TRICLADiDA PALUDiCOU

INVERTEBRATE ZOOLOGY
SMITHSOMAN IMSTITUTiON

ilP.

i'

New

Series, No.

184 (Vol. 46, Part

Price

4).

FEBRIIAEY,

1903.

THE

QUARTERLY JOURNAL
OF

MICROSCOPICAL SCIENCE.
EDITKI)

E.

RAY LANKESTEK,

HY

M.A., LL.D., E.R.S.,

SOKABV FELLOW OP EXETER COLLEGE OXPOB D COH B ESPONDK > T OK THE INSTITUTE OK PBAXCI
AND OK TUB IMPEBIAL ACADEMY OK SCIENCES OK ST. PETERSBURG, AND OK THE ACADKMV
KORKIGN MEUBBR OK TUE BOAL BOHEMIAN
OK SCIENCES OK PUILADELPIIIA
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AND OK TUK AMERICAN ACADEMY OK ARTS AND SCIENCES OK BOSTON
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CAMBRIDGE PHILOSOPHICAL SOCIETY, AND OK THE ROYAL
PHYSICAL SOCIETY OP EUINBUBOH HONORARY
MEMBER OK THE BIOLOGICAL SOCIETY
,

OK Paris;
DIRECTOR OP TUB NATURAL HISTORY DEPARTMENTS OK THE BRITISH MUSEUM; LATE KULLERIAN
PBOKBSSOR OK PHYSIOLOGY IN THE ROYAL INSTITUTION OK GREAT BRITAIN.

WITH THE CO-OPERATION OF

ADAM SEDGWICK,

M.A.,

F.R.S.,

KELLOW AND TUTOR OK TRINITY COLLEGE, CAMBRIDGE;

W.

F. E.

WELDON,

M.A., F.R.S.,

LINACHE PROFESSOR OF COMPARATIVE ANATOMY AND KELLOW OK MEUTON COLLEGE, OXFORD:

LATE KELLOW OF ST. JOHN's COLLEGE, CAMBRIDGE;

AND

SYDNEY

HICKSON,

J.

BEYER PROtKSSOU OP ZOOLOGY

IN

M.A.,

F.R.S.,

THE OWENS COLLEGE, MANCHESTER.

WITH LITHOGRAPHIC PLATES AND ENGRAVINGS ON WOOD.

LONDON:
J.

&

A.

CHURCHILL,

7.

GREAT MARLBOROUGH STREET.

1903.

Adliird

and

.Son,]

[London and Dorkinc

CONTENTS OF

No.

184.-New

Series.

MEMOIRS
PAGE

The Movements and Reactions of Fresh-water Plauarians a Study

By flAyMOND Peael, Ph.D., Instructor in
in Animal Behaviour.
:

Zoology in the University of Michigan,

Ann

Michigan,

Arbor,

509

U.S.A

Part IV. On the Central Complex

By A. T. Mastekman, M.A.,
dl>aus dodecalophus, McI.
D.Sc, Lecturer on Zoology, School of Medicine, Edinburgh. (With

Ofl the Diplochorda.

of Cepliaifl-

715

Plates 33, 33)

On Hypurgon Skeati,
Ascidians.

By Igerna B.

New Genus

and Species of Compound

Sollas, B.Sc.Lond.

J.

(With Plates
729

34,35)

Ehlers, and of a New Variety

some Observations on the Post-larval Stages.
-737
H. AsHWORTH, D.Sc. (With Plates 36, 37).

The Anatomy

of

Arenicola ass mills,

i

of the Species, with

By

J.

Title, Contents, and Index.

-7||-,

iniuUiUsuH rauuui^uui \^

QUARTERLY JOURNAL

MICROSCOPICAL SCIENCE.
EDITED BY

B.

RAY LANKESTER,

M.A., LL.D., P.R.S.,

EXETER COM.EGE, OXFOUD COR R KSPON nENT OF THE INSTITUTE OF FRANCE

AND OF THE IMPERIAL ACADEMY OF SCIENCES OF ST. PETERSBURG, AND OF THE ACADKMV
OF SCIENCES OF PHILADELPHIA; FOREIGN MEMUKR OF THE ROVAI. BOHEMIAN
SOCIETY OF SCIENCES, AND OF THE ACADEMY OF THE I.INCEI OF ROME,
AND OF THH AMERICAN ACADEMY OF ARTS AND SCIENCES OF BOSTON
ASSOCIATE OF THE ROYAI. ACADEMY OF BELGIUM; HONORARY MEMBER
YORK ACADEMY OF SCIENCES, AND OF THE
<1F THE NEW
CAMBRIDGE PHIl.OSOPHICAI. SOCIETY, AND OF THE ROYAL
PHYSICAL SttCIETY OF EDINBURGH HONORAR Y
MEMBER OF THE BIOLOGICAL SOCIETY

ilONORARV FRl.I.OW

Ol'

OF Paris;
nlHECTOR nr the natural history departments of the BRITISH MUSEUM; LATE FULLERIAN
PROFESSOR OF PHYSIOLOGY IN THE ROYAL INSTITUTION OF GREAT BRITAIN.

WITH THE CO-OPEUATION OF

ADAM SEDGWICK,

M.A.,

F.R.S.,

FELLOW AND TUTOR OF TRINITY COLLEGE, CAMBRIDGE;

W.

F. R.

WELDON,

M.A., F.R.S.,

LINACKE PROFESSOR OF COMPARATIVE ANATOMY AND FELLOW OF MERTON COLLEGE, OXFORD;

LATE FELLOW OF ST. JOHN's COLLEGE, CAMBRIDGE;

SYDNEY

HICKSON,

J.

M.A.,

F.R.S.,

BEYER PROFESSOR OF ZOOLOGY IN THE OWENS COLLEGE, MANCHESTKK.

VOLUME
3i5litlj

Ififljogrnpljit

46. New Series.

|!lales

anb

(L-ncirRbings

on

3!5loob.

LONDON:
J.

&

A.

CHURCHILL,

7,

GREAT MARLBOROUGH STREET.

1903.

INVERTEBRATE ZOOLOGY
SMITHSOMIAM mSTiTUTiOAl

CONTENTS.
CONTENTS OF

No. 181, N.S., JULY, 1902.

MEMOIRS
PAGE

On

a Free-swimmiiif^

By Arthur Dendy, D.Sc, F.L.S., rrofessor

et sp.
in tlie

HydroiH, Pelaf^ohydra mirabilis,

New

Canterbury College, University of

Plates 1 and 2)

Studies in the Retina.

of Biology

(With

Zealand.

-1

n. ;en.

Parts III, IV, and V, with Summary.

Henry M. Bernard, M.A.Cantab.

(From the

By

Biological

(With

Laboratories of the Eoyal College of Science, London.)

3 5)

Plates

25

Haliotis tuberculata,

Notes on the Relations of the Kidneys in

By II. J. Fleure, B.Sc, U.C.W., Aberystwyth.
etc.

(With
'^^

Plate 6)

Notes on the Development

of

Paludina vivipara,

with special

reference to the Urino-genital Organs and Theories of Gasteropod

(With Plates 79)
Torsion. By Isabella M. Drummond.

97

Is

Chemotaxis a Factor

Animals?
strator in

the

Fertilisation

the

Eggs of

By A. H. Reginald Buller, B.Sc, Ph.D., DemonBotany

CONTENTS OE
MEMOIRS

in

of

at the University of

No. 182, N.S.,

Birmingham

SEPTEMBER,

.145

1902.

Maturation of the

Ovum

in

H. Bryce, M.A., M.D.

Echinus esculent us.

(With Plates 1012)

Studies on the Arachnid Entosternite.

Plates 13 and 14)

By R.

I.

By Thomas
.

PococK.

.177
(With
22J

CONTENTS.

IV

PAGE
Morphology of the Cheilostomata. By Sidney P. Harmer,
(With Plates 15 18)
.263
Sc.D., F.U.S.

On

tlie

On

the Development of Sagitta

the Adult.

with Notes on the Anatomy of

By L. Doncaster.

CONTENTS OF

(With Plates 1921)

No. 183, N.S.,

DECEMBER,

351

1902.

MEMOIRS
On

By William A. Haswell, M.A.,

a Cestode from Cestracion.

(With Plates 2224)

D.Sc., F.R.S.

The Development

of

velopment of the Skin and

Kerr.

Lepidosiren par ado xa.

Derivatives.

its

(With Plates 2528)

Part
By

III.

J.

.399
De-

Graham
,

.417

The Metamorphosis of Corystes Cassivelaunus (Pennant).

By Robert Gurney, B.A.(Oxon.), P.Z.S. fWith Plates
29" 31)
Artificial

Parthenogenesis

Thomas H. Bryce

CONTENTS OF

......
and

Fertilisation

No. 184, N.S.,

Review.

FEBRUARY,

461

By
479

1903.

MEMOIRS:
The Movements and Reactions
Animal Behaviour.

in

Zoology

in

the

of

Presh-water Planarians; a Study

By Raymond Pearl, Ph.D., Instructor

University of Michigan, Ann Arbor, Michigan,

in

U.S.A

509

On the Diplochorda. Part IV. On the Central Complex of

Cephalodiscus dodecalophus, McI. By A. T. Masterman,

...

M.A., D.Sc., Lecturer on Zoology, School of Medicine, Edinburgh.

(With Plates 32 and 33)

On Hypurgon Skeati,
Ascidians.

By Igerna B.

new Genus and Species

J.

Sollas, B.Sc.Lond.

of

(With Plates

34 and 35)

The Anatomy

715

Compound
729

of

Arenicola assimilis,

Ehlers, and of a

New

Variety of the Species, with some Observations on the Postlarval Stages.

By

J.

H. Asiiwortii, D.Sc.

and 37)

(With Plates 36
737

Title, Index,

and Contents.

->

fc>r,

'

MOVKM?]NTS, ETC, OF riJESFI-WATER rr,AN A 1(1 ANS.

';i>tx

509

The Movements and Reactions of Freshwater

Planarians a Study in Animal Behaviour.'
:

By

Kayiiioiid PcnrI, Pli.D.

(rnsfniRlnr

Zonlotry in

in

tiic

Univcisily of Mirliipjan,

Ann Arbor,

Michigan, U.S.A.)

Contents.

A.

Tntkodtjction

B.

l^ESUME OF LlTERATUKE
I.

JI.

Morplioiogical and Systematic

logical

Pliy si

Material
Habits and Naturae History
I. Occurrence and Distribution

c.

D.

II.

III.

52G

Activities

527

a.

Sensitivity

h.

Secretion of

c.

Periods of Activity and Rest

527

Mncus

529

d.

Formation of Collections

e.

Movement on

532

533

Surface Film

534.

Food

535

IV. Defecation
V.

Summary

of

537

Factors

in

JJcliaviour

538

Normal Motor Activities

I. Locomotor Movements

E.

a.

'

539
539

Gliding

539

1.

Rate of Gliding Movement

545

2.

Direction

548

Contributions from the Zoological Laboratory, University of Michigan,

Ann Arbor, Michigan, No.

VOL. 46, I'ART 4.

58.

NKW

SERIES.

LL

RAYMOND

510

b.

c.
(1.

Crawl in J? Movement
Direction

2.

Stimuli which induce Crawlino:

Movement on

Movements

b.

Extension of the

c.

Rest

d.

The

Descrii)tion of Reactions

555

555
556

.557

566
570
576

....
.

Reactions to Strong Stimuli

j8.

Reactions to Wc:ik Stimuli

Head Region

Body

Reactions to Strong Stimuli

jS.

Reactions to

Weak

Stimuli

588

589

S.

Reactions to Stimuli applied to the Posterior Region

4.

Reactions to Stimulation of the Ventral Surface

5.

Reactions

6.

Reactions to

7.

The

dure

Body

of

Specimens

Resting

.592

The General Features

.

the

of

by

Operative

Hindrance

Reactions

.595

given

to

Proce-

Movement

to

595

597

Mechanical
.

.600

The Mechanism of the Reactions

1. The Relation of the Brain to the Reactions
2. The Neuro-muscular Mechanism
Features in the General Behaviour of the Organism which

602

602
606

the Reactions to Mechanical Stimuli explain

619

.....

Summary
Reactions to Food and Chemical Stimuli
a.

594

Mechanical

to

.....
.

Slimuli

Effect of Mechanical

Stimuli

II.

578
582

.588

o.

Stimuli

/.

577
577

Reactions to Stimuli applied to the Middle Region of

of the

e.

576

.576

Reactions to Stimuli applied to the

the

f/.

.....

b.

c.

Movement

Mechanical Stimuli

Methods

2.

Effect of Operations on

a.

1.

Formation of Collections

to

'

Reactions

553

....

Body
Body

Contraction of the

551
552

of Tiiclads to those of other

Non-locomotor Movements
a.

548

Eeactions to Stimuli
T.

PAGE

.550

the Surface Film

Relation of the

i.

p.

....

1.

Forms
TI.

PEAKIi

Food Reactions
1. Food Reactions of Specimens
2. Summary of Food Reactions

after Operations
,

.623

623

624
637

640

MOVKMKNTS,

ETC., OF FEI']SH-WATRR TLANARJANS.

511
page

b.

Reactions to Chemical Stimuli

1.

ITT.

Cliemotaxis

643

Reactions to Localised Chemical Stimuli

G43

a.

Methods

043

j3.

Results

646

Summary

2.

General

3.

Unlocalised Action of Chemical

657
669

Thifjmotaxis and the Ri^htinj? Reaction

670

a.

Thiii;motaxis

670

b.

The Riifhtinj^ Reaction

The ]\[echauism of the Reaction

676

Summary

684

c.

673

TV. Elect rot ax is

685

a.

Methods

685

b.

liesults

685

c.

Mechanism

d.

Summary

V. Tieaction to

of the Reactions

690
695

Desiccation

695

VT. Rlieotaxis
G.

H.

697

General Summary and Di.scussion of Results

List of Literature

A.

The present study has

698

7T0

Introduction.
for its purpose the analysis of the

l)ehaviour of the coinniou fresh-water planarian into

ponent factors.

It

is

well

known

we have

little

organisiiis.

aside

that,

researches of a few investigators on a small

number

its

com-

from

the

of forms,

knowledge of the behaviour of lower

coming to be realised, too, tliat knowledge

detailed

It is

what an animal does is just as important in the general

life phenomena as a knowledge of how it is constructed, or how it develops.
But it must be admitted that

of

study of

until quite recent times the study of the activities of living

things

was

much

neglected

field

Origin of Species

in

biology.

The

gave the biological

pendulum a swing towards the study of phylogeny, from
which it is onl}' just beginning to return.
As a consequence of this concentration of interest on other
subjects, we possess an accurate and full knowledge of the

publication of the

'

'

RAYMOND PRARL.

512

The beliaviour of the

activities of very few lower organisms.
Protozoa has been quite fully described and analysed by tlie
Avork of Verworn ('89) and Jennings ('97, '99, '99a, '99&,
'99r,

00,

OOff,

00?^,

In the earlier work

of

:00c, 01, Jennings and Moore :02).

Yerworn the general features of most
:

of the reactions of the Protozoa are described, special attention

In

The reactions

being paid to the rhizopods.

Jiifnsoi-ia

the

of

the

have been very thoroughly worked out by Jennings.

case

of

the

In-fusoria

we now know

exactly

the

mechanism of the renction to a large number of stimuli. The

reactions and general behaviour in the case of two groups of
echinoderms are quite thoronghly known from the early work
of Preyer {'86, '87) on the starfish and the recent brilliant
work of von Uexkiill ('96, '96ff, '99, 00, :00ft) on the seaThese few instances are the only ones in the
urchin.
literature where the inovements and reactions of an organism,
or group of organisms, have been investigated in any comprehensive " monographic " way. There is a great body of
:

literature dealing with isolated leactions in a variety of forms,

but the thorough investigation of the activities of animals in
a way comparable to that in which their morphology has

been investigated remains in large degree yet to be done.

It appeared highly desirable that this sort of knowledge be
extended, and it was with this idea in mind that this work
was undertaken. The form used, Planaria, was chosen for
several reasons.

paradigm

In the

work

for

(ui

first

place,

already well known.

standpoint

is

or more of

its

it

has come to be a sort of

regeneration, and

its

biology from that

Furthermore,

in

some one

an almost universally distributed

form and can always be obtained in quantities. Finally, and
particularly, it is a representative of an animal type about
whose activities we know only the most general facts. It is
species

it

is

a symmetrical aquatic organism of low organisation, and

behaviour

is

rathei' complicated.

ing a detailed knowledge of the

symmc'trical,

when

it is

The importance
.activities

its

of possess-

of a bilaterally

free-moving, low organism will

be apparent

considered that such an organism has never been

MOVEMENTS,

ETC.,

OF FI.'ESll-WATEi; PLANA IMANS.

513

made the subject of sucli a study. Tlie behaviour of typically

uusymmetrical organisms, the Infusoria, has been analysed,
as has also that of some radially symmetrical animals, and in
is found to be a very close iuterrelationbetween the general form of the body and the reactions.
To investigate, then, iu a comprehensive way the activities
of a bilaterally symmetrical organism standing low in the
animal series was the purpose of this work.
The most
general problem which presents itself is the establishment

both cases there

sliip

of the animal's position in the objective psychogenetic series.

Are

its activities relatively

simple or are they complex

Do

under the same general type as those of the

Infusoria or those of the higher organisms, or do they occupy
an intermediate position ? Another general problem of importance is whether there is any marked correlation between
the behaviour and the form of the body, such as has been
found to obtain in so marked a degree in the case of the
Infusoria and the rotifers (vide Jennings, loc. cit.). We
have in the tlat-worm a symmetrical animal; are its reactions
they

fall

?
Besides these broad fundamental
problems there are, of course, a large number of subsidiary
questions which readily suggest themselves iu connection
with a work of this sort.
These need not be specifically
mentioned here, but will be brought out in the course of

of a symmetrical type

the paper.

As to the scope of the work as actually done, the following

may be said: The general '^ natural history" of the animal
was studied as completely as possible. All the normal movements were studied iu detail. The reactions to mechanical
stimuli; the food reactions and reactions to chemicals in

general; electrutaxis

thigmotaxis; rheotaxis; the righting

reaction; the reaction of cut

and regenerating pieces

and

hydrotaxis and the reactions during desiccation were investigated.

No work was done

on the phototaxis or thermotaxis.

study of the phototaxis was omitted for two reasons;

first

on account of the fact that during the progress of this investigation Parker and Burnett (: 00) reported their work on

514

KAYIMOND

PEAllL.

the sumo subject^ and furthermore on account of lack of

As

opportunity.

made

a result of some

incidental

observations

appeared that it
extend the Avork of Parker and

during- the course of this work,

it

lias

would be profitable to
Burnett, and this, together with a study of the thermotaxis, I
hope to be able to do iu the future. Another field for further
work is afforded iu the study of the reactions of regenerating
individuals.
As this subject did not fall immediately into
the general plan of this work, but comparatively little attention has been g'iven to it, yet the work done gives much
promise of important results to be gained by more extended
investigations.

So

far

as

possible

the

thei-e

is

details

much

not as

much need

in physiological

of the recent

work

movements

and

It is not easy to see

why

details

reactions will be described fully.

for a

work
in

as

the

of

complete knowledge of
in

morphological, yet in

comparative physiology only the

most general results are reported. To gain a knowledge of

the details one must do the work over again. While such
more or less general papers are easy to read, and put the main
results in such a form as to be easily accessible, yet it is
believed by the writer that the solid foundations of comparative physiology and psychology must consist of detailed
" fine " work, just as has been the case in morphology. It
seems to the writer that the tendency to abandon the detailed
descriptive method in favour of the extreme experimental
method iu biological work is unfortunate. Both ways of
working are methods of getting at the truth, and, as proven
by their results, both are good methods. The current notion
of the sufliciency of the experimental method to the exclusion
of others is not only an evident exaggeration oi the facts in
the case, but, in the opinion of the writer, the exclusive use
of the " crucial-experiment "

ments and reactions

of

method

organisms has

in

in

work upon the movesome cases hindered

rather than hel[)ed us to gain a clear understanding of the

])henomena.

The importance

of close observational

work

in

the study of animal behaviour has been strongly emphasised

MOVEMI'JNTS, KTC,

FBE8H-\VATER I'LANAKIANS.

Ob'

The aim

recently by Wliittmm ('09).

in

the present

515

work

has been to get as extensive and detailed a knowledge as

possible of the behaviour of the organism

by

direct observa-

tion before resorting to experiments.

At

this point I wish to

acknowledge

my

indebtedness to

work has been

Jennings, under whose general

the officials of the laboratory in which this

done.

To Professor H.

S.

oversight this investigation has been prosecuted, I wish to

extend

my

heartfelt thanks

for

freely giving advice, suggestion,

Any adequate

measurable value.
ness to him

is

impossible.

his uniform kindness in

and kindly criticism

expression of

my

of im-

iudebted-

further wish to express

my

Jacob Reighard for the numerous

facilities which I have enjoyed
during my stay in his
laboratory, and for his kindly interest, which has made work
Finally, I desire to acknowledge my inthei'e a pleasure.

thanks

Professor

to

debtedness to Professor F. C. Newcombe, of the Botanical

Department

of

the

University

of

Michigan,

for

many

valuable suggestions and advice.

B.

Resume op LrrERATURE.

But little has been done on the physiology of the movements or on the psychology of the Turbellaria, and, as in
the case of most of the literature dealing with these subjects,
what has been done has been in comparatively recent years.
Investigators of the old " natural-history " school which
flourished before the time when Darwin's work changed the
course of zoology seem not to have given much attention to
planarians, while the later systematists and morphologists
for the most part carefully avoided any reference to the
activities of the forms which they studied.
I.

Among

Morphological and Systematic.

the papers devoted primarily to the systenuitic or

morphological treatment of the group, there are occasional

references to jioints in the behaviour of the oi-ganisms which

RAYMOND PEARL.

516

importance from the present standpoint. Aiuon

such references the following may be noted
Moseley ('74), in a paper concerned principally with the
anatomy and histology of the land plaiiarians, devotes a sec-

are of

He com-

tion to a discussion of the habits of these forms.

ments on the ''avoidance of light" (negative phototaxis) of

hind and aquatic planarians, and discusses the habitat and

He reaches the conclusion that all

food of the animals.
plauariaus are carnivorous, but gives no account of the
method of feeding. He quotes Rolleston as having found
Planaria torva and Deudrocoelum lacteura in a
dish in which had been placed a freshly killed earthworm
" crowded on to the worm's body and soon sucked all the
hajmoglobin out of it, leaving it white and pulpy." Brief

that

mention

made

is

of

the habit of the land planarians

of

mucous thread and hanging from it as a mollusc

method of movement of Bipalium with
the head raised and waved from side to side as the animal

secreting a

Finally, the

does.

proceeds
is

is

described.

bibliography of previous literature

given.

In another paper Moseley

('77,

pp.

273, 271)

account of the movements and general habits of

11

aV

in

a,

a Brazilian species.

gives an

Geoplaua

This species was found to keep

shaded and moderately lighted places.

The

direction of

the ciliary currents was tested by placing small bits of paper

on the surface of the body, and

was

it

was found that when the

movement the

effective beat of the

on the anterior part of the dorsal surface was forward
and outward, while on the posterior portion of the dorsal
sui'face the beating was backward and outward.
The

aninuil

in active

cilia

currents on the ventral surface were always straight back-

ward.

The author concludes that the function

on the dorsal surface

When

is

to

of the cilia

quickly remove foreign bodies.

the organism was at rest there was no

movement

of

"the animal moves to a large extent by

muscular action, the body alternately contracting and expiinding during motion.
When moving it lifted its anterior
the dorsal cilia;

517

MOVEMENTS, EIC, OF FUESII-WATEU PLANARIANS.

extfoinity

or see

its

was cut

and moved

offceiij

"

way/'

off

When

it

aud ffo as

to

it"

to fee

the anterior extremity of the body

the remainder of the animal seemed

still to

move

with definite purpose, avoiding obstacles and retreating- from

end was raised and thrust in various

the light; while the cut

directions as

to search for

if

In a brief note Zacharias

an object on which to climb."

mentions the occurrence

('88)

of

terrestris between the lamellic of Agaricus

the slow
Particular points mentioned are
deliciosus.
movement^ characterised by the raised anterior end, aud the
hanging by a mucous thread after passing over the edge of
a glass plate. Light stimulation of the anterior end with a
needle induces a very strong contraction of the whole body.
(Jamble ('03), in a systematic paper on marine 'i\xrbel-

Geodesmus

the movements of a number of

and triclads.
Lang ('84), in his monograph on the polyclads, devotes a
chapter to the habits, movements, and natural history of this
group of })lanarians (loc. cit., pp. Goi Gil). While not
done particularly from the physiological standpoint and not
laria,

describes

briefly

species of rhabdocujles

treating

the

experimentally, this

subject

numerous valuable observations.

contains

w(jrk

Points especially treated

method of feeding,
swimming, copulation,
respiration, regeneration, growth, and duration of life.
The

are the habitat, colouration, food and

movements,

defecation,

details iu the

monograph

including

behaviour of the polyclads recorded

will

in

this

be discussed later in connection with the

points on which they have direct bearing.

The most important paper dealing with the movements

and general behaviour

of planarians

to find in the literature

is

is

which

I Iiave

that of Lehuert ('91).

been able
This work

principally devoted to an account of the biology of three

fornjs

of

keweuse

land
var.

planarians,

viridis,

viz.

and

Bipalium kewense,

B.

Geodesmus bilineatus.

Besides the work on these land forms, Lehnert also made

some comparative studies on several fresh-water dendroceeles
aud rhabdocoeles. He gives au admirably full and detailed

RAYMOND PEAKL.

518
account

by far
phenomena in the Hterature.
of the movement to the mucous

the inoveiueuts of laud plauariaus; in fact,

of

the best description of these

In this account the rehition

secretion from the ventral surface of the

in

The

great detail.

movement

body

is

brought out

principal factors in producing the

the case of the land planarians he gives as

in

movement on the

ventral surface, [h] rhythmical

waves passing longitudinally over the ventral
surface, (f) secretion of slime, and (d) snake-like movements
A comparison with the movements of
of the whole body.
other planarians (fresh-water) is made. In this connection
it may be mentioned that Lehnert considered rhythmical contraction waves passing along the ventral surface of the
(a) ciliary

contnictiou

animal

be a factor

to

in

(Dendrocujlum

planarians

movement of fresh-water
lacteum, PI an aria poly-

the

chroa, and Polycelis tenuis). This I am unable to confirm from observations on the planarians which I have
studied.
This point will be discussed more fully later. The
food and the method of taking food in case of the land
planarians, Lehnert worked out very thoroughly. They were
found to be carnivorous, and in the case of Bipalium the
pharynx was ca])able of being stretched over a large j)iece of
earthworm, so that it resembled a very thin transparent skin
covering

it.

ment, e.g.

'JMie

air,

relations to other phases of the environ-

water, temperature, light, solid bodies,

etc.,

are described very briefly.

Easpail

('93), in

a brief note, mentions the feeding of a

planarian.

Van Dnyne

('06)

mentions

briefly

heteromorphic forms of PI an aria

that the

parts

of

two-headed

movements

the

torva

individuals

of

He

found

moved

inde-

(?).

pendently of each other, and that each piece Avould move

away from the other until they had completely torn apart.
Willey ('07 j, in a brief note, describes the structure of a

remarkable asymmetrical planarian, for which he proposes

name Hetero])lana, having the left side of the

body almost completely atrophied.
Regarding the locothe generic

M0VI':MENT8, etc., of laiESll-WATEIi rJ,ANAUlANS.

519

motion of this reiiuirkable form he says, '^lu llctorophma

the locomotion is usually conducted in a somewhat
one-sided fashion/' and he furthermore figures the animal
It is to be regretted
as constantly moving* towards the left.
is made to how this form reacts to stimuli,
would be of great interest to know whether the reactions
are asymmetrical, and in general how they compare with
the normal planariau type.
A series of papers by Morgan ('08, 00, 01) contains
numerous references of importance on the movements of cut
and regenerated specimens of various fresh-water planarians.
He finds ('98), in conlirmation of van Dnyue, that in twoheaded individuals each head tends to move in its own proper
direction.
In the case of a hetex'omorphic form with two

that no reference
as

it

lieads pointed in opposite directions, this likewise held true

but one component being stronger this determined the move-

ment

The lack

of the whole.

of

movement

In his

of cut pieces is also noted.

the readiness with which " Planaria, sp. "

maculata

in

certain forms

00 paper Morgan notes

take food, although no account

and Planaria
is

given of the

method of the feeding reaction. An interesting observation,

and one of considerable theoretical importance, is also reported in

tliis

paper.

In an individual split longitudinally in

the median line from the posterior end forward,

in which the
two parts were united only by a small connecting band of
tissue at the anterior end, it often appeared " as though these
pieces would pull apart, but as soon as the tension on the
connecting band becomes too strong, the rest of the piece, by
a sort of adaptive response, ceases pulling in its former
direction."
In the most recent ])aper cited (:01) Morgan
corrects a statement of Bardeen
regarding the feeding of
Planaria. It is maintained (and 1 may mention at this
'-^

point that

my own

observations agree entirely with those of

Morgan) that Planaria

1

Later

'

To be reviewed

iileiitiiied

'^

responds

by Woodwoitli as
later.

I'l

freely" to food sub-

anuria lugubris.

RAYMOND

520

PEAIIL.

stances uot actually iu contact with

This point will be

it.

discussed in detail later.

Lillie (:01) bring's

out the fact that cut posterior parts of

the body of Dendrocoelum lucteum show very little

movement^ and in general fail to give the typical reaction to
light after removal of the brain.
Filially^

there are accounts of

natural

the

history

and

habits of various planarians in numerous "natural histories"

and text-books. As such accounts are for the most part
brief and of no great significance from our standpoint, they
will not

be referred to iu
II.

The

detail.

Physiological.

literature dealing with the

planarians from a purely

Furthermore, for
is very meagre.
most part it deals only with special phases of the
physiology of these organisms, there being very little work
attempting to bring the behaviour of planarians into relation
physiological standpoint
the

with that of other forms.

The most important work dealing experimentally with the

physiology of the movements of flatworms which I have
found is that of Loeb ('Df). The purpose of his work was to
determine

in

how

far the reactions of

such low organisms as

worms were dependent upon the brain. The planarians used

Thy sano zoo n brocchii, and Planaria torva. In

were

Thysanozoon he found

that if the animal were quickly cut

two pieces transversely with a sharp scalpel or scissors
undisturbed, while the
the anterior piece crawled on
The concluposterior piece showed no further movement.
sion is then drawn that " Die Spontaneitiit der Progressivbewegungen ist also bei Thysanozoon eine Fuuktion des
Gehirus." This form shows no definite ''geotropic " reac-

into

tion,

but crawls about with the axes of the body forming any

The very strong

angles with the direction of gravitation.
" stereotropism " (thigmotaxis) of the ventral side of Thysanozoon, which always tends
solid

body,

is

noted.

to

keep

This reaction

is

contact with a
found to be indein

MOVRMENTS,

OF FRESn-WATER PLANARTANS.

ETC.,

521

There was fonud to be co-ordination

and posterior pieces of a worm in
wliich tlie Lateral Longitudinal nerves had been cut, but in
which a narrow connecting strip of tissue had been left
between the pieces. In P. torva Loeb states that posterior
])arts of the bod}' which have been separated by a transverse
cut fioni all connection with the brain crawl " ebenso ninnter
The reaction of this form to
weiter, wie die orale Halfte."
changes of light intensity is discussed in considerable detail,
it being shown that in strong
light the organism is stimulated to active movement, while in the shade it remains quiet
or moves very slowly.
This was found to occur as well in
decapitated as in nortnal woi-ms. The " stereotropic " reacIn concluding, the
tion in this form is also mentioned.
author holds that in worms there is no ''associative Gediichtniss," and hence no consciousness.
These results have been
recently incorporated without essential change into a larger
work (Loeb 00).
In an earlier paper Loeb ('9o) fii-st described the reactions
to light of PI an aria torva.
These results were incorporated without essential change into the '94 pnper menpendent of the
l)et\veen

lirain.

the anterior

tioned above.

Hesse

('97), in

his

morphological studies on the eyes of

flat-worms, devotes a section to the subject of the reactions

to light of

Euplanaria gonocephala and Dendrocoelum

lacteum.

His results are confirmatory of Loeb's, nothing

of particular significance being added.

Steiner {'^S) found that posterior pieces of Planaria

Neapolitana (=:8tylochus ])ilidium, Lang) separated
from the brain by a transverse cut would move about freelyafter recovery from the operation.
to

move

is

conditioned

b}^

He

believes this ability

the presence of ganglion-cells in

other parts of the body than the brain (along the lateral
nerve-cords)

Parker and Burnett

00)

have recently made a thorough

study, using very careful experimental methods, and treating

the

results

statistically,

of

the

reactions

of

Planaria

EAYMOND

522

gonocepliala

to

PEART;.

This form moves away from the

light.

The amount

of directive influence was

was found that specimens without eyes^ i.e.
in which the anterior end had been cut off, react in much the
same way to light as do normal individuals, 'Mn that they
have a tendency to turn away from the course when directed
towards the source of light, and to keep in it when directed
awa}' from the source, though with less precision, and often
Furthermore,
to less extent, than planarians with eyes."
figures are given showing that planarians from which the
anterior end has been cut off move more slowly than normal
This is thought to bo due to the absence of the
animals.

source of the light.

measured.

It

eyes.

The most extensive paper dealing with the physiology

planarians

is

that of Bardeen (:01).

This paper

is

of

mainly

devoted to a stud}' of regeneration in Planaria niaculatn,

but before enteriiig upon the discussion of this subject the
author devotes considerable space to an account of
anatoni}'^

and physiology

of the organism.

the

In the section

devoted to physiology, the author discusses, under the

caption " Environmental Activities," sensation, movement,

and the central nervous system.

The author makes the
and
obviously
incorrect,
remarkable,
statement that the
planarian is sensitive only to light and contact.
A ver}' few
inconclusive experiments having reference to thigmotaxis,
geotaxis (?) and hj-drotaxis, are reported. 'The statement is
made that specimens " would remain unmoved by the presence
close l)y their side of a piece of fresh snail, a food

prized by them."

"swimming" and
Avhat he calls the "

Two

much

forms of movement are described

crawling.
The author's description of
swimming " movement will be discussed

Brief and very general statements

later in this paper.
regarding the reactions to mechanical stimuli are presented.
Under the heading " Internal activities " are discussed
deglutition, food dispersion, defecation,

rather loose and liy]iothetical Avay.

followintj;;

and respiration,

Tiie

in a

author makes the

contribution regarding exci-etion in

Planaria:

MOVKMKNTS,
''Excretion

is

ETC.,

OF FRES[T-WATER PLANARFANS.

523

carried in part through the intestines by the

act of defecation

in part

it

is

doubtless carried on by an

excretory system opening on the sui'face."

more detailed

discussion of various points raised by Bardeen will be entered

work on which they

have direct bearing.
A second paper by the same author (Bardeen^ Ola)
describes briefly the normal food reactions of Planaria, and
shows that a decapitated specimen will not find food materinl
in a dish, although one such a specimen could " be made to
eat if it were placed on its back on a slide in a small drop of
Under the conditions mentioned the pharynx is
water.
usually protruded, and will engulf bits of food placed in the
mouth." An experiment was performed in which the pai't
of the head in front of the eyes was cut off. Such specijnens,
from which merely the tip of the head had been removed, reIt is also shown that specimens
acted normally to food.
fi'om which the part of the body posterior to the pharynx has
Regarding the
been removed feed like normal worms.
method by which planarians find food in their immediate
vicinity, Bardeen says (p. 176), ''It is difficult to determine
the source of the impulse which gives rise to this purposeful
activity.
It is possible that the auricular appendages here
act as delicate organs capable of stimulation by slight
currents in the water set up by the minute organisms that
prey at once upon the flesh of the dead snail." Experiments
to be reported in the course of the present paper show, I
think, that the mechanical and chemical stimuli given by food
are the ones which affect planarians.
into in connection with the parts of this

c.

Material.

The following species have been principally used in this

Planaria maculata, Leidy
Planaria gonocephala, Duges Planaria dorotocephala, Woodworth.^
Of these P. dorotocephala and P. maculata have been
study

'

Excellent figures and descriptions of

lislied

by Woodwoitli, '97.

lljese

three species have been pub-

RAYMOND PEAKL.

524

most used, both on account of their abundance and, furthermore, because P. dorotocephala is a form particularly
very active, and
after being disturbed continues in movement longer than
either P. macnlata or P. gonocephala, as has ali-eady
favourable for the study of reactions.

It is

I have found
been noted by Woodworth CIoc. cit., p. 7).
also that it moves faster than either of the other two species.

There is a general precisiou and jiositiveness of i-esponse in

behaviour which make it es]iecia]ly favourable for experimental work. A large number of experiments have been
made with a view to determining whether there was any
difference in the i-oactions of these three species, but no

its

essential difference has

is

the same in

been found.

Whatever

all cases.

The form

of the reactions

differences there are are

differences of degree, such as would be conditioned by the

relative sluggishness

and

activity.

Certain forms of reaction to mechanical stimuli, and to

chemical

stimuli,

more

rather

are

dorotocephala than

easily

induced

in

P.

in either of the others, yet, as will

be

be given, under the proper

This being the case, and
conditions, by the other species.

shown

later, these I'eactions will

since P.

used

dorotocephala

in this

work,

it will

as the type form, and

it

was, for reasons noted above, most

be employed throughout the paper

will be

understood,

when

statement of the species, that P. dorotocephala

there
is

is

no

the form

meant.

No

account of the anatomy of these forms

will be given
has been very fully treated in other readily
The most important papers dealing with
accessible papers.

here, because

it

morphology of the fresh-water triclads are those of

Lang ('81, '81a), Kennel ('88), Chichkoff ('92),
and Woodworth ('91 and '97).
Besides the species mentioned above, on which the most of
the work was done, a number of observations and experiments have been made on several other species of triclads
and rhabdocoeles. The other triclad most frequentlj' met,
and whose reactions have been found to agree closely Avith
the

Jijiina ('84),

MOVEMENTS,

OF FRKSH-WATER PLANAllIANS.

ETC.,

Plauaria,

of tlio species of

tliose

is

525

a form Avliich agrees

witli tlie description of

Dendrococluni lacteum, except

respect to the colour.

This form

is

The colouring

light grey to nearly black.

in

usually coloured from a

is

uniform.

In

found any deviation from this

typical coloration, and in that case there Avas a band of
black pigment extending the whole length of the body
along the mid-dorsal line. In width tliis band occupied about
one third the whole width of the body. The margins were a
only one specimen have

pure white, without the faintest trace of pigmentation.

This

specimen was kept under observation for some time, and

belonged to the same species

at once as beingtransitional between the ordinary white to cream-coloured
Dend rocoelum lacteum, and the grey form found about
Ann Arbor. Being in some doubt as to the true taxonomic
position of this grey species, I shall refer to it throughout
this paper as Dendrocoelum, sp.
Besides the forms mentioned, several undetermined triclads
have been collected and worked with to some extent, but as
no new factors presented themselves in their reactions they

there was no doubt that

will not

it

The specimen struck one

as the grey form.

be considered in this papei'.

I have identified asMesostoma
found rather commonly in certain

Alargei'habdocoele, which

personatum,

0. Schm.,

about

localities

woi'k on this

have studied
Schui.

No

Ann

is

I have done some

Another rhabdoco?le whose actions I
some extent is Stenostoma leucops, O.

Ai-bor in the spring.

form.
to

detailed

investigation of the behaviour of the

vhabdocosles was made,

Init as

opportunity offered they were

used for comparison.

The methods used

in

experimentation

will

be given under

the separate headings dealing with the reactions.

T).

Habits^ and Natural History.

In the course of more tlian two years' study of planarians

'

In

tills

section the

activities of the

word

" habit " will be

used to signify merely

tliose

organisms which are frcquontly ol)served to occur under

VOL. 46, PART 4.

NEW

SERIES.

M M

RAYMOND PEARL.

526

numerous observations have been made on tlieir general

natural llistor3^
It is thought desirable to present a general
account of this here for two reasons first, because there is no
:

adequate discussion of the natural history of the fresh-Avater

triclads in the literature; and furthermore, because it will
bring out prominently
seeking an explanation.

problems with which

I.

the

phenomena

for

In other words,

this study has

had

it

which we are
present the

will

to do.

Occurrence and Distribution.

The species of Planaria (maculata, dorotocephala, and

gonocephala) used in this study have been collected mainly
from the Huron River near Ann Arbor. They are found, for
the most part, on the under surfaces of stones in places where
the current is of moderate swiftness, and the substrate on
which the stones rest is rather soft. They are also found
among the fronds of such water plants as Ceratophyl lum
and El odea, although less abundantly than on stones. I
have obtained these species only very I'arely in collections
fi'om ponds and small pools of stagnant water.
They appear
to be, in general, much more abundant in shallow water than
in deep.

Rhabdocoeles I have found in great abundance in small

ponds and pools of stagnant water, and, with the exception
of Stenostoma leucops, almost never in running water.
Dendrocoelum, sp., is also much more abundant in stagnant
Avater than in streams.

There

is

Planaria

no marked seasonal distribution of the species of

studied.
They appear to be slightly more abun-

dant in the

fall than in the spring. I have found no evidence

any migration into deep water during the winter in the
case of these forms, as has been described by Child (: 01, pp.
978 981) as occurring in Stichostemma. The seasonal

of

distribution of

well marked,

conditions, williout necessarily implying Uie same idea as that embraced

term " habit " as used by the psychologists.

iiat.ui-al

in tlie

Dendrocoel um,sp., appears to be

MOVEMENTS,

OF PHESH-WATKB PLANAEIANS.

ETC.,

individuals being found in considerably greater

527

numbers

the spring than at any other time in the year so far as

in

my

This seems to be true also of the rhab-

observations go.
docoeles.

Relatively the most abundant species of planarians in this

region aie

numbers

Planaria dorotocephala and maculata.

The

two are about equal, with a slight advantage

in favour of P. dorotocephala.
Next in abundance I have
found to be P. gonocephalaj but this is considerably below
the other two. Finally comes Dendrocoelum, sp., which I
have never found in an abundance to be compared with the
species of Planaria.
of these

II.

The movements

Activities.

of planarians will be discussed in detail in

it is desired to take up here
and relations to the environment
under the general subject of natural

a later section of the paper,^ but

certain general activities

which properly

fall

history.

The
a.

first of

these subjects

Sensitivity.

is

The flat-worm

is

extremely sensitive to

Among the different stimuli which produce specific reactions, and to which we must therefore
conclude it is sensitive, are the following
Mechanical
disturbances of the general environment (shaking, jarring,
a variety of stimuli.

movement

of

Avater,

etc.),

contact

stimulation), chemical changes

in

(localised

mechanical

the environment

(in

the

widest sense, including food substances), light, the electric

current, desiccation, a current of water,
Its

extreme

sensitivity,

which makes

slight changes in the environment,

'

It

may be slated

lieie, for

and heat.
it

responsive to very

mny bo shown by

the conveuience of

llie

a very

reader before reacliinjf

movements, that the progressive movements of

triclads are of two sorts.
These are (a) gliding movements, in which there is
little or no muscular action
and (6) crawling movement?, in which the motion
is effected by muscular contractions involving the wiiole body.
Tlie crawling
the

full

discussion of the

has some general resemblances to the method of progression observed in a

leech of the genus

Clepsine.

528

EAYWOND PEAEL.

simple experiment.

If a dish containing

specimens not

any wa}^ stirred up by rough handling, but gliding along the

bottom, be jarred ever so slightly, every individual will
If only a
instantly stop, contract, and remain immovable.
single jar

pause.

is

If in

thing.

momentary
more strikingly the same
a dish containing water to a depth of not more

given, the

worms will

start after only a

further experiment shows

cm. a single specimen gliding quietly is

is touched to the surface of the water
above or to one side of it, it will be seen, if closely watched,
to give the same momentary pause and partial contraction.
If the needle is pushed down through the water towards the
than

to

1-5

selected, and a needle

any but the quietest and gentlest way the contracted

Only at the moment when quietness in
the surroundings intervenes again will movement be resumed.
I have frequently tried to introduce a needle close beside the

worm

in

state will continue.

animal without causing this momentary pause. With a layer

of water not over a centimetre in depth covering the worm I
have not been able to do this, except in rare instances.
After the point

is

once through the surface film

it

may be

brought nearer the worm without causing a persistence of the

contraction, provided it is advanced in line with itself, i. e.
not slid up laterally. In order to observe this extreme sensitiveness to disturbance of the water one must take care that
the animals have not been violently disturbed just previously.

Any marked

disturbance or persistent, more or less violent

stimulation puts the animals in a condition which may be

called, for lack of a better term, " excited." Such a condition
is

characterised by increased rapidity of

movement and

in-

creased general activity, and in this condition the animals do

not give the " finer " responses, that is, responses to weak

stimuli.

I shall

have occasion to discuss

this

matter in more

detail later.

This marked sensitivity and

its

associated behaviour are

remarkably similar to Avhat has been found by Whitman ('99)

He has further
to obtain in the case of the leech Clepsine.
pointed out that lack of attention to this extreme sensitivity.

MOVEMENTS,
which

ETC.,

OF EKESH-WATEIi PLANAHIAN.S.

529

apparently quite generally distributed aniono- lower

may be an important source of error in work on

is

organisms,

the behaviour.

Regarding the statement of Bardeen (: 01, p. 14) that he

does not find that Planaria " is sensitive to anything but light
and contact," nothing need be said here. The detailed
accounts of the reactions of the organism to a variety of
stimuli

which

iollow

in

this

paper are

themselves a

in

sufficient criticism.

h. Secretion of Mucus.
There is secreted at all times
over the surface of planarians a sticky slime, apparently of
the nature of mucus.
This secretion is increased when the

animal is irritated, and is under normal conditions more

abundant on the ventral than on the dorsal side. If a
needle or fine glass rod is touched several times on the
surface of the body its end becomes covered with this
secretion.
For this reason it is necessary in applyin<)^
localised mechanical stimuli to wipe the

of the needle frequently,

Similarly,

the edge

iP

one

is

mucus

off

the end

order to obtain good results.

using a sharp scalpel to cut the animals, if
in

with the surface of the body an}'

length of time before the decisive cut is made, the edge will
become so coated with mucus that a clean cut is impossible;
is left

in contact

from under the knife.

it leaves behind
a more
or less heavy string of this mucus, so that if several specimens are placed iu a clean glass dish the bottom will, in
a short time, become covered with a network of interlacing
instead, the animal will slip

When

the animal moves about

mucus threads. The same phenomenon occurs in other

Turbellaria and among the Nemerteans (cf. Child,
(Jl,
and Wilson, 00). The threads when first secreted are so
:

transparent as to be invisible, but in larger quantities they

appear opalescent, and may be picked out of the dish with
forceps.

The function of
the body

attach

this secretion iu
to

purpose occur widely

locomotion

the substrate.

among

is

evidently to

Secretions for such a

aquatic organisms.

RAYMOND PEARL.

530

The mucus

also

undoubtedly plays au important part

in

the attachment of the animal to the under side oE the surface

film.
When the worm leaves the surface film in open water,

when it cannot reach any solid body, it hangs by the

mucus thread in much the same way that a terrestrial
mollusc, like the common slug, does when it passes through
i.

e.

This observation I
the air from a higher to a lower point.
have made many times, though generally in an indirect way.

As has been
first

mucus thread is
when a worm leaves the

said before, the

secreted, so that

invisible

when

surface film

it

seems to glide freely through the water. If, however, one

passes a needle horizontally through the Avater immediately
above the posterior end of a worm which has just left the
surface film, it will be seen that at a certain point (where the
needle strikes the thread) the end of the worm will be jerked
Furthermore, one may with care pick up the
with forceps and raise the whole
worm, provided the attempt is made before the anterior end
reaches the bottom. I have seen specimens of P. maculata

to one side.
invisible

mucus thread

back upon the thread after going a part of the way

down to the bottom, and again regain a position on the
The same thing is frequently done by slugs.
surface film.

craAvl

When

the animal has fully reached the bottom, connection

with the thread which has served to suspend

is

it

in the

water

usually broken by several sharp jerks of the posterior end

body from side to side.

The relation of the organism to this slimy secretion is
much the same in the land plauarians, according to the

of the

observations of Lehnert

('91).

He

distinguishes in case of

these forms " Kriechfaden,^' " Briickenfaden,'' and " Gleitenfaden " formed fi'om the slime, the distinction being based
The
on the relation of the thread to the surroundings.
" Kriechfaden " are the threads left behind as the organism
"
moves over a continuous solid body, and the " Gleitenfaden
are the threads on

which

the

animal

hangs

throuo-h the air from a hio-her to a lower level.

forms of threads

have found

in case

in

passing

Both these

of the fresh-water

MOVEMENTS,

ETC.,

OF FRESn-WATER I'LANAIilANS.

531

"

The

Bruckenfadcn," wliicli are formed by the

hiiid plauarians when they pass from one solid body to
another at about the same level, I have never observed in
case of fresh-water planarians, tiiough I see no reason why
under proper conditions they would not be formed. Lehnert
plauarians.

says, " Die

(loc. cit., p. 17)

Landformen ihren
also noted that

Wasserplanarien bilden wie die

Kriecli-, Briicken-,

und Gleitfaden."

Poly cells tenuis was

He

able to crawl back

upon a mucus thread after passing for some distance down

over

it.

Nothing- like the formation of "cysts" froui this mucus,

such as Child

01, pp.

(:

989 to 993), found

Stichostemma, has been observed

Its only biological significance in these

to

in

the case of

in the case of planarians.

forms

is

in relation

movetuent, as pointed out above.

In connection with the subject of

be well

to point

mucus

secretion

it

may

out the tenacity of the attachment of the

fiat-worm to the bottom.

be found in attempting to

It will

dislodge the animal that the extreme anterior end and the

extreme posterior end stick very firmly to the substrate.

Whether this holding is the result of a sucker-like action of
the ends of the body, or is due merely to the stickiness of the
mucus, I have been unable to decide. It is easily possible
that the muscles could be so contracted as to form out of
either end of the body a practical sucker, but whether this is

done or not

it

is

impossible to say.

Woodworth

described a permanent anterior adhesive disc in

('97)

has

Dendro-

coelum lacteum, but considers that this "is not a true

sucker, nor does the animal employ its anterior end for the
purpose of attachment to any greater degree than the
posterior or lateral margins of

surface

of

which

its

body, along the ventral

numerous mucus glauds have their

it is the margins and posterior end that

openings. In truth,
adhere more firmly to a support often when the animal is
forcibly removed from the sides of the aquarium the parts of
the margin or the posterior end will adhere so firmly to the
;

glass that the

points

of

attachment

are

drawn out

into

P^'^

532

KAYiMOND PEARL.

digitate processes."

I incliue to

the view that in

Plan aria

the mucus which attaches the organism to the support,

although it must be said that the jippearance is at times

ib is

strikingly as

if

posterior ends acted as

the anterior and

suckers.
c.

Periods

of

There

Activity and Pest.

is

in the

case of freshly collected planarians a certain periodicity in

First, there is the rather marked difference
amount of activit}^ in the night aud day. It has been
stated by a number of investigators that planarians were
probably nocturnal in their habits, i. e. more active at night

the activities.
in the

This can easily be seen to be the case

than during the day.

in the

following way

In a dish

of planarians, together with

tophyllum,

usually

seen during the day.

number
material like Cera-

containing a large

some plant

comparatively few specimens will be

Nearly

all will

be in among the fronds

of the plant material in a quiet condition.

comes

(in case of

however, one

winter days) there has been two and a half or

more hours

of darkness, a large

number

found on the sides and bottom

movement. Again, one will frequently

will be

and bottom

of the

of these will

among

aquarium dish

have disappeared,

in

are

the specimens

find in the
all

at rest.

or,

the plants, where they

of

of the dish in active

that the specimens are scattered about

in

If,

into the laboratory at 8 p.m. or later at night, so that

over

morning
tlie

sides

By noon numy

other words, gone

protected from the

light.

Besides this day and night periodicity there

is another
during the day,
at any rate, they seem to be incapable of continuing
movement more than a certain, not very great, length of

fact that

may be mentioned;

Then

this is that

must intervene. Thus one may

which has been moving about come to rest,
and after a length of time, varying from a comparatively few
minutes to several hours, it Avill start into spontaneous movement again, and repeat the Avhole cycle over and over. It
seems that the periods of ({uiet are really for the })urpose of

time.

a period of rest

see a specimen

MOVKMKN'I'S,
resting,

i.

e.

E'I'O.,

OF FRKSll- WATKi;

the aniiua-l seems to

own movements.

Tin's

is

])e

533

I'l.ANAK lANS.

(luickly fatig-ued

indicated by the fact that

by its
one

it'

up a specimen, and sets it into activity again just as

soon as it comes to rest, the periods of spontaneous activity
will become progressively shorter, until finally the wcn-m will
only move a very short distance before coming to rest again.
The periods of activity are longer and more frequent in
P. dorotocephala than in any of the other species I have

stirs

studied.

Formation

d.

of Collections.

Tlu'rt; is a

well-marked

tendency for specimens of planarians to form well-defined

groups or collections when they come to rest on an open

surface like the bottom or sides of a glass dish, or on the
under side oC rocks, under natural conditions. Of course.

C.^

I'lG.

1.

Diagram

bliowiiig

he

apjjcaraiice

of

a culled ion of

ifbUiig

jilaiiariaiis.

this

is

in part a result of tlieir reaction to ligld, as

noted by Loeb

('04).

has been

Besides this there seems to be some

other factor at work, for in the same dish one fre([uently

finds several localised collections

diameter

from one

in different parts of the dish.

to

two inches

in

In these collections

may be closely packed t(.)gether, and with

some specimens overla]iping and lying partly over others, yet
in the species I have studied a looser arrangement of the
character shown in fig. 1 is the more usual one.
On the
under surface of stones such groups arc frequently seen
two or three nniy bi^ ft)uud on the same medium-sized stone.
the specimens

In this case light as a factor cannot be present, since the

couditions of

all

with reference to this stimulus are equal.

RAYMOND PEARL.

534

We

have, then, here a case of what appears superficially to

be " social instinct."

Movement on Surface

e.

flat-worms and a

number

Film.

As

is

well

known,

of other animals frequently

move

about on the under side of the surface film at the top of the
water.
On account of the flexibility of the support, motion
under these conditions is very slow, and usually, after having

been on the surface film for a short time, the worm will
loosen its hold and pass down to the bottom in the way which
has been described above.
The worms do not remain
customarily in the angles formed by the surface film with the
side of the dish, as does Stichostemma (Child, :01), but
instead pass out at once on to the free surface. Further, the
flat-worms never push through the fllm at the side of the

and pass up out

dish

The occurrence

of the water as the

nemertean does.

on the surface

film is not the

of planarians

result of any thigmotactic reaction (using thigmotactic in the

sense ordinarily understood), but

simple reflex act, and

is tlie

is

brought about by a

result of the configuration of the

surface of the water and the side of the dish.

brought out

in

more

detail later.

It is

This will be
probable that fresh-

water planarians, in their normal habitat, very rarely take up

a position on the surface fihu.
Among other organisms
(Entomostraca, Hydra,

etc.)

this habit

probably has a much

than in planarians (cf.

When on the surface film the
Scourfield, 'Of,
00, :01).
worm behaves in nearly every respect as it does when on the
bottom. The head is frequently raised (with reference to
the worm) and waved about in the water just as occurs in
greater

biological

significance

That the situation is a more or less

abnormal one, however, is shown by the fact that, so far as I
have observed, the worni never comes to rest on the surface
the normal movement.

film,

but instead, always keeps in active movement

leaves

till

it

it.

The means by which the animals maintain

their position on
undoubtedly the mucous secreThis is very sticky, and holds
tion from the ventral surface.

the under side of the film

is

ETC., OF FRESU-W/Vl'ER

MOVEMENTS,

PLANA HIANS.

535

animal to the film, the surface tension being sufficiently

great to support a considerably greater weight than that of
tlio

flat- worm.

note in this connection that the land

It is interesting to

move about on the top

planarians are able to

film

water to

of

extent

a limited

The immediate means

p. IG).

(cf.

of the surface

Lehnert,

loc.

cit.,

of support here, as in the case

of the fresh-water planarians,

is

the mucous secretion.

of the

mucus

thread described above apparently does not take place

if it is

The leaving
possible for the

way.

Before

surface film

of the

same

result to be accomplished in any other

worm

occurs the

it

by means

usually

the

stretches

and turns it in all direcwith

something solid the
If it comes in contact
tions.
anterior end becomes attached to this, and pulls the posterior
end of the body away from the film. If nothing solid is
anterior end

down

within reach the

into the water,

worm

Avill

usually, after a time, drop

down

on a mucus thread as described.

III.

Food.

Planarians will take almost any sort of animal food very

have used mainly,

readily,

crushed

pieces

Planorbis, etc.
shell and placed

in

the feeding experiments,

fresh-water molluscs, such as

of

One

of these molluscs,

a dish centainiug a large

in

Physa,

removed from the

number

of

planarians, will, in a short time, be literally covered with the

worms

feeding.

If a

worm

is

gently lifted

off

the

])ile

the

greatly stretched pharynx will bo brought into view. The

worms will eat any other kind of animal tissue (fresh meat,
parts of insects, pieces of fresh-water worms, etc.), so far
as I have observed, tlu; only condition being that the meat
must be fresh. As will be shown latei", the juices from the

food act as chemical stimuli, so that

tissue be crushed or bruised so that

the water.

though

still

pai'tially

able to

it

its

is

necessary that the

juices can escape into

crushed specimen of PI an aria, even

move about,

will

be seized upon and eaten

RAYMOND

536

PEAlil;.

as quickly as any other food.

specimens thus eaten.

patience, to

make

posterior end of

It

is,

liavu several

in fact, possible,

tiiues seeu

with a

little

a specimen eat a small piece cut olf the

own body

This eating of each other

have observed, unless an individual
is bruised so that some of the tissue underlying the epidermis
is exposed.
Under these conditions juices escape from the
body and act ;is stimuli on the other worms. Under normal
conditions contact of one individual with another does not
start the feeding reaction, Avhich is a purely reflex phenomenon, cispablo of being started only by a certain set of
stimuli.
Promiscuous cannibalism, such as Child (:Ul)
suspects to occur among individuals of Stichostem ma, I
have seen no evidence for among the Turbellaria.^
In the feeding the worm lies fully 'distended, with the
posterioi' two thirds of the body on the meat, or whatever
else is being used for food
the pharynx is extruded,
its

does not occur, so far as

frequently to nearly half the length of the body, audits end

attached to the meat.

end

of the

worm

is

the piece of food

is

During the feeding the very anterior

attached to the bottom of the dish, provided

is

make

not so large as to

this impossible.

Besides the animal food which the worms will take so

some extent, feed on

have not been able to induce
the typical food reactions (to be discussed later) with vegeThe evidence for the statement that
table material.
food
is
used by planarians is of two sorts (o)
vegetable
specimens a,re frequently found extended on the stalks of
water plants, with the pharynx extruded and attached to the
stalks; and [b) the faeces which have been observed immediately after defecation have been found to consist largely of
It would appear, however, that
finely divided plant tissue.
readily, they also normally, probably to

vegetable mattei', although

'

Biirdccu

ones.

Ill

upon the

(:

01, A,

p.

176) says, "Stroiii,^ planaiiaiis oficu

such instances the strong individual

boi]y of tlie

weak one, usually near

al

tiic

\n-c.y

upon weak

its

pharynx somewliere

liead."

I iiavc never seen

laches

even the largest specimens eat smaller ones unless these latter were bruised
in

some way.

M(WrL\[KN'l'.S,

KTC, OF FRRST[-\VATKIi PLANARTAXS.

veg'ofcable food is not alone sufficient to

good condition^

for

specimens kept

plenty of living plant

material,

in

keep

tlie

animals

The food

is

is

in

an aqnarinm dish with

on which they stay the

greater part of the time dni'ing the day, will steadily

smaller niiless animal food

53/

grow

given them.
rate softoiied, and

in part digested, or at an}'

physically changed ontside the body.

piece of mollusc,

on which a nnmber of worms have been feeding for some

time, has a white, Huffy appearance, similar to that of meat
'J'his is ajiparently brought
after partial gastric digestion.
about by a secretion poured out of the end of the pharynx.
The necessity for some such action is apparent, because the

flat-worm has no teeth or other means of separating a portion

tough fibrous tissue off from a mass so that it can

This can be done, liowever, if the mass is first
softened and partially dissolved.
There are certain other
of ordinary

be swallowed.

evidences that a secretion

is

during the feeding process.

poured out from the pharynx

These will be taken up in

ano-ther connection.

After the worms have fed undisturbed for a certain length

of time they will leave the meat, and, after a short period of

come
The worms

activity,

time

(at

least

continually

to rest.

are able to live for a considerable length of

two months)

grow smaller

without food, although

dui-ing this time.

the}-

This marked

decrease in size while starving has been noted by several

observers, and especially studied by Lillie (:00).
This

author finds that the decrease in size

simplification

of structure

sort of

accompanied by a
"development back-

is

ward," such as has been described by Patten

normal embryos of Limulus.

('06)

for ab-

IV. Defecation.

The process of defecation has been observed by Bai-deen

The process consists of three or four general con-

(:0l).

tractions

involving

the

whole

body,

during

which

the

RAYMOND PEARL.

538

coutents of the intestine can be seen to be in rapid motion.

Soen

after the beginning of the contractions, Avhich are in

character different from any other of the

movements

of the

have observed, and which cannot be adequately

described, the intestinal debris is shot out of the pharynx.

body which

The
out

force of the expulsion

of the pharjaix.

times

is

so great that the f^ces spread

the water some considerable distance from the opening

in

apparently

have observed the process only a few

it

occurs only at infrequent intervals.

Summary

V.

From

of

Factors

in

Behaviour.

the above sketch the behaviour of the flat-worm can

be seen to be of considerable complexity.

show many variations

animal

in character, rate,

and

The movements
direction.
The

shows apparent preferences for certain situations

while avoiding others.

It reacts to a variety of stimuli in

ways which, on the whole, further its preservation and wellbeing perhaps as well as if guided by careful thought. It
chooses its food, taking certain things and passing by others.
It forms gatherings of a sort which apparently indicate that
the flat-worm prefers to be in the company of his fellows

seems to have something of "social

instinct."
On the whole, as the further analysis will show,
it fits itself to its environment by its activities in a way
which would not be discreditable to a being possessed of
considerable powers of reasoning.
Our problem now is to analyse, as far as possible, this
complex behaviour into its component factors. Each activity
will be taken up in detail and subjected to thorough scrutiu}',
to determine its essential nature, and whether it may not be
With this analysis comresolved into simple components.
pleted, it will be possible to assign the organism a definite
in

words,

other

position

in

the

it

objective

psychological scale.

internal psychological factors

objective

criterion

we

purpose of the paper

is

shall

those

of

With the

which there

not attempt to

to furnish the data

deal.

is

no

The

which may be

MOVRMENTS,

ETC., OP

FRKSH-WATKR TLANARIANS.

539

obtained by an objective study of the phenomena precisely

what these imply as to internal factors would doubtless be a
:

subject of dispute

E.

among

psychologists of different schools.

Normal Motor

Activities.

Under this heading will be included all the purely motor

This will include the
phenomena of the organism.
movements (without refei-ence to special reaction to stimuli),
the coming to rest^ and the general resting condition of the
organism.

The movements

natui*ally

fall

two categories;

into

locomotor movements, and

[h)

movements

and expansions and the

as contractions

I.

like.

Locomotor Movements.

As has been mentioned above

(p. 19), thei-e

are two sorts

of locomotor movements, the gliding and the crawling.

gliding
slips

ripple

is

(a)

non-locomotor, including such

The

the smooth, even motion by which the flat-worm

about over surfaces Avithout showing a perceptible

of muscular movement.
This is the characteristic

movement when the organism is not particularly stirred up.

The crawling is the characteristic movement when the animal
or has been recentl}', strongly stimulated.
muscular movement.
is,

a.

ming " by Bardeen

in all of

my

a purely

The movement which I have called gliding

the same as that which has been called " swim-

Gliding.

is apparentl}''

It is

(loc. cit., p. 15),

yet

it

must be stated that

observations on a very large

number

of plana-

have never seen anything corresponding to some of

the details which this author mentions in this movement.
In the first place, he speaks of the worms moving progresrians I

when not in contact with a solid body, i. e. of a

movement freely through the water. This I am unable to
sively

understand, as I have never seen the slightest indication of

the organism moving

freel}'^

in the

water without contact

with a solid body or something which served the purpose of

RAYMOND rEARL.

540
a solid

(viz.

The

the surface film).

only possible exception

passage of the animals from the surface film

to the bottom on a string- of mucus, as described above.
to this is the

Furthermore, so far as

can find

in

the literature, no one

else has ever seen a fresh- water triclad

swim

freely

body

in

through

the water.

The movement takes place

witli tlie

contact with

There is,
between the ventral surface of the body and the
surface on which it is moving, the thin layer of mucus which
Ifc
is in this mucus layer
is constantly being secreted.
rather than the free water that the cilia beat.
This gliding movement is, so far as I have been able to
ascertain, brought about by the action of the cilia on the
ventral surface.
There may also be some very slight
muscular movement of the ventral body-wall comparable to
that in the foot of a mollusc like Physa, which assists in the
a surface either of a solid or of the surface film.
of course,

locomotion
exists at

but in the case of the flat-worm this factor,

all, is

vei*y insignificant.

Only

in a

if it

few instances

been able to satisfy myself that any such movement

was taking place, and then it did not have the characteristic

have

rhythm seen

has any effect at all

must be an extremely slight one.
The cilia beat strongly backward, i, e. towards the posterior end of the bod3\
I have not been able to induce any
reversal of the ciliary beats in these ventral cilia.
Bardeen
(loc. cit., p. 15) states that when the head is suddenly drawn
back from some object the movement of the cilia on the
antero-lateral margin of the head is reversed, and further
suggests that " this i-eversed action may possibly be set up
by the mechanical friction of the water." It would appear
that the suggestion is the correct explanation, and that this
in a mollusc.

on the gliding movement

is

If this factor

ifc

not a true reversal of effective beat.

The

cilia

which are mainly

effective in

producing the

movement are distributed on the ventral surface of

There is a band down the
the body, as shown in Fig. 2.

gliding

centre of the body, which widens out at the anterior end so

MOVEMENTS,

ETC., OF

as to cover nearly

tlic

FRESH-WATKR TLAXAIUAN.^.

Avliolc of

541

the ventral surface of the

The beat is the strongest down the median line of

band, and diminishes in intensity towards either edge

head.
this

until at the mai'gins there is

no ciliary movement at

all.

At

the anterior end the cilia near the side of the head beat

backwards and
line, so that

at the

same

tiaie

inward towards the median

the currents take the course indicated

arrows in that region in Fig.

The

2.

by the
and action
powdered indigo

distribution

made out by stirring finely

and then either directly observing the ciliary
action on these suspended particles as the animal glided on
the surface film, or by indirectly observing it in a mirror
placed below the bottom of the glass dish in which the
worms were. Both of these methods gave the same results,

of these cilia were

in the water,

Dingrani of the ventral surface of Plaiiaria, to sliow the

Fig. 2.
distribution of cilia. The stijjpled area is that, which bears cilia.

The arrows
pharynx

is

indicate the direction of the ciliary currents.

(The

omitted for the sake of clearness.)

and showed very

clearly the distribution

of the

effective

cilia.

have found no evidence of

on the dorsal
head there
are cilia, but in other parts of the body, either on the dorsal
surface or the edges, I have found no evidence of their
I

surface of the body.

presence.
of a

This

worm
is

in

ciliary action

Around the margins

Particles of indigo dropped on the dorsal surface

will

remain

in the

same place

of

is

for hours at a time.

striking contrast to the conditions in the land

planariaus as described by Moseley

surface

of the

thickly covered Avith

cilia,

('77), Avhere

the dorsal

which serve the purpose

keeping the body freed of foreign matter.

In the gliding movement the head

is

raised slightly from

the bottom so as to form an angle with the rest of the body.

VOL. 46, PART 4.

NEW SERIES.

NN

RAYMOND PEARL.

542
This position

is

shown

in

Fig.

head is
and apparently its elevation

o.

As

will

be brought out

a particularly sensitive portion of the bod}',

later, the

is

related to

its

sensory function,

head into close relation with

The head is not held in a fixed
a large environmental field.
raised position, but is in constant though slight movement
whenever the animal as a whole is moving. These "feeling"
movements ('Hastende Bewegungen ") of the head are very
characteristic.
The head as a whole is raised and lowered,
and turned from side to side, while at the same time the
antero-lateral margins are moved up and down and extended
and retracted. These "feeling" movements of the head
region are usually very slight, and escape notice except
When the organism is much
under close observation.
stirred up, however, tliey may become quite apparent.
in that

it

practically brings the

Their purpose

is

evidently to increase the chances of re-

ceiving stimuli, so that any stimulus in the neighbourhood

Fig. 3.

may be

Diuj^rammalic

quickly

side view of a gliding plaiiaiiaii.

received.

Constantly

surfaces are presented to the environment.

acts in
is

movement

different

sensory

The head region

as a single great tentacle-like organ which

constantly testing the environment as the animal pro-

At the same time

tlie auricles are fully extended and

do not think that this marked sensory activity
functions so much for the protection of the organism against
harmful environmental influences as it does to give prompt
for example, stimuli due to the
notice of useful stimuli,
presence of food material. In general it would not appear
that such an organism as the flat-worm runs as great i*isk of
elimination from enemies as it does from not finding food
In the ctenophore Mnemiopmaterial for its own support.
sis Leidyi, whose reactions I have studioLl, no trace was
observed of a reaction adapted to the purpose of getting the
organism out of dangerous surroundings, but its only specific

ceeds.

raised.

MOVEMENTS,
reaction

ETC., OF FRRSH-AVATEIJ

one which would

bring

PLANARIANS.

543

it

towards any food

material which miglit be encountered.^

Wliile^ as will be

shown
which

is

later,
is

there

adapted

is

in the

case of

Plauaria a

reaction

to getting the organism, out of danger, yet

weak a stimulus as is the food

evidently for the purpose of receiving
stimuli of the lowest intensity that the "feeling" move-

it is

not called forth by so

reaction,

and

it

is

ments are adapted.

In addition to the slight "feeling movements" of the
head, described in the preceding paragraph, the organism
frequently in the course of its gliding raises the whole
anterior part of the body off the bottom
in the water.

The portion

of

and waves

the body so raised

it

about

may

in-

whole anterior half.

The gliding is usually
entirely stopped or very much decreased in rate while these
waving movements are taking place. The head is swept
from one side to the other and raised high in the water,
covering a considerable area. This movement is also undoubtedly for sensory purposes.
clude

the

In the gliding movement the body back of the head is kept

an approximately straight line ; that is, there is no sinuous

in

bending of the body such as

Stichostemma
movement

of

(Child, loc.

is

cit.,

the earthworm.

observed, for example, in

p. 981), or at

Furthermore,

times in the

have never
observed any regular undulation of the margins of the body
during movements such as take place in case of many poljI

e. g.,
Leptoplana tremellaris (cf. Lang, '84).
Bardeen (loc. cit., p. 15) seems to imply that such motions
occur, and are an aid in the locomotion, but I am unable to
confirm this statement. There are, of course, slight movements and changes of contour of the margins of the body,
but they are not of a prominence or character to warrant
thinking that they in any way contribute to the propulsion
In fact, it seems more probable that they are
of the animal.
in part passive results of the motion of the whole body, and

clads,

'

brief preliminary account, of tlie reactions of

publisiied in

'

Science,'

N.

S., vol. xii,

No. 311,

Mncniiopsis

pp. 927, 02S,

00.

has been

RAYMOND PEARL.

544

in part tlic expression of local

changes in the tonic contrac-

tion of the muscles.

In the gliding

movement the body

is

the surface along which the animal

in close contact

with

When

moving.

is

an

animal passes from the resting condition into movement one

can see the body lengthen and flatten so as to hug the
By observing with a compound microscope an
surface.
animal gliding along the vertical side of a dish so that the

brought sharply into view, the closeness of the conmargin of the body with the surface can be well
Furthermoi'e, in specimens in which the posterior part
seen.
of the body has been split longitudinally in the middle line
to a point just behind the head, it is found that the half of
the body which is determining the direction of the moveedge

is

tact of the

ment

always in close contact with the surface, while the

other half only lightly touches it.
It would appear from all the observations which have been
given that the gliding movement is brought about in the
is

The ventral surface of the body constantly

mucus in greater or less quantity. This mucus can
be shown experimentally to be very sticky immediately after
As it is secreted under normal
it is secreted into the water.

following Avay

secretes

conditions

animal

is

it

immediately sticks to the surface on which the

reposing.

Thus there

will

the animal and the surface on which

mucus which

is

be constantly between
it

moving a

is

adherent to the substrate.

the lowest part of this

mucus layer where

layer of

We

can think of

is

stuck to the

it

upper layers which

In
this
upper
layer of the
the
animal.
with
contact
in
are
and
constantly
pushing the
beating
are
mucus the cilia
surface as of denser consistency than

its

animal forward. Of course, what really takes place is that

the cilia are pushing the secreted mucus backward, but as
this layer of
it is

mucus becomes

fixed to

secreted, the practical result

ahead.

'IMiis

relation

side view of a gliding

is

shown

worm

is

tiie

substrate as soon as

that the animal

in Fig. 4.
is

on the ventral side of the organism

the substrate
;

and

13

the

is

pushed

represents a
;

the cilia

mucus

secre-

MOVEMENTS,
tion,

ness.

represented

OF FRESII-WATER PLANARIANS.

disproporfcioiiately

exaggerated iu

545

tliick-

This sticks to the surface of the substrate, and the

backward beating
].

KTCI.,

of the cilia drives tlio

Rate of Gliding Movement.

marked

worm

ahead.

There

difference in the rate of the gliding

case of the species of l^lanaria studied.

no

is

very

movement

On

in

the whole,

specimens of P. dorotocephala move more rajjidly than do

those of the other two species, but there arc large individual
Active specimens of Dendrodifferences in this matter.

coelum, sp., move much faster than any other planarians I

have observed. Large specimens of this form will sometimes
glide along with simply amazing rapidity, not showing the
slightest tremor of the surface of the body.

As

to the

absolute rate of the crawling, some statistics

yi^jy///f//Y///r7T//Tf/rr/r>J!:r//7Tt/^//-///:^/^7/,:

Fig.

4.
Diagram Iu show \\\v. niecliauism of t.lic gliding movenient. A
represents a specimen of I'laiiaria seen from t,lic side
W, the layer
of nniciis secreted by tiie animal.
(This layer is represented as
greatly exaggerated iu lliickness in |)roportioii lo the animal.)
C,
cilia.
I), the subslrafe.
The arrow indicates tJic direction in which
the organism is moving.
For further explanation see text.
;

have been collected and will be presented. The statistics

were obtained in the following way
A paper was ruled into
centimetre squares; over this was placed a Hat Petri dish
:

containing the Avorm to be tested.

of P.

maculata were

Normal

active specimens

used, and nothing was put into the

dish but fresh clean water.

at night,

and the source

The experiments were performed

of

illumination

Avas

a 16-candle

power electric light enclosed within a ground glass globe.

This lamp was 35 cm. above and 35 cm. distant iu a horizontal direction from the centre of the dish, so that the light

struck the animal at an angle of approximately 45 on

dorsal surface.

The worm was allowed

to get into

its

an even.

RAYMOND PEARL.

546
uoruial glitle^

and then

come around, as it usually would in

was headed in the direction of the

to

sliort time, so that it

light.

Then the time which

it

took the

worm

to glide three

centimetres was taken by means of a stop-watch.

animal started crawling, or abruptly changed

The average

the trial Avas ruled out.

second determined in this

individuals

is

1'34.

is

trials

on two

considerably higher than

those obtaiued by I'arker and Burnett

cephala.

If the

direction,

rate in millimetres per

way from twenty

This rate

its

(:

00)

for P.

In that form they found a rate of 1"04

gono-

mm, per

moving toward a horizontal light; 1'12 mm. per second when movement was away
from a horizontal light, and 1'08 mm. nev second when the
animals were moving under a vertical light. There seems to
be a well-marked correlation between the size of individual
and the rate of gliding', as would be expected on general
grounds, and is apparent from merely qualitative observations
on the movement. One of the specimens from which observations were taken was 11 mm. long when extended, and its
rate of gliding was 1"48 mm. per second; while the other
specimen, which was only G mm. long when extended, showed
a correspondingly slower rate of l'2-3 mm. per second.
The
statistics are, of course, very meagre, and are not offered for
any other purpose than to give a concrete idea of the approximate rate of the gliding movement. A thorough quantitative
study of this matter of the rate of movement in planarians
and other related organisms, and of the effect of different
agents on the rate, would, I believe, be very interesting, and
second

in

might lead

the

case of

individuals

to valuable results.

hope

to

be able to make

such a study at some future time.

Lelinert

(loc. cit., p. 17)

gives a table of the rate of move-

ment (presumably

in

rate of the gliding

movement)

the case of the fresh-water forms the

lie gives no account of

how

of several species of flat-worms,

the data were collected, but his

may be inserted here for the sake of comparison. His

Bipalium kewense and B. kewense viridis are
Usual rate, 1 to 1"33 mm. per second; occasional rate,
values

rates for

MOVEMKNTS,
l"8o
for

ETC., OF FI{ESH-WATElt I'LANARIANS.

mm. per second. Tliis agrees

maculata given above
s

mus

L i 1 i n e a t u s and

considerably slower (0"5

mm.

1*33
.(1"66

2"5

very closely with the rate

P.

G eo d

per second

mm.

rates

for

a c t e u ni are

per second and 0"75

respectively).

1"83 mm. per second), Planaria

mm. per

His

(lo4).

D e u d r o c oo

O'GG

547

Poly eel is

tenuis

polycliroa

(2*16

mm. per second), and

mm. per second) show a

second, exceptionally 3*33

Mesostomum tetragonum

(2*GG

markedly faster rate than the forms I have studied.

Regarding the effect of different agencies on the rate of the
gliding movement no special study lias been made, and I can
Such a study
only report a few incidental observations.
should be made by exact quantitative methods, and this I
have not had the opportunity to do. What the effect of light
on the rate is, it seems to me, impossible to say with entire
certainty.
Cole and myself^ have found that light of great
intensity (that obtained from a projection lantern with an
electric arc

as its source of illumination)

causes a definite

increase in the rate of gliding, but this increase has not been

The

measured.
to

answer

results oE

Parker and Burnett do not help us

this question of the effect of the intensity of light

on gliding, as they are concerned only with the direction of

its rays.
The well-known phenomenon of " Unterschiedsenipiindlichkeit " for light which Planaria shows (Loeb,
'93, et al.)
would indicate that increased light causes
increased rapidity of movement.
The electric current causes
a very marked diminution in the rate of gliding

in the weakest
which affect the organism at all. The effects of
chemicals on the rate of gliding are not altogether uniform.

intensities

Solutions of

all

chemicals tried with this point

in

view,

when

above a certain strength, caused a marked diminution in the

rate of the gliding, or else an entire inhibition of it, and the
The action of
substitution of some other form of movement.

weak
weak

Very
AVeak sugar solutions
as rate of movement was cou-

solutions varied with the different substances.

acids slightly increased the rate.

had no observable

effect so far
'

Unpubliahed observations.

RAYMOND PEARL.

548

Copper sulphate causes an entire inhibition of the

in moderately weak solutions, even when
Weak mechanical stimuli
these are not immediately fatal.
applied at the posterior end of the body cause a slight
increase in the rate of gliding, but this is not marked, as any
decided stimulus in this region of the body causes the crawl-

cerned.

gliding*

movement

ing motion to supervene.

It is probable that the various agents affect the rate merely
by causing changes in the general tonus of the animal. There
is much evidence to support the view that the rate of gliding

an individual

of

is

a direct function of

its

tonic condition.

Thus in the resting condition, which is characterised by a

general lowering of the tonus, as will be brought out later,
there is little, if any, ciliary movement. Again, after operations Avhicli result in lowering the tonus, the gliding

is

very

slow or entirely absent.

Direction.

2.

my
to

The direction

of the gliding

observations go, always forward.

make the animal

with

the

st em

ma.

finding of
It, of

glide backward.

Child

01)

(:

in

is,

so far as

have never been able

This is in agreement
the case of St idle-

course, indicates that the effective beat of

In the case of the planarians

cilia cannot be reversed.
on which this work was principally done, a lateral change
of direction of movement is not brought about by the
stronger beating of the cilia on one side. In other words,
when the animal turns to one side it does so by a muscular
the

bending of the body in that direction, and not by ciliary

In an undetermined species of triclad, however, I
action.
found that the most usual method of turning towards one side
was by the stronger beating of the cilia on the opposite side
of the body.
As an individual was gliding along the bottom
of the aquarium dish it would swerve off at an angle to its
former course without bending its body in the slightest
observable degree.
b.

Crawling Movement.

motor activity, the

muscular movement.

The

crawling

second

movement,

It takes place only

form
is

of

loco-

distinctly

when

the animal

MOVEMENTS,

Las beeu stimulated

The crawling
is

movement
which

is

tlic

is

much

less

always induced when the posterior end of

The

strongly stimulated.

are as follows

half

is

gliding.

If

worm

touched with a needle the

immediately contracts longi-

body

the

of

characteristics of the

the posterior end of a

gliding smoothly along

posterior

and

certain ways,

in

frequent occurrence than

the body

549

OF FUHSII-WATEK FLANAUIANS.

ETC.,

is

an instant later the anterior end stretches out far

Then there is a longiin front and fastens to the substrate.
tudinal contraction which begins just back of the head and
tudinally

draws forward the

and as this comes forward and
gets a hold on the surface on which the animal is crawling,
the anterior end is again extended far in front and attached
This, of course, at once

runs posteriorly.

posterior part of the body,

This process

to the substrate.
settles

down

tially in

repeated until the animal

It consists essen-

into the regular glide again.

a stretching out of the head followed by a pulling

body forward by an active muscular contraction.

the

of

When

is

the

tion of the

animal

is

very

body posterior

part in the crawling after the

fact, the posterior half

of

stimulated

strongly

to the
first

the

the

por-

pharynx usually takes no

general contraction.

body

In

may even be held

between the
head and the origin of the pharynx is actively expanding and
contracting and sending the body ahead. These strong expansions and contractions of the anterior end which make up
slightly raised off the bottom, while the region

the

crawling movement

may

each other in rapid

follow

succession as described above, or there

may be a

considerable

between one contraction and the next. In this interval the body as a whole keeps moving ahead as a result of

interval

the ciliary action

that

is,

the gliding

movement continues

during the crawling, so that the latter may be regarded as

an additional movement for the purpose of advancing the
animal faster than the gliding alone can do it. Tiie crawling
may take place, however, with the ciliary beat entirely
stopped.

The duration

of the crawling-

movement

after

it is

induced

RAYMOND PEARL.

550
is

usually rather short.

single strong stimulus at the

such, for example, as given by

through the body
not usually cause

posterior end of the body

is

will
running a needle
more than three of the strong contractions of the crawling
nioveraentj and then the animal will relapse into the usual
glide.
The limits in this matter I found to be from a single
contraction and expansion as a minimum up to six or seven

as a

maximum.

stimulus only.

This

By

is,

o course, in response to a single

repeating the stimuli the animal

may be

made to continue the crawling indefiuitel3^

The effective rate of this form of progression is faster than
The crawling rate of one of
that of the gliding movement.
the worms used for the measurement of the rate of gliding
(the specimen 11 mm. long) was measured in the same way
The worm was stimulated Avith a
as was the gliding rate.
needle at its posterior end just enough to keep it crawling,
The
i. e.
prevent it from settling into the regular glide.
average rate of the crawling was found to be I'GO mm. per
second. Merely qualitative observation shows that the worm
gets along somewhat faster in the crawling than in the
gliding.
1.

Direction.

The

crawling

may take

place so as to

The backinduced by very strong stimulation of the

It does not alwa3's occur tiven
anterior region of the body.
after such stimulation, there being apparently some incarry the animal either forward or backward.

ward crawling

is

dividual differences
factor

which

among

partial desiccation.

allowed to dry,

it

One
backward crawling is

the specimens in this respect.

will call forth persistent

If the dorsal surface of the

will

organism

is

attempt to crawl backward violently.

of the backward crawling is jnst the reverse

when the animal moves forward. The
which
obtains
of that
then
posterior end is extended and fastened to the bottom
a wave of contraction, starting in this case from the posterior
end, draws the remainder of the animal backwards, and then
The backward crawlthe posterior end is again extended.

The mechanism

ing

is

usually induced

when

the

worm

is

excessively stimu-

MOVEMENTS,

OF FliliSH-WATER

ETC.,

lated or iujurcd at

tliu

anterior end.

always occurs when the head

induced

in

cut

is

551

J'L.\NAUIA?sS.

This moveineut almost

off,

aud may usually be

such decapitated specimens for a considerable

period after the operation by stimulating the anterior end.

The backward
ment forward.

crawling*

posterior end

is

is

not so rapid as the same move-

The reason

for this appears to be that the

unable to take so firm a hold upon the

The backward

bottom as does the anterior end.

crawling"

is

coming to
a backward

usually not very long continued, the animal soon

rest.

The

inability of the

animal to glide

in

course, be noted
this connection.
Strong chemical stimulation of the anterior end will cause
Ijight, so
the backward movement to appear in some cases.
far as I have observed, will not, nor will the electric current.
.There is considerable variation as to the appearance of this
backward crawling. Some individuals cannot be induced to
do it at all, or only in a very slight degree, while others will

direction should, of

in

crawl backward for considerable distances after injury to the

anterior end.

It

appears to be a complex of reflexes which

under normal circumstances

is inhibited, and only appears

any pronounced way under comparatively rare conditions. It is not, as might be expected, a method ordinarily
used by the organism to get out of danger. This is one of
the cases quite frequently met where an organism has among
its available assets, so to speak, a reaction which is well
adapted to a certain end, but of which use is not nuxde at all,

in

or but very
2.

little.

Stimuli which induce Crawling.

It

may be

said

any strong stimulus applied to the

posterior portion of the organism causes the forward crawling movement to ap])ear. Mechanical and strong chemical
in general that almost

stimuli applied in this region will do this.

Light, either of

ordinary intensity, or of such high intensity as that from an

arc light, so far as

ing movement.

have observed,

The

electric

greatly diminishes the rate.

will

not cause the crawl-

current does cause

Any

operative treatment

example, cutting the body in two in the middle

it,

but

as, for

almost

in-

RAYMOND PEAUL.

552

variably causes the portion iu front of the cut to advance

by

the crawling movement, and, as has been mentioned in the

preceding section, at the same time frequently causes the
posterior piece to crawl backward.

There

is

no reason to

suppose that the operative procedure acts in this respect in

any other way than merely as a strong mechanical stimulus
Other stimuli which induce
applied at the point of the cut.
the backward crawling have been taken up in the preceding
section.
c.

Movement on

surface film

on account
the animal

Motion

is

slower in rate than that on the bottom, largely

of the greater flexibility of the surface

is

on the

practically confined to the gliding movement.

is

This gliding

the Surface Film,

While the mechanism

moving.

on which

of the

move-

ment is the same in the two cases, the surface film is elastic,
and does not give so firm a basis as does a solid body. The
effect of this elasticity of the film is very well seen when the
animal attempts to change its course and turn to one side.
The film offers little resistance to the posterior end, so that
this

cannot easily serve as a fixed point for the anterior part

Furthermore, in case the anterior end is left

to turn about.
in contact

with the film when the turn

usually the case, there

is

almost as

much

turning of the anterior part as there

is

is

attempted, as

is

resistance against this

resistance to hold the

The consequence is
that the worm is unable to change its direction of movement
quickly when on the film, and it has to go through a succession of muscular twists and jerks towards one side before the
posterior end fixed as a pivot support.

result

is

attained.

co-ordinated crawling

have not been able

movements

in a

to

worm

induce well

while on the

The preliminary contraction of the posterior

when that region is stimulated, but
the subsequent stretching out of the head and drawing up of
I have tried stimulating
the body does not usually follow.
surface film.

part of the body occurs

both the exposed ventral surface of the animal and the dorsal
surface from below, but neither method is effective.
The
reason for this

is

probably to be found again

in the elasticity

MOVEMENTS,
of

tlie

ETC.,

The

film.

OF FRKSII-WATEK TLANARIANS.

iiiiterior

end

553

unable to get any firm

is

may be drawn
Furthermore, similar resistance is offered to the
stretching- of the head forward as to the turning of it
towards one side.
attaclunenb so

tliat

tlie

rest

the body

of

forward.

When

the animal

gliding on the surface film the same

is

raising of the head (with reference to the

worm, of course:
lowering with reference to the centre of the
earth) and waving it about in the water occurs as under
in this case, a

normal circumstances. In some cases two thirds or three

fourths of the whole body will be thus raised and waved
about, ex;tending itself to its utmost capacity, and apparently
seeking some solid body on which to attach itself. In these
cases only a small portion of the very posterior end of the

body

be

will

in

left

contact Avith the film to support the

whole.

On coming

to the side of the dish

surface film, the

turns

down

worm almost

when

gliding on the

invariably leaves the film and

The

the side of the dish.

reaction which

is

the

cause of this and of the organisms passing from the side of

the dish on to the film will be brought out later.

Relation of Movements of Triclads to those of

other Forms. In respect to their movements, the triclads
studied occupy a somewhat intermediate position between
certain othei* groups.
The rhabdocccles are in general
characterised by free movements in the water, brought
about by cilia covering the whole body. Their movement
d.

general

in

features

Infusoria.

among

is

not at

On

activity.

polyclads
is

Turbellaria

the

movement

is

resembles

that

all

is

or very

little

the other hand, the

found in the case of

movement

of

the

the

liolotrichous

Stenostoma leucops.

is

margins

The

dependent upon muscular

movement

characteristically muscular.

the

of

type showing well this class of movement

many

of the

An example

of this

of

Leptoplana tremellaris, where

largely effected by the rhythmical beating

(cf.

movement has become

Lang,

'84).

In

fact,

this

form

of

so well developed in these animals

RAYMOND PEARL.

554

we may think

that

locomotor organs.

movement

the

of

tlie

margins of the body as special'

if any part in

Ciliary action plays little

of sucli a form.

It is to

be noted, however,

that both the rhabdocoeles and the polyclads are capable of

performing true swimming movements,

in the Avater

i.e.

movements

withont contact witli any solid body.

fresh-water triclads, especially

of

the

free

In the

Plannria

genera

and Dendrocoelum, the

cilia have become much diminished

comparison with the rhabdocoeles, and are restricted to a
portion of the ventral surface only.
Consequently they are

in

not numerous and strong enough to support and

move

the

disproportionately heavier body freely through the water.

The movement of the cilia merely serves in these forms to

propel the body while insufficient to support its weight.
Consequently we find the principal form of movement to be
On the. other
a gliding over the surfaces of solid bodies.^
hand, the fresh-water triclads have not attained the liigh

development of muscular locomotion which the polyclads

There is a purely muscular movement in their case,
but it is not by far the most important form of locomotion,
and is not so highly developed as is that of the polyclads.
Evidently, then, the fresh-water ti-iclads seem to form a
transitional stage in respect to locomotor phenomena between
the rhabdocoeles on the one hand, where purely ciliary
locomotion obtains, and the polyclads on the other hand,
where we find the locomotion largely if not entirely
muscular. Whether this has any phylogenetic significance

have.

is

not certain.

The land planarians

occup)^ a position very similar to that

the fresh-water forms so far as

of
'

ment

do not

their

movements

forms of move-

wisli to imply, in tills discussion of tlie different

as related to the

number and

distribution

of tiie cilia, any belief that

structure gave rise to function or function to structure.

I wish merely to

point out the evident correlation wiiich exists in the matter.

most probable that structure and function changed together

in

many

otlier similar cases, positive

evidence

are

is

It
;

seems to me

but

in this, as

lacking, and consequently

attempts to settle the phylogenetic development of the plienomena would

appear to be fruitless.

MOVEMRNTS, KTC, OP FRESH-WATER PI-ANARTANS.

555

There is, however, a notewortliy difference

between the two groups. In tlie movements of tlie hind
planarians the muscular factors (rhythmical wave motion of
concerned.

the ventral surface, and. snake-like

movements

of the

whole

body) are more important relatively to the ciliary component

than in the fresh- water forms. In these land planarians
there

evidently

is

beginning-

the

of

the

characteristic

rhythmical wave motion of the part of the body in contact

with the substrate, Avhich reaches

highest development

its

in

the case of the Molhiscn.

Non- Loco motor Movements.^

II.

Under non-locomotor movements will be included the

phenomena of contraction, expansion, " feeling movements,"
movements of the pharynx, etc. The purpose of discussing
these phenomena, which are not immediately included in the
is to give an account of their mechanism
which may be referred to in succeeding portions of the paper.
These movements are the physiological foundations on which
the locomotor movements and the reactions are based, and it
is necessary to determine their mechanism in order to bring

general standpoint,

the analysis of the behaviour to completion.

a.

Contraction of the Body. By the term

when applied to forms like the

tion of the body,"

usually meant the sliortening of the

is

the flat-worm this

movement

is

'^contrac-

flat-worm,

body lengthwise.

brouglit about by

traction of the longitudinal muscle-fibres.

tlie

In
con-

It ma}' involve

Most frequently only

the whole body or only a portion of it.
a part of the body contracts longitudinally after stimulation
thus,

if

middle

the anterior end

line,

is

rather strongly stimulated in the

the resulting contraction will usually involve only

the anterior third of the body.

'

III

discussing

lie

In this longitudinal contrac-

nius-culaluic I liave usetl tlirouiiliout,

of Jijima (loc. cit.), in wliose paper a very

he fouiul.

have identified

of muscle-fibres:

full

in sections of V.

tlie

iiomcnciature

(iescriptiou of this

niaculata

system

will

the following groups

() outer longitudinal, (il circular, (c) oblique (?), (d) inner

loiiL;itudinal, (f) dorso-veiilral,

(/) transverse.

EAYMOXI> PEARL.

556

tion all the sets of muscle-fibres other than the longitudinal

must be relaxed completely, because as the animal shortens

it grows broader and thicker, which would be impossible if
the ring, or transverse or dorso-ventral musculature, also contracted.

The longitudinal musculature

is

apparently better

developed and more effective on the ventral side of the body

than on the dorsal, because after very strong stimulation of
the anterior end there is a well-marked tendency for the
middle portion of the body to be raised and the head somewhat curled in under it. This relation is shown diaoramThis curliug under of the head does not
matically in Fig. 5.
specific reaction, but, on the contrary, merely
an expression of the fact that the ventral musculature is
capable of shortening its side of the body more in maximal
contraction than the dorsal side. Jijima (loc. cit., p. 378)

appear to be a

Fig.

5.

Diagram

showiiif^ the

appearance in side view of a

niaxinialiy contracted planarian.

finds,

from a histological study

of the musculature, that the

bundles of fibres in the main longitudinal muscle layer are

thicker on the ventral than on the dorsal side.
b.

Extension

of the

Body.

The mechanisms by which

brought about are

probably very different in different groups. In the case of
the flat-worm extension is produced by the contraction of the
circular muscular layers surrounding the body, and of the
transverse and dorso-ventral systems of musculature. Probably
extension

of

a soft-bodied animal

also the oblique musculature,

when

is

by its
The mechanical

present, assists,

contraction, in the extension of the body.

necessity for extension of the bodj^, after contraction of these

muscles,
sake,

be

is

readily apparent.

considered

If

a cylinder,

the body, for simplicity's

contraction

of

circular

MOVEMENTS,

ETC., OF

FRESH-WATER PLAXARIAXS.

557

muscles must cause it to lengthen, wliile witli the form of

body which really exists in the flat-worm, the contraction of
the well-developed dorso-ventral fibres must bring about the
flattening seen in the fully extended gliding auimal.

Probably by far the most important sets of muscle-fibres

producing the general extension are the dorso-ventral and
circular.
It is to be noted that contractions of any of the
for

may

sets of fibres

take place in localised regions, producing

extensions or contractions of that region, according to the set

affected.
The sensory, or " feeling " movements of the head
are brought about in this way.

The
of the

extension,

pharynx

is

and probably also in large part the extrusion

brought about by contraction of its well-

developed circular musculature.

In the case of the marine mollusc,

A ply si a li niacin a,
Jordan (: 01, pp. 11 15) has recently shown that extension is
brought about in an entirely different manner. It results
from passage of fluid from vesicles in the skin into the spaces

in

the

When
into

body parenchyma when the body muscles are

the animal conti^acts this fluid

the vesicles, which

swollen

then,

when

relaxed.

(blood) is pressed out

become very much extended and

the muscles are relaxed, the elasticity of

the walls of the vesicles forces the fluid back into the body,
and thus causes its extension. As a result of this method of
expansion it is possible to kill the animal fully extended by

the use of such poisons as cocaine.

It is not

unusual to consider the fully extended condition

of such an organism as a flat-worm as one of approximate

relaxation.

Instead of

this, it is, in fact,

a certain part of the musculature

tonic contraction.
it is

impossible to

is

a condition in which

in a state of well-marked

This furnishes a reason for the fact that

animals in a completely extended

kill these

condition by the use of poisons which tend to produce a

relaxation of the muscles.

Under these circumstances the

animals take on the typical relaxed form, which

from that of extension.

Rest. Inasmuch as a very large,

is

quite

different
c.

VOL. 46, PART 4.

NEW

SERIES.

if

not the larger

O

RAYMOND PEAKL.

558
portion of the

life

of a planavian

will be well to discuss

rest, it

is

tliis

spent in a condition of

matter ; and it may, perthe general heading of

taken up under
although really the opposite of activity.
The appearance of the worm when resting is, as has already
been mentioned, quite different from its appearance in the

haps,

best be

''activities,"

The body is shorter, wider, and thicker.

The ordinary contour of the head is almost entirely lost, and
active condition.

sharply pointed anterior end of a form like P.

dorotocephala, the end is evenly rounded. The auricles
disappear almost entirely, and their position is indicated only
in place of the

by the

difference in the pigmentation at that part of the

dorsal surface.

j^iG.

The

lateral

edges of the body frequently have

sliowing the typical appearance of a resting planarian.

bounds approximately tlie area covered in tlie
" testing movements " wliicii precede the coming to rest (cf. text).

G. Diagram

Tlie

dolAed

line

a wavy line instead of the straight one of the active condiThe anterior end of the body is in contact with the
tion.

bottom, and not raised as in movement. The general appearance of a resting planarian is shosvn in Fig. G.
The coming to rest of a gliding animal is usuall}' done in a
First, the animal glides more and
very characteristic way.
more slowly for some distance before reaching the point at
which it will finally stop. The distance before reaching the

stopping place in which the worm glides appreciably slower

usually not
is not however, in most cases very considerable
own
length.
is to be
its
It
times
three
or
two
than
more

noted that this slower gliding which precedes the coming to

MOVEMENTS,
rest

PRESH-WATKR PLANARIANS.

ETC., OP

559

not in form or rate distinguishable from the other

is

worm which is not followed by

In other words, a specimen may glide slowly for a long
time without stopping, so that one cannot prophesy with
certainty from the rate of movement whether the specimen is
soon coming to rest or not. The coming to rest is practically
always preceded by a period of slower gliding, but all slow
gliding is not immediately followed by rest. After a brief
slow gliding motion of the

rest.

period of this slower gliding the

the posterior half of the

same

The

position.

worm suddenly

body remains fixed

anterior half of the body

about over the bottom from side to

touched frequently

and

in precisely the
is

slowly

moved

the head being

bottom or any other solid object in

The anterior part in this "feeling"

to the

the neighbourhood.

movement moves about

the posterior part as a fixed point,

the latter very rarely changing

its

The thoroughness of

stopped.

side,

stops,

it has once
"testing" of the sur-

position after

this

roundings by the sensory anterior end varies much in different

cases, but in practically all cases one can see some indication
of

it.

have

in

some instances seen

it

done very thoroughly,

so that the whole surroundings within a radius of 3

were gone over.

to

complete

rest,

Finally,

when

this

is

mm.

done the animal comes

and assumes the typical relaxed condition

shown in Fig. 6. The apparent significance of the " testing "

movements at the time of stopping is that it is apiece of proThe worm examines the surroundings
tective behaviour.
before coming to rest, to see if there is anything dangerous
(either

of

solid

nature or

immediate neighbourhood.
is

the

true

one,

a harmful chemical) in

Whether

the

or not this explanation

and further, whether natural

selection

developed this reaction for protective purposes, seems to

me

to be very doubtful, for reasons brought out in anotlier place

and 543). In some cases I have seen the Avorms

by simply stopping without any appreciable trace
of the " feeling " movements, but this is not the usual procedure. In coming to rest in one of the collections already mentioned, the '^feeling" movements are usually very well marked.
(cf.

pp. 542

come

to rest

RAYMOND PEARL.

560

There
to

come

is

a "well-marked tendency for the planarians studied

to rest in such a

way that the

long* axis of the

body

forms a right angle, or nearly a right angle, with the lines

of the force of gravitation. The cases in which the organisms
come to rest with the long axis forming an angle of less than
thirty degrees with the line of gravitation are rather few.

Of course, when they come to rest on the bottom, the angle

formed is approximately ninety degrees. A large number of
observations on individuals which came to rest on the sides
of dishes of various shapes have given the general result
stated above.
There are, of course, exceptions, but there is,

Fig.

7-

Diagram

showing

(lie

positions taken by planarians

coming

to

rest iu a dish (see text).

making due allowance

tendency to a horiThis is the only

behaviour of the organism which bears any resemblance to
a geotactic reaction. Another tendency, less marked than
the former, is for the animals to come to rest in the angle
formed by the sides and bottom of the dish. Not only do
specimens come to rest lying directly in the angle, as shown
at A in fig. 7, but also, and more frequently, they lie in such
a position that a part of the body is on the side of the dish
and a part on the bottom, as shown at B, fig. 7.
In this
after

for these, a

zontal position as the position of rest.

position the anininl usuall}' lios obliquely rather than at right

MOVEMENTS,

ETC.,

OF FIil!]SH-WATEli PLANARIANS.

5G1

angles to the line of the junction of the side and bottom of

The animals usually come

dish.

tlio

to vest in this position

have been gliding on the side of the dish. When

they come to rest from movement on the bottom of the dish,
a position frequently taken is that shown in C, fig. 7, where
only the very anterior end is in contact with the side. This
coming to rest in the angle of a dish is apparently a reaction
which agrees with those usually called thigmotactic reactions.
But it is not, as has been stated by several writers, due to a
tendency to get more of the body in contact with something
after they

solid^

than

is in

such contact under usual conditions; for in

the case of an organism like a flatworm,

any more of the surface of the body

solid
flat,

in

when

it is

as in B.

either

to

shown in Fig.
the same amount

bent, as

There

is

The ventral surface

case.

is

it

in contact

8,

A, than

of

it is

the flat-worm

is

8.

with a

when

of surface in contact

L
Fig.

impossible for

be

Diagrammatic cross-sect ion of

an angle

and B, on a

])laiiaiiaii at rest

A,

in

i)!ane surface.

strongly positively thigmotactic under

all

circumstances, and

the dorsal surface negatively thigmotactic, but this does not

help us understand
in angles.

why

the animal comes to rest frequently

This behaviour of the flat-worm in dishes

is

due to

the same sort of reaction as that which causes them to

come

on unevennosses on rocks, and also causes the same

phenomenon in a more marked degree in the case of Litto rest

torina, as recently described by

common

Mitsukuri

factor in the reaction in all cases

body

is

(:01).

The

that difl^erent

brought into such positions that they

form unusual angles with each other. Since this phenomenon
is distinctly different from any embraced by the term thigmotaxis as used in its true sense, it seems desirable that it be
parts of the

are

EAYMOND PEARL.

562

given a specific name.

term gouiotaxis.^

When

would propose for

a flat-worm starts from

nature of the movement,

e.

i.

this reaction the

a resting condition

tlie

whether gliding or crawling,

depends in large measure on the intensity of the stimulus

If the resting animal is rather strongly
which starts it.
stimulated it will start at once into a crawling movement,

which changes

to gliding after three or four, or fewer, con-

not again renewed.

tractions, ])rovided the stimulus

is

possible by the use of a very

weak stimulus

It is

to start the

resting animal off at once into the gliding movement, or with

only the faintest indication of a single crawling contraction.

When

the animal

spontaneously

starts

usually begins at oace with the glide.

movement

into

When

it

starting spon-

taneously the glide is usually preceded by some of the

" feeling " movements of the head end, such as precede the

coming

to rest.

The purpose

as in the former case.

Any

of these is evidently the

same

sort of strong stimulus will start

the resting animal into movement.

The physiological condition

of the resting animal

is,

as has

the
been mentioned, one
apparently
commuscuUir systems of the body are in an
This is evidenced by the form of
pletely relaxed condition.
the resting animal, which differs from that of one in movement, in being shorter, wider, and thicker, and in not showing

already

of

relaxation.

All

of

such features as the auricles, or the pointed tip of the head.

This relaxed condition is evidently one of lowered tonus, as

may be determined by

simple observation.

Stimuli of an

which would cause a marked reaction in an individual in an active condition, will produce no effect on a
This point has been tested with a variety of
resting animal.
stimuli, including- mechanical, chemical, food, etc., and the
intensity

markedly loAver tonus of the resting animal is very evident.

The reactions which are produced, provided the stimulus is
made strong enough to be just effective, are weak. Of
course,

if

the stimulus
'

is

above a certain strength, or

From

ywri'a

angle.

is

con-

MOVEMENTS,
tinned for some

ETC.,
tiinOj

OF FRESH-WATER PLANARIANS.
tlic

ciuiinal

generally

b(!couiu

will

563

This condition of
up and glide or erawl away.
lowered tonus in the resting animal reminds one of the conThere, as in
ditions found in sleep in the higher animals.
this case, the general sensory and muscular tonus is greatly
reduced, and there seems to be no good reason why the
resting condition of these lo.wer organisms may not be conThe two things appear to be
sidered and called '' sleep."
fundamentally the same physiologically, and would appear
Furthermore, there is no
to serve the same purpose.
apparent reason why the lower organisms should nut have as
stirred

great a need as the

higher for periods of rest or sleep,

during which the anabolic processes are in considerable
The fact that some lower
excess over the katabolic.

organisms

are

balanced

so

physiologically

apparently do not require such periods of rest

that
is

they

not con-

must be similarly
the writer, there has been

clusive evidence that other low organisms

balanced.

So

comparatively

far as
little

is

known

to

attention paid to the physiological con-

dition of lower organisms during different phases of their

activities.

An

animal which

is

not moving

to be in a " resting condition," Avhen in

as

in

Clepsine, the quiet

animal

heightened rather than lowered

is

tonus

in
(cf.

is

loosely said

nuiny instances,
a

condition

Whitman,

of
loc.

cit.).

As was noted

ia the section

on " Natural History," the

periods of activity of PI an aria are separated by periods of

The time spent in the restrest of greater or less length.

ing condition, at least during the daytime, is considerably
greater, on the average, than that spent in movement.
Probably, however, this is reversed during the night, when
the activity is greater than during the day. This periodicity
in the activity is just

what would be expected

if

there

is

necessity for rest at intervals as in the higher animals.

The causes which immediately induce the coming to rest

The principal cause, as has been
considered.
indicated above, is that the animal becomes fatigued by

may now be

RAYMOND PEARL.

564

movement, and

As
in

general tonus becomes lower and lower.

its

a result of this

it

must remain relaxed

order that recovery

may

take place.

of the activity of the animal

certain point

it

its

for a certain time

When

in the course

general tonus gets below a

coming

stops, the actual process of

to rest

being a more or less gradual one.

A strong piece of
evidence in favour of this view is the fact already given in
the section on " Natural History/' namely, that if the
animal is stirred up and made to start moving again immediately after coming to rest each time,
that the periods of activity

it

will

be found

become progressively

shorter.

Furthermore, when the general physiological condition of

the organisms is weakened by keeping them for a time in
the laboratory,

it

is

found that the periods of rest become

progressively longer in proportion to the periods of activity.

The general " predisposing condition " to the coming to rest

then probably a lower tonus due to fatigue. The immediate causes determining the exact place chosen are of
three sorts. First, and probably most important of these, is
is

the intensity of the light.

It is well

known

that planarians

tend to come to rest in regions of comparatively low light

having been first noted by Loeb ('93),

"
him
Unterschiedsempfiudlichkeit."
and
by
This
factor seems to bo the most important of any in determining
the region in which the animals come to rest, both under
experimental conditions and in the natural habitat.
In
aquarium dishes placed close to a window, and containing
considerable plant material, the worms will be found resting
practically always in the half of the dish away from the
The largest number of individuals will be
window.
entangled in the plant material, and usually for the most

intensity, the reaction

called

part invisible

and bottom

while of those specimens resting on the sides

of the dish the greatest

number

will

be found in

such places that there are heavy masses of plant material

between them and the window. A few will come to rest far
around on the sides of the dish where the glass itself cuts off
some of the light. This last position has been mentioned by

MOVEMENTS,
Loeb

ETC.,

OF FUESH-WATER PLANARIANS.

565

as the oue most ffequently taken by plauarians in a

dish containing' only water.

Tiiis

behaviour towards light

an absolutely precise reaction. Many times

during- experiments I have seen specimens come to rest in
but in
the vei-y lightest parts of the dish and remain there
general this reaction will cause most of the animals to gather
is

not, however,

in shaded areas.
It is probably the principal factor in
causing the animals to take positions beneath stones in their

natural habitat.

The second immediate cause

animals shall come to rest

is

in

determiuiug where the

the goniotaxis mentioned above.

an animal in the proper physiological condition of reduced

tonus comes to an unevenness in the surface on which it is
moving, it will in most cases come to rest there. This, again,
If

not sufficiently so as to

make

is

not a very precise reaction

it

possible to predict beforehand where any given individual

will stop.

In this case, just as in the case of light,

much

depends on the animal's physiological condition, and when in

the proper condition they may come to rest on a perfectly
smooth surface. Thus in a dish individuals will always be
found at rest on the smooth sides and bottom, yet there is a
distinctly marked tendency, when the animals are put under
experimental conditions and closely observed, for them to

come

to rest in the

angle of the dish.

This reaction probably

also plays a considerable part in the habit of coming' to rest

among

the branches and leaves of the plant material.

the natural habitat

them

may be
is

it

is

to take positions

In

undoubtedly the factor which causes

on the uneven parts of stones.

It

that the immediate cause of the stopping in this case

the increased

i-esi

stance

to

movement afforded by

the

unevenness of the surface. This, acting on an animal in a

fatigued condition, might give the necessary stimulus fur
the stopping.

The third factor in doterminiug where the animals shall

come to rest is one about which I am doubtful. There seems
to be some evidence, from the behavioui" of the animals
themselves, that in the foimatiou of the groups or collections

RAYMOND PEAUL.

566
previously mentioned,

That

kinosis.

is

(pjj.

say,

to

533, 534) tliere

a sort of cliemo-

is

chemical

presence of some

the

water causes the animals to stop. The

evidence for this factor will be taken up with the discussion
It probably does not play
of the formation of collections.
substance in the

any part

determining where a single individual shall come

in

to rest outside of a collection.

must be emphasised that all of these three factors are

in importance as compared with the physiological
condition of the animal, which may be said to prepare it for
An active animal, in which the tonus is at
the resting state.
or near the maximum, will pass through regions of low
illumination, uneven surface, or collections of other individuals without stopping.
Only when the animal is in the
It

secondary

right general condition do these factors

come

in to determine

the precise point where the stop shall be made.

Formation

of Collections. Since the formation of

dependent on the animals coming to rest in a
certain area, it ma,y properly be taken up in this section.
The collections are fairly well-defined groups of from six or
eight up to twenty or more individuals. The general appear1.

collections

is

ance of such a gi'oup

is

shown

in Fig. 1.

The

individuals

composing it have no definite orientation, but are scattered

about with the anterior ends directed in whatever way they
happened to be pointed when the individuals stopped. The
distance separating the individuals varies
cases.

In some cases

may be

or again

it

may be

as

much

much

in different

as a half-centimetre,

the Avidth of an individual

worm

or

less.

This formation of collections of this sort might be considered

the result of a " social instinct " by animal psychologists of
the Binet school.

simple reactions

Actually,

taken

in

it

appears to be due to two

conjunction

with

the

physiological condition of the individuals composing

first of

these reactions

individuals
certain

come

to

is

a comparatively

degree of illumination,

condition

of

That

that to light.

if

is

to say,

restricted

they are in

reduced tonus, they stop.

general
it.

The
when

area of
a

certain

Those which are

MOVEMENTS,

liTC,

OF FRESH-WATER FLANARIANS.

567

very active pass on through the region, bnt necessarily in

course of some time several individuals will have stopped,

and a group will have been begun. Wlieu once started

another reaction apparently enters to assist in enlarging it.
This reaction appears to be due to some chemical substance,
and belongs to the class of reactions which Eiio-elmann has
suggested should be called "kinetic,'^

in tliis case chemowould appear that planarians excrete or secrete

some cliemical substance towards which they are themselves
positively chemotactic, and which also causes them to come

kinesis.

It

to rest.
When several individuals remain quiet in a small
area this substance, of course, accumulates and affects other
individuals passing.
That some such a substance is sepa-

rated from the bodies of the animals is evidenced by two

phenomena. First, in the case of the food reaction, which
will be taken up in detail later, it is found that after one or
two individuals have attached themselves to a piece of food
material and begun feeding the mass of food and planarians
is a much more effective stimulus to positive chemotaxis than
is the same food substance alone, even though it may have
remained in the water a greater length of time. The "zone
of iufluence " (vide infra, p. 626) of the food and feeding

individuals together

is

much wider than

that of the food

Specimens are affected at a greater distance from

the food and react more sharply.
As a result of this, dense
aggregations of planarians will be formed in a comparatively
short time after the first two or three individuals have found
a bit of food. As there is no reason to suppose that the
alone.

of the food itself is different in the two cases, we

must conclude that the greater effectiveness of the food and
feeding indivitluals is duo to some chemical substance coming
from the organisms themselves.

action

The second

line of

evidence for the existence of a reaction

to a chemical in the formation of collections

is found in the
behaviour of specimens coming near a group of individuals
resting on the bottom of a dish.
When some distance away

from the outer boundary of such a group a gliding animal

RAYMOND PEARL.

568
will

frequently be seen

to

give a well-defined

positive

and turn towards the group. The reaction is of

precisely the same character as that given by the organism
to weak chemicals (to be described later), and the behaviour
reaction,

convinces one observing

it

that the specimen

stimulated

is

by some chemical diffusing out from the group. After

turning towards the group the specimen will glide into it
and usually coiiie to rest, in the manner which has been
described above.

What

the nature of the chemical substance present in the

region about the groups

is, I have not been able to discover.

Neither rosolic acid nor methyl orange is discoloured by it.

Whatever
state.

its

nature,

it

must be

an extremely diluted

in

This seems evident for two reasons

does not affect delicate indicators

first,

does not have any effect on active specimens of

A large

number

because

and second, because

it
it

Plan aria.

of experiments have been

performed to test
this latter point, but always with the same result.
Unless
the individuals were in the proper predisposing condition of
lowered tonus, they would pass by or through groups of
other individuals without giving any reaction.
Attempts to produce, artificially, collections of planarians
in chemicals have been unsuccessful.
I have tried various
solutions (such as sugar, weak alkalies, etc.) to which the
organisms showed a well marked positive chemotaxis when
tested by other methods, but have not been able to get any
formation of collections in them. The animals would give
the positive reaction on coming to the edge of the diffusing
chemical and pass into it, but would not come to rest. This
failure to produce collections artificially is not surprising
when one considers the number of conditions necessary for
the production of the desired result.
The organism must be
in just the right physiological condition, the chemical must
be of a certain concentration, and finally, it must be located
in an area of

a certain

light intensity.

almost or quite impossible to

fulfil

the same time in an experiment.

all

It

is

practically

these conditions at

MOVEMENTS, ETC, OP FRESH-WATER PLANARTANS.

The

569

collection seems to be due

simply to the direct effect of the chemical on the org-anism.
'J'here is no evidence that the animals are held in the group
coniing- to rest

in

the

as a result of a negative reaction to the surrounding water,

as

the case in the collections formed by the infusoria

Jennings, '99, h). The method of formation of collections

is

(cf.

in chemicals in the case of the infusoria

is

as follows

Specimens swimming about at random come to the edges of

If, for example, the
drops of chemicals purely by chance.
happens
to
a
weak
acid,
chemical
be
the specimens will pass
into the drop without giving any reaction.
When, however,
they reach the opposite edge of the drop and attempt to pass
from the chemical back into the water they are stimulated,
and give their usual motor reaction. This turns them back
into the drop, in which they are, as it Avere, '' caught in a
As a consequence of this method of reaction a very
trap."
dense collection will be formed in a shoit time. With the
flat-worm the case

is

very different in that an active

dividual frequently passes into and

out

of

in-

one of these

showing the faintest trace of a reaction

The only way in which any stopping in the

collections without

on either

side.

brought about by a chemical is by a chemokinetic

The fundamental difference in the reactions of the
two groups of organisms on which this difference in the matter
of forming collections is based will be brought out in the
section on the reactions to chemicals.
region

is

reaction.

To sum

up, the

formation of collections of individuals

seems to be due, in the first instance, to the tendency of the

organisms to come to rest in areas of a certain degree of
intensity of light, and in a lesser degree to a tendency to
turn towards and come to rest in

areas containing some

substance secreted or excreted by the worms themselves.

prerequisite in the formation of collection, as in the

coming

under any circumstances, is a proper physiological

condition of reduced tonus.
There does not appear to bo any special biological significance to this tendency of the animals to collect in gi'oups.
to rest

RAYMOND PEARL.

570

The behavionr

is

not of auy evident benefit to the organisms,

where it is apparently closely

connected with the obtaining of food. On the contraiy, it
seems to be, at least potentially, a harmful thing, because
any accident or enemy would affect a number of individuals
rather than a single one when they were so collected.
as

it is

d.

in the case of infusoria,

The Effect

Operations on Movement.

of

It

may

be well to put together in one place the results which

were obtained with reference to the movements from animals

which had been cut in various ways. From these we can

form some idea of the relation of the nervous system of the
planarian to its movements.^
The immediate effect of any operation is that of a very
strong mechanical stimulus applied to the same part of the

Fig.

9.

Operation

Tlie heavy str.night, line indicates tlie cut

diaj^rani.

For results see

made.

body, and the sort of movement

text.

resulting in each

piece

depends on the position of the cut. The details of this immediate effect will be described in connection with other
mechanical stimuli. What concerns us here is the permanent
after-effect of operations on the movements.
We can best
by
taking
up
matter
this
some
specific
get at
cases.
^

All the operations

with the specimen

were performed with

in a dish of

water.

a sharp scalpel,

In some cases the

most cases

in

worm was

to a drop of water on a soft board for the cuttins', but in

all

transferred

cases where

immediate observations were wanted, the operations were performed

dishes used for

t,lie

experiments.

on planarians arises from

rest

tiie fact

The only
that

if

difficulty in

in tlie

performing operations

the edge of the knife

is

allowed to

on the surface of the body for even a very short time before the cut

made,

it

will

any clean cut

without

become covered with the sticky slime from the

is

impossible.

iicnct rating.

The edge

will

slip

off

the

aninuil,

and

back of the

is

tiien

worm

MOVKMENTS,
If a

cut

planai-ian is

9,

FRESH-WATRR TLANArjAN?.

scjiiai-ely

across

571

the body in

tlie

distance behind the head, as indicated

I'egion a short

Fig.

ETC., OF

the anterior piece will continue

to

move

in

after tlie

the same rate as the whole

After the immediate effect of the operapast the glide is its ordinary movement, and it will go

operation

approximately

at

animal did before.

tion

is

al)out the dish

At

and behave

the outstart

in

general like a whole individual.

of activity

its pei-iods

about as in a normal individual,

movement

of spontaneous

On

is

and

or, in

rest are distributed

other words,

its

power

not inipnired, at least for a time.

the other hand, the posterior piece comparatively soon

comes

Its gliding movement is

become longer and longer in
activity.
Its power of spon-

to rest after the operation.

slower, and the periods of rest

compai'ison with the periods of

taneous movement becomes very greatly diminished within a
comparatively short time after the operation, and it remains

Fig. 10.

Diagrammatic side view of

a decapitated specimen performing

the gliding movement.

in the relaxed resting condition during the greater pnrt of

the time spent in the process of regeneration.

When

this

posterior piece does glide about soon after the operation

anterior end

higher than

is

is

the head of a normal

This

circumstances.
''feeling"

usually raised off the bottom

movements

but instead this end

If,

instead of

made back

is

is

shown

in

flat-

Fig.

worm under
10.

its

considerabl}-

similar

There are no

of the anterior end of such a piece,

held very

making the cut

stiffly in

the raised position.

so close behind the head,

it is

the middle region of the body, the anterior

in

piece behaves as before,

e. like the normal animal.

The
moves slower thau did the corresponding piece in the previous experiment, and it comes to
rest sooner after the operation, and remains quiet longer.
In the same way cuts may be made nearer and nearer the
the posterior piece will move more and more
posterior end

posterior piece, however,

i.

RAYMOND PEAEL.

572

At the same time the

anterior pieces will appear more and more like the normal.
In both sets of pieces the crawling movement may be induced by proper stimulation of the posterior ends.
01)lique transverse cuts produce the same results as do
and come

slowly,

sooner.

to rest

B
Fig.

Heavy

11. Operation diagram.

For results see

The same laws hold

direct ones.

In case a strip

pieces.

is

lines indicate the cuts

as to the

make any

further

movements

of the

cut from the side of the body, as

in Fig. 11, the smaller piece

shown

made.

text.

progressive

curls np,

movements,

and does not

although

it

remains alive, and will eventually regenerate in most cases.

The main part B contracts on the cut side, and hence becomes

curved

move

in that direction

about, but

at a

after the operation.

It

somewhat slower gliding

is

able to

rate than

normal, and in a path curved towards the cut side. In case

a worm is slit down the middle line at the anterior end, as in
Fig.

12,

it

is

able to glide, but at a

slower rate than

normal. It performs the crawling movement in response

to stimulation at the posterior end, and each half of the head
performs feeling movements independently of the other half.

Fig.

made.

An

The heavy line indicates the cut

For results see text.

12. Operation diagram.

individual

slit

up

in the

middle

line

from the posterior

end, as in Fig. 13, glides at approximately the normal rate,

provided the cut is not carried too far forward. If the cut

extends into

the

mediately slow^er.

head region the gliding becomes imSuch a specimen performs the crawling

MOVEMENTS,

KTO., OF

FKESH-WATEK PLANAKIANS.

573

nioveineut upon stimulation of the posterior end of either

piece, but in a peculiar way, whicli will be described later.

Putting

all

these results together,

we

see that there

is

general tendency for animals on which operations have been

performed to glide at a slower rate than normal. In some
of the pieces this tendency is very slight, and frequently
hardly noticeable. In others the movement is very much
In all cases the periods of rest are
longer during the time of regeneration than normally. Tiiis
tendenc}^ for the animals to remain quiet during regeneraslower than normal.

tion increases up to a certain point as regeneration proceeds.

piece of a planarian njay be quite active for three or four

hours after the operation, while during the following three

or four days it will scarcely move at all. After the regeneration

is

practically complete the

again approximately as

JflG. 13.

it

Operation diagram.
For

worm

will

begin to move about

normally does.

The heavy

During the

line indicates the cut

re-

made.

results see text.

generating process the anterior pieces, bearing an uninjured

head, are

much more

posterior parts.

inclined to

These

move about than

latter usually

are the

remain entirely quiet

during regeneration.
This behaviour of the posterior parts during regeneration
appears to be distinctly purposive, and to belong to the class

phenomena

called regulatory.
The general tonus of these
immediately lowered by the operation, and consequently they keep quiet.
Yet at the same time the processes
of morphallaxis and, in many cases, growth begin at once,

of

pieces

is

and proceed very vigorously

we consider

till

the

missing parts are

worm

or part of a worm
has at the beginning a certain sum-total of energy available

restored.

If

for all activities, including

and

all its

that the

movement, growth, morphallaxis,

it would appear that

other vital processes, then

VOL. 46, PART 4.

NEW

SKRIES.

P P

RAYMOND PEARL.

574

the performance of any single set of activities in excess must

cause a corresponding diminution in other activities.
is

exactly what

we

While the processes concerned

planarian.

are at their masimiuu activity^

the

amount

of

movement.

part of the energy which

is

This

find to be the case with the regenerating

It

is

we get

in regeneration

a decided reduction in

would seem, then, that a large

expended in movement

ordinarily

used after operatiou or injury in the processes of regene-

As

ration.

the regeneration

completion more and

and used in movement. This
nears

more energy is available for,

would seem to be a sort of "energy regulation." The
behaviour is evidently further beueficial in the case of the
posterior pieces, because their anterior ends are very insensitive as compared with the head of the normal animal, and
about they would certainly be more apt
if they moved
in the long run to get into difficulties than if they remained
quiet.
It

may be

well in closing the section to point out the

relation of the nervous system

(:00)

has

maintained

planarian, for instance

that

to

"if

the movements.

we

divide

Planaria torva,

Loeb

a fresh-water

transversely, the

posterior half, that has no brain, crawls just as well as the

oral half.

Spontaneity in Planaria torva

no means a function of the brain."

sentence we understand "glide"

first

ment

is

"move

If

is,

therefore,

by "crawl"

by

in the

be meant, the statenot strictly accurate. The posterior pieces do not

just as well as the oral" (anterior), but, as has
to

already been brought out, more slowly.

For a very short

time after the operation the statement would in some cases

be correct, but it certainly would not be twenty-four hours
later,

according to

all

the observations I have been able to

As for the spontaneity of the movement, that also becomes very much lowered with the loss of
The very
the brain, as I have attempted to show above.
much lessened activity of posterior pieces of planarians has

make on

the subject.

been mentioned by

From my own

Lillie

(:

01, pp. 182, loo).

observations

it

seems clear that the principal

MOVEMIiNTS, ETC., OF FliESH- WATI'JIi

I'LANAIil ANS.

575

function of the brain of PI an aria with reference to move-

ments

is

That

to maintain the tonus of the ciliary system.

neither the crawling nor the gliding movements are specific

functions of the central nervous system

both sorts of movement

my

Yet

all

loss

of

may

is

evident, because

take place after

removal.

its

observations tend to show that after injury to or

the brain the gliding

immediately, markedly slower.

movement becomes, almost

This relation

is

especially

by the experiments noted above on splitting

the animal longitudinally from the anterior and the posterior
ends.
In the one case the gliding movement becomes at
well indicated

once distinctly slower, while in the other case there

is

slight difference in the rate, evidently conditioned

only a

by the

few of the cilia can get a hold,

The force of the
so to speak, so that they can function.
argument will be impressed if one glances at the relative si/e
of the cuts in Figs. 12 and 13, and then remembers that the

fact that only comparatively

rate of gliding of the specimen figured in Fig. 13

With

than that of the one in Fig. 12.

is

faster

the co-ordination of

movements, including the crawling, the ceutral nervous

system has very little to do in the case of Planaria. With
regard to the spontaneity of movement
in

how

far the brain functions.

It

is

it is difficult

to decide

certain that regenerat-

ing antei'ior pieces show more spontaneous movement than

do posterior pieces, yet the anterior pieces are behind the

normal worm in this respect. The brain probably plays
some part in the perforuiance of normal spontaneous movements, but, as has been pointed

out, in

these

operation

experiments the whole matter is very defiuitely related to

the regenerative process, and loss of substance plays nearly,
if not quite as great a part as loss of nervous system.

Summarising, we

may

1. For the performance

movements the brain is not specifiThese movements are normally co-ordinated

say that

of the crawling or gliding

caliy necessary.
in the
2.

absence of the brain.

The maintenance

of

the ciliary system

the tonus of

(which produces the gliding movement)

is

a specific function

EAYMOND PEARL.

576
of tlie brain,

and

most important function so

further, its

is,

movement is concerned.
The brain plays a certain part

far as
3.

in the production

of

spontaneous movements.

Reactions to Stimuli.

F.

I.

Since

Eeactions to Mechanical Stimuli.

reactions which

the

mechanical stimuli are

given by Planaria to
foundation on which

are

in a sense the

it may be well to
working out the
thoroughly
consider them
reactions to mechanical stimuli we have a very definite clue
to practically all the animal's behaviour.
For rough, general work with mechanical
a. Methods.

the reactions to other stimuli are based,

first.

After

may be used

stimuli a needle or a sharp-pointed scalpel

the stimulating agent.

For the

ising the stimulus, I at first

tubing drawn out

work

finer

as

in sharply local-

made use

of pieces of glass

fineness.

This method was

to capillary

was too stiff to admit

the body under some circumstances.

not, however, satisfactory, as the glass

of reaching all points of

Furthermore,
the end,

this stiffness, together

made

it

with the sharpness of

almost impossible to give the animal a

moderately strong stimulus without wounding it. A far

better plan was found to be to fasten with sealing-wax a
moderately
tubing,

the

stiff

piece

latter

to

of

human

serve

as

hair to a piece of glass

a handle.

With such an

arrangement the stiffness of the stimulating point can be

Danger of
varied by varying the length of the hair.
wounding the animal is avoided, yet repeated strong stimuli
may be given, while, further, the flexibility of the hair makes
it possible to stimulate the animal at any point and from any
desired direction.

An

annoying

difficulty in

connection with this work was

the clinging of the slimy secretion of the body to the point

Once coated with this slime the
used for stimulating.

MOVEMENTS,

KTO.,

OF FRESH-WATER FLANARTANS.

sharpest point will slide

the

off

body without

givinc^

577

any

effective stimulation.
h.

Description of Reactions. The

by taking up

be described

order

in

reactions can best

the

typical

results"

following stimulation of the different parts of the body.

Stimulation of Head Region.

1.

If

planariau

gliding along on the bottom of a dish be touched with a

needle on one side of the head,

it Avill,

under normal circumhead and anterior

stances, in the majority of cases, turn the

one fourth of the body away from the side stimulated, and
continue gliding along in the

turning of the anterior end.

or, as

will

we may

call it for

always be given

strong.

There

is

new path determined by the

away" reaction,

This 'burning

economy
if

the

of words, negative reaction,

stimulus

is

made

sufficiently

a certain intensity of stimulation below

which the negative reaction may or may not be produced,

depending on the physiological condition of the individual,
but above which it always occurs. If, again, a normally
gliding planarian be selected for stimulation, and this time
the stimulating point (preferably something finer and more
flexible than a needle) be touched very lightly to the
edges of the sides of the head or the auricles, we get,
provided the specimen is in the proper physiological condition, a very gi*aceful and striking reaction, quite different
from that obtained in the former case. This time the flatworm will stop for the briefest instant, turn the head and a little
of the anterior end of the body towards the side stimulated,
and at the same time raise the head from the bottom, until
finally the tip of the head points exactly towards the point
from which the stimulus came, and then glide forward in that
This "turning towards" or positive reaction is
direction.
given only in response to very weak mechanical stimuli, and
then only when other conditions are favourable. It is a very
and characteristic reaction when it does appear.

precise

Having outlined
ical

the two main reactions following mechan-

stimulation in the head region,

sider each of

them

in

more

detail.

we may proceed

to con-

RAYMOND PEARL.

578
a.

Reactions

to

Strong Stimnli. Tlie

negative reac-

tion is the characteristic reaction given to all strong stimuli,

whether mechanical or of some other sort. It is, further, the

same tjpe of reaction which most organisms with fairly welldifferentiated reactions give in response to strong stimulation.
It

takes the animal away from what might be a dangerous

object.

In

Planar

a the portion of the body which takes part

turning away varies with the strength of the stimulus

in the

to a certain degree.

Stimuli just strong enough to call forth

the negative reaction will cause onl}^ the head to be turned

The

away.

first

turn away of the definite reaction never in-

Diagram

sliowing the form of the nen;ative reaction to mccliaiiA shows the position just, bel'oie the slimuhis is
applied, and P> the position after the reaction.

FiG, 14.

ical

st.iinuli.

eludes any of the

body back

of the pharynx, so far as I

have

observed, except in the case of very strong and repeated

stimuli.

In the typical and tn^st often observed form of the

negative reaction the portion of the body which turns away

is

that anterior to a point about halfway between the level

of the eyes

shown
is

and the point

in Fig. 14.

With

of origin of the

pharynx.

This

is

stronger stimuli the poiut of turning

farther back on the body.

The number
in the

of degrees through which the head

negative reaction

is

depends on the intensity

turned
of

the

MOVEMENTS,

OP FRESH-WATKR PLANARFANS.

579

Witli stimuli just effective iu calliug- forth

stimulus.

reaction the turn

liead

ETC.,

is

only slight, and since

it

end the direction of movement of the whole animal

The amount

be scarcely changed at

all.

anterior end

from 30

There

is

typically

the

affects only the

ma}'^

of turning of the

to 40.

the negative reaction a pause at the instant of

stimulation, pi-eceding the turning away.
The first effect of
is in

to cause the animal to stop its relatively rapid

the stimulus

is

movement.

This pause

may be

on the other hand, may lengthen

interval
istic

when the stimulus

is

almost

so slight as to be

weak

stimuli, or,

a quite

noticeable

imperceptible in the case of comparatively

to

very strong.

It is a character-

feature of both the positive and negative reactions of

and

is evidently duo merely to the fact that before

something involving a change of motion) the
former movement must stop.
The effect of localisation of mechanical stimuli in the head
region may next be considered. As has already been men-

planarians,

a reaction

tioned,

(i.

e.

stimulation of the sides of the head produces the

positive or negative reaction according to the intensity of the

stimulus.

There are no special regions of

specific

organs connected with either of these reactions.

tive response is given after strong stimulation of

one side or the other of the head and, so far as

to observe, just as decidedly after stimulation of
of another.

It

is

of interest to

any part of
it is

possible

one part as

know what happens

after

median line. It is very

difficult to get a stimulus exactly in the median line, but one
may come very near it by stimulating the dorsal surface of
the liead in the region between the eyes. The reaction produced is a longitudinal contraction of the anterior part of the
body, drawing the head back away from the stimulus. The
head is then turned to one side or the other as in the usual
negntive reaction, and the animal starts ahead again in the
new direction. The side towards which the turn is made
that is, there is no
after median stimulation is indeterminate
tendency to turn in more cases towards one side than towards
stimulation

of

the head

sense-

The nega-

iu

the

EAYMOND PEARL.

580

been found by Prandsen (: 01) to be the

This is what would be expected in the
case with Li max.
case of the flat-worm, because it is a perfectly bilaterally symProbably what actually determines which
metrical organism.
the

ofclier,

as has

way the organism

shall turn after

attempted median stimula-

tion is the fact that the stimulus really acts a little to one

side

or the other, and the turning

is

really the negative

reaction.

end of a worm with

moderately strong mechanical stimuli its reactions may be
In the beginning of such a series of stimulation
modified.
the worm turns away farther and farther from each succeeding stimulus, at the same time remaining at the same
place in the dish, i. e. not making any progressive move-

By

repeatedl}^ stimulating the anterior

make

This process tends to

ments.
circle

away from the

fixed

point.

It

stimulus, about

the animal describe a

its

posterior end as a

never completely describes a

ever, but after several stimuli

howwhich it

circle,

have been given, to

responded progressively more vigorously, it finally

jerks back with a strong longitudinal contraction, and turns
the anterior end through a considerable arc, so that it points
has

in

an entirely different direction.

This final strong reaction

end towards the

in other words,
which
the
stimulation
is
coming,
or,
from
side
in an exactly opposite direction to that of the previous
This reaction appears as if, after the animal has
reactions.
tried in vain to get away from an uncomfortable stimulus by
in the majority of cases turns the anterior

its

ordinary reaction,

a wild jump in the oppo-

it finally tries

This curious change

in the i-eactions induced

have observed many times.
It indicates the effect of the organism as a whole on its
As von Uexkiill (: 00, p. 73) has well brought
refiexes.
out, we must consider that in the case of a higher organism,
site direction.

by a repetition

of strong stimuli I

like a dog, the animal

moves

organism whose activities are

urchin it is really the " legs"

move the animal.

its

legs,

reflex
(i.

e.

while with a lower

for example,

the sea-

locomotor organs) which

In the flat-woi-m the movements of the

MOVEMENTS,
whole organism

are

in such

yet

reflexes,

OP FRESH-WATER PLANA1?TANS.

ETC.,

determined

by

exceptional

cases

definite

as

581

stereotyped

one

the

just

described the organism as a whole takes control, and does

something quite different from what the normal reflex fitted

to the case wonld accomplish.
Very sti'ong mechanical stimulation of the anterior end,
such as to

wound

the

much more

animal, causes a very

vigorous reaction than the ordinary negative one, and of a

slightly different form.
The animal contracts strongly longitudinally, and, as a result of the heavier musculature on the

ventral surface, curls the head in under the body.

Then the
turned to one side through a larger angle
usually the case, and the worm straightens out in this

anterior end

than

is

is

The point of importance to be noted in this

maximal stimuli is the curling under of the head.
The turn away from the side stimulated frequently is so great

new

direction.

reaction to

as to turn the animal squarely about, so that

dii'ection opposite to that before stimulation.

effect of

maximal

stimuli just described, they

it

heads in the
Besides this

may

also pro-

duce a change in the movement from gliding to crawling.

The crawling does not usually follow stimulation of the
head end of the body, but it is possible in some cases to produce it by very strong stimulation here. I have also been
able in a normal animal to induce crawling backward by
very strong and continued stimulation of the anterior end of
This backward crawling, when it occurs, is of the
same character as the same movement in a forward direction,
the body.

except that

all

described above

the
(cf.

factors
p. 551).

are

reversed.

It is

It

much more

has

been

easily pro-

duced after certain operative procedures, and in connection

them further details regarding it will be brought out.

Avitli

The negative reaction, o. that to strong stimuli, is given

more frequently than any other in the course of the activity
of the individual, and apparently does not depend on the
i.

presence of an 3^ special ph^^siological condition. It is given

in response to stimuli covering a wide range of intensit}^.

The lower liminal value

of the stimulus producing

it

(there

RAYMOND PRARL.

582
is

some extent with the

'Julius
in some
which would ordinarily

appai'ently no upper limit) varies to

condition

physiological

of

the

individual.

specimens at certain times stimuli

produce a rather strong negative reaction will call forth
nothing but the positive reaction. This condition is only a
transitory one, and the reason for it seems to be a heightened
Specimens exhibiting this
tonic condition of the animal.
relation to rather strong stimuli are always very active, and

move about with great

anterior end of the

rapidity,

frequently

body and waving

it

raising

the

about through the

water as they glide along. Persistent strong stimulation

of the organism rapidly changes the general physiological
condition.

This

is

not more true of stimulation applied to

the head region than of strong mechanical stimulation of any

The animal becomes '^ stirred up "genepart of the body.
rally,

moves about with increased rapidity, its sensitiveness

becomes diminished, and it will give only the

to stimuli

negative response to stimulation of the anterior end.

change

in the

physiological

result of continued stimulation of

experiments,

is

any

This

the animal as a

condition of

sort, as in a series of

a matter of great practical importance in

One may get totally different appearances from an individual Avhich has been " stirred
up " from what are seen in the case of one which is in the

connection with reaction work.

normal condition. This is only one of a number of factors

which must be taken into account in woi'k on the reactions
and behaviour of an organism if one is to obtain trustworthy
It is almost an absolute necessity that one should
results.
become familiar, or perhaps better intimate, with an organism,
so that he knows it in something the same way that he
knows a person, before he can hope to get at even an approximation of the truth regarding its behaviour.
The positive reacj3. Reactions to Weak Stimuli.

tion

is

the characteristic reaction given to

all

weak

stimuli.

an orienting reaction in the sense that it brings the

anterior end of the animal in a position such that it points

It is

approximately towards

the

source

of

the

stimulus.

On

MOViaiENTS, ETC., OF FRESH-AVATRR PLANARFANS.

account of
strength

it-^

of

583

fineness of adjustment witli refiM-ence to tlie

the

and

stinuilus

condition of the animal,

general

tlie

pli^'siological

a response which might

be
very easily overlooked in a superficial examination of the

As

behaviour.

the

is

it

worm

gives

positive

this

reaction

in

response to a gentle stimulus, turning the head towards the

source of stimulation, and at the same time raising
gives one the impression that

such

behaviour

the

animal psychologists.
fact that

if

would

it

is

it

be called by some

doubtless

This impression

the head does not

it,

seeking something, and

come

enhanced by the

is

contact with the

in

first reaction, the animal advances

from which the stimulus came, with the
anterior part of the body raised and waving from side to

stimulating object at the

in

the

direction

side in the water.

As has been mentioned,

adjusted

physiologically.

reaction

the
In

the

very

is

majority

of

delicately

the

cases

animal must be in a comparatively quiet condition, that is,

not " stirred up ^' or excited, and gliding smoothly at the
ordinary rate, in order that the reaction

may appear

at

The stimulus must ordinarily be very weak, and given

not to disturb the animal by abruptly changing the
rounding conditions.

by the use

It is possible to

of a needle or scalpel point

all.

so as

sur-

produce the reaction

if

sufficient care is

taken, but better results are obtained by the use of a hair

The point should be lightly touched

edge or dorsal surface of the head, and then quickly
drawn a short distance away. Even when all these precautions are taken one may fail to produce the characteristic
response.
I have frequently found that the same specimen
which at one time would give the positive reaction in a very
as the stimulating point.

to the

definite

and

characteristic

way

head could not be made to show

shows how closely it depends
conditions.

to every light touch

on the

This
a few hours
on general physiological
On the other hand, specimens will frequently
it

later.

be found that for short periods of time (two or three hours)

can hardly be induced to give any other response to median-

RAYMOND PEARL.

584

ical sfciraulation of tlie head.

Stimuli strongs enong'h to be far
above the usual upper limiual value for this reaction will call
it forth.
Such specimens show the reaction in a much more
pronounced type than is usually the case. After a stimulus
lias been given they will turn towards it, and if the source is

not touched immediately they will remain in the same spot

waving the head about the region from which the stimulus
came, at the same time stretcliing the anterior end of the
body far out in all directions, precisely as if in search of the
Usually this

stimulating body.

condition

hypei'sensitive

passes off in a short time, and the animals behave again in a

more normal fashion. It was thought that possibly this

condition was due to hunger, but experiments^ devised to
was not the case.
We can only say that it is due to some intimate physiological condition, the exact nature of which we do not know.
Another fact which may be mentioned in this connection is
that sometimes a specimen in normal condition will give the
test

question

this

indicated that this

positive reactioti in response to a certain strength of stimulus

only a part of the time.

Other

trials I'esult in entire indiffer-

ence on the part of the organism.

possible

to

give

Of course,

it

is

not

the same

mechanical stimuli always of

strength, yet with the closest possible approximation to this

by an experienced operator, some

the animal in any

way except

of the trials will not affect

to cause a slight local con-

traction at the point on the head

glides

along

without

any

in

rate

Altogether we must conchide that the reaction

is

The worm

stimulated.

change

or

direction.

is

one which

very closely dependent on the existence of certain definite

internal conditions as well as the external ones.

The

typical course of the reaction

is,

as has

been described,

momentary pause, followed by a turning of the head

towards the stimulus, accompanied by a raising of the antefirst

rior part of the body.

many

there are

From

variations.

this typical

The

form of the reaction

raising of the anterior end

from the bottom just before and during the time

'

it is

See section on " Reactions to Fooii and Chemicals."

being

MOVEMENTS,

OE FRESH- WATEli PLANAttlANS.

ET(J.,

585

turned towards the source of the stimulus may be entirely

omitted.
In this case the head is swept around towards the
stimulus without being any further i-aised from the bottom

The duration

than in the ordinary glide.

immediately

great variation.
ceptible, the

stimuhition

following
It

may be

worm sweeping

pause

the

of

subject

likewise

is

to

so diminished as to be imper-

the anterior end around through

the water without any change in the rate of the glide.

The

amount of the turn varies with the point of application of the

stimulus, the head being turned just far

enough

This orieuta-

the direction from which the stimulus comes.

tion,

we may

if

stimulus

is

so call

it, is

to point in

If the

generally quite exact.

near the middle line on the edge of the head the

turn will be only through a few degrees, while

are touched

it

will

amount

if

the auricles

This fact indicates

to nearly 90.

the remarkably well-developed co-ordination of the reaction.

There

is

a great deal of variation with regard to what takes

place after the turn has been made, and the anterior end
directed towards the stimulus.

removed immediately

is

If the stimulating point is

after stimulation, so that the animal

by means of the first reaction, a normal

lower the head and continue gliding in
As has been mentioned, however, in
the new direction.
some cases a specimen will continue " feeling" about in the
does not touch

specimen

it

will usually

locality for

some

time.

If the stimulating point

is

held in

been given,
the first reaction will in most cases bring the head into
In this event the animal usually moves the
contact with it.

about

its

original position after the stimulus

tip of the

head about over the hair

luis

(or other point)

for a

short time, and then drops back to the bottom and continues
gliding.

In other cases

it

will clasp the anterior

end about

the hair (as in the feeding reaction to be described later),

and then in a moment start gliding up over it. When this

happens the hair or needle may be moved about in the water
or even lifted out of it, and the animal will not let go its
hold and drop

animal

off.

If the needle

will in a short

time glide

is

held

down

however, the
and proceed on

fjuiet,

off

it

EAYMOND PEARL.

586
its
is

way along the bottom.

This behaviour when the animal

able to reach the stimulating- object

Avhicli

is

evidently the action

most frequently occurs in natural environmental con-

ditions.

With

reg'ard to the localisation of the stimulus producing"

may say that I have been able to produce it

by proper stimulation of any part of tlie edge or dorsal
surface of the head region under favourable circumstances.
It seems to be moi-e certainly produced
that is, in a larger
number of cases by stimulation of the auricles than of any
other part of the head, and it may be that in this is to be
found an indication of the chief function of these sense-organ
bearing structures. At any rate, this is the only indication
of a special function for them which I have been able to
this reaction, I

discover.

The

positive reaction given in

response to light

stimulation of the dorsal surface of the head

somewhat

is

necessarily

from the typical reaction which has been

described.
In this case there can be no turning towards one
side, because if this were done the liead would not be directed
towards the source of the stimulus.
Instead, what takes
place is this the head is sharply raised and twisted, so as
to form a part of a spiral in the region posterior to the head.
This brings the anterior end into a position pointing towards
the source of the stimulus, and at the same time the ventral
surface is brought around so as to be, in most instances, the
tirst portion of the body
to touch the stimulating point.
different

This reaction, following stimulation of the dorsal surface of

the head,

is

not an easy one to obtain.

best in producing

it in

sensitive condition

With regard
call forth

have succeeded

the case of individuals in the hyper-

mentioned above.

to the strength

of the stimulus necessary to

the positive reaction, only very relative statements

may be made.

Unfortunately

Ave

have no method of measur-

ing the intensity of such weak mechanical

used in work on lower organisms.

stimuli as are

Our only idea

of the

strength of the stimulus must come from the reaction of the

organism

itself.

It

must

suffice to say,

regarding the reaction

MOVKMENTS,

ETC., OF

FRESH-WATER I'LANARIANS.

587

uirIlt discussion, that iu au auinial in a condition of liypor-

have been able to produce the reaction by the

weakest stimuli whicli I was practically able to give. Under
normal conditions of sensitivenes it takes a slightly stronger
No absolute value can be given for the upper
stimulus.
limen of the reaction, beyond which it docs not appear, but

seusitivity I

gives place

to

negative

the

may be made

This value varies

reaction.

The general statement

greatly with different individuals.

that the positive reaction

is

the characteristic

response to stimuli of very low intensity, and

its

production

very closely dependent on the proper gradation in the

This dependence is so close that it is possible to
intensity.
is

obtain a part of both the negative and positive responses

combined

in the

same reaction by

tlie

use of a stimulus of

have been able in a few very favourby

a single stimulus a pronounced
produce
able cases to
raising of the head, such as is characteristic of the positive
reaction, followed by a turning away from the source of the
stimulus. Now the raising of the bead is no part of the typical
negative reaction, and, furthermore, was done in the very
characteristic way in which it occurs in the positive reaction.
The stimulus whicli produced it was evidently about intermediate in intensity between what, in the case of tlie particular animal used, would have called forth either the positive
or negative reaction, as the case might be. This experiment
shows in a very striking and conclusive way that in both
the positive and negative reactions we are dealing with a
complex of reflexes, since here a part of one of the reactions
This point will be
is associated with a part of the other.
alluded to again in another connection, and its significance
the proper intensity.

more fully pointed out.

The evident purposeful character
plainly apparent.

It is

of the positive reaction

is

a reaction admirably suited, on the

whole, to bring the organism into contact with beneficial

things, such as food, etc.

It

seems

to

me

that

it

nnist be

by

far the most important reaction of the animal in the struggle

for existence.

In the

conditions

under which planariaus

RAYMOND PEARL.

588

a reaction which gets

live

it

foodj or helps to^

is

of far

greater importance for the survival of the individual than a

reaction which takes
tion

out of dauger;

can show, the dangers

move over

It does not

as

it

is

known,

it

it

for, so far as

observa-

encounters are relatively few.

large areas of territory, and, so far

does not furnish a considerable part of the

food supply of any other organism.

struggle for existence would

seem

Altogether

its

chief

to consist in obtaining

subsistence for itself, and for this the positive reaction to

mechanical stimuli would appear to be an important aid.
As will be shown later, the food reaction proper consists

largely of this same response.

We may

now pass to a consideration of the

Reactions to Stimuli applied to the Middle
Region of the Body. I use the term "middle region of
2.

the body " to distinguish that portion extending from the

of the head to about the middle of the
pharynx. The separation of the body behind the head into
a " middle '^ and a " posterior " region is based entirely on
physiological considerations, and is not defined morpho-

posterior border

logically.
a.

Reactions to Strong Stimuli.

Strong mechanical

stimulation of the middle region of the body along the lateral

edges causes, in the

body

in the

contraction

is

well

the head region.

especially

if

first

instance, a local contraction of the

immediate region of the stimulus.

marked; much more

This local

distinct than that in

If the stimulus is sufficiently strong,

the stimulating point

is

and

applied to the edge

from above rather than from the side, the previous gliding
movement will be changed to crawling. This will continue
for a brief interval, usually from two to four crawling contractions being given; then the animal will relapse again
into the glide, provided the stimulus is not repeated.
In
the case of a strong stimulus applied

middle region of

the

several times repeated,

body, especially

to the
if

the

side of the

stimulus

is

we get the negative reaction

turning away from the side stimulated

just as in ihe similar

MOVEIFENTS, ETC., OF FRESH-WATER FLANAIJIANS.

case in

head region.

tlie

The nearer

end of

more easily
the negative reaction he produced, while back in the

the middle region the stimulus

will

to the anterior

589

pharyngeal region
cases.

In

all

it

is

applied, the

follows even strong stimulation in fewer

cases where this reaction

movement

is

not produced the

unchanged by mechanical
direction of
stimulation, or the anterior end may be brought around
vei'y slightly
i*esult of

towards

is

either

the side stimulated as a mechanical

the local contraction on that side.

Hy repeating
summa-

the strong stimuli on one side of the middle region,

tion effects similar to those described

when

the head

is

above as taking place

similarly treated are not produced.

animal crawls faster and faster away from the stimulus.

movement

The
Its

changed, but usually not more than

We see here evidence of precise
thirty to forty degrees.
response to localisation of stimulus. Stimulation of the head
direction of

is

turn to one side, and, in case the

very strong, to contract longitudinally strongly
As we go back from tiie head in the
before doing so.
middle region of the body, the tendency to crawl rapidly
causes the animal to

stimulus

is

ahead away from the stimulus increases. At the same time

the turning away from the stimulus becomes less and less
marked the farther back it is applied. In no case do we get
any strong retraction of the anterior end, which, in case of
stimulation of the middle region of the bod}^, would tend to
bring the animal, or at any rate a part of it, into further
contact with the stimulus.

Strong stimulation applied to the dorsal surface of the

middle region of the body causes the animal to change from
This change of the form of motion
the glide to the crawl.

may be regarded

as a specific reaction in response to strong

stimulation of the middle or posterior regions of the body.

Stimulation of the dorsal surface of the middle region does

not change the direction of the movement unless the stimulus

is applied near the lateral margin, in which case it may
cause the negative reaction, as mentioned above.
/3.

Reactions to

VOL. 4G, PAUT 4.

Weak
NEW

Stimuli.

SEUIES.

Weak

media nical
Q Q

RAYMOND PEARL.

590
stimulation of

tlie

sides of the

middle region

of the

body

at the
causes, in the first instance, a small local contraction
direction
general
place stimulated, without any effect on the
movement of the whole organism. Under favourable
of

circumstances, however,
quite different result

it

is

frequently possible to get a

of a weak stimulus on the

by the use

margins of this region. Specimens of Planaria may

reaction
be induced to give the characteristic positive
as the
back
far
as
point
of
a
described above by stimulation
it
inducing
stimulus
The
middle of the pharyngeal region.
prowill
which
as that
is of the same intensity and character
duce the same result in the head region.
Some experiments bearing on this point will be reported
theoretical
in full, on account of their important bearing on
were to
experiments
These
later.
up
taken
questions to be

lateral

Diagram to show the portion of tlie side of the body {a I)

Fig. 15.
within wliich weak stimulation produces the positive reaction.
test the effects of

weak

stimuli on the sides of the middle

region of the body, especially with reference to the relation

of the physiological condition of the individual to its

The

reactions.

specimens

P.

of

stimulated was

experiments

were

maculata, and

performed

the region

that from a to h in Fig.

15.

of

on

large

body

the

Most

of the

and
The stimuli were given by
the origin of the pharynx.
moving the point of a fine scalpel along the bottom of the
dish till it came into contact with the margin of the bod}^.
In this way no general disturbance was produced. The

stimuli were applied in the region between the auricles

attempt was made to make

possible the

the

stimuli

same intensity each time.

of

The

as nearly as
results

were

classed as positive, negative, or indifferent, according as the

MOVEMENTS,

ETC.,

OF FRESH-WATER PLANARIANS.

591

specimens gave the usual positive or negative reaction, or

kept their course without change, showing only the local con-

The

first series which I will report was on an

which was in a condition of great excitation,
moving about with more than normal rapidity, and generally

traction.

individual

''

The

stirred up."

this

results of twenty-three stimulations on

specimen were

Expt.

Positive responses

I.

JNegative responses

Indifferent responses

experiment

similar

1\
p.

J.-

Tvr

0,'-

Specimen
^

22J

another

with

individual

entirely normal unexcited state, gliding at a

gave the following results

Expt.

II.

The

in

moderate

an

rate,

Positive responses

20^

AT
iJNegative
responses

o
2

Indifferent responses

ISJ

Specimen
^

in

normal

striking preponderance of the positive reaction in the

case o the

unexcited individual

it

notable.

is

individual used in Experiment II was

poking

in state of

The same

" stirred

now

up" by

violently about the dish with a needle for about

minutes. It was then allowed to settle into a glide

which was at a more rapid rate than normal, and another
series of stimulations was made, with the following results

five

Expt. II r. Positive responses

XT

iNegative
responses
T :i-cc
Indinerent responses

^1
o

o
8;

i.

Specimen

in

condi-

'
.

tion of excitation.

Here, again, the indifferent responses are in excess, and

there are practically no positive reactions. The specimen
was again " stirred up " in the same way as before, and

another series taken.

Expt. IV. Indifferent responses
1

(trials

to 11 inclusive)

Specimen

Positive responses (trials 12

and 13)
Negative responses

(0)

in state of

excitation.

RAYMOND PEARL.

592

The specimen was ag-nin stirred np in the same

another series taken, with the following resnlts

way and

Expt. Y. Indifferent responses

1 to

(trials

9 inclusiv^e)

Specimen

Positive responses (trials 10

and 11)
Negative responses

The

(0)

positive responses in all these experiments were very

definite
in

in state of

excitation.

and

many

characteristic.

other series of

recorded in detail.
in order for the

have obtained the same results

experiments, which need not be

The experiments show very

clearly that

animal to give positive responses to weak

it be in an unexcited condition.
have also an important bearing on the question
of the mechanism of the positive response, in that they show
conclusively that the reaction does not depend on the stimu-

stinn;li it is

These

necessary that

resnlts

lation of special sense organs located in the

head regions

alone.

Weak

mechanical stimulation of the dorsal snrface in the

middle region of the body

is

usually without any effect other

than the causing of a slight local contraction at the point

If any specific effect on the whole animal is
stimulated.
produced,

it

is

merely a change from the gliding-

to

the

crawling movement^ such as results from strong stimulation

in the

same region.

Reactions to Stimuli applied to the Posterior

Region of the Body. By '^posterior region of the body
I mean that part of the body from the pharyngeal region to
3.

the posterior end.

'^

This region

is

not sharply marked

physiologically from the middle region, and

to say in

any given individual

at just

what

it

is

off

impossible

level the

demar-

be found. The physiological distinction between

the two regions is founded on the fact that it is possible by
unilateral stimulation of the middle region of the body to
cation will

produce a change in the direction of the movement of

the animal as a whole, while in case of the posterior region.

MOVEMENTS,

ETC., OE

FUESH-WATEK I'LANAKIANS.

On

as will be showily this canuot be done.

will not

bo necessary

in

tin's

593

uccounfc

it

the description of the reactions to

sliarply distinguish between the effects of stimulation of the

margins and of the dorsal surface^ as has been done in the

previous cases.

Strong mechanical stimulation of the posterior region of

the flat-worm produces as a specific reaction an immediate
change from the gliding to the crawling movement. The
direction of the crawling is the same as that of the gliding;
that is to say, the worm keeps on in a straight lino, taking
itself directly and in the quickest possible way away from
the

The duration

stimulus.

of

crawling movement

the

following stimulation of the posterior region varies with the

and the physiological conmost

usual number of the
dition of the specimen.
The
strong, crawling contraction waves following strong stimularelative intensity of the stimulus

tion

is

We

three or four.

may

number than

get a smaller

and very frequently do, but iu the species studied I

have very rarely seen more than four of the general con-

this,

tractions following a single stimulus.

This

evidently

is

all

that would be necessary under normal circumstances, since

the animal a

four of these strong contractions will carry

considerable distance ahead, and probably out of reach of

the stimulating agent.

The weaker the stimulus

is,

the fewer

are the contractions and the shorter the distance crawled.

In some individuals
the crawling

it is

at times almost impossible to induce

movement except by repeated

Such specimens

will

stimulation.

merely draw up the posterior end in a

single crawling contraction after stimulation, and then immediately relapse into the glide. If a strong stimulus is
repeatedly given at the posterior end the crawling is con-

tinued,
effect of

becoming more and more

rapid.

This

is

the only

continued stimulation in this region, there being no

effect corresponding to that produced by stimu-

summation

lating the anterior eud.

No

different effect

is

produced by

stimulating the margins of the posterior region of the body

from what takes place when the point stimulated lies near

RAYMOND PEAUL.

594
the middle

any part

line.

There

is

unilateral stimulation

no turning towards or away of

of any special effect of

The lack

of the body.

not surprising, for the reason that

is

rapid movement in a forward direction will get the animal

away from harmful stimuli affecting this region, in the long
Further, there would be
run, more quickly than any other.
no advantage in the production of a positive reaction by
stimuli at the posterior end.

If

we think

of these reactions

by natural selection there would

be no possibility of such a reaction having arisen, for the
reason that practically any favourable stimulus would be
encountered by the anterior end before it possibly could be
by the posterior. Very weak mechanical stimulation of the
posterior end of the body causes only a local contraction at
as having been developed

the point stimulated.

Reactions to Stimulation of the Ventral Sur-

4,

face.

In

the descriptions of the reactions to

up

stimuli

this

to

mechanical

we have been considering

point

stimuli

applied to the dorsal surface and to the margins of the body.

It

may be

well to describe briefly

what the reactions

in

response to localised stimulation of the ventral surface are.

This matter can best be tested when the animal is moving on
the under side of the
that this
testing

surface film, with

ventral side

its

It

reactions to ventral stimulation, but, as a matter

of fact, the conditions are

and

its

might be supposed before the trial was made

habit of the animal would afford ideal conditions for

uppermost.

anything but

elasticity of the surface film

makes

ideal.
it

The

flexibility

almost impossible

touch it with a stimulating point anywhere within a radius

of a centimetre about a planarian without causing the animal
to be jerked bodily to one side or the other, quite sharply
to

This is, of course, a mere

distance.
which takes place with lifeless bodies also.
Furthermore, as has been mentioned in an earlier section, it
appears to be very difficult for planarians to quickly change
the direction of their movement when on the surface film (as

and

for

some

mechanical

is

little

effect,

necessary in reacting to stimuli).

On

account of these

MOVEMENTS,
conditions

ETC., OF

very

FRESH-WATER PLANARTANS.

595

any certain and trustworthy results from the stimulation of the ventral surface.
My results have been as follows: strong stimulation of the
anterior end on one side of the middle line causes the
it

is

difficult

to got

when

negative reaction just as

the stimulus

is

corresponding point on the dorsal surface.

reasons the response
is

is

not as extensive as

on a solid, but there seems no doubt of

positive reaction to

weak

applied at a

For mechanical

when

its

the animal

character.

The

stimuli I have not been able to

produce in any certainly recognisable form in response to

stimulation of the ventral surface, but

think this negative

due probably to the external conditions, and not to a

real failure of the organism to react.
Strong stimulation of
the posterior end of the body causes the gliding to change
to the crawling just as under other conditions.
Very
strong mechanical stimulation of the ventral surface of the
body causes the animal to let go its hold and pass down
result

is

to the bottom.
5.

Reactions of Kestin g S pecimens

Stimuli.
to

weak

stimuli which arc

definite reaction

The stimulus

to

Median ical

resting specimen gives no response whatever

is

when

the

still

strong enough to produce a

worm

is

in

the active condition.

simply below the threshold of the resting

To stronger

animal's sensitiveness.

stimuli

the

reactions

correspond in form with those given by the active animal,

but are less pronounced. For example, rather strong stimulation at the anterior end induces a weak negative reaction
similar stimulation of the posterior end sets the animal off
Strong stimulation of any part of
into the crawling motion.
the body besides producing the characteristic reaction for
that region (that
cases start the

the case

if

is

the negative reaction) will also in most

animal into movement.

the stimulus

times repeated.

is

of sufficient strength, or is several

As would be expected from

tiveness of the resting flat-worm,

forth from
G.

it

This will always be

it

is

the low sensi-

impossible to call

any positive reaction.

Reactions to Stimuli given by Operative Pro-

RAYMOND PEARL.

596

cedure.

Evidently

wlieu

a plauarian

induces a strong stimulation,

cut

is

wliicli is of

the

cutting

tlie

same kind as

induced by ordinary mechanical stimuli, only much

more intense. The immediate effects of operations may then
be taken up in this section.
If we take first the typical case given by cutting the
animal transversely in two in the region between the posterior border of the head and the origin of the pharynx, and
make the cut by a single stroke of a sharp scalpel, we find
that the effect on the anterior piece is precisely the same as
that of an ordinary strong mechanical stimulation of the
same place. That is, this piece merely changes from the
gliding to the crawling movement, and after giving three or
that

crawling

four
glide.

This

coiitractions

is

settles

down again

into

the

the same result essentially as that obtained

by Norman (: 00) and earlier by Loeb ('94 and 00). In the

behaviour of the posterior piece in this experiment under
discussion there is a great deal of variation.
In about 70
per cent, of all cases in which I have observed the results of
:

such an operation, the posterior piece crawled backwards

In the remainder of the cases the

as a result of the cut.

piece

either

stayed

in

the

violently, or else glided ahead.

ward crawling when

this

same place and contracted

The amount of the back-

occurs

varies

from a

greatly,

short distance involving only one longitudinal crawling contraction to several times the length of the

ment

worm, the move-

lasting in this latter case for over a minute.

that this backward crawling

and distinct form

above a certain

it

is

size.

may appear

In order

well-marked
necessary that the posterior piece be

Very small

in a

posterior

pieces

after

operation usually remain quiet.

cut so

made

as to split the anterior

the middle line in most cases causes the

wards just as does a transverse

cut.

end of the body in

to crawl back-

worm

In some cases

this, as

well as other operations, merely causes the animal to contract

and squirm about at the same

posterior end of the body in the middle
violently

place.

Splitting the

line causes the

parts

MOVEMENTS,
on either side of

is,

worm

as a whole starts crawling ahead

from

changes

it

597

cut to give violent lougitudiual con-

tlio

tractions, while the

that

FRESH-WATKll PLANAUIANS.

ETC., OF

the

gliding

to

crawling

the

movement.
Oblique cuts produce essentially the same effects as would
trausvei'se cuts in the same part of the body, i. e. forward
crawling of the anterior piece, and usually backward crawling
This

of the posterior piece.

is

true unless the cuts are very

form very acute angles with the sagittal

In such cases the effects produced more
the body.

oblique, so as to

plane of

nearly resemble

obtained in

those

splitting of the body.

If the

body

two parts longitudinally, there

gressive

movement

is

complete longitudinal
is

split

completely into

usually very

of either piece afterwards.

little

The

pro-

pieces

contract strongly on the cut sides very soon after the operation

is

which

performed, so that they take on the form of a bow,

many instances becomes a nearly complete circle.

in

This being the case, any progressive movement, either by

Cuts
is nearly or
quite impossible.

gliding or crawling,

involving only

produce,

if

small

portion

of

one side of the body

in the anterior region, the characteristic negative

reaction given to other strong mechanical stimuli, while

if

in

the posterior region they cause the crawling ahead.

Cuts made on the resting animal produce essentially the

Unilateral cuts
effects as on the gliding specimen.

same

have the same

7.

effect in

The Effect

ment.

coclum,

series of

of

sp.,

producing the negative reaction.

Mechanical Hindrance

experiments was performed on

MoveDendro-

to

with reference to the behaviour of the animal

when progressive movement was made

impossible, and yet

the animal was stimulated strongly at the same time.

These

conditions can be realised by thrusting a needle through the

centre of the body from above, and then holding
position.

The

results

of

this

according to the portion of

In case the hindrance
needle was thrust.
region of the body,

e.

it

fixed in

procedure varied somewhat,

the body through which the
is

in the posterior

g. at a point just behind the posterior

EAYMOND PEARL.

598

end of the pharynx^ the

effect

thnistiiig iu of the needle

is

immediately following the

a strong longitudinal contraction

whole body. After this first strong contraction the

animal remains perfectly quiet in the contracted form for a
varying length of time (in some cases as long as five minutes,
of the

but usually less). After this period of quiet a series of

rhythmical waves of contraction pass longitudinally over the
still

contracted

evidently to

The purpose

body.

loosen

of

these

restraining object by

the

waves

is

making the

hole in the body through which it passes lai'ger.

This is the
same behaviour that I have observed in the deposition of the

This process of rhythmical longitudinal contrac-

large egg.
tion

is

continued for a time; then the animal stretches to

its

extreme lengthy attaches the anterior end to the substrate,

and attempts to crawl away. The movement of the anterior
end is precisely the same as in crawling. The animal turns
and twists and struggles violently in this attempt to crawl
away, and the cilia beat strongly. If the needle occupies a
position near the edge of the body this first struggle will
usually be sufficient to tear the body loose from the needle, so
that the animal may then move ahead freely. Such specimens
will, of course, have a large jagged wound in one side of the
body, which, however, closes in and heals in a short time. In
case the first struggle of the extended animal to craAvl ahead
is not effective, that is if the needle is too far in towards the
centre of the body to make the tearing out possible, the
animal, after continuing the struggle for a time, contracts

strongly longitudinally and goes through the whole series of

stages

of

quiet,

rhythmical,

attempted crawling again.

first

and succeeding

the

animal

is

longitudinal

The only

series of trials

contracted

strongly

become shorter with each

In case the needle

is

is

contraction

and

difference between the

that the stages in which

longitudinally

tend to

repetition.

thrust through the body in front of

the pharynx, the strong longitudinal contraction appears as

before,

and

is

followed after some time by an extension of the

part in front of the needle, while the rest of the body re-

MOVEMENTS,

OP FliESH-WATEU PLANARIANS.

ETC.,

599

muins quiet iiiid coutracted.

This short anterior region,
hardly more than the head, goes through the
crawling movements, but on account of its small size is very

including-

'

body away from the needle

experiments I have never seen any
getting free from a needle put through the

ineffective so far as pulling tiie

is

concerned.

worm
body

In

succeed in

my

in this position.

This general behaviour of the animal in response to restraint

of

movement

is

very interesting, especially in the cases where

at the posterior end, as showing the relation

between the behaviour and the capability of regenerating.
The organism tears itself loose from a restraining body with
entire nonchalance, as it were, and its confidence is well
founded because no permanent harm comes from the action.
The lost and wounded parts are regenerated and healed in a
short time.
The behaviour takes advantage of the ability to
regenerate.
Whether the form of behaviour (pulling away
from restraining objects) or the power of regeneration and
reparation appear in the organism first we cauuot say, for
either might very well follow, in a more or less remote causal
connection, the other. What we do know is that at present
there is a very nice condition of mutual adaptation between
the two things.
The effect of the hindrance of a rather light weight at the
posterior end of a worm is to induce the crawling movement.

the restraint

is

This can be seen in case the animal

is

feeding on a small

piece of food material, and, as frequently happens, starts into

movement before the pharynx

is withdrawn.
The piece of
food attached to the end of the pharynx is dragged along
behind, and the movement is the crawling. Frequently, also,

in feeding

experiments pieces of food

posterior end of the

of t])e body,

worm by means

will

of the

and these have the same

get stuck to the

mucous secretion

effect in

inducing the

crawliug movement.

Having now obtained a descriptive basis we may pass

some general features of these reactions.

discussion of

may

first

take up

to

We

RAYMOND PEARL.

600

c. The
General Features
Mechanical Stimuli. From

the Reactions to
above description it
appears that the nature of the reactions to mechanical stimuli
depends upon several factors. These are
of

the

The intensity of the stimulus.

The localisation of the stimulus.
3. The physiological condition of the organism.
The reactions given may be of several different
1.

2.

pending on the factors mentioned above.

as follows

kinds, de-

These are chiefly

The resting individual may begin locomotion.

2. The gliding movement may be changed to the crawling
movement.
3. The forward movement may be transformed to movement backward.
4. The animal may turn away from the source of the
1.

stimulus (the "negative " reaction).

may turn towards

5. The animal
stimulus (the "positive " reaction).

source

the

named

It is evident that the reactions last

the

of

the negative

and positive reactions are the most important and most

It is of the
interesting from the theoretical standpoint.
greatest interest to note that these two qualitatively opposite
reactions are induced merely by differing intensities of
stiumli, the stimuli

being otherwise identical throughout.

be noted further that the positive and negative

It is to

reactions have the characteristics of purely reflex acts.

reaction

Each

has a perfectly definite and characteristic form.

While, in some cases, which of the two reactions will be

given in response to a particular stimulus depends on the
physiological condition

ot"

the organism, yet

it

is

practically

always either one or the other of the typical reactions. Only

very rarely do we get any deviation from the type forms,
and in such cases the reaction is evidently a combination of
easily recognisable

components

of the

two typical complexes

of reflexes.

These

two reactions are

evidently

not

single

simple

ETC., OF

MOVEMENTS,

FRESH-WATKR PLANARTANS.

601

reflexes, but arc coiii])lexes of several simple refJex acts.

may be

It

com-

well to present in tabular form the different

ponents in each of these reactions, indicating- by the position

in the table the relations of the parts.

Component Phases
with

Stimuli,
E. e

g'

of the Reactions to Mechanical

special reference to the Head

on

Nkoativk.

rosiTivi';.

A. Moiueutary

of pre-

stopjjiiii,'

A. Same as

in i)OMliv('.

movement.

to as

Referred
" pause " or " lusita-

tiou "

ill

vious

description.

B. Longiludinal

ext.ensioii of the

anterior end

Amount

extent.

less

greater

to

B. Longitudinal contraction of ante-

or

rior

end

of

intensity.

de-

appear more

Usually distinctly noticeable.

longer in duration.

stimulated.

that
is

viz.

C. Turning

defined by the [josilion of

the
lus,

source

of

as

in

less

make A
ar.d

one side,

positive.

viz.

Defined

stimulated.

No

sharp

" orientation."

slimu-

the

towards

not

that

side

Tliis

to

pronounced

pends on previous extension.

C. Turning towards one side,

or

greater

Tends

Sharp

not structurally.

"orientation."

C. Raising

of anterior end.

This

takes place at the same time

as C.

D. Movement towards
Direction
position

end

Time

stimulus.

determined
taken

l)y

1).

by

anterior

Movement away from

Direction

stimulus.

determined

as

in

positive,

at termination of C.

relations arc indicated by vertical position in the table.

occurring at the same time are included

Components

in braces.

components before D may be considered as a

and thus there are in one case four and in the
other case three simple reflexes which go to make up the
whole reaction. That these reactions are composites of the
distinct parts is evidenced, lirst, by direct observation of the
reactions themselves ; and second, by the fact that it is

Each

of the

sinofle reflex,

RAYMOND PEARL.

602
possible

by varying

tlie

parts

of

strength of

tlie

stimulus to produce

without

the

remainder, and, furthermore, that a part of one reaction

may

only

in

certain

the

whole

reaction

be combined with a part of

rare instance
587).

(v. sup., p.

the other

the Reactions. A question which

is of the greatest iniportauce in all work on the reactions of
organisms is, what is the mechanism of the reaction ? In
the case of the flat-worm this becomes, what is the neuromuscular mechanism of the reactions ? Very little direct
evidence bearing on this question can be obtained from the
Taking the positive and negative
reactions themselves.
d.

Mechanism

of

reactions as they occur, there are several different sets of

muscles and of nerve connections by means of which they

might conceivably be brought about. The best evidence on
the question

is

the indirect evidence from operation experi-

ments, in which parts of the

mechanism are injured or

removed.

first

Relation of the Brain to the Reactions. The

specific problem which may be taken up may be stated

thus

1.

is

the brain necessary

normal reactions

to

for the performance of the

mechanical stimuli

Or, in other words,

from which the brain has been removed

This question can be answered
react normally to stimuli ?
from the study of specimens which have been cut in two
transversely, and consequently we may proceed at once to a
description of the reactions of the pieces resulting from such
an operation. A typical specimen is cut in two transversely
at the level of a point about halfway between the head and
the origin of the pharynx, as shown in Fig. IG. As has been
mentioned above, the cut itself acts as a strong mechanical
will a plauarian

stimulus, and the immediate effect of the operation is to set

both pieces crawling, the anterior one ahead and the
posterior one usually backward.
If now the pieces are allowed some hours to recover from
the immediate effect of the operation, and then stimulation is
tried, the following results are

obtained

With the anterior

MOVEMENTS,

ETC., OF FREST[-WATER PLANARIAXS.

piece A, containing

tlie bi'ain,

603

the results are entirely similar

normal animal. Stron<if

head causes the negative
I'eaction, weak stimulation of the same sort the positive
Stimulation at the posterior end causes the crawlreaction.
ing movement to appear, and altogether the appearances are
essentially the same as in the normal complete specimen.
The posterior piece B (lacking the brain) behaves in a
somewhat different manner. If the anterior end of this piece
is given a stimulus of moderate intensity anywhere on the

to

those obtained in case of

stimulation

unilateral

tlie

the

of

cut surface the piece will usually start crawling straight

backwards.
after

This

is

the operation,

almost always true for a short time

is especially well shown in such

and

specimens as started crawling backwards as a result of the

AVhen from twenty-four to forty-eight hours have
cut.
elapsed after the operation this tendency of posterior pieces

Fig,

Heavy

1G. Operation diagram.

line indicates cut.

backward on stimulation

of the anterior end begins

marked, and, as regeneration proceeds, finally
In many such posterior pieces I have been able
disappears.
to produce this backward crawling in a very pronounced
form, and of comparatively long duration (three or four
The chai*acter of the movement has
minutes at a time).
been described above. If the stimulus is applied to one side
or the other of the anterior end of such a posterior piece,

to crawl
to

grow

less

instead of squarely against the cut surface, a well-marked

negative
turns

reaction

is

produced

away from the stimulus

The reaction

is

character

the

as

very

definite,

normal

difference to be observed

is

that

just as a

is, the anterior end

whole animal would.

and of precisely the same

negative

reaction.

The

only

that in proportion to the strength

of the stimulus the reaction is not so

pronounced as

in the

RAYMOND PEARL.

604

being due to the generally lowered tonus

have not been able to obtain any positive
reaction (i. e. turning towards the stimulus) in sucli a
posterior piece after operation.
Stimuli which are at all
This experiment
effective produce the negative response.
has been tried many times, but always with the same result
the positive reaction never appears. If the posterior end of
such a posterior cut piece is stimulated the crawling movement is produced just as in case of the normal complete
animal. As has been noted in connection with the move-

uormal animal^

in

such a piece.

ment, there

is

this
I

a general reduction of tonus in the posterior

pieces resulting from transverse cuts.

This low tonus in-

not only the motor functions, resulting in slower

movement, but also to a less extent the sensory functions.
Such a piece is somewhat less sensitive to mechanical stimuli
The cut surface is more sensitive to
than normally.
mechanical stimuli than any other part.
Now it will be seen from the above description of the
reactions of a piece from which the brain has been removed,
that the most striking difference in the behaviour of such a
piece from that of a normal animal is to be found in the
volves

absence of the positive reaction.

There are three conceivable

possibilities as to the cause of

the absence of the positive reaction in pieces from which the

head has been removed. First, the positive reaction might

be due to the stimulation of certain sense organs which are
removed by the operation. But this is decisively negatived
by the fact that in an entire worm stimulation of points
posterior to the level of the cut removing the anterior end
will cause the positive reaction.

Second, it might be conceived that the reaction is brought

about by a special localised muscular mechanism, which is
removed or destroyed by the cut. But there is no evidence
of the existence of such a

mechanism

and

furthei-, it will

be

shown later that the ordinary musculature of the body, which

is

of course uninjured in

brino-

about the reaction.

the posterior part,

is

sufficient to

JJOVEMUNTS, ETC., OF l'liE8il-\VATEK PLANAUl ANS.

605

might in sonic way be a

which is removed by the
operation.
As the evidence seems to be decisive against the
first two possibilities this seems
probably true. Is this
because the brain contains a special " centre " whose function
it is to produce the reaction ?
There is no reason to think of the reaction as a function of
the brain in the sense that that organ forms a centre which
Finally, the

positive

specific function

of

the

veuctiou
brain,

On

originates the impulses Avliich cause the reaction.

contrary,

seems much more probable that the

it

the

loss of the

brain causes the loss of reaction for the following reason.

It has

been shown that removal of the brain causes a general

lowering of the tonus of the organism, and further that the

appearance of the reaction in a normal animal is closely

dependent on the tonic condition of the organism. Probably,
then, the chief reason for the non-appearance of the positive
reaction in posterior pieces

is

that in these the conditions of

general tonus are so changed by the loss of the brain that the
reaction

is

too sluggish to give

is

This being the case,

possible

Expressing

no longer possible.

the animal

to

it

the

it

in another way,

positive

would be expected that

it

response.

might be

induce the positive reaction in a decapitated

specimen provided the tonus were raised in some way. As a

matter of fact, as will be shown later, positive reactions to

certain chemical stimuli have been observed in a few cases

(cf. p.

is

649).

In

its

form and mechanism the positive reaction

not directly dependent upon the brain.

Summing up

the evidence on the relation of the brain to

the reactions of the flat-worm,

reactions to mechanical stimuli

it

may be

said that all the

shown by the normal animal,

with the single exception of the positive reaction, are given

by specimens from which the brain has been removed.

The relation of the brain to the positive reaction is, in large
be obtained, an indirect one, viz.
of the proper tonic
Thus far there is no evidence of
conditions of the organism.
any special " centre " functions of the brain, similar to those
part, so far as evidence can
it

is

necessary for the maintenance

VOL. 46, PART 4.

NEW SERIES.

R K

RAYMOND PEARL.

606
supposed

exist in the cortical centres, for example, of a

to

mammal.
2.

The Neuro-muscular Mecliauism.

In the negative

reaction to mechanical stimuli the anterior end of the body

is

turned sharply away from the source of the stimulation,

it is equally sharply turned

while in the positive reaction

These relations immediately suggest the

towards the source.

following questions

Is the negative reaction the result of a

crossed impulse, which, originating at the point stimulated,

body and causes the

crosses over to the other side of the

contraction of the longitudinal muscles on that side, thus

producing the turning away from the stimulus ? What is the

course of the nerve impulse which produces the positive
What sets of muscles are concerned in the proreaction ?
duction of each reaction

The
first.

discussion of the negative reaction

If

the

nervous

impulse

may be taken up

pi'oduciug

this

reaction

body to produce a contraction on the side

opposite from the stimulus, the experiment cited in the
section above shows that this crossing cannot occur entirely
in the brain^ but must also occur in some part of the body
or at any rate, be capable of so doing
posterior to the brain
in a quite normal fashion immediately after removal of the
brain.
In this experiment where the body has been cut
in two behind the brain, the posterior piece performs the
negative reaction in a quite normal way immediately after
crosses

the

the operation.

This experiment

may be

carried

farther,

and the animal cut in two transversely in places nearer

and nearer to the posterior end of the body. In all of these
cases, until the piece becomes too small to show definite
movements of any sort, the negative reaction may be obtained
by strong unilateral stimulation. This shows conclusively,
then, that

if

the negative reaction

crossed reflex, there must be

all

cross-commissures which are at

is

all

])reviously

to

do.

be considered a

times ready to bring about

in co-ordinated perfection a result with

had anything

to

along the body a series of

which they have never

This conclusion seems in-

MOVEMENTS,

ETC., OF

because,

ovitablo

sfciniuhitioii

as

FUESH-WATER TEANAUIANS.

has

been

shown

above,

of the posterior region of the

body

607

uiiihitei'al

in a

normal

individual does not cause the negative reaction, but instead

merely causes the animal to move ahead, faster by crawling.

paths for the crossing of impulses which are so

If these

immediately effective after the operation are present in the

uninjured specimen, one would expect the reaction to be of
quite a different character

from what actually occurs.

stimulus applied near the posterior end would naturally cross

over at once and produce a bending on the opposite side at

the same level.

Or the stimulus might

entire op])osite side Avould bo affected

diffuse, so that the

and the worm would

But as a matter of

become uniformly curved on that side.

fact we find that the turning affects only the anterior portion
of the body.

If it is

urged that after operation the crossing

of impulses takes place through the general protoplasm the

difficulties encountered are no less, for it must be shown how
passage of an impulse through the protoplasm to cause a

perfectly well co-ordinated reaction can appear so quickly

produce such perfect results at once.

If tested

and

immediately

after the operation, before the general lowering of tonus

felt,

is

the reaction time for the negative response of a posterior

piece of the body will not differ appreciably from that of a

normal worm.

Now, according

to the

views of the advocates

of the theory that after operations involving loss of nervous

tissue,

impulses

protoplasm,

it is

may be

conducted through the general

is always at first

held that such conduction

It would
must be true.

appreciably slower than in nervous tissue.

seem on purely a priori grounds that

Thus

it is

this

also

seen that there are serious objections to the view

that the negative reaction

is

the result of a contraction on the

body opposite to that stimulated that is, that it is

a crossed reflex. The question now arises, if the reaction is
not produced in this way, in what other way can it be
side of the

produced

Evidently

it

is

quite possible that the anterior

away from the stimulus by a

lengthening of the side stimulated, quite as well as by a

part of the body can be turned

RAYMOND PEAWL

608

sliorteuing or contraction of the opposite side.

We

may now
away

consider the evidence as to whether or not the turning

is

actually due to a lengthening of the side stimulated.

Very

little

observation

evidence can be obtained regarding this from

the normal moving animal, because the

of

in the turning would be the same

whether it were due to a shortening of one side or a
lengthening of the other. The results from certain sorts of
operation, however, give definite evidence on the question.
A specimen split longitudinally in the posterior end, as
shown in Fig. 17, a, and the cut was extended forward to the
Several days were
posterior border of the head region.

general appearance

Fig. 17.

B.

Operation diagram,

h.

For further explanation see

Showing

text.

side

supported on
is omitted

(Tiie pliarynx

for the salve of clearness.)

allowed for recovery from the shock of the operation, care

being taken to prevent the two parts from growing together
again.

By

this time the cut

edges had healed well, and the

specimen was in good condition for experimentation. The

results of mechanical stimulation were as follows: strong
stimulation of the head or anterior part of the body on either
side caused the negative reaction; the anterior end turned

But it was possible to tell in this

stimulus.
or halves of the body were
pieces
of
the
two
which
case
turning.
It could be seen clearly
the
producing
effective in
that the half stimulated, immediately on stimulation, flattened

away from the

MOVEMENTS,
out

slig-litly

ETC., OF

FRESH-WATER PLANARTANS.

ventrally, thus bringing the ventral cilia in close

contact with the bottom, as

is

At the same time

working.

GOO

necessary for their effective

it

lengthened along

its

outer

thus forcing the anterior end around towards the side

opposite from the stimulus.
That the '' side opposite " had
side,

nothing to do with the turning could be observed

in

many

cases directly, for this side (B) would remain in an almost

entirely relaxed condition after the stimulus

was given, and

not get any effective hold on the bottom so that

affect the

movement.

It

was further

possible

it

by a

could
little

be
by it, as shown in Fig. 17, h. If
with such conditions the worm was stimulated rather strongly
on the A side of the head, it gave a strong negative reaction,
the poiiit about which the turn was made being as far back
as X.
Evidently with part B up on the dorsal surface of A,
and consequently having no hold on the bottom, it could
have no effect in the reaction. The reaction must have been
due to the side A alone. The same thing could be shown by
manipulation to get the piece

laid

over on

so as to

practically entirely supported

very gently lifting on a needle the side

in

so that

it svas

contact with the bottom, and then stimulating A,

not

when

again the negative reaction occurred. This experiment I

have repeated with variations many times, but always with
the same result, showing that the side stimulated is the
effective one in producing the turning.
It

may be mentioned

here that the effect of strongly

stimulating the posterior end of either of the two pieces of a

specimen

slit in

this

way was

to cause a local contraction of

the piece stimulated, and a crawling

portion of the body in front of the

movement
slit.

of the short

This crawling was

not very effective, since so small a portion took part in it,

but it is of interest to note that what crawling appeared

involved only the uncut part of the body.

It being established that the side stimulated produces the
turning, the question may be raised, how, supposing in these
longitudinally split individuals that this side does produce

the reaction,

is

it

known

that

it

does this by lengthening

EAYMOND PEAEL.

GIO
along
its

its

outer margin rather than by actively contracting on

inner cut margin

may be answered by

This question

operative experiments of a different character.

If the side

stimulated, acting independently, produces the reaction by

lengthening on

its

own outer

side,

then an isolated longitu-

dinal half of the body ought to be able to give only one

reaction wherever stimulated, or, in other words, it ought
always to turn towards the same side. Furthermore, such a
piece ought always to turn towards the cut edge, since onl}'

on the side opposite to

the necessary

this has

muscles

circular

it

for

margin possessing

extension

(vide

sup.,

On the other hand, if the contrai'y view is

pp. 556, 557).
correct, that the turning away is due to contraction of the
longitudinal muscles on the side opposite that stimulated,

Fig.

18. Showing

the appearance of a longitudinal half of a

phinarian wlien at rest.

then such an isolated longitudinal half of the body ought to

be able to turn either way, according

to the localisation of

the stimulus, since there are longitudinal muscle-fibres along

the cut edge as well as along the other.

We

may determine

two views is correct.

impossible to get any clear evidence

from experiments which

of these

Unfortunately, it is
on this point from entirely separated longitudinal halves of
When a planarian is split in two lengthwise
tiie Avorm.

each of the pieces immediately becomes strongly contracted

longitudinally on the cut side, the apparent purpose of this
reaction being to reduce the exposed surface at once to a

After this strong contraction has taken place,

giving the piece the form shown in Fig. 18, no further
pi'ogressive movement can take place, and the general tonus

minimum.

MOVEMENTS,

ETC., OF FUl'^Sn-WATER

PLANARIANS.

Oil

becomes immediately very much lowered. In view of these

results
it is impossible to get any very trustworthy
from the stimulation of such a piece.
There is another operation, however, which, while it does
not isolate completely two longitudinal halves of the body,
facts

does

yet

separate

reacting parts,
splitting-

ot'

the

into

namely,

worm

longitudinal halves

the

essential

head regions. This is the

middle line for a short distance

the

in the

back from the anterior end, as shown

operation the two anterior pieces

in Fig, 12.

move about

After this

violently

and

independently for a time, taking all the various positions

shown in Fig-, ]{). The animal soon recovers from the immc-

FiG. 19.

Diagram sliowing

tlie

different positions taken by the

components resulting from longitudinal

two

splitting of the liead.

diate effects of the opei-ation, glides about in a normal way,

only at a rather slow rate, and responds well to stimuli.

The

much
than when the

anterior piece keeps comparatively straight, there being

less
split

tendency

to contraction

on the cut side

extends the whole length of the body.

The reactions

of

such a specimen to mechanical stimuli are as follows. To

stimuli applied at the posterior end along the sides of the

body the reactions are precisely the same as those already

normal individual. Stimulation in the
regions a a (Fig. 20) of moderate or strong intensity produces
the negative reaction. The organism turns away from the
described for the

RAYMOND PEARL.

612

side stimulated quite as

promptly and

does a normal specimen.

If

21^) are stimulated in the

used for

in the

same way as
and B (Fig.

the cut edges

same way

the specimen will

this)

stimulus.

now

may best be
always turn towards the
(a

needle

This can best be brought out by describing a

which a series of fifty stimulations in the

and B were made on a favourable individual cut in

typical case in

regions

way. In thirty-nine of the reactions the animal turned

towards the stimulated side. That is, if the stimulus was
this

applied at
a

while

if

the animal turned in the direction of the arrow

B was the

direction of the arrow

trials

stimulated edge the reaction was in the

In eight of the remaining eleven

b.

the reaction was indifferent.

Fig. 20.

Operation diagram.

stimulation and then started

The animal stopped

at

See text.

moving straight ahead again,

stimulus evidently having been ineffective so far as

the

special reaction
fifty

is

concerned.

In only three cases out of

did the specimen turn away from the stimulus.

required the greatest

Since

manipulation to give the

stimulus to one cut edge without touching the other side,

it

cai-e

in

especially in view of the fact that the animal

was moving

all

seems very probable that in these three cases a

stimulus was accidentally given to the side wliich it was not
intended to stimulate.
The same general result of turning
the time,

'

After

shown

it

tills

operation the two parts of the head usually take the position

in this figin-e after the first

tion have ceased.

spasmodic movements following the opera-

MOVEMENTS,

FRESH-WATER PLANARIANP.

ErO., OF

613

towards the stimulus when applied to the cut edge was

obtained in several other series with this same specimen, and

many
will
if

times with other specimens similarly mutilated.

be seen that this

the turning

lated.

away

is

is

It

the result which would be expected

due

to

lengthening of the side stimu-

Stimulation of either side of the cut portions, inner

or outer, causes turning in the

same

direction,

and that

Diagram

to ineclianical stimuli and

case of a specimen in which tlie head
has been split longitudinally. For further explanation see text.

Fig. 21.

to

show Uie reactions

tlieir niecliaiiisms in tlie

is the one in which turniug would be caused provided each piece did actively lengthen on its outer side.
There seems to be no reason whatever, if the turning away

direction

were due to contraction of the side opposite that stimulated,

why the specimen should not turn away from stimuli applied
This it does not do. There seems
to the cut inner edges.
to

be no escape, then, from the conclusion that the turning

EAYMOND PEARL.

614

uway from the stimulus (negative

reaction)

due

is

to

lengtliening of the side stimuhated.

It

may

possibly be objected to the last experiment that

the impulse from a stimulation

at,

for example,

(Fig. 21)

took the path indicated by the dotted line in that figure^ and

caused a contraction on the left side of the body, so that

really the observed turning was the result of a contraction on

To

the side opposite that stimnlated.

this objection

it

may

be answered that by stimulating different points along the

edge B it is possible to cause the point about which the
turn occurs as a pivot to be located anywhere along the line x y.
It is very evident that contraction of muscles in the region N
can have nothing whatever to do with turning of the right
So this objection is without force.
piece about the point x.

As the

process of regeneration of a cut longitudinal half of

from the
made, and it is consequently possible to obtain specimens in which the regeneration of the missing half of the body has produced only a very
small amount of new tissue, and which are at the same time
nearly straight in outline and able to make progressive movements. The reactions of such partially regenerated specimens are of importance as throwing light on the normal
the body goes on^ the piece will

curved form

mechanism

of

straighten out

takes after the cut

it

the

specimen of this

reactions.

soT't

is

The

may be described

reactions
in detail.

of

a typical

On October

10th, 1901, a small piece of the anterior end of a specimen of

The piece was cut as nearly as

shown in Fig. 22, a. On October IGth
A narrow strip
the piece had the form shown iu Fig. 22, h.
side,
and the forinaright
of new tissue had formed down the
tion of the outline of the head and of the right eye was just
beginning. At this time the reactions of the specimen were
Stimuli applied at y caused the head to turn
as follows.
sharply away from the stimulus (typical negative reaction).
This reaction was quite like that given by a normal individual
Stimulation at x, however,
stimulated in the same way.
produced no trace whatever of a negative reaction. On
P.

maculata was

isolated.

possible in the form

MOVEMENTS,

ETC., OF

FRESH-WATER PLANARIANS.

615

stimu];ition at this point the speciinon contracted longitiuli-

and then staited moving ahead again in exactly the

same direction in whicli it was going before stimulation. It
was impossible to indncc any turning away following stimulanally,

tion of the side

Now

is

it

,r,

this

specimen

times.

comes very near

being an isolated longitudinal half-planarian.

to

structures of the original one half are

is

many

althongh this was tried

evident that

only a very

the line of

new

little

new

of the other side of

tissue,

down

All the

and there
the body produced in
present,

the originally cut edge.

In this

and the
muscle layers are not well formed. It Avas brought out above
(p. GIO) that an isolated half of the body ought to be able to
tissue there is probably very little differentiation,

Fig. 22.
in

o.

Operation diagram,

Diagram

a.

The new

b.

tissue

Piece wliich regenerated from

is indicated by stippling.

give only one reaction, or, in other words, ought to be able to

turn the body in only one direction in response to stimulation,

provided this turning

side.

men

We

is

due

to

an extension of the stimulated

find precisely this result in the regenerating speci-

just discussed.

It

turns away from stimuli applied at y

all the muscles necessary for

because on that side are present

extension just as in a normal animal.

from stimulation of the side x because

It
it

does not turn away

has not the necessary

On the view that the

on the side opposite that
stimulated, there is no reason why stimulation at .r should not
cause the animal to turn away from the stimulus, because the
opposite side ((/) has all its muscular mechanisms intact.
muscles for extension on that side.

turning away

is

due

to contraction

RAYMOND PEARL.

616

The reason why the specimen in this last expei'iment does

when stimulated on the side x,

not turn towards the stimulus

is

apparently because the regeneration has proceeded only far

to produce just enough new tissue to form the

beginning of a new side to the body. This new side receives
the stimulus and is sufficiently potent to determine the reaction of the whole (the straight longitudinal contraction),
but is lacking in the mechanism necessai-y to produce its own
proper reaction, the negative reaction.
On the other hand,

enough

in the case of the individual with the split anterior end,

each

piece turns towards the stimulus after stimulation of the cut

edge because here only one half the organism is present

either to be stimulated or to react; there is not even the
beginning of the formation of a new side along the cut edge.
Putting all the evidence together, I think it must be regarded as demonstrated that the turning away from the
stimulus in the negative reaction to mechanical stimuli is due
to an extension of the side of the body stimulated.
This
extension is brought about by the contraction of the circular
and dorso -ventral muscle-fibres probably also assisted by
the transverse and oblique systems of fibres in the region
stimulated.
This reaction is a simple reflex act involving

only the side stimulated.

response

is

concerned,

identical, but

halves.

is

The normal organism, so far

composed

to be considered as

in a certain

as this
of

two

sense independent longitudinal

Thus, representing these halves diagrammatically,

as in Pig. 23, a, the evidence presented indicates that stimulation of one side of the

worm,

as

A, causes a reaction

in that

side, and, so far as essential features of the directive reactions

go, only in that side.

The movements

of half

after its

stimulation determine and, in fact, cause the reaction of the

Furthermore, these longitudinal halves retain

their individuality as halves if they are isolated from each
other.
A separated half-worm (longitudinal) reacts as a
half-worm, just as it did when in connection with the other
half in the body, and not, as might perhaps be expected on
a priori grounds, as a whole worm. It reacts as a whole

whole animal.

MOVEMENTS,

worm

ET(!.,

OF FIlESH-WATEIi PLANAKIANS.

only ufier a uuw half

cut edge.

Tlie

various

lias

been regenerated along

its

change from the

those as a whole worm can be

stages

reactions as a half-worm to

617

in

the

The new tissue

followed step by step as regeneration proceeds.

formed along the cut edge very quickly takes on some of the
Wiien only a narrow

functions of a side.

formed

it

has been

strip

serves for the reception of the stimulus, and hence

stops the reaction of the opposite side, as in the experiment

last discussed.

may be made

to

To make the meaning more clear, reference

diagrams h and c of Fig. 2o. In h is repre-

sented, in a straightened position, the half

of a

normal worm

A
Fig. 23.
of

Diagrams
Planaria

tliereto.

to
to

show the relations of the halves of the body

the reception of st.imuli, ant] the reactions

See account

in text.

(Tlie

pharynx

is

omitted for

the sake of clearness.)

immediately after being separated from the other

c

half,

while

represents the same half after regeneration has begun and

a strip of new tissue has been formed

Now

of the piece to turn

own proper

the cut edge.

towards the stimulus, i. c. to give its

(cf. experiment given above on

negative reaction

slitting anterior end).

side

down

stimulation of the cut edge of h causes the anterior end

that

is

This

is

because in this case

stimulated, although along

Stimulation along the right-liand edge of

its

it

is

inner edge.

does not cause

the turning towards the stimulus, because in order that this

RAYMOND PKARL.

618

may

take place

would be necessary

it

proper negative reaction.

its

It

for the side

to give

cannot do this because

it

is

not directly stimulated, but the new very small side A is

stimulated.
This side may not have the necessary muscles to
give

negative

described above,

reaction

3'et

may

itself

as

receive

the

in

the

experiment

stimulus

and

so

from reacting. Another way of expressing this same fact is by saying that in regenerating
longitudinal halves of planarians the physiological middle
line remains at the line of the former cut edge for some time
after regeneration has begun. ^
In connection Avith this
indirectly prevent

discussion of the reactions of half-animals

it is

greatly to be

that Willey ('97) did not get any data on the

regretted

remarkable form Heteroplana. In this

form we have a natural " half-planarian," or very nearly that.
reactions of the

One

side

seems

It

is

so greatly atrophied as to be practically absent.

me very probable that this organism would

in much the same way that a longitudinally

to

to stimuli

specimen of

Planaria, which had begun

to

react
split

regenerate,

does.
I

do not wish

it

to

be understood from the analysis of the

negative reaction which has been given that I intend to

maintain that in this reaction the side opposite that stimulated never contracts longitudinally.
this, especially in cases of

It

probably often does

very strong stimulation which cause

a general excitation and reaction of the whole body.

I have
merely wished to show that the fundamental basis of the

negative reaction

is

the extension of the side stimulated.

seems to me quite possible that

may be shown by

it

It

close

analysis in other cases that supposedly crossed reflexes are

not fundamentally such at

We

may now

the

of
'

all.

pass to a brief consideration of the mechanism

positive

1 liavc rc'coids in

reaction

my

of

the

planarian

notes of cxpcrinientb

to

mechanical

wliicli bliow tliat in llie

case of

oblique cuts the physiological middle line remains at the cut edge until after
the

new head

is

well formed in (he

fcpace forbids detailed

new

tissue on tlie obli(|ue edge.

desciipUon of these expeiiinenls here.

Lack of

MOVEMENTS,
stimuli.

ETC., OF

FRESH-WATER PLANARIANS.

As has been sLowu above^ removal

of

tlio

G19

anterior

end of the body containing the brain causes the disappearance

and

probably due rather

to the lowering of tonus than to the removal of any special
centre having the causation of this reaction as its function.
Additional evidence on this view that lowering of the tonus
of this positive reaction,

is

this result

is

the chief cause of the disappearance of the reaction

is

found

in the fact that other injuries to the head, such as longitudinal

splitting,

which produce a lowering of the general tonus,

also

cause the disappearance of the positive reaction.

This very close dependence of the reaction on the general

tonic conditions of the organism

makes

its

analysis difficult,

seems most probable that its mechanism is as follows

a light stimulus, when the organism is in a certain definite
tonic condition, sets off a reaction involving (1) an e(|ual
bilateral contraction of the circular musculature, producing
the extension of the body
(2) a contraction of the longitudinal musculature of the side stimulated, })roducing the
but

it

turning towards the stimulus (this the definitive part of the

reaction);

and

(3)

contraction

of

the

dorsal

longitudinal

musculature, producing the raising of the anterior end.

this reaction the sides

In

do not act independently, but there

is

and finely co-ordinated reaction of the

organism as a whole, depending for its existence on an
entirely normal physiological condition.
It is to be noted,
a delicately balanced

however, that the definitive part of the reaction, namely, the

turning, is a response of the side of the body stimulated.
This point

is

one of fundamental importance for the general

theory of the reactions.

The mechanism

of

the

other

stimuli are evidently very simple.

which must be considered as the

reactions

to

mechanical

The crawling movement,

specific reaction to

mechanical

stimulation of the posterior region of the body,

is due to
rhythmical contraction of the longitudinal musculature. The
only other reactions to mechanical stimulation are local con-

tractions,
e.

whose mechanism is evident.

Uehaviour
in the General

Features

of

the

RAYMOND PEAKL.

620

Organism

which

Stimuli explain.

Reactions to Mechanical
much of the behaviour of pla-

the

That

narians in their natural surroundings

actions above described

Among

them.

is

the result of the re-

is

very evident to any one watching

specific features of this sort in

reactions play a part

may be mentioned

the

which these
escape from

enemies or harmful surroundings, the getting of food

(to

be

discussed in detail later), the localities chosen for coming to

behaviour on meeting solid obstacles in the path of

All
movement^ the passing on to the surface film, etc.
of these need not be discussed specifically, as their relations
will be evident enough on a moment's thought, but the last
two deserve special mention.
The behaviour of planarians on meeting solid bodies in
their path in the course of movement is entirely made up of

rest, the

reactions to mechanical stimuli.

as follows

The behaviour

detail

in

is

If a gliding specimen meets squarely head-on an

obstruction of considerable size, so that

it

cannot glide over

without changing to some extent the position of

its long
an instant^ raise the head, let it drop down
till it touches the obstruction again, and then glide directly
up on to and over the solid body. This behaviour is invariable, so far as my observations go, if the worm meets the
It is at once seen to be merely a
obstruction squarely.
special case of the usual reaction to a weak mechanical
it

axis,

it

will stop

stimulus, characterised

behaviour

is

by the raising

of

the

head.

The

evidently purposeful in the long run, because

it

organism up on to food material just as well as

If the gliding worm meets the obstrucindifferent bodies.
tion obliquely the behaviour depends in large part on the
physical nature of the object. If it is food material, or something else of a rather soft and yielding texture as, for
example, another planarian, the worm will immediately raise
the head, turn it towards the object, and crawl up over it.
will take the

This behaviour

is

evidently the typical positive reaction to a

weak mechanical stimulus.

case of this positive

which

i-eaction,

special
1

and rather curious

have twice observed,

MOVI'IMENTS,

OF FIfKSIT-WATKT?

KIT'.,

C21

I'LAN AKIANS.

appeared wlien two ?-periin(Mis o-liditiQ* alono-, wifli the nnterior

ends slightly r:iised in tlu^ iioi'inal inaiiiicv, met liead-on.
Both were siinnltaiieonsly stimulated to the positive reaction
and raised the anterior ends, and then let them di'op ai>-ain.
As they came down the two ventral surfaces were l)roni;ht
squarely together in the way shown in Fig. 2t; then each
started gliding

movement

up the ventral surface of the other.

changing form

ns a result of the constantly

body, the ventral surfaces slipjiod

the two

worms went on

off

their way.

fi*om

When

In a
of the

one another and

the obstruction

a hard body, as a piece of glass, the specimen meeting

obliquely usually turns the head away slightly at the

is
it

first

contact (negative reaction), and then glides along parallel to

the edge of the body for a distance.
it

again with the side of the head,

If
it

negative reaction and turns away again.

Fig. 2i:

Side view

of

two

it

happens

to touch

frequently gives the

After the solid body

plaiiariaiis starting: to glide

up on Uie

ventral surfaces of cacli other.

has been touched several times, however, the positive reaction

is usually given, and the worm passes at once up on to the
solid body. This behaviour is shown in Fig. 2h. The precise
form of the behaviour on meeting obliquely a solid body in
the path varies considerably with the general physiological

condition of the individual.

first

touch

In case

it is

much

excited, the

induce a strong negative reaction, and tlie

turn away and pass out of the neighbourhood.

will

individual will

In the cases whei'e the final positive reaction is preceded by

two or three negative ones, it would seem as if repetition of
what must be an almost identical stimulus causes it to l)ecome in effect weaker. Leaving aside all variations in the
exact character of the behaviour on meeting a solid, the
important point to be brought out is that all this behaviour is
based on the simple reactions to mechanical stimuli. The
VOL. 46, PART 4.

NEW

SERIES.

S S

622

I^AYMONII TKARL.

different factors.

any o-jven case depends on several

These are the position of the animal with

reference

obstruction,

exact behaviour in

to

the

the physicnl

and the physiological

obstruction^

nature of the

condition,

whether

of

greater or less excitation.

So, again, with reference to the habit of the animal of

moving about on the surface film, a problem is presented.

When a specimen, gliding up the side of a dish, touches its
anterior end to the surface film at the point where the latter
joins the glass,

it

immediately gives a characteristic positive

any other weak

reaction, precisely like that in response to

mechanical stimulus.

The head

is

raised and turned towards

the side from Avhich the stimulus came_, and then dropped

and fi are successive stafifes in the reactions of

PI an aria on meeting obliquely an obstacle in its path. Tlie hcav)^

Fig. 25.

1, 2, 3, 4, 5,

straight line represents the obstacle.

As a consequence of this reaction, the head end

comes to rest on the under side of the surface film at a point
The
some little distance out from the side of the dish.
ventral surface of the anterior end of the body flattens out on
the surface film, and the animal glides out on to the film,
following the direction determined by the reaction of the anterior end. Thus it is seen that the going on to the surface film is

again.

only a special case of a response to a

i.

e.

weak mechanical

stimulus,

the positive reaction, the film itself acting as the stimulant.

The leaving
the dish

is

of the surface film

and passing down the side of

evidently also due to the same positive reaction.

MOVEMENTS,

ETC., OF

FHESH-WATKR TLANAHIANS.

623

There ai-e a nninbcr of other points in the general behaviour

which are directly rehited to the reactions to nieclianical
stimuli, wliich \Yill be taken up later in connection with the
other reactions,

Summary. Before

/.

next subject,

it

may be

on to a discussion of the
summarise briefly the chief
the effect of mechanical stimuli on

passino-

well

findings with reference to

to

planarians.

has been shown that the planarian responds

It

nigli

manner

perfect

meclianical stimuli.

the

to

localisation

and

in a well-

intensity'

of

turns awa}- from strong stimuli (in

Tt

the long run harmful) applied to the side of the body turns
towards weak stimuli (in the long run beneficial, almost never
;

harmful)
to the

it

crawls rapidly away from strong stimuli applied

backs and turns away from similar

posterior end;

strong stinmli applied at the anterior end.

It has been shown, further, that these reactions have
the characteristics of reflex actions, complex,
still

it

is

all

true, but

reflexes.

The mechanisms
are unilateral, and
Discussion

mechanical

of

of the reactions to unilateral stimulation

the side stimulated.

lie in

the

implications

these

of

stimulation, with reference to the

results

on

psychology of

the organism and the general theories regarding the reactions

of oro-anisms to stimuli,

is

deferred

till

the results from other

sorts of stimuli are in hand.

II.

Reactions

to

Food and Chemical Stimuli.

Evidently one of the most important factors in the sum

any aquatic organism is its reactions

total of the activities of

to

chemical

substances.

Its

ability

to

receive

chemical

prime importance in its

habitat such an
natural
in
its
for
struggle for existence,
encountering
constantly
aquatic organism must be almost
Some of these may be harmdifferent chemical substances.
stimuli and react

ful

and some

to

them must be

beneficial,

and

it

of

would seem that

if

a species

is

RAYMOND

624

PEAIJL.

must have some sort of reaction

whereby they may avoid the harmful and take advautnge of

to survive, its individuals

the beneficial.

In the case of planarians, the reactions to

chemicals seem to be of about equal importance with the reactions to contact stimuli in the general activities.

Since the

reactions to food substances are a special case of the reactions

to chemicals in general, they
a.

Food Reactions

may be

The

discussed

first.

nature of the things used as

food by fresh-water planarians has been discussed already in

the section on

'^

Natural Histor}-," and hence need not detain

us here.

typical case of the food reactions to a bit of crushed

Fig. 2G.
Diaj^ram sliowiiif^ the successive stages in tlie normal food
reaction of PI an aria.
A represents a small bit of meat.

mollusc

may

account.^

first

be described, to serve as a basis for the

body of Physa which has just

If a piece of the

been extracted from the

of a pair of forceps

is

shell

and crushed between the points

placed in a small dish containing a

number of active planarians, it will result from chance alone

that some of the flat-worms must in course of time pass near
For a very short time

the food material.

^

The food reactions

(:01,a).

of

Planaria have been

after the food has

briefly

described by Bardeen

MOVEMENTS,

625

ETC., OF EIJESlI-WATEl; I'LANAEIAMS.

been placed iu the dish specimeus may pass very near it

within two or three luilhmetres without being affected in
any way. They simply glide straight by as if there were no
food there.
After a few minutes have passed, however, it
will be found that a worm coming near the food is affected
Its behaviour is as
in
a very characteristic manner.
follows:
When within about three or four millimetres of
the piece of meat (Fig. -0, o) it stops abruptl}', raises the
head, and turns it towards the food (Fig. 2G, h). As the head
is raised and turned the gliding is resumed, and the head
being almost immediately lowered, the movement is directly
towards the food. Thus fur the reaction is evidently
precisely like the positive reaction to weak mechanical

stimuli,

and so we may speak

the stimulus differs.

Tig.

27.

Diagrammalic
*'

is

(Fig. 26, c).

view

in

siiie

gripping "

it

such

ol'

view of
a bit of

as

apparently

to

show

tlie

A.

to

Fig. 27) are as follows

piece

Planaria
fooil,

flattens

down over the very edge

whole

as the positive reaction to

it

same

down upon it, and, if the connnike it possible, " grips " it
The details of this " gripping " (shown in side

touches the food

figuration

of

in the two cases, though

^Vhen the anterior end of the head

food, the reaction being the

provided

squeezes

it

The

anterior

end

closes

of the piece of food, or over the

it

is

small

by contraction

enough,
of

and

then

the longitudinal

muscles on the ventral surface of the head, 'i'lie action is

very characteristic, and evidently forms an integral part of
Its probable function will be
the normal food reaction.

brought out Inter. While it is taking place the worm as a

whole stops its progressive movement and remains quiet.
After the " gripping" has continued for some time the worm
It passes forward
starts gliding ahead up on to the food.
the opening for the extrusion of the
till the point where
pharynx is located is approximately over the place pre-

KAYMOND PEARL.

626

viously "g-i'ipped" (Fig. 26,(1).

Then the

truded and feeding begins (Fig. 26,

worm

voluntarily

After

e).

is

ex-

time

tlie

pliai'ynx
ti

leaves the food and glides off over the

bottom.

Having described the typical case of a food reaction,, we

take up some of the more important variations from the

may

and describe the various phases

tyjoe,

the reaction in

in

greater detail.

Starting with the very beginning of the reaction, it may

be said that the distance from the food at which any effect
on the planarian is produced varies greatly, as is to be
expected.
This distance, of course, depends on the extent
which the juices or chemicals of the food have diffused from
it.
When a piece of meat is first put into the water

specimens will pass very close to it without being stimulated.

In fact, if a specimen finds a piece of food within three or
four minutes after it is put into the dish, it will usually have
done so as a result of accidentally coming in contact with it.

As has been brought out above, when

gliding

worm

touches anything of a rather jaeldiug texture, like food,

it

iuimediately gives the positive reaction and passes up over

it.

This plays an important

])art

in the getting of food,

because, as I have found in experiments, unless the food

crushed and pressed with forceps the juices

slowly,

and

for

some time

specimens

will

is

diffuse rather

not

give the

positive reaction unless they actually touch the food.

On

the other hand, after the food has been in the water for

some time,

so that diffusion has taken place, the distance at

may be affected becomes quite considerable.

have seen specimens gliding by a small piece of meat at
a distance of Ig cm. from it give the positive reaction and
turn towards it. At greater distances than this food is not
Avhich specimens
I

effective,

according to

my

observations.

The distance from

which a given specimen will give the positive

reaction and go towards it depends also on the })hysiological
Specimens in a state of general
condition of the individual.
observed, go closely by
frequently
have
I
excitation will, as
food

at

move:men'1',s, etc.,

of fkesh-wa'ier planaiiians.

627

a jDJece of food

without tiiruiug towards it, while other

specimens in a more normal condition will give the positive

reaction

some distance from

After the

first

it.

specimen has begun feeding on a piece of

material the zone of influence of that piece becomes almost

immediately widened appreciably.

ing specimens increases the area

which

As

the

number

of feed-

the surrounding water

in

becomes corres])ondingly greater. This

many cases, as an illustration
Several pieces of crushed snail were put in a

affects others

phenomenon
will indicate.

dish with a

is

very striking in

number

of

In

])lanarians.

short

time

about the dish had come near to one

had given the positive reaction and begun
feeding.
At almost the same time another of the pieces of
food had " attracted " another specimen. The other bits of
food were quite similar in every way to these two, and lay in
the dish not far from them.
Yet at the end of fifteen
minutes the two pieces by which the first two worms had
been affected were completely covered with feeding

specimen

in gliding

of these pieces,

specimens, Avhile the remaining pieces of food, with a single

exception,' did not have a specimen on them.

This increase

a stimulus must be due

in the effectiveness of the food as

more

chemical substance from it.

due either to some secretion of
the feeding animals or to some change which they induce in
the food. It is probably due to a combination of these two
That a digestive secretion is poured out through
factors.
the pharynx of the feeding worm is well known, and clearly
shown by the appearance of a piece of food on which a
specimen has been feeding.
The surface of the meat is
turned white, and rendered very soft and almost flocculent.
the

to

diffusion

of

Apparently the increase

It is

is

probable that this digestive secretion acts as a positive

chemotactic stimulus to other worms, and that coupled with

this there
itself

is

an increased diffusion of juices from the food

caused by the changes which

The reaction which

'

One

is

piece ruiUicsl lemuved

it is

caused by
fruiii

undergoing.

this chemical stimulus

the ollicib luul

;i

single specimen on

it.

KAYMOND PEARL

628
fi'uin

the food

is

evidently esseutially the same thiug' as the

weak

positive reaction given to

sists in

uiecliauical stimuli.

It con-

a turning of the anterior end of the body towards

There is no reason for supposing

any way diiferent from that of tlie
same reaction to mechanical stimuli, and hence this need not
be further discussed here. A question of prime importance

the source of the stimulus.

that

mechanism

its

is

in

with regard to this positive reaction in response to chemical

stimuli, Avhicli

was not taken up before,

with reference to the stimulus,

how

words,

how well localised,

?

or, in

other

precisely does the anterior end point towards the

source of the stimulus,

in

this case food

clear-cut orienting response

may be

is

the reaction

is

said that

when

the

worm

is

Have we here

In answer

to this

problem

a
it

only a short distance

from the food the response is very precise. The anterior

end is brought by the first positive reaction so as to point
exactly towards the meat, and as the worm glides ahead it
never misses it. This is true where the specimen is near
enough (usually within three quarters of its own length), so
that the stimulus which reaches it is a fairly strong one.
In case the worm is stimulated near the edge of a large
diffusion area when the stimulus is very weak, the first
reaction may not suffice to direct the animal straight towards
In this case the behaviour is usually like that
the food.
shown in Fig. 28, in which the line B, B, B, represents the
effective margin of the diffusion area of the piece of food A.
(By " effective nuirgin " is meant the line outside of which no
effect
first

is

produced by the food on passing specimens.)

reaction which the

sion area (Fig. 28,

and

worm
2) is

proceeds on the new path into

towards the food.

it is

The

gives on reaching this diffu-

weak
tliis

positive one.
area, but

It

then

not directly

After a short time, however (Fig. 28,

again stimulated to a

})ositive

reaction" (4).

3),

This time

both the stimulus and the reaction are stronger than before,

and the worm

diilusion,

but

is

directed more nearly towards the centre of

still

not exactly.

When

it

food again (5) another positive reaction (0)

gets oj^posite the

is

given, and this

aJOVEMENTS, ETC., UF FUESU-WATEU I'LANAlilANiS.

time, since the stiuiulus
is

is

a rather slruug' one, the reaction

a very precise ouO; and the subsequent

movement

the animal directly io the food (Fig. 28, 7).

is

typical for this sort of stimulation, but

component phases, depeudiug- on the

stimulus

the distance from the

food

Diagram

carries

This heluiviour

may vary

in its

relative strengtli of the

when

first

stimulated.

Fig. 28.

629

showiiij; llic rcaclioiis of

Plaiiaiia

l,u

food

(A)

from which juices liavc l)i:cii diirusiiii^ into ho walcr for some time.
B, B, B, represent the ell'ectivc marj^in of the (liH'usiDii area of the
food A.
1, 2, 3, i, 5, 6, and 7 are successive positions talvcn by
the organism.
t

two or as many as four positive reactions niay be

to the food.
This shows
clearly that with reference to chemical stimuli, the precision

Thus

eithei"

necessary to bring the animal

of localisation

of the positive reaction decreases as the in-

Indeed, I have observed

nu unlocalised positive reaction, although

tensity of the stimulus iliiuinishes.

what

is L'viileutly

030

EAYI\IOND PEARL

The behaviour was as follows

larfro diffusion area had beeu formed^ and a specimen was
stimulated to a weak positive reaction at a distance of about
twice its own length from the food (Fig. 29, 1).
It passed
this

seems piiradoxical.

into the diffusion area, but did not give another positive reac-

/ B

Fig. 29. Showing the reaction of ii plauariiin to a very weak food

stimulus.
Letters as in l"ig. 28.

tion when opposite the food, but instead glided by and away
from it. When it had gone some distance in this direction
it stopped and gave a very clear and characteristic positive

reaction, so far as

the

form of the reaction indicated, but

with the turn away from instead of lowai'ds the centre of

MOVEMENTS,

There was no doubt of

cliifiisioii.

reaction

usual

ETC., OF FUESH-WATI-:!; PI-ANA l.'IANH.

llie

631
of the

elianioter

the head was raised and the body turned in the

manner

of the positive reaction, whicli

one can never

become familiar with it.

The
specimen kept on in the path determined by this last
reaction (Fig. 29, 4), and passed entirely out of the region of
the food.
Evidently in this the Avorni was stimulated very
weakly by a chemical, and the stimulus was nearly as strong
on one side of the body as on the other, and when the reflex
was set off it was on the wrong side of the body. This is
not the usual result of weak stimulation, and has been
observed in only two cases, but it serves very well to show
the decrease of the power of localisation when the stimulus
mistake

is

after once having-

very weak.

When,

as

fi'equeutly ha])})ens, the

food exactly head-on, the

worm approaches

the

reaction usually consists merely

of that portion of the reflex expressed in the raising of the

head, while the

the food.

worm keeps on

in its straight path

The head may be waved from

but the general direction of motion

till it

reaches

side to side slightly,

The

not changed.

is

action evidently corresponds to the positive reaction following

weak mechanical stimulation

of the dorsal surface of the

in the middle line, as described above.

ever, I

head

In some cases, how-

have observed very active and hungry specimens of

sp., which were going straight towards the

Dendrococlum,

and turn
away from the

food, give a complete positive reaction

and

start off in a

new

direction

to

one side

food.

This,

however, of course brought the specimen at once into a

where the stimulus was acting unilaterally, and it

again gave a positive reaction, this time heading it again
for the food, which it usually reached without further
reaction,
liut in some cases I have observed the specimen
position

give so strong a reaction

as

to

taken

be

almost directly

away from the food by the subsequent movement, and,

fail

to reach

Avere

''wild"

passing out of the area of diffusion,

Specimens behaving
general reactions.

in

this

way

it

in

at

all.

their

The responses were very vigorous, but

IJAYMOND PEARL.

632

not localised with refeieucc to

tlio

sLiiuulus with the

utjiial

precision.

of

The ''gripping" of the food substance by the anterior part

the worm is a very characteristic feature of the normal

form depends on the configuration

body "gripped." In its most typical
form, where the food material is in the form of a cylinder, or
approximately such, the action reminds one of the action of
the human hand in grasping a stick.
The tip of the head
closes over the material in the same way that the fingers do,
while the region just behind the auricles bears the same
food reaction.

Its exact

of the food or other

relation as does the proximal part of the palm, just in front

of the wrist, in grasping.

After the head has been placed

over the material

way

can be seen to

contract

rather strongly, and thus literally squeeze the food.

In case

in

this

it

the surface contour of the food does not admit of this reflex

being carried out in

its

typical form, as close an approxima-

made as possible. To compare again with the

human hand, when the surface is flat, or forms the surface of
tion to this

is

a cylinder of large radius, the ventral surface of the head

pressed closely to

it,

below the surface,

the tip attempting to dip

in,

as

it

is

were,

same way that a man "claws"

in just the

with his finger tips in attempting to

obtain

liold

on a

similarly configured body, too large for complete grasping.

While the "gripping"

is

in general a very characteristic

feature of the food reaction,

The cause

may be omitted

it

in

rather

where it
occurs, or any laws governing the matter, I have not been
able to discover.
A necessary accompaniment of the
"gripping" of the food is the cessation of the forward
exceptional cases.

movement
food

ot"

is first

omission

This pause

when

touched by the anterior end and before the

it,

the

worm

occurs in practically every case, whether

the grijipmg accompanies

of

the

the animal as a whole.

passes up on to

is,

for

it

or not.

course, considerably greater

when it is absent.
The function of the "gripping"

The length
when the

of the pause
" gi'ipping "

occurs than

of the food material before

MOVEMENTS, KTC, OF
feeding begins

regard

to

not inmicdiately apparent, but I

is

purpose

to think its

is

am

inelined

to intimately test the substance with

availability as

its

G33

I'K'KSTT-WATKi; I'LANAinAXS.

Some evidence on

food.

point and further discussion regnrding

will

it

tliis

be introduced

later.

After the preliminar}- pause nnd ^^ gripping" of the food

the worm glides up on to it to begin active feeding,
l^he
position taken

by the worm brings out a very nice correlation

In a ver}' large number of cases (certainly over

in reflexes.

75 per cent., so far as

my

observations have gone) the

worm

advances over
the pharyngeal opening is
exactly over the place where the first "gripping" occurred,
and there the pharynx is extruded iind feeding begins.
the

food

until

Diagram showing great extension of the pharynx. Tlie

stippled area represents food substance on wliirh the phmarian

Fic. 30.

is rest

When

the

in?.

worm

reaches this position the posterior

the body relaxes and takes

istic of

on

the resting specimen.

and becomes
the opening

in

it

The pharynx

attaclied very quick!}'.

extended, and

than when

the

the body-wall

it

As

it

are

of

becomes

is

thrust out,

passes out through

usuall}' considerai)ly

diameter becomes correspondingly smaller

in the pharyngeal sac.
It ma}' or may not

its
is

attach to the food directly beneath the body.

ditions

i^art

appearance chai-acter-

favourable

it

usually

does,

and

When

con-

consequently

cannot be seen on looking down on the animal from above.

On

the other hand, I have frequently seen

it

stretched out

and attached some little distance to one side of the body, as

shown in Fig, 30. The stinnilus, causing the extrusion of the

RAYMOND

634
pliarynXj

is

rFAKT,.

the contact of food or other solid body with

tlie

pharyngeal region of the ventral surface, together with an

appropriate chemical stimulus. The pharynx is not extruded
until the

animal gets up on to the food so that the opening' of

is in direct contact with it.

This can be
demonstrated by direct observation by the use of a very
small piece of food material and a plane mirror placed
beneath the glass dish in which the specimen is moving. By

the pharyngeal sac

end

lifting gently the posterior

also

be seen that the pharynx

the food.

The most striking

of the

is

body on a needle

not extruded before

can

it

it is

over

illustration of the correlation in

the reaction which brings about the extrusion of the pharynx

when

just over the food, is to be seen when a specimen

nemertean Stichosterama asensoriatum is used as
food, and the long axis of the planarian and of the nemertean
are at right angles to each other.
After first " gripping "
it is

of the

Fig. 31.

Diagrammatic

Icngiludinal sccHon of a planarian feeding

on a nemertean (shown in cross-section at

).

the nemertean the planarian glides along over

it

until

the

pharyngeal opening is just above it, and then pauses, and the
pharynx is extruded and attached ( and h, Fig. 31). These
facts strongly indicate that the effective stimulus for pharyn-

geal extrusion

is

received, at least in part, in the pharyn-

That it is necessary for both contact

and chemical stimuli to act to produce the extrusion of
the phaiynx may be shown by experiments on specimens
geal region

itself.

gliding on the surface film ventral side uppermost.

If, with
such a specimen, a chemical known to produce under other

conditions extrusion of the pharynx,

is

allowed to come in

contact with the pharyngeal region, there

is no result.
Of
experiment proper precautions
were taken not to disturb the animal by allowing the solution
to drop upon it. Another demonstration of the same fact that
a chemical stimulus alone does not suffice to cause extrusion

course in performing

this

MOVEMENTS,
of tlio

pharynx

solutions, such

V.TC, OF FltRSTT-WATRK PLANARTANS.

that specimens

is

as

sn^^ar solutions,

635

immersed in favonvahle
do not show tliis phe-

nomenon. 'IMiat mechanical stimulation alone does not suffice

is demonstrated by the fact that planarians pass over and rest
on other planarians without extruding- the phai-jnx, althoug'h
the consistency of their l)odies
that of the animals

used

used as food frequently

if

as

is

In

much
fact,

the same as

they

will

l)e

wounded

so as to afford

The stimulation

of tlie anterior

they

the proper cliemicfd stimulus.

evident!}'

food.
ai"e

end of the body by the food seems also to be necessai"y before

pharyngeal extrusion takes place. The data on this point will
be presented later in connection with operation experiments.
The appearance of the body on the food is quite characAs mentioned above, when the pharynx is extruded
teristic.
forward, movement stops, and the posterior part of the body

The anterior third of the

body, however, keeps in movement during a considerable
The head is waved
part of the time the specimen is feeding.

becomes more or

less

relaxed.

side, and touched to the food or the

bottom of the dish here and there. It keeps its characteristic extended form to a greater or less degree.
A favourite
position is for the anterior third or half of the body to lie on
the bottom and move about, while the posterior part lies up
on the food. This is the position most frequently seen in
specimens feeding on a rather small piece of meat. Wiieu
the anterior end gets on the bottom it gives every appearance, in many cases, of attempting to glide away, and being
only restrained by the attachment of the pharynx to the
In other cases, however, the anterior end remains
food.
The importance of the attempted movement will be
quiet.
brought out later. As has been mentioned above, the flatworm is able to move off and drag the food still attached to
In the fastening of
the extruded pharynx along behind it.
tiie food to the body in this case the sticky slime undoubtedly
assists the pharynx.
After the food has been softened by the digestive juices,
it is taken into the body through the pharynx.

about from side to

T^AYMOND TEARL.

636

worm

After the
time

it

will

Ims been feeding for

certnin

leno-tli

of

detach the phaiynx and spontaneously move

off

n,

from the food, the pharynx: being withdrawn again into its
The length of time after the beginning of the feeding
sac.
at which this takes place varies very greatly in different
I have observed a specimen which fed on a piece of
cases.
mollusc for as long as an hour and thirty minutes, while in
other cases the worm may stay on the food only ten minutes,
Judging from the rate at which food is taken
or even less.
np while the animal is feeding during the day, and from the
fact that pieces of meat left in the dish overnight are almost
entirely consumed by morning, it would appear that much of
the

time during the night

is

spent in

material available for the purpose

anterior end of the feeding
tiveness

stimuli,

to

worm

a feeding

feeding when any

While the

at hand.

retains its normal sensi-

nevertheless

it

stimulation to induce

is

requires

worm

considerable

to leave

the

food.

which would ordinarily set all resting

specimens into rapid movement, has little or no effect on
feeding specimens. If a worm is suddenly pulled off a piece
of meat on which it is feeding a very good view of the
extruded pharynx may usually be had, as this organ is
retracted somewhat slowly when torn from food in this way.
So far as I have been able to discover, the presence of
food in the immediate neighbourhood of a resting planarian
has no effect upon it. Apparently the stimulus afforded by
crushed meat is not sufficiently strong to produce a response
from such an individual. The following experiment copied
from my notes will show this.

Shaking

May
snail

of the dish,

14th,

1901, 3.10

was placed

resting specimen.

3.30 p.m.
4.5 p.m.

p.m.

piece of freshly crushed

mm. distant from the anterior end

No reaction or other effect produced.

Worm in same

No change.

dentally started into

of a

position as before.

(At this time the

movement and

worm was

acci-

the experiment conse-

quently ended.)

This lack of effect of food on resting specimens

may be

MOVEMENTS,
tlio

OF FRESH-WATET^ PT-ANAIHANS.

ETC.,

reason for the statement of Bnrdeen

C37

p 522) '' that

which liaJ been kept in pure rain water for a week or

worms

(loc. cit.,

two, and were thus in a hungry condition, would remain

unmoved by the presence close by their side of a piece of

fresh snail, a food much prized by them,"
1
Food Reactions of Specimens after Opera.

For the purpose of throwing" light on the general

mechanism of the food reaction, experiments were tried on

tions.

specimens cut in different ways.

It

is

unfortunately very

from practical reasons to get many certain results

different

from these experiments. Many of the results are negative,

and hence not entirely conclusive. Since, however, some
important facts have been brought out by these experiments,
they will be described.

The

which

operation

first

will

be discussed

cutting the animal in two transversely.

made

in the region in

resulting piece, after

shock

On coming
it

somewhat from the

show the following reac-

into the zone of diffusion about a piece of

gives the positive reaction just as a normal

On reaching

does, and turns towards the food.

the meat

animal

that of

has recovered

effect of the operation, will

tion.

meat

it

is

such a cut is
front of the pharynx, the anterior
If

its

it

worm

the edge of

behaviour

is again like that of the normal

" grips " the food, and then passes
At this point appears the striking difference

stops, usually

on over it.
between the behaviour of this anterior piece, which, it must
be remembered, has no pharynx, and the behaviour of the
entire

worm.

glides up over

The
it,

anterior piece after gripping

and without the

the

food

slightest change, even in

down off of it on the other side.

There is not the slightest indication of any stopping for the
pharynx to be extruded.
the rate of gliding, passes

If the transverse cut is

pharynx is included
behave with reference
It will stop

The

in

made

farther back, so that the

the anterior piece, this will then

to food quite as

a normal animal does.

on the food and extrude the pharynx.

posterior pieces resulting from transverse cuts do not

VOL. 40, PART 4.

NEW

3EKIES.

TT

RAYMOND TEAEL.

638

give any definite food reaction, so far as I have been able to

ascertain, until they have been regenerated to some considerable extent.

has been cut

Posterior pieces from which only the head

will glide

by

pieces of snail on which other

wornas are feeding, without giving the slightest reaction.^

In experiments so arranged that the gliding posterior piece

would just touch with its anterior end the edge of a piece of
This same arrangement -with a
food, it gave no reaction.
normal w^orm practically never fails to call forth the positive
Posterior
reaction and bring the worm up on to the food.
pieces placed gently on pieces of food material do not extrude
the pharynx and start feeding, but immediately glide down
from it and over the bottom of the dish. These experiments
with posterior pieces have been tried many times and under
varied conditions, in the hope that some sort of positive
This is
results might be obtained, but never with success.
After a new head
true for three days after the operation.
has been fairly well formed the animal will react to food
The behaviour of one of these posterior pieces on
again.
touching with the anterior end a piece of food is very
The cut
strikingly different from that of a normal animal.
piece, if it touches with the front or sides of the anterior end
the smallest shred of food material, or any other substance,
gives a well-marked negative reaction, and goes in a new
direction away from the obstruction.
It does not, as a rule,
crawl up over anything which it meets squarely " head-on,"
but instead turns away.
Thinking that possibly the pharynx might play a more or
loss independent part in the normal food reaction, i. e., that
it might have a set of reflexes of its own, not determined by
the rest of the body, I tried experiments with the isolated

pharynx removed entire from the body. Such an isolated

pharynx will remain alive for a considerable period, and
respond to stimulation. When first removed from the body
'

Bardeen

(:

01, a) has sliowii

that,

if

the transverse cut

is

in the region in

front of the eyes the posterior piece (comprising in this case nearly the

worm)

will react

normally to food.

whole

MOVEMENTS,
it

contracts

KTf'.,

OE FIJESTT-WATER PLANA IITANS.

in

I'lijtliinically

a longitndinnl

time, and then comes to rest at about

in the body.

its

039

direction

normal

for n

Icuotli wlien

Mechanical stimnlation causes merely

tudinal contraction, Avhile the presence of food near

it

lono'i-

has no

meat placed Avithin a

had
no effect upon it
millimetre of such an isolated pharynx
1 have tried layino- the isolated
in the course of an honr.
pharynx directly on pieces of meat to see if there would be
any tendency for the end of the orc^an to attach itself as it
normally does.
This was not done, nor was any other

effect

whatever.

Freslily crushed snail

definite reaction produced.

These operation experiments show, so far as they pfo, that

(i.e., the
(1) The presence of the pharynx in the body
take
food)
has
nothing
functional ability to
to do with determining- the reaction of the anterior end of the bod}^ to food
stimuli.

The anterior part

of the

body gives the same

re-

action to food in every case, without regard to whether so

doing actually puts the animal in a position to get food oi'

The i*eaction is only purposive under certain circumstances; when changed conditions make it no longer purposive, no adaptive change in the behaviour of the anterior end
not.

This shows clearly

occurs.

how

little

basis there

sidering the behaviour towards food as

nature of intelligent behaviour.

The stopping

(2)

circumstances

is

due

of the

worm on

for con-

is

anything

of

the

the food under normal

to the posterior half of the

body, not the

The behaviour of the anterior cut piece in gliding

directly over the food is what one might be led to expect from
the behaviour of the same part of the body under normal
As described above^ it was seen that the
circumstances.
anterior.

anterior end of the normal individual gives every appearance

of attempting to continue

part

is

feeding, and

is

moving forward while the

posterior

only prevented from doing this by the

mechanical hindrance of the attached pharynx. In a sense,

we may consider that in a large degree the work of tlie
anterior end of the body with reference to feeding

when

it

gets the animal up on to the food.

is

over

RAYMOND PEARL.

640
(3)

head.

The reception

of the food stimulus

In other words, the head

is

is

a function of

tlie

the only part of the body

capable of receiving very Aveak chemical stimuli.

(4)

far as

Decapitated specimens do not extrude the pharynx, so

my

observations go, even though the proper normal

Presumably the
we have

stimuli are given the pharyngeal region.

brain

is

the necessary organ in this connection, as

already seen that the sense organs concerned with the act of extrusion are not those of the head, but of the pharyngeal region.

Bardeen

(:

01, a, p. 178) states that

extending the pharynx and

removal of the head.

'^

the simple reflexes of

of swallowing are preserved after

found, by repeated

the headless pieces could usually be

trials,

made

to

that one of

eat

if

it

was

back on a slide in a small drop of water. Under

the conditions mentioned the pharynx is usually protruded,
and will engulf bits of food placed in the mouth." Regarding this conclusion, I can only say that in a large number of
experiments with decapitated specimens I have never been
able to induce extrusion of the pharynx, under conditions
approximating as closely as possible to the normal. I do not
placed on

its

wish to affirm that the decapitated planarian cannot extrude

the pharynx, but merely that it does not when placed in
situations which normally produce pharynx extrusion.

The pharynx is not an independent organ in its reactions,

since, when separated from the body, it does not react with
reference to the localisation of the stimulus, as it does when
(5)

normally connected with the remainder of the body.^

It is shown above
2. Summary of Food Eeactions.

that planarians have a very definite and characteristic set of

reactions to food substances which enable them to become

aware of the presence of food, and

find

it.

The importance

of these reactions in the life of the individual can hardly be

over-estimated.

While planarians,

like

many other lower

organisms, can live for a considerable time without food, yet

The question
in the long run they must, of course, have it.
^

Evidence on

this latter point will be

the reaction to chemicals.

brought forward

in

connection with

MOVEMENTS, KTC, UV FRESH-WATI'iR PLANAiaANS.

how

of

a lower orguiiiisiu gvia

its

641

food, taking' advantage of

the good and rejecting the bad, and thus apparently choosing
one thing from several, is one of the most interesting and

important in comparative psychology.

The food reaction

of planarians consists of

an extremely

which may be

set into action

well co-ordinated set of reflexes,

and second,
two sorts, first, chemical
Both sorts of stimuli are, of course, given by
the food. The first and most important of till the reflexes in
the food reaction is the turning of the head towards the
source of stimulation, followed by movement in that direction.
This is the reaction which enables the animal to find food.
Evidently it is the same thing exactly as what has been

by

stimuli

of

mechanical.

described as the positive reaction to mechanical stimuli; or,

in other

words, the positive reaction to mechanical stimuli is

Its primary

only a special case of the general food reaction.

function

which

is

evidently the getting of food, whatever the stimulus

calls it forth.

for a food response following

be found in the fact that it most

The reason

mechanical stimulation

is

frequently happens that

to

many

things

(e. g.,

whole animals)

which are available for food are not emitting chemical substances into the water in sufficient quantity to cause an
effective stimulus.

If the planarian did not

give a positive

reaction after contact with such bodies they would be missed,

and no advantage taken

of

them

By

as food.

positively to Aveak mechanical stimuli

the

animal

reacting
is

in a

position to take advantage of the presence of food of all sorts,

whether it is in condition such as to diffuse chemical subThis fact that the animals
stances through the water or not.
react to food substances as a result of mechanical stimulation
affords a possible explanation of the "gripping " phase of the

general response.

The purpose

of this " gripping "

may be

to bring the sense organs of the head, which are sensitive to

chemical stimuli, into very close contact with the substance
in order to determine whether it possesses the chemical

characteristics of food.

In other words,

"tasting" reaction, which

is

this

made necessary by

reaction

is

the fact that

RAYMOND PEAKL.

642

the organism turus toward

all

bodies of a certain physical

texture under most circumstances.

Tlie active squeezing of

the material in the "gripping" undoubtedly helps to press

out to the surface any juices which may be in the material.

In closing the section on food reactions it may be well to

give a sort of general picture of the whole behaviour of
fresh-Avater planarians towards food.

The method by which

the planarian finds material suitable for food is as follows

1. Chemical substances diffusing from food come in con:

head region

tact with the sensitive

of the planarian

or

The movino- animal touches with the head some

stance, and as a result of either of these two

soft sub-

sorts of

stimulation
2.

The organism

gives

positive

towards the source of the stimulus.

most cases, and

precisely localised in

part of the whole food behaviour.

previously described
3.

When

result of

(v. sup., p.

Its

reaction,

i.

is

turns

e.

This reaction

is

mechanism has been

619).

the anterior end squarely touches the food as a

this reaction

it

typically closes tightly over

giving what I have called the ''gripping'^ reaction.

reaction

ment

is

very

the most essential

evidently a very

for the purpose

much

specialised feeling

it,

This

move-

of closely testing the chemical nature

produced by a contraction of the ventral

While it is taking
longitudinal muscles of the head region.
place progressive motion ceases.
4. After this pause the worm glides over the piece of food
till the opening of the pharyngeal sac lies over or nearly
of material.

It is

over the place " gripped/' and there the posterior part stops
and the pharynx is extruded and attached to the food. The
factors determining the place

extruded are

(a)

where the pharynx

shall

be

the stimulation of the ventral surface of

the body in the pharyngeal region of the food (pure reflex

and [h) the presence of the brain, which probably

factor),

acts as a co-ordinating centre for this reaction.

5.

digestive Huid

and the food

is

is

poured out through the pharynx,

partly digested before being taken up.

MOVEMENTS,
6.

FRESH-WATEK PLANARIANS.

ETC., OF

The softened food

is

643

taken into the body through the

phaiynx.
7.

The animal spontaneously

stops feeding after a certain

time.

The
food

question

now

bottom

is at

arises, if the

in the

chemical stimulus, what

stance causing

it ?

normal process of getting

majority of cases a reaction to a

is

the uatui*e of the chemical sub-

Can the same response be induced by

the use of different inorganic and organic chemicals

Is

there any relation between chemical composition and the

intensity or form of the reaction

number
be had

To answer these and a

them recourse must
the nuture and concentra?

of other questions arising out of

experiments in Avhich

to

tion of the chemicals affecting the organisms

trolled.

All

the

experiments of this kind

may be
I

will

con-

group

together under the heading

h.

Keactions to Chemical ^Stimuli

1.

Reactions

to

Localised

Che mot axis.

Chemical Stimuli.

This particular phase of the general subject of the effects of

chemicals may be considered first, since it is most closely
related to what has preceded on the food reactions.
The
plan of the experiments was to try the effect of a series of
substances

when applied to restricted areas of the body. A

number of chemicals were used to include

sufficiently large

representatives from each of the main groups of substances

which have been found to have marked effects on organisms.
The method Avhich was found to give the
a. Methods.
most satisfactory results in the application of localised
chemical stimuli was the use of a capillary tube filled with
the solution whose effects it was desired to test. The form
of the tube used is shown in Fig. 32.
The tubes were 10 to
15 cm. long, and were made from glass tubing of about
2'5 mm. internal diameter. Each end was drawn to capillary
fineness, and then broken off so as to give an opening of the
desired size.
The opening at the upper end was made

644

RAYMOND PEARL.

slightly larger than that at the lower,

which was used in

The tube was filled with solution by
suction.
The rate of diffusion can be regulated by changing
the sizes of the openings, and can be determined for each
tube from the rate at which the fluid sinks at the upper end
giving the stiniulas.

of the tube.
Considerable experimenting is necessary in
order to get the best rate of diffusion for work on planarians.
Since the animal is moving rather rapidly while the stimulus
is

being applied

diffusion or the

it

necessary to have reasonably rapid

is

worm

will not react at all, or not for so long

a time after the stimulation has begun that one cannot be

certain

of

the

results.

It

is

easily

possible to get the

capillary so fine that no results can be obtained.

other hand,

when

it is

On

the

too large the solution affects too large

a portion of the body at one time, and furthermore, as will

be shown

later,

organism.

This, of course, introduces a possible source of

Fig.

may

cause a rheotactic

32. Glass tube used

serious error.

It

in

reaction

of

the

giving localised chemical stimuli.

can be avoided by frequent and proper

control experiments.
It will

be well to describe in advance the conduct of a

and the precautions taken, so that it may

not be necessary to repeat these details in the account of
each experiment. Six to ten normal active planarians were

typical experiment

taken from the aquarium dish and put in a Petri dish of

about 10 cm. diameter, in freshly drawn, filtered tap water.
Enough water was put in the dish to give a depth of about

Two or three of the capillary tubes with different

openings were filled with the tost solution.
These
tubes were all tested before a final experimental series was
begun, and usually only one which had been found to allow
1

cm.

sized

was used. In some cases,

varying degrees of sensitiveness among the
different specimens made it necessary to use for some indiffusion at the satisfactory rate

however,

MOVEMENTS,

ETC., OF

645

FRESH-WATER PEANARIANS.

dividuals capillaries of faster or slower rates than what

may

be called the standard. After preliminary experiments to

determine the relative sensitiveness of the different parts of
the body to chemicals, attention was devoted almost entirely
to stimulation of the head region, and consequently in tlie
experiments which

will

be reported

first

the stimulus was

The
head, unless otherwise stated.
of
])oint
the
place
to
was
stimulus
applying
the
method of
the
from
mm.)
(about
2
the capillary tube a short distance

applied only to

the

place on the body to be stimulated.

The animal was stimu-

it was gliding along in the normal way, and hence

was necessary to move the capillary tube at the same rate
the animal moved in order to keep it opposite the same point

lated as
it

in case the reaction

was put into

was not given

at the instant the capillary

place, which, of course, almost never happens.

move the tube along

as the

With

worm

glides so as to keep the relative position of the two

little

practice one can

almost identically the same.

Just as soon as a reaction had

been obtained with a given specimen the capillary tube was

removed from the water, so as to permit as little as possible
of the chemical to get into the water surrounding the
organism. After a series with any substance, the worms
were transferred at once to a dish of fresh water before
beginning another series. Further, in any long series, when
for any reason it might be supposed that the water was
contaminated with the chemical to an extent
worms were transferred to
another dish of fresh water. All through the course of an
experiment frequent control tests were made by trying the
effect on the worms of the water surrounding thein when
diffusing out from the same tube used previously for the
After each experiment the tubes were thoroughly
chemical.
rinsed by drawing distilled water back and forth through

becoming

sufficient to affect the results, the

them many times. The tubes were

and new ones substituted.

also frequently discarded

RAYMOND PEARL.

646

The following substances were used

in the

experiments

/-Nitric

Mineral acids

Hydrochloric

ISu Iphuric
Oxalic

Organic acids

Citric

Formic

fSodium hydrate
(Sodium carbonate

Alkalies

Salts of heavy metals

fCopper sulphate
(Zinc sulphate

Other

Sodium chloride
Sodium bromide
salts

Potassium chloride

Masfuesium chloride
Cane-sugar.
Distilled water.

Since distilled water was found to have a decided effect in

producing a reaction^ the solutions were prepared iu both
In case of any
distilled water and in filtered tap water.
doubt; as with very dilute solutions, the effects of the solutions prepared in each sort of water were tested and

compared.
Since only qualitative results were desired, and for the
practical reason of greater convenience; perceutage rather

than molecular solutions were used.

The results are, in a way, so remarkable that
/3. Results.

they will be presented in some detail.

Mineral Acids.
Nitric

(sp.

gr.

1*42),

strong negative reaction.

This

solution causes

If applied to the

head region the

i per cent.

animal turns away from the side stimulated immediately; and

If the stimulus is long continued the animal
strongly.
writhes and twists about violently.

Stimulation of the posterior region causes the part where

MOVEMENTS,

OF FRESn-AVATEIi PLANA RIANS.

ETC.,

647

the solution strikes to contract very violently, and the whole

crawling ahead

animal to start
is

very injurious, and

the individual.
essentially as

its

if

It will

This concentration

rapidly.

action

continued^ quickly kills

is

be noted that

its effects

are the same

those of strong mechanical stimuli applied to

the same parts of the body.

y*y

per cent, and

per cent.

^^^

per cent.

The animal

is

Kesnlts

the same as in 4

and extensively

not as quickly

injured by these solutions as by the former.

noted

that

with these comparatively strong

It is to

solutions

be
the

reaction time after stimulation of the posterior end of the

body

so slow that this part of the

is

body

is

permanently

injured or destroyed before the animal gets away.

per cent.

-^^

In

some cases a well-marked positive reby stimulation of the head region with
The head would turn towards the mouth

action was caused

this

solution.

of the pipette in the characteristic fashion of the food reaction,

weak mechanical stimuli. In other inwas weakly negative, while still

In cases where there was
other specimens were indifferent.
or the reaction to

dividuals the reaction given

an indifferent reaction there was a

side of the head stimulated.
g*g-

per cent.

Clearly

marked

majority of cases after the

time.

local contraction of the

positive

stimulus

reaction

in large

has acted for some

This solution never caused the negative reaction.

Some

individuals were, in a few cases, indifferent to this solution.

This solution

too

is

weak

to start a

resting specimen into

movement.
Y^y-

per cunt, and weaker.

Indifferent

reactions or

weak

positive.

This acid appears to be a very strong stimulus for the

negative reaction in concentrations down to j,, P<-'i" cent.,
while below that

it

is

a rather ineffective stimulus, and the

when induced is positive.

Hydrochloric, j\j per cent. Strong negative

reaction

There

is

individuals

nuticuablc
to

turn

in

some cases a tendency

very

slightly

reaction.

for

some

towards the source of

RAYMOND PKARL.

648
stimulation

before

giving'

tlie

strong

negative reaction.

Stimulation of the anterior end of a decapitated specimen

caused a slow negative reaction with long reaction time.

This solution causes the change from the glide to the crawl

when

applied to the posterior end of a normal worm.

yV per cent. Negative reaction; rather weaker than with

preceding solution. With this solution one specimen would

came
and then

turn towards the source of the stimulus until the head

into the strong acid near the

mouth

of the pipette,

give the sharp negative reaction.

Specimen A gave positive reaction in every

specimen B in about 50 per cent, of all cases, while
iu the remainder of trials gave weak negative.
Other
specimens negative reaction.
^L per cent. Specimen A as in preceding case. Specimen
:f

J per cent.

case

gave positive reaction in about 90 per cent, of

Other specimens weakly negative reactions.
Y^jj

per cent.

"grip"

All

it

to the

end of the capillary and

with the anterior end as in the food

moment they

After holding on for a

let

when continued.

reaction.

go and give a sharp

negative reaction, indicating that the stimulus

strong

trials.

specimens give well-marked positive

They glide up

reaction.

all

is

still

too

This behaviour will indicate the

machine-like character of the positive reaction.

^Q
action.

per cent.

In

Remainder

To give an idea

the majority of cases

indifferent

re-

positive.

of the

dependence

of the reactions to

chemicals on the physiological condition of

following series of experiments with

per cent, and weaker concentrations

HCl
may be

tlic

organism, the

in solutions of

described.

y-^^^

It is

understood that these experiments were carried out

on different animals from those just given.
Specimens
per cent. No sharp positive reaction.
y-g^ij
Avill give a weak negative reaction if the opening of the
capillary is held very near the head.
In most cases reactions
to be

are indifferent.
^TU" P^^* cent.

One

specimen gives positive reaction and

MOVEMENTS,
jrroos

whole food rcnction on

tlirono'li

The remainder
cTU

P^^"

still

cent.

649

ETC., OF FRESIT-WATEK PLANARTANS.

end of

tlio

tube.

give weak negative reactions.

Reactions

same as

essentially the

in

^r

per cent.

At this point this series was discontinued. It shows that

any absolnte concentration for a chemical solution which will
cause all planarians to give the positive reaction cannot be

How

assigned.

a given individual will

react

concentration of chemical depends almost,

if

to

given

not quite as

it does upon the solution.

Sulphuric, j\j- per cent, and -^ per cent. Caused immediate and violent reaction.
Decapitated worm reacts like

much, on the individual as

This

normal.
jij

per cent.

of cases.

is

evidently a very strong stimulus.

Caused strong negative

One specimen reacted

tube being held some

first

reaction in majority

as follows

distance

the

capillary

away from the head,

gave a well-marked positive

reaction.

it

On coming

the mouth of the tube it

began strong convulsive contractions (evidently on account

into the stronger solution near

stimulation).

of too sti'ong

same spot, and after

and swept it about
mained this way for
removed immediately

Ifc

remained, however, at the

a few minutes extruded the

pharynx

The specimen reThe tube was, of course,

the first positive reaction was

over the bottom.

some

time.

after

decapitated specimen in one case gave

a very
distinct positive reaction to this solution, the tube being
held some distance away from the specimen.
giy per cent.
Negative reaction. Decapitated specimen
given.

gave

reaction

positive

the posterior

end

once.

Tliis

solution,

applied

of the body, induces the crawling

to

move-

ment.
Too"

P^'^'

cent.

Negative reaction.

tracts into a ball

when stimulated with

Positive

Isolated pharynx conthis solution.

Remainder
pharynx as in ji^ per cent.
With
^^j; per cent., y'^t^tt pcr cent., and ttttto pci' cent.
these solutions the reactions were for the most part negative.
j^jj

per cent.

negative.

Same

reaction in one case.

result with

EAYMOND PEARL.

650

In a few cases positive responses were produced^ but not

regu]ixv]y.
psi'

otVtt

cent.

Positive

reaction

in

all

cases.

The

whole food response was produced in case the end of the

tube was left in position. The worms '^gripped" it, glided

up on

to

it,

and extruded the pharynx,

in

many

the latter up into the lumen of the tube.

cases running"

Anterior piece,

resulting from cutting animal in two transversel}^, acts like

worm (positive reaction), but less strongly. Decapitated

worm gave no response. In order to make sure that in this

whole

was the extremely diluted acid which was producing

the result, numerous controls with distilled water and culture
water and fresh tap water were tried on the same specimens, in alternation with trials with the acid. With tap
water and culture water the specimens were indifferent
but with the acid solution (-g-yVo P^" cent.) mixed in either
tap water or distilled water they gave a well-marked positive
This showed clearly that the results were due to
reaction.
case

it

the acid.

Summary. With the three mineral acids tested it was

found that to concentrations above a certain point the specimens always gave the negative reaction, while to concentraThe
tions below this point the positive reaction was given.
absolute value of this " critical point " varies widely with
different

The

individuals.

same as that

behaviour

essentially

is

the

in response to mechanical stimulation, viz. to

strong stimuli the negative reaction

is

given, to

weak the

positive.

Organic Acids.
Oxalic, i per cent, and yL per cent.
This solution affords a very

reaction.

and quickly

kills

the

specimen.

The

Sharp

negative

strong

stimulus

negative reaction

very violent when once induced, but several speciuiens

were killed before they turned away. There was noticeis

able

a slight

instant

it

tendenc}^ to turn towards the

was perceived, and before

this could

stimulus the

be replaced by

MOVEMKNTS,
tlie

ETC., OF

FRESH-WATKIJ PLANAT^IANS.

negative reaction the specimens were

651

nearly or quite

killed.

per cent.

-^^

Convulsive

by sharp

negative reaction in

In one case stimulation was

majority of cases.

followed

by extrusion

positive reaction, succeeded

great

tlie

of the

pharynx.
trials

give positive

reaction,

A few specimens on some

and then go into convulsive

per cent, and J^ per cent.

-^jy

movements as they get into stronger solution.

Remainder negative.
^i- per cent. Positive and weak negaT^TT P^^' cent, and

twisting

about equally divided.

tive reactions
UTir P^^'

becoming
actions

are very

full

and

y^rg-Tv

cent.

P^i'

weak when given

In the cases where there

solutions.

the

cent,

Positive

proportionately more numerous.

response

is

Negative

response

the mouth of the tube, but do not grip

re-

these

to

a positive reaction

given; the specimens go

not

is

in

reactions

it

up to

nor extrude the

pharynx.

Y5Vn P^^' cent. With this solution all but one specimen
give the positive reaction. Specimens will follow the end of
the pipette about the dish

by a

if it is

moved

Specimens

series of positive reactions.

This

slowly.

is

done

will give the

complete food reaction on the end of the tube.

Citric, 2 per cent.
1

per cent.

Less

to positive in
Y*^

some

Strong negative reactions.

marked negative

reactions.

Tendency

cases.

Positive reactions
Indifferent.

per cent.

in nearly all cases.

Re-

mainder indifferent.
YX)

per cent.

Citric acid in

weak

solutions seems to be a ver}^ ineffective

pronounced reactions of any kind.

and yV V^^ cent. Prompt and de-

sort of stimulus, not causing

Formic, i per

cent,

Causes a resting worm to give

a weak negative reaction of the anterior end, but does not
start the whole animal into movement, provided the tube is

cided negative reaction.

withdrawn

after the first reaction

is

obtained.

EAYMOND PEARL.

652

Negative reaction, but decidedly less proTTjy per cent.

Does not
nounced than with preceding concentrations.
cause any movement whatever in resting specimen.
per cent. Negative reaction, less strong than in preNoticeable
vious cases. In some cases positive reaction.
tendency to give slight positive reaction just before the

definite negative response.

g\p

per cent.

Well-marked positive response.

Summary. The same

conclusions are to be

drawn from

the experiments on organic acids as from those on mineral

acids, viz. that to strong concentrations of a given substance
is

given, while

cause a positive response.

Oxalic acid

the
that

negative reaction

it

appears to furnish in

all

weak concentrations
is

rather peculiar in

concentrations a stimulus of

the proper quality to induce the positive response, but

the

same time excessively harmful

in

is

at

any above the weakest

solutions.

Al kalies.

Sodium Hydrate,
cent.

Immediate

i per

cent.,

j^jy

strong negative

per cent,,and ^V

response.

P^i"

Specimens

per cent, the reaction

turn away very sharply. In
slightly weaker than in the other two.

is

J^ per cent. Negative reaction. Stimulus applied to posterior end of body is sufficiently strong to cause crawling
movement.

Weaker negative reaction.

resting animal into movement.
Weak negative reaction. Ineffective

yL per cent.
strong to start
jjrjj-

Sufficiently

per cent.

resting

worm and on

end

posterior

of

body

of

on

moving

specimen.

In one
-^^ per cent. A^ery weak negative response.
specimen sharp positive reaction performs whole food reaction on the end of the tube.
Positive reactions from all specimens. The
-^^-^ per cent.
;

complete food reaction

is

given.

MOVEMENTS, KTC, OP FRESH-WATIOR PLANA RTANS.

To

653

solutions below this concentration the organisms

are

either indifferent or, in a few cases, weakly positive.

Sodium Carbonate,

Rather weak negative

per cent.

-i-

reaction,
y\)

Majority
Well-marked

per cent.

Remainder

of all reactions positive.

weakly negative.
Ya per cent.

The specimens can be

positive reaction in

led around the dish

all

by moving

cases.

tube

tlio

slowly.

Below

concentration

this

different or

Summary.

A caustic

alkaline reaction

in

alkali

(NaoCOg)

(NaOH) and

produce

in-

a salt of strong

essentially

the

same

weak, positive.

Salts of

Heavy

Metals.

Copper Sulphate,
trials the

were either

In strong solutions they cause negative

results as the acids.

reactions

the reactions

weakly positive.

animals

the stimulus

all

per cent.
At the very first
-^jj
turned sharply and immediately towards
reaction),

(positive

but the solution was so

strong as to throw the animal into convulsions, when the

head came very near the mouth of the tube. Subsequent
trials

produced the negative response,

All specimens give positive reaction.
-jjy per cent.
is brought up to the tube, and the worm glides up

and
The head

over the latter.

With
marked

all

concentrations of

CuSO^

there

is

a very well-

local' contraction of that part of the bod}^

which

is

stimulated.

In another series of experiments with this same substance,

the strongest concentration which would produce the positive

reaction in

all

cases

Zinc Sulphate,

was ^^ per cent.

and jjy per cent.
-g-

Sharp

and imme-

diate negative reaction,

jLy

in

per cent. Negative reaction, but less pronounced than

former cases.
and -Jy per cent. Specimens stop when stimulated,
-^-^y

VOL. 40, PART 4.

NKW

SElllES.

U U

RAYMOND

654

PEAR];.

and wave the anterior end about in the water, first

away from and then towards the source of stimulation.
As the head comes nearer to the end of the tube, where the
solution is strongest, it is more strongly stimulated, and
gives a definite negative reaction.

weaker zone again

once more.

If the

it

is

tube

As

gets out into the

it

stimulated to a positive
is

now removed

reaction

the specimen

will,

some cases, after a short time turn sharply towards the

In other
place where it was, and move in that direction.
in

cases the negative reaction finally predominates.

not in-

It

frequently happens that in the earlier part of this reaction the

only moves very

away from the stimulus,

anterior end

slightly

slightly

so that the

be fixed in one position.

first sight, to

towards,

The

or

very

body seems,

at

planarian, in this

strenuous reaction, probably comes as near to the hypothecated behaviour of the famous "Buridan's ass" as anything
is

ever likely to in actual practice.

y-iy-

per cent.

One specimen

gave clearly marked positive

Others as in the preceding solutions

reaction in every case.

(tV P^* cent,
TT^jy

and

per cent.

-^-^j

per cent.).

Well-marked positive

reaction.

Specimens

give complete typical food reaction.

In one case, with a small worm, I was able to produce

craAvling in a

backward

end
per cent. ZnSO^.

of the anterior
-yL

Summary.

The

heavy metals are

in

in

direction by continuous stimulation

the middle line of the body with

results

from solutions

of

salts of

two

accord with those obtained with other

chemicals.

Other

Sodium Chloride, i

Salts.

per cent, and yV

tive reaction; distinct, but not

P^'^'

as strongly

cent.

Nega-

marked

as the

negative reaction to strong acids.

^\y

per cent.

Weak negative reactions

reactions in about equal numbers.

no response whatever.

Many

and weak positive

of the tiials produce

MOVEMENTS,
-^^

per cent.

No

ETC., OF

Weak

EKESII-WATRR PLANARIANS.

positive

negative reactions.

reactions

The

in

655

nearly every

complete food
reflex I have not been able to induce with sodium chh^ride.
Concentrations below this do not produce any definite recase.

typical,

action.

In general, NaCl

is

a very ineffective stimulus

to

narians, either to the positive or the negative reaction.

tilled

water

is

pla-

Dis-

a considerably stronger stimulus to the positive

reaction.

Sodium Bromide,
negative reaction in

per

cent.

Weak

but

distinct

all cases.

per cent.
Well-marked positive reaction in nil cases.
I"
Complete normal food reaction is produced.
Potassium Chloride, 2 per cent. The animals usually
react in a pecnliar way to this and stronger solutions of KCl.
When stimulated they stop, turn the anterior end either
slightly towards or slightly awny from the source of stimulation, and then stay in the same place and squirm and twist
the body.
In some cases there is a well-marked negative

reaction.

i per cent. Some specimens give negative reactions

the fii'st few trials
afterwards give definite positive
responses, ns do other specimens in all cases.
In one case
the specimen gave marked positive reaction, and after the
head was turned towai'ds the stimulus, remained quiet in the
same position as long as the chemical acted.
per cent. All specimens give positive reaction or
y*y
are indifferent.
The whole food reaction took place on the
end of the tube.
In this experiment it could be clearly
demonstrated that the })harynx is positively chemotactic to
this substance.
It is probably positively chemotactic to all
substances which induce the preceding portions of the feeding reaction. If, after the pharynx had been extruded, the
tube was turned about so that the ventral surface of the
animal could be seen, and the posterior part of the body was
moved with a needle, so as to change the position of the
pharynx with reference to the mouth of the tube, it could be

in

RAYMOND PEAEL.

656

seen that tins organ bent directly towards the mouth of the
The pharynx oriented itself with reference to the
capillary.
issuing chemical.

The

cases in which specimens were "indifferent" to this

solution (i.e. did not give either the positive or negative reaction) were evidently not due to the fact that the animal

was not stimulated^ hul, on the contrary, that it was stimuThis

lated about equally to negative and positive responses.
"
when
indifferent."
behaviour
restless
by
their
was indicated
While the animal as a Avhole moves in a straight line, the
head constantly moves slightly towards and away from the
Evidently the solution is not quite strong enough
stimulus.
to induce a definite negative reaction, nor quite weak enough
to cause a clear positive response.
^jj-

per cent.,

per cent., and

j\j-

-^j^

per cent.

Distinct

posi-

tive reaction in all cases.

-jjrjj

per cent.

indifferent.

Positive

The "

reactions

indifference "

in

some

cases,

now due

is

to

mainly

lack

of

stimulation.

Below

yiy- per

cent. I

have been unable

to get definite

responses of any sort with KOI.

Magnesium
negative reaction.

Chloride, i per
In some cases a

cent.

Usually

sharp

slight turn towards the

stimulus preceded the negative response, nnd in some few

other trials the aniaial was indifferent.

-i-

per cent.

Weaker

clear positive reaction.

negative

No

reaction.

In

one case

local contraction of the region

stimulated is caused by this chemical.

Jy per cent. Positive reaction in

all

cases.

Complete

food reaction could be induced.

_i_-

per cent.

Weak positive reaction

or indifferent.

NaCl, NaBr, KCl, and MgCU

Summary. To the
the planarians react as to other chemicals, by giving the
negative response to strong concentrations and the positive
to weak.
Cane-sugar, Sugar solutions, in all concentrations
above j^ per cent., so far as I have been able to discover.
salts

MOVEMENT^;, ETC., OF

057

PLANAUlANt!.

l-'liliSlI-WATEU

cause well-marked positive reactions in all cases.

This is
the ODly chemical which I have found that causes only one of
the reactions.

Distilled AV'ator. To distilled water applied by the

method the organisms give a well-marked positive
reaction in all cases.
That the reactions to very dilute solutions of chemicals were not due to the distilled water in cases
where this was used as the solvent, rather than to the chemical
itself, was proven in the following way
Parallel experiments
were performed, using tap water as a solvent, and in every
cnpillary

case the same reaction was given to the tap-water solution as

to that in distilled water.

would not react

the same tube.
2.

to clear tap

General Summary.

effects of localised

At the same time the specimens

water applied in the same way by

Putting

all

the results on the

chemical stimuli together,

we

are forced to

the somewhat remarkable conclusion that practically all substances are both '' attractive '^ and " repellent " to planarians.
Evidently, then, the chemical composition of a substance
not of the

importance in determining

first

shall react to

it

how

but, on the contrary, its concentration

important matter.

To weak solutions

of

is

the individuals
is

the

any chemical the

animals give positive responses, while to strong solutions they

give negative.

Between the behaviour towards chemical stimuli and

towards mechanical stimuli there is a very close parallelism,
or, perhaps better, identity, which is evidently something of
fundamental importance.
clearly

it

may

In

order to bring this out more

be well to arrange in tabular form the results

of the study of the reactions to these

two

stimuli.

658

RAYMOND PEAKL.

MOVEMIiNTS, ETC., OF FRESH-WATKIi PLANA inAN.S.

effects of ions

and the conclusions based on very

tative results with chemicals.

Two

659

fine (jii;inti-

such series of experi-

ments as those quoted above from HCl and CuSO^ indicate

what would be the worth of the assig'nment of an absolute
value for the concentration of either of these two substances
which would produce the positive reaction in planarians.
Such instances might be multiplied, and they serve to bring
out the fact, apparently so frequently lost sight of, that what
an organism will do when stimulated is quite as much a
function of the physiological condition of the organism itself
at the time as

it is

of the stimulus.

comparison of these results with those of Yerkes

on the reactions of
author finds that

G on ion emus

of

is

though

there

characteristic food reaction,

which

is
is

much

(:

interest.

02)

This

well-marked and

given in response to

food substances, whether in solid or liquid form, yet this

by other chemicals. It is stated

were tried in all concentrations
for the special purpose of detei-mining whether the food
reaction might not depend upon intensity rather than quality
This was not found to be the case. We must,
of stimulus.

reaction cannot be induced

that a

number

of chemicals

then, conclude that

its

Gonionemus

psychic development than

is

is

a stage farther along in

the flat-worm, for the

medusa

reacts with reference to the quality as well as to the in-

and location of the stimulus, while with the fhit-worm

the intensity and location of the stimulus are by fai- the most
tensity

important factors. It is necessary in the case of the flatworm, to be sure, that there be mechanical and chemical
stimuli acting together in order to produce the complex of
reflexes

forming the complete food reaction, thus indicating

some relation to quality of stimulus. But for the production

of what is, in one sense, the most important phase of the reaction, the turning towards the source of stimulation,

quality of the stimulus

With an

is

the

not significant.

understanding of the method of reaction to

localised chemical stimuli, a

problems present themselves.

number
While

of interesting special
it

will not

be possible

RAYMOND PEARL.

6G0
to take

up

all of

important ones

tliem in this paper, a few of the specially

may be

considered.

One such important general question which

problem

arises

Do

of orientation to diffusing chemicals.

orient themselves along radial lines of diffusion

is

the

planarians

and proceed

towards the centre of diffusion ? It would seem that in the

case of such a perfectly bilaterally symmetrical organism as
PI an aria, if anywhere, Loeb's theory of orientation ought to
This theory accounts for orientation by supposing that when an organism is stimulated unilaterally its

hold good.

motor organs are caused

to act either

more strongly

or

more

weakly, as the case may be, on that side than on the other.
This results in bringing the long axis of the body parallel

with the lines of action of the stimulus; and then, since

symmetrical points on either side of the body must be equally

stimulated, the organism moves in a straight line towards or

away from the stimulus.

most stimuli

Jennings has shown

this theory of orientation

(:

01) that for

does not hold in the

case of the Infusoria.

From

the

account

of

the

reactions

chemical stimuli which has been given,

seen that there

is

in this

case,

to

of
it

planarians to

will

be at once

some degree

at least,

With weak chemical stimuli the

an orienting reaction.
head turns towards the stimulus in such a way as to point
the anterior end very directly towards the source of stimulaIt might be thought that this marked a pure orientation.
tion, but it must be remembered that the organisms turn the
head just as precisely towards the point from which a weak
mechanical stimulus comes.
exactly

the

same

thing.

The two reactions are evidently

However, a single mechanical

stimulus can hardly be considered a directive stimulus of

the sort which induces an orientation, such as, for example,

the electric current.

The orientation

to the constant current

is

of unicellular

organisms

the purest type of an orienting

response, however, and the most characteristic thing about

is

it

that the organism, after having the anterior end turned

towards one of the poles, keeps the long axis of the

MOVEMENTS,

body

ETC.,

G61

FKESlI-WATEll I'LANAKIANS.

01-'

action of the stimu-

pai'iiUol to the lines of

This movement of the animal in a constant rehition to

a constantly acting stimulus is^ as I understand it, the fundamental criterion of an orientation according to the theory
lus.

above mentioned. Now if we lind, as has been shown above

to be the case, that the organism gives precisely the same
reaction to a chemical unilaterally applied as
single

weak mechanical stimulus

we can

doubtful whether

it

docs to a

similarly applied,

consider that there

is

seems

it

such an

orientation in the case of the chemical, even though the

is

directed very precisely towards

On

it.

head

the contrary,

it

seems apparent that we are dealing here with a well coordinated motor reflex only such as, for example, the reflex

of a frog's hind leg,

which brings

its

foot very exactly to the

point stimulated on the side of the body.

crucial test of this point

may be obtained by submitting

the animals to the action of some chemical to which they are

known

when

to give the positive reaction

only arranging the experiment so that

Under

large area.

positive orientation,

these conditions,
it

ought

it is

applied locally,

is

diffusing over a

it

if

the oi'ganism shows

move along the lines of diffuTo test this matter

the form shown in Fig. 33, I.

to

sion straight up to the source of diffusion.

I constructed a trough

On

a plate of glass

of

A was

fastened the trough B, which was

cut from a block of paraffin.

trough were 50

mm. x

50

The

mm. x

internal dimensions of this

5

mm. Only

the sides were

of paraffin, the glass plate serving as the bottom.

hollow

was cut in one end of the trough, and a glass tube D, about
4 cm. long, was fastened into it in an upright position. Then
from the point x on the inside of the trough a fine needle was
thrust through the paraffin

previously cut
the device
use

is

in

the wall.

shown

till it

came out

in Fig. 33, II.

the apparatus the trough

into the hollow

sectional view of this part of

was

When
filled

it

was desired

to

with filtered tap

water and a number of planarians placed in it. Then into

the tube D was introduced a certain amount of the solution
whose effects were to be tested. By varying the amount of

RAYMOND

062

PEAUIi.

the solution introduced, the rate of

diffusion

its

into the Wciter could be very nicely controlled.

was

thoroughly

tested,

and

apparatus

the

through x

This matter
in

sense

calibrated by the use of coloured solutions before the actual

experiments were begun.

considerable

number

of experiments

were

diffusion trough, with the following results

tried with this

In no case was

A typical
there any observable orientation of the organisms.
experiment will illustrate what actually took place.

per cent, solution of NaoCO., which

method always produces a sharp

Fig.

by the

capillary

positive reaction, was put

DifTusion trough used for testing tlie reactions of planarians

A, A. Glass base plate. B, B, B. Paraffin
C. Cavity of
trough,
Point of opening of diffusion tube.
x.
trough in which the specimens are placed. D. Tube in whicii the
II. Enlarged sectional view of the
solution to be tested is placed.
end of the trough bearing the diffusion tube. Lettering as in I.
1)3.

I.

to diffusing cliemicals.

into the tube

in

sufficient

quantity to give a diffusion of

had been diffusing for some time (by

test with coloured solutions long enough to reach the middle
of the trough) specimens were introduced at the end C. They
Some
started gliding about in random directions at once.
remained
nearer
passed diagonally up to the end D others
while still others went up on the paraffin sides to
the end
the end D. None 'went straight towards .p after they had
come into the region where the chemical had diffused. No
moderate

rate.

After

it

reaction of any sort was given in the course of the passage

MOVEiM LINTS, KTC, OF Flib]SH-WATEli I'LANAKIANS.

towards the end

in

(i.

e.

this

weak

in

positive reaction

x) at some point in
amount that it did not

turn slightly towards

was so small

lu some few

the majority of cases.

instances an individual would give a

turn the animal directly towards

x.

663

its

course, but

in

most cases

Further, the direction

of movement was frequently changed considerably, and turned

away from x after this weak positive response. In other
words, the animals moved about in the trough practically at

random, giving only slight reactions

the area of diffusion.

end

Many

in a

few cases while

in

of the individuals, after reaching

of the trough, turned

around and went back

to

the

other end again, just as they would have done provided no

chemical had been present.

Other specimens would glide

Only these

across the trough on the paraffin of the end D.

specimens showed any definite response to the chemical.

AVhen they came within the length of their own bodies from
the opening x they gave a well-marked positive reaction and
went to X.
Having arrived there, they explored and
"gripped" the edge of the hole with the head, and then
extruded the pharynx. The pharynx was usually stretched
up into the diffusion opening, and the worm proceeded to
feed for a time on NaoCOo.
These experiments were repeated many times with a
variety of chemicals of various concentrations, and diffusing at
various rates.
It was very certain in all cases that there was
no definite orientation along lines of diffusing ions. When
the organism by chance came near the diffusion opening x, it
would give a positive reaction if the solution was of the proper
concentration, and then proceed to give the complete food
reaction over the hole, bat there was no continued orientation.
There Avas a similar absence of a negative orienting
response when strong solutions of acids were used. In this
case the animals stayed at end C of the trough, but this was
because when, in the course of their random movements, they
struck the diffusing chemical where it was of sufficient concentration, they gave the usual negative reaction, turning the
anterior ends about 30"^ away, and starting off on the courses

RAYMOND PEARL.

G6^
SO defiued.

again

the}'

It"

they came in contact with the strong solution

In no case did they turn

repeated the reaction.

squarely around with their heads directly

away from x and

the long axis parallel to the lines of diffusion.

would be unprofitable to further multiply accounts of

all led to
the same result.
No
definite orientation occurred, but only the positive and negawith random movements.
tive motor reflexes coupled
Whether, as some maintain, we have in these positive and
negative reactions the " Diuge an sich " of orientations is a
It

these experiments, since

question for the metaphysician rather than the physiologist

The objective reality of the matter is that in the

behaviour of planariaus towards chemicals there is no orientation in the lines of diffusing ions, i. e. no phenomenon like
to decide.

the orientation of

Paramecium

Another problem

to the electric current.

importance in connection with the reactions of the organisms to chemicals has to do with the
formation

formed

of

collections

of

individuals.

in certain chemicals, as is the case

Infusoria as described by Jennings

forth,

Paramecia

will

As

Are

collections

with certain of the

this

author has set

form dense aggregations

in

drops of

weak acids, introduced into

The method by which this is done is as
the culture water.
Individuals swimming about at random strike the
follows:
of
acid
by chance and pass into it without giving
drop
any reaction; when, however, they come to the opposite
side of the drop, and start to pass from it to the water again,
they are stimulated and give their characteristic motor reThis
action (jerk back and turn towards the aboral side).
reaction turns them back into the drop, which forms, as it

various chemicals, particularly

were, a trap for

aggregation

is

all

that enter

formed.

known

This

it.

In a short time a dense

is

almost the only method of

from orientation, which will

produce collections of organisms in chemicals. Its essential
feature is not the getting of the organisms into the chemical,
active reaction,

this

them

aside

being purely a matter of chance, but the holding of

in the chemical after they have entered it, by what

MOYKMENTS,
amounts

to a

ETC.,

OF FRESTI-WAl'KR rLANAlUAXR.

665

negative reaction to the suvronnding water.

is, can we get any such formation

of
by the retention of those specimens wliicli have
entered an area by cliance in the case of Planaria ?
This problem was attacked in a nnmber of different ways,
but the clearest results could be obtained by the " two-drop "
method of Massart. Two drops of fluid of e(|ual siz(^ are
placed near each other on a slide, and a narrow connecting
band is made between the two by drawing some of the fluid
One of them was usually of culture
across with a needle.
water, while the other was of the solution to be tested.
Now
evidently, if the animals form collections b}' the " motor
reflex " method, they ought to pass into the drop of solution
without any reaction, but when they attempt to pass back
into the water drop they should be stimulated to a negative
reaction and thus turned back.
An experiment with a solution to which the animal gives a
sharp positive reaction may first be reported. One of the
drops was tap water, and the other was 1 per cent, sugar
solution, to which the specimens gave a strong positive retSeveral small planarians were put into the water
action.
They glided rapidly about this drop, and soon one
drop.
came up to the bridge connecting the water with the sugar.
It was headed sti*aight for the sugar drop, and passed over
into it without any reaction whatever.
Up to this point the
behaviour is like that of the Infusoria towards the acid drop.
This specimen circled about in the sugar drop, and after a
time became directed towards the connection between the
sugar and water, and passed back into the water drop without giving the faintest trace of a reaction of any sort.
All
the specimens passed back and forth betw^een the two drops
without giving any reaction, except in some cases a weak
positive one.
The conditions under which a positive reaction
is given are that a specimen should come more or less transversely across one end of the connecting bridge, as shown in

Tlie question, then,

collections

Fig. 34.

It

then usually gives a weak positive reaction and

It may do this on

turns slightly towards the other drop.

EAYMOND PEARL.

666

passing either from the water to the sugar or vice versa.

When

in

sugar solution

it

gives a positive reaction to tap

method or

as just

from this experiment, that

collec-

water, whether applied by the capillary tube

described.

It is evident,

tions are not

are by

formed by planarians

Infusoria.

in the

same way that they

The animals are not negative

to

the

To
was repeated

surrounding water after they have been in the solution.

test

and

verify this conclusiou the experiment

with solutions of diiferent substances.

case of

all

It

was found that

in

substances in concentrations to which the animals

gave a positive response when stimulated by the capillary

method, the specimens would pass back and forth from water
If solutions were
to solution and vice versTi, indifferently.
reaction
concentrations
to
which
a
negative
was given
in
used

Fig. 34.

Diat^ram showing the arrangement of " two-drop " experiment with chemicals.

when stimulation was by the

capillary method, the specimens

merely stayed in the water drop. When they came to the

boundary line of the strong solution they gave the negative
This immediately
reaction, and hence stayed in the water.
raises

the question,

why would

collection of the planarians

there not be a permanent

formed

in a

drop

of^

a substance

which they give the positive reaction, provided they were

First put in a drop of some substance to which they were
strongly negative ? There is evidently no theoretical reason
why this should not take place, but there is an important
This is that any solution which would cause a
practical one.

to

negative reaction, under these circumstances, will, so far as I

have found, also seriously modify the animals' movements, if

MOVEMENTS,

lOTO.,

667

OF ERE,S1I-WAT1<:R PLANA RIANS.

They will simply squirm about autl

movements, and hence not get into
the drop of substance to wliich they are positive. But it is
quite possible that by making a long enough scries of experiments on this point, one might get a solution just strong
enough to cause a negative reaction, and in which the
organisms would still move well.
We would then get a
The important thing, howcollection in the positive drop.
ever, is that to the water in which they live tlie animals do
not, under any circumstances, give a negative reaction, and
hence under normal conditions no collections can be found by
a "motor reflex" method.
It may be well, before leaving this subject, to point out the
fundamental physiological difference betAveen the Infusoria
and the planariaus, on which the difference in the behaviour
towards chemicals is based. It is that in the case of the
Infusoria there is but one form of reaction (the "motor reflex"
turn towards a structurally defined side) regardless of whether
the stimulus is strong or weak, while in the case of the
they are immersed

make no

in

it.

progressive

planarian there

is

stimuli from that

a qualitatively different reaction to strong

which

is

stimulated at
it

it

is

it

is

the in-

apparently not

what reason we do not know).

all (for

attempts to pass from acid to water

which must be
it

When

given to weak.

fusorian passes into the drop of acid

When

given a stimulus

weak one, yet

and is, as a
With the planarian any slight

in the nature of things a rather

responds with the only reaction

consequence, kept in the acid.

it

has,

in environmental conditions gives a weak stimulus,

and the specimen turns towards the source of stimulation.
This serves, together with random movements, to get it into
the drop of solution but when it strikes again the water,
which again must furnish a weak stimulus, it gives the same
The ability
positive reaction and passes out into the water.
to differentiate in the reactions between the strong and weak

change

stimuli

gives

the

organism a

far

greater

range

in

its

activities.

Another problem which

is

of ititerest in connection with

RAYMOND PEARL.

668
food and

clieiiiical

reactions

is

the relation of the condition

of tlie organism as regards liuiiger to

its

reactions to stimuli.

might be supposed that an individual which had not had food

for some time would be more apt to give the positive reaction
to a given stimulus than one which had just fed.
To test this point parallel experiments were instituted with
It

specimeus allowed to feed till they left the food spontaneously

about three hours before the experiments, and specimens
which had been kept for three weeks in a dish of clear water.
NaBi- was used as the stimulating solution, and Avas applied
by the capillary method. The specimens chosen were of the
same species, P. dorotocephala, and as nearly as possible
of the same size. The only difference which could be detected
between the fed and the unfed animals in their behaviour
towards a f per cent, solution of NaBr was that the unfed
animals gave the whole food reaction on the end of the
capillary tube, while the recently fed specimens only went so
far as to give the positive reaction, and touched the end of
the tube with the anterior end of tlie head. They did not
"grip" it and pass up on to it, as did the others. In the
main point at which I was working, namely, the giving of
the definite positive reaction, there was no discoverable
One set
difference between the fed and unfed specimens.
reaction
just
promptly
and
the
as
decidedly
as
did the
gave
Next a weaker solution, y\y per cent., was tried.
other.
With this solution about 50 per cent, of the specimens in
ordinary condition give a v/eak positive reaction, and 50 per
cent, are indifferent. This concentration, being about on the
border line betAveen that which affords no stimulus at all and
that Avhicli

is

a definite stimulus for the positive reaction,

ought to bring out any differences which may exist between

fed and unfed individuals in the sensitivity to stimuli for the
As a matter of fact, no difference in the
positive reaction.
behaviour of the two sets was to be observed. One gave a
well-marked positive reaction in as many cases as did the
In some instances the reaction time of the fed
other.
specimens seemed to be slightly greater than that of the

MOVEMKNTS, KTC, OK FRESH-WATKR PLANARIANS.

669

unfed, but this was neither

marked nor of general occurThis experiment was afterwards repeated with other
specimens, and with sugar as the stimuhiSj with essentially

rence.

the same results,

have also repeatedly tried stimulating

specimens which had just ceased
feeding, and in these cases found no certain difference
between their behaviour and that of specimens whicli had
not been fed for some time, with regard to the giving of the
positive reaction. It would appear, then, that so far as tlie
with various

solutions

giving of the positive response to

amount

weak

stimuli

of food the animal has previously

consequence.

The

failure of fully fed

is

had

concerned, the

of very little
specimens to give the
is

feeding reaction on the end of the capillary tube indicates

full

that

changes induced by recent feeding

performance of the food-taking rather than the

physiological

the

affect the

food-seeking reflexes.
3.

Unlocalised Action of Chemicals.

Fig. 35.

An extensive

Diagram showing tlicfonu of crawling niovcnient, exhibited

by PI

11

aria

wlieii

placed in 10 per cent. NaCl.

experiments on the effects of immersing planarians

was performed, but as the results threw
but comparatively little light on the general nature of the
series of

in various solutions

behavioui', they will be reported only briefly.

any strong solution causes marked changes

ments.

The gliding

disappears.

is

Immersion in
the move-

in

made very much slower

NaCl a peculiar form

In 10 per cent.

or entirely
of crawling

Very pronounced contraction waves pass over the

body longitudinally, giving it the aj)pearance shown in
Fig. 35.
In 2 per cent. CuSO.j^ the animals make no progressive movements, but wave the head violently from side to

appears.

side.
In strong solutions of acids the worms squirm violently
without making any effective progressive movements. In all
these strong solutions the sensitiveness to all stimuli is

VOL. 46, PART 4.

NEW

SERIES.

XX

RAYMOND PEARL.

670

This can best be shown with mechanical

greatly diminished.

In strong solutions of NaCl (10 per cent.) the

animals make no attempt to right themselves if placed with
their dorsal surfaces down. Another peculiar effect of strong
stimulation.

NaCl

solutions of

This organ

is

to

is

solutions

example^

and extended to a much

Immersion of the animal in

thrust out of the body

greater length than

weak

cause the extrusion of the pharynx.

usual.

is

cause

that

per cent, sugar

movements, but when

the

has

capillary tube method.

contact with water

as a

weak

III.

reaction

no definite

in these solutions the

the positive reaction to tap water

by the

positive

effect

as^

for

on the

animals will give

when the latter is applied

Under such circumstances

a slight environmental change, and acts

is

stimulus.

Tliigmotaxis and the Righting Reaction.

Tliigmotaxis. If a specimen of Planaria is turned

over and placed dorsal side down on the bottom, it will
immediately right itself. This is done by a very characteristic
reaction, and is one of the first things to attract the attention
of one studying the behaviour of the organism. Loeb ('94, pp.
a.

251

252)

held that the righting reaction in the poly clad

Thysauozoon was due

to

the

negative and positive

tliig-

and ventral surfaces

The evidence offered for this view was that
respectively.
when the thigmotactic relations of these two surfaces were
reversed, the animal reacted strongly, and tliat this result
could not be due to any effect of gravitation, since the animal
assumed all possible relations to gravity, and kept them for
It seemed to me desirable to
considerable periods of time.
get, if possible, some further evidence on this subject, and to
work out the mechanism of the righting reaction.
motaxis

{" stereotropism ") of the dorsal

That the dorsal surface of the animal is negatively thigmotactic is certain, and can be shown in other ways than by
laying the animal on its dorsal surface. For example, if a
piece of cover-glass be gently laid on the dorsal surface of

MOVLIWENTS, ETO., OF FRESH-WATER PLANAKIANS.

either a resting or a moving' specimen,

move out from under

Furtlier,

it.

if

it

will

671

very promptly

crevices are arranged of

this

form

and

letting the two opposite corners rest

by supporting cover-glasses at two corners,

on the bottom of the
specimens will not go into them. The moment the

dish,

dorsal

(^^r"^)

touches

surface

the

cover-glass above,

begins to react violently, changing

the

worm

direction of movement,
and goes out from under the cover.
With the existence of an apparent negative thigmotaxis of
the dorsal surface established, however, there still arises the
question as to whether this is the sole cause which induces the
its

itself.
The following experiment
throws light on this point
A specimen is placed ventral
side up on a dry spatula in the air, and then the spatula is
placed just beneath the surface of the Avater in a tall jar or

inverted animal to right

large test-tube

and quickly pulled out from under the worm,

so that the latter starts falling through the water in an in-

Another Avay in which the worm may be

up is by holding it on a scalpel
point above the water, and then dropping it beneath the
verted position.

started falling ventral side

surface

in

the

desired

down
when the animal

ventral side

way

as

described later).
itself it

itself

again.

Before

position.

dropped any great distance

righting reaction, and turn

it

itself

This
is

the

will give the

is

worm has

characteristic

over so as to bring the

done

in precisely the

same

inverted on the bottom (to be

After the falling animal has thus righted

may again

give the same reaction, and thus turn

over so that the dorsal side

cases I have seen a

worm

is

down

again.

In a few

after righting itself the first time

keep right side up during the remainder of the fall. The

most usual behaviour is for the animal to keep giving the
righting reaction all the time that it is falling, although this
does not, of course, keep it all the time with the same side
uppermost.
I have performed a large number of these
dropping experiments in which the animals were started in
both upright and inverted positions, and in all cases they
gave the righting reacting one or more (usually more) times

RAYMOND TEARL.

672

before reacliing the bottom, provided the distauce through

which the drop was made was greater than 7 10 cm. This
result seems to indicate that there is something more concerned in the righting reaction than the negative thigmo-

taxis of the dorsal surface for the following reasons

dorsal surface

ment
the

is

(2) it is

case

(1)

the

not in contact with any solid of this experi-

in contact with water onl}', just as

when

the animal

right

is

up.

side

is

normally

It

may be

objected that the experiment

is

result of the falling, there

is

an increased water-pressure on

the dorsal surface, and

this

not conclusive, because, as a

may

act

as

thigmotactic

This objection is met by two different facts.

animal gives the righting response in some cases
while falling ventral side down, under which circumstances
there can be no increased pressure on the dorsal surface.
Second, if a stream of water from a pipette is directly
squarely against the dorsal surface of a worm normally
stimulus.

First, the

gliding about on the bottom the righting reaction

induced, regardless of the force of the stream.

this

is

not

Evidently

stream of water against the dorsal surface produces a

on the dorsal surface similar to that when the

and if the righting reaction in the falling
is due to increase of pressure on the dorsal surface, we might
suppose that some indication of it would be produced in this
case.
As a matter of fact it is not. We must conclude,
pressure

animal

is

falling,

then, that the righting reaction

is

due, at least in very large

some other cause than the negative thigmotaxis of

This is indicated also by the fact that
when solid bodies are laid on the back of a specimen in its
normal position, the reaction which is caused is not the
righting action, as would be expected if the latter were
due solely to the negative thigmotaxis of the dorsal surface.
The righting reaction is clearly not due to gravitation, since
part, to

the dorsal surface.

the flat-worms

move on the

surface downward.

surface

film

with the dorsal

This leaves, as the only factor to which

the reaction can be due, the positive thigmotaxis of the

ventral surface.

am

convinced that

it is

to this factor that

MOVEMENTS,

ETC.,

OF FRKSH-WATER PLANARFANS.

G73

While the negative thigniotaxis

some part in the reaction, it is, as
the experiments described above show^ a comparatively unimportant factor. The specific relation of these two factors
the reaction

is

chiefly due.

of the dorsal surface plays

to the definite righting reaction will be

next section, in

Avliich

the

brought out

and mechanism

form

in the

of

this

reaction will be set forth.

The Righting lieaction. The righting reaction is

a very characteristic piece of behavioui', and can best be
described in a single phrase by saying that when the animal
is placed on its back it throws itself into a spiral in such a
h.

way that

tlie

ventral surface of the head

with the bottom.

comes

into contact

This ventral surface then attaches

itself to

the bottom by means of the mucous secretion, and starts

gliding ahead. As it goes forward it unwinds the remainder
of the spiral, as each successive posterior part of the ventral

Fig. 36.

Showinpr

tlie

form taken by Planaria

in the righting reaction,

surface comes into full contact with the bottom.

The form

of this spiral just after the ventral surface of the

head has

come

into contact with the

bottom

is

shown

in Fig. 36.

The

thrown very quickly after the dorsal surface touches

the bottom, and usually includes the whole length of the
body at once. However, by observing a specimen in which
it takes place a little more slowly than usual, it can be seen
spiral is

that the

movement

is

started at the anterior end,

with, for example, the right side of the head, this

under, while at the same time the

left

side

is

13eginning
is

turned

raised.

This,

head region down,

and at the same time makes a twist in the body, just back of
the head. In some cases this is the only twist that is made,
while in others another similar twist is thrown in the body
farther back.
As the anterior end after it is righted glides
of course, brings the ventral surface of the

RAYMOND PEARL.

674

nnwouiul by the raised edge of each

to the bottom as soon as
Thus, of course,
it is in a position where this is possible.
when the animal has traversed a distance equal to its own
length it will have come entirely into the normal position
again. The reaction is really a rotation of the body on its
long axis through 180. The mechanism of the turning is
ahead, the spiral

is

down and attaching

twist dropping

such that only a part of the body rotates at a time, first

the anterior end, then the portion next behind that, and so
on,

till

This rotation by

the whole animal has turned over.

sections, as

it

were, causes the spiral form which the animal

takes on in the reaction.

The number
forming

of turns into

which the body

the spiral varies Avith the

is

thrown

in

length of the individual,

some extent witli its physiological conThere may be only a half-turn in the whole body, or
there may be one complete turn or, again, one and a half
turns; or, finally, as many as two complete turns in the body.
One complete or nearly complete turn, as shown in Fig. 36,
In large individuals
is the usual form of the reaction.
and

appai'ently to

dition.

more twisting
that

is

is

frequently seen.

all

the twisting

absolutely essential for the righting of the specimen

the half-turn given by the

is

Evidently

turning of the anterior end

ventral side down.

The determination
thrown,

or,

in other

of the direction in

which the spiral

is

words, the side of the body towards

which the anterior end turns in order to get right side up,
was for some time a very puzzling problem. A collection of
statistics on the matter showed that the anterior end twisted
towards the right and towards the left in an approximately
number of cases. This is precisely the result which would
be expected if the matter were due to chance only, but the
reaction did not give the appearance of being a chance
Finally, the determining factor was found to be the
matter.
^

relation of the dorsal surface to the bottom.

'

Tn

tlie

towards

figure (Fig. oH)

tlie left..

tlie

worm

is

repiTseiifed willi

cross-section

(lie

spiral

tlu-owii

MOVEMENTS,

RTO.,

OF FRESM-WATRIl PLANARTANS.

G75

body of Plan aria has tlie form sliowii in Fig. 37. It

is convex in outline on the dorsal side, and nearly straight on
tlie ventral.
As a consequence of this shape of the dorsal
surface the animal when placed in an inverted position very
seldom lies exactly on the mid-dorsal line, and if it does at
first it almost immediately tips over to one side or the other,
so that its cross-section has the relation to the bottom shown
in Fig. 37, B and C.
It h then found that the side of the
body which is in contact with the bottom determines in
which direction the spiral shall be thrown.
If the right
of

tlie

is down the right side of the head

nnder towards the left and the left side will be
raised up over towards the right, or, in other words, the
head as a whole will rotate from I'ight to left, i. e. in a

side of the dorsal surface

will turn

Dorsal

Vfntral

B
Fig.

37- Diagranimalic cross-section of Plaiiaria t,o sliow the contact

relations of the dorsal surface of the body to Uie substrate in llie
case of a specimen in an inverted position.

counter-clockwise direction.
surface of the body

down

If the left side of the dorsal

head will
from left to right. This relation may bo made out
easily by direct observation in all cases where the reaction is
is

at the beginning, the

rotate

not too rapid.

The righting reaction is a fnirly rapid one. The head is

turned over and the spiral thrown in the case of a normal individual almost immediately

The length

when the

dorsal surface touches

which it takes a specimen to

get completely righted evidently depends on the length of
the body, because the longer spiral which must be unwound,
the more the time which must be taken. The following
figures will bring out this relation between the size of the individual and the time taken in righting.
In ten trials with
the solid.

of time

RAYMOND PEARL.

676

an
P.

but

active

large

specimen

(about

12

mm.

long)

of

dorotocepliala the average time taken to regain com-

normal position after being inverted was 8'68

seconds.
With a small specimen (5'5mra. long) the average
time taken in righting in ten trials was 5*22 seconds. The
time taken in the reaction also depends, of course, on
pletely the

Thus

the general physiological condition of the animal.

ten trials with a sluggish specimen, approximately 9

long (thus shorter than the

in

mm.

specimen mentioned), the

average time taken in regaining- the normal position was
first

10'90 seconds.

The thigmotactic

may be

irritability

modified or reduced

in several ways, and, as a consequence, the righting reaction

will disappear entirely or in part.
One of these cases has
been mentioned above (p. 670) where it was shown that a
specimen placed on its back in a 10 per cent, solution of
NaCl makes no attempt to right itself. Similarly a specimen
put in an inverted position on a dry surface, care being taken
that no water surrounds the animal, will not give the righting
reaction.
In both of these cases the specimens are able to
move.
The Mechanism of the E-eaction. It is a very
difiicult matter to determine exactly the muscular mechanism

of this righting reaction, since

ment, and

the formation of the spiral

as will
it

is

it is

ordinarily done in

such a complicated move-

most essential feature

Furthermore,
appear from the operation experiments to be described,
is

its

so very quickly.

almost impossible to devise crucial experiments of a

character which will demonstrate what the mechanism

What

I shall do, then, will

be to present a tentative explana-

the mechanism of the reaction, together with the

tion of

evidence for

it

which

have been able

that the view to be presented

careful study of the

worms, and

is

of

to obtain.

body

may

say

in

normal and operated

a correct explanation.

the righting reaction

of the

the result of a long and

phenomena both

I believe that it is

The mechanism
The half

follows

is.

is

probably as

of an inverted specimen

which

MOVEMENTS, KTC, OF FRESH-WATEE PLANA RIANS.

677

in contact with the bottom extends (by the mechanism

previously described, pp. 556, 557) in response to the stimulus
given by the contact of the dorsal surface of that side of the
is

body with the bottom. At the same time the opposite half of
the body, by active muscular contraction, keeps its length
the same. Thus any bending of the body away from the
side stimulated as in the ordinai-y negative reaction

vented, or, in other words, the long axis

is

pre-

kept straight by
the opposite side maintaining actively its normal length.
Now the necessary mechanical result of keeping one side of
a flexible system at a constant length while the other side
lengthens must be that the lengthening side will be thrown
into a series of waves.
In other words, it is mechanically
is

impossible for the lengthening side to keep

the same plane.

Furthermore,

if

its

in such

whole edge in
a system

it

is

possible for rotation about a longitudinal axis to occur, the

system will be thrown into a spiral of the form which the

Again, as soon as
one side of such a system under elongating stress changes its
level with reference to the remainder of the system, and thus
starts the formation of the spiral, the long axis of the
planarian takes in the righting reaction.

system

(i.

e.

the centre of the spiral) will keep itself straight.

Any

further force elongating one side will merely throw the

spiral into tighter coils without having any tendency to bend
its

long axis.

This fact

is

of importance in the case of the

planarian where the maintenance of the

initial

straightness

done by the opposite side of the body. Of

course, a symmetrical spiral cannot be formed unless the two
edges are of equal length, but the moment the spiral of the
planarian is started all necessity for one side keeping a conIt must be kept in mind, however, as
stant length ceases.
has been indicated above, that the force which produces the
spiral must act on one side only, and hence the side of the
of the long axis

is

the movement must be

by the other in the spiral formation after
The direction in which the spiral shall
this has once begun.
turn will evidently not be determined by the mere lengthen-

planarian opposite that initiating

moved

passively

RAYMOND

678

PEAT?L.

The determinant

ing of one side of the body.

of this

is

evidently a difference of tension on the uppei^ and lower sides,

the spiral turning towards the side of greatest tension,^ This
greatest tension

evidently, then, in the normal reaction on

is

the dorsal surface, as

since that

we should expect on

a priori grounds,

the part directly stimulated.

is

up, the spiral righting reaction of the planarian,

as I have worked it out, is due to an elongation of that side

To sum

of the

body whose dorsal surface

is

in contact

with the

solid,

while the opposite side of the body actively maintains its

As the elongation occurs the various parts
original length.
of the

body rotate

freely about its long axis,

and hence the

The

spiral turns
whole worm takes on
(i.
so
as to bring
e.
towards the dorsal surface in every case
the ventral surface of the head down), as a result of the
greater tension of the dorsal musculature on the elongating-

the spiral form.

side.

The

reaction

is

thus seen to be of almost the same cha-

racter as the ordinary negative reaction to strong mechanical

primary reaction is an extension of the

The difference between the two is that in

stimuli, in that the

side stimulated.

one case there is a bending of the longitudinal axis of the

body, while in the other there is a rotation about this axis.
On the view just given of the mechanism of the righting
reaction the specific parts played by the positive and negative
thigmotaxis of the ventral and dorsal surfaces are evident.
The positive thigmotaxis of the ventral surface is the primary
cause of the whole reaction, and
factor of the two, as

is

evidently the stronger

shown by the experiments

of laying

on the dorsal surface of the animal when in a

normal position. It will be recalled that such treatment does
Further
not call forth the specific righting reaction.
solid bodies

evidence of this same thing

'

fied

The statements

is

found

in the fact that speci-

as to tlie mechanical principles of a spiral have been veri-

with different sorts of models, including plastic clay, rubber bands, etc.

Lack of space

will

anyone can verily

not permit the enumeration of these experiments

for iiimself the various

statements with very

in detail,

little

but

trouble.

MOVEMENTS,

OF FRRSII-WATER TLANARTANS.

ETC.,

G79

mens will remain in the normal position on tlio bottom of a

clisli when there is a layer of plant debris a half-centimetre
in thickness above them, and necessarily in contact with the
The negative thigmotaxis of the dorsal
dorsal surface.
surface plays
in

its

part in the righting reaction in determining

which direction the turning shall take place.

It has so far been shown that the view of the mechanism

of the righting reaction presented

mechanical principles

is

in accord with all the

necessary to produce the

observed
an examination
of the evidence that this mechanism is the one which actually
brings about the reaction. This evidence is obtained from
experiments with worms on which operations have been per-

results.

formed.

The

attention

Obviously,

if

may now be turned

to

the mechanism described

by which the reaction

is

the one

produced, any operation which

destroys or throws out of working order any essential part
of the

mechanism

is

will cause the typical reaction to disappear,

or be greatly modified.

We may

first

consider the reactions of the pieces resulting

from cutting the animal in two transversely in the middle of

the body. It is found that each of the pieces resulting from
such a cut will perform the righting reaction in the typical
manner. The spiral is formed, but there is usually only one
half-turn of the body, i. e. just enough to bring the anterior
This then attaches itself to the
end ventral side down.
bottom and starts gliding*, unwinding the spiral just as under
normal circumstances. There is observable the same relation between the side of the body, which is in contact with
the bottom and the direction of the turn as in the normal inThe only striking diiference in the behaviour of
dividual.
the anterior and posterior pieces is that the reaction time of
the former

is

much

shorter

than that of the

latter.

The

anterior piece rights itself practically as quickly as does the

normal animal, while the posterior piece took in one series of

experiments 1 minute and 38*1 seconds (average of ten trials)
for

complete righting.

This

slower

righting

reaction

is

another expression of the generally lowered tonus of such

RAYMOND PEARL.

680
posterior pieces.

By varying

tlio

position of

tlie

cuts, seg-

body of various lengtlis may be obtained. All

of these, which are about 1-|- mm. in length, will usually right
themselves by as close an approximation to the typical spiral
The side of
reaction as is possible under the circumstances.
the body which is lowest can be seen to elongate in these
very short pieces, and just enough of a twist is found to
bring the ventral surface of one corner of the anterior end
Of course, no complete spiral
into contact with the bottom.
ments

of the

can be found in such short pieces. Their reaction time is

very slow.
Next, experiments were tried with the pieces resulting

from

splitting longitudinally anterior halves of

worms

in the

These pieces had the form shown in Fig. 38.

Evidently such pieces have only a half of the mechanism
necessary for the performance of the spiral righting reaction,
middle

line.

Fig. 38.

Operation diagram

(see text).

according to the view given above, and therefore should not

be able to give the typical response. They have one complete side which may elongate, but they have no other side
to keep the middle line straight, and so make the elongation
eifective in forming a spiral. Such pieces, when placed with the
dorsal surface down, reacted immediately by bending strongly
towards the cut side, i. e., so that the concavity was on the
cut side.
This was kept up for a time, the animal squirming
about violently, but it was finally replaced by another reaction.
The ventral longitudinal muscles contracted strongly, and
raised the anterior end of the piece well up from the bottom
(shown in side view in Fig. 39, a)
After a strong raising contraction the piece would extend and settle back again. Then
.

after a time the raising

noticeable that the piece

was repeated, and it soon became

was rising higher each time and

MOVEMimTS,
settling

back

ETC., OF

loss after

FRMSH-WATER PI,ANARIANS.

each

681

Successive stages of this

trial.

it worked up till
and then the next
contraction caused it to fall over of its own weight and come
down right side up (/). The sticky mucous secretion at the
posterior end was undoubtedly what held the piece up after
each successive trial. This behaviour, as described, was

rising are sliown in Fig. 89,

it

h, c, d.

Finally,

stood directly on the posterior end

[e],

uniform in all the trials.

The behaviour of these pieces brings out several points of
importance. First, it is to be noticed that no trace of the
typical spiral righting reaction is to be seen yet, on the other
hand, we find the pieces bending strongly towards the cut
side when first inverted, which is just the ell'ect Avhich would
;

a^

.cdUL

f
Fig.

3'J.

Diagram

the nietliod of riirliliiig adopted

of the pieces shown in Eig. 38.
sliowiiif;

l)y

one

be produced by the lengthening of the stimulated side in the

normal righting reaction, provided, as actually obtains in this
case, there was no opposite side to keep the long axis of
the

piece

straight.

Tlius

which would be expected

if

we get

precisely

the

result

the view given of the mechanism

of the reaction is the correct one.

Another fact that is
brought out by this experiment is the apparent adaptation
shown. When the animal is unable to give the usual reaction
for righting itseK it very quickly reacts in an entirely
different way, but attains the same end result.
A worm was cut so as to give a piece of the form shown at
A in Fig. 40. This piece was placed in an inverted position

EAYMOND

682

and

its

reactions observed.

PEAEI;.

Evidently, so far as injury of

the mechanism by the operation

is

in essentially the

in the previous expeimnent.

of the body.

ably at

is

concerned, such a piece

same condition as the pieces described

It

has only one complete side

The piece when inverted scpiirmed about

first,

but gave no indication whatever

consider-

of the

normal

spiral reaction. In a short time the violent movements ceased,

and a notch was noticed in about the middle of the uncut
edge (cf. Fig. 40, h). This soon grew larger, and extended
more and more towards the ends of the piece, as shown in

l'"iG.

ileavy lines indicate tiic cuts.

0, c,
iO.
a. Operation diagram,
and d. Successive stages in tlie righting reaction of the piece A of
diagram a. e and /. Cross-sections through A at two successive
See text for further explanation.
stages in the rigliting process.

and d. By close observation the cause of this appearance

was found to be that the thin mobile edge was folding
under and attaching its ventral surface along the bottom. A
cross-section through the worm at this stage had the outline
shown in e. As soon as a considerable portion of the edge
had so folded under and become attached, the piece gave
a series of strong contractions and literally " flopped " over
A stage in
the attached edge and came down right side up.
This behaviour
this process is shown in cross-section in /.
was so peculiar, and at the same time precise, that the
c

MOVEMENTS,

ETC.,

OF FUBSH-WATEll I'LANAUIANS.

683

experiment was repeated many times on this piece and on

Tlie same method of" righting
others cut in the same way.
After the first few times the turn
was always observed.
is

made

in this

way;

it

is

done more

each succes-

(piickly at

sive trial.

This experiment leads to the same conclusion regarding

the mechanism of the righting reaction as did the previous
one.

It affords

lation

in

position)

another and more striking example of regu-

The piece
by a reaction which
reactions.

the

attains

end

(normal

undoubtedly never had

it

occasion to practise before.

Isolated longitudinal halves of the

body react

in the

same

way as did the piece described in the |)receding e.\})erinient.

They right themselves by folding under the edge, and then,
by violent contraction, drawing the rest of the body up over
There is no trace of the spiral righting reaction.
A specimen cut in the manner shown in Fig. 41 shows a

it.

Tig. 41.

Operation diagram

very peculiar righting reaction.

down

(see text).

When

the portion posterior to the

placed dorsal side

median longitudinal

immediately gives the spiral righting reaction, and drags

The process is slow but
very characteristic, so that there is no doubt of the nature of
slit

the two ])assive anterior pieces over.

the reaction.
of a

This

worm where

sIioavs

that in that part of a single piece

the necessary mechanism

is

present

the spiral righting reaction, while in other parts

it

we

get

does not

appear.

The same

point can be brought out by splitting a

worm

longitudinally from the posterior end up to a point near the

head.

The complete anterior part

of such specimens gives

the normal spiral reaction, while the posterior parts remain

passive so far as this reaction

considerable

number

is

concerned.

of different experiments

were per-

EAYMOND PEAEL.

684

formed for
operations,

tlic

purpose of testing the righting reactions after

bring out anything

but since none of them

from the results already given, they will

not be reported here. But it may be said in general, that all
the experiments gave the same results with reference to the
mechanism of the reaction, namely, that so long as the
mechanism described above was intact the typical spiral rewhen this mechanism was destroyed or
action was given
injured the reaction was not given, but the animal, if it
righted itself at all, did it by a different method.
When the animal falls freely in the water the righting reaction is induced because the ventral surface is no longer in
contact with a solid. There is no reason for thinking that
the mechanism of the reaction in this case is any different
from what it is when the animal is placed in an inverted
The direction in which the spiral is
position on the bottom.
thrown in the case of the falling animal is probably determined by slight differences of pressure on the two sides of the
body.
The flat-worm is positively thigmotactic on
c. Summary.
its ventral surface, and negatively thigmotactic on its dorsal
different in principle

surface.

As

a result of this

it

gives a characteristic righting

reaction whenever the normal relations of either surface are

changed.

body

This righting reaction consists in throwing the

face of the anterior end

cases except

The

way as to bring the ventral surdown into contact with a solid (in all

into a spiral in such a

when the animal

anterior

end

starts

is

gliding

dropped into free water).

and unwinds the spiral,

The thigmotactic reaction

thus righting the whole body.
may be modified by chemical and other stimuli. All the
evidence shows that the spiral righting reaction is due to a
lengthening of the side whose dorsal surface is in contact
with a solid, while the other side of the body keeps the long
axis straight.

The

direction

of the

turn in the spiral

is

determined by the side of the body which is in contact with

the solid. This reaction is thus seen to be closely related to
the negative reaction to mechanical and chemical stimuli, so

MOVKMKNTS,
far

iiun'linnisni

;is

nurinal

OF PRKSII-WATKIJ I'l.AXAElANS.

ETC.,

Cut

ooiict'Tiiod.

is

ineclianisin

for

tlic

destroyed, rig-ht themselves

in

])ioc{'s,

G85

wliicli

in

tlic

been
various ways, thus showing a
rio'litiny

reaction

lias

sort of regulation in reactions.

lA'^.

El ec trot axis.

In view of the sliarp and precise reactions of ]^l;inarians to

other

stinnili,

it

oljjects for tlie

was thought

tliat

they would furnish excellent

study of electnjtaxis,

Init nnfortnnntely this is

Their reactions to the constant current are nf)t
clear-cnt, since the specimens become wholly or partially

not the case.

paralysed in a very short time after the current begins to

act, and as a consequence the reactions become feeble and in-

For tho sake of completeness, however, and since

some facts of importance are brought out, the experiments on

distinct.

this subject will

a.

Methods.

be briefly reported.

The

following methods were

used:

The

constant current used was obtained from the general lighting

and reduced to the proper intensity

by interposed resistance.
This apjiaratus for getting the
current I have described fully elsewhere (: 00, 01), so that
it need not detain us here.
In the circuit a rheostat was
circuit of the University,

inserted for regulating

tlie

strength of the current.

unpolarisable brush electrodes were used.

Ordinary

The specimens

were placed either in a trough with clay ends, to serve as

poles, and with paraffin sides of 5 mm. depth, or else on a
slide under a cover supported by several layers of moistened
filter-paper.
These filter-paper ends then serve as the poles
of the preparation, the brushes of the electrodes being laid

upon them.
in this

The layer

of

which the specimens were

was approximately 2'5 mm. in
were obtained by both the trough

water

in

sort of a preparation

thickness.

Identical results

and the filter-paper method, but since tho latter is the lu^ater
and generally more satisfactory method, it was used almost
entirely in preference to the trough.
h.

Results.

The

typical

result

of

the action of

the

current on specimens in such a position that the long axis of

VOL. 46, PAKT 4.

NEW

SERIES.

RAYMOND PEARL.

686
the body

is

approximately at

riglit

angles to the direction of

first.
If a number of specimens are g-liding- about at the normal rate, and a current of
from Aveak to medium intensity is made through the prepara-

the cnrrent, niny be described

of all the specimens is to stop their

forward motion, turn towards the kathode, and start crawling
very slowly towards this pole. The orientation towards the
kathode is at the first trial usually rather precise. The whole
animal gets squarely into line with the current and moves

tion, the first reaction

slowly towards this pole.

anode end of the body, in

While the current

is

acting the

this case the posterior end,

remains

3 +
d
Pjg. 42.

Diagram showing;

lie

PI an aria.
and d. Suc-

typical electiofaclic reaction of

the moment of
cessive phases of the reaction.

a. Position at

making the current,

b, c,

rather strongly contracted, presenting the same appearance

as

when mechanically

stimulated.

Movement

a short time after the current begins acting.

comes

to rest,

occurs only for

The worm soon

and further stimulation serves only

to cause

contraction of various parts of the body without producing

any progressive movement.

The successive stages

shown

typical reaction to the constant current are

of the first
in Fig. 42.

In succeeding experiments on a given individual, and in

many

cases with the very

first

experiment, the reaction

is

IMOYM.MKNTS, KTC, OF FRKSII-WATER PLANAKIANS.

much

less ]')rononiicod.

moment

at the

'I'lie

aiiiinal in

687

the transverse position,

of makino- tlie current, will

sim])lv tnrn

the

somewhat toward the kathcxh' niid

tlien stop.
Reversal of the current causes the head to swing
a short distance towards the new kathode, and then stop
again.
The oinentation becomes less and less precise the
longer the current acts. The position most fre([uently tnken
by a speciuien after it has been submitted to the action of the
current for a short time is shown in Fig. 4^, where it is seen
anterior ]iart of the body

that the orientation of even the anterior end

is not very
such cases the specimen remains perfectly
quiet after the first turn towards the kathode until the

In

precise.

current

is

all

reversed or broken.

The behaviour described is that which is typical for currents

of medium to fairly Aveak intensities.
With verj' weak
currents no striking effect is produced.
With a current

+
Fig. 43.

Diagram showing

partial orientation of

Plaiiaria

to the

constant, current.

which

is

just stong

enough

Paramoccium towards

to cause a general

the kathode, the

planarian gliding at right angles to the current

some cases a very

moment

slight turn of the

movement

of

only eifect on a
is

to cause in

head towards the kathode

making. The specimen does not stop the

gliding movement, and is not forced into any orientation, but
may give a slight turning reaction, which changes its course
from one squarely at right angles to the current to one
turned a little diagonally towards the kathode.
In many
at the

of

cases such a current produces no effect whatever.

strong currents the planarian stops at the

With very

moment

of

jerks the anterior end around towards the kathode

less,

and then

curls up into the

result of verv strono- contraction

form shown

making,
more or

in Fig. 44, as

of the ventral

lonsjitudinal

RAYMONP

688
musculature^ and dies.

PEARFi.

have never been able

to

produce

disintegration on the anode side Avith any current strength at

my

where disintegration
in a specimen
pharynx
behind the

disposal except in a single

began

in the region just

strongly curled up in the

way

case,

described.

In case the long axis of the planarian

is

parallel with the

V
Fig. 44.

Diagrammatic

side

view of a planarian subjecteil

to the action

of a very strong constant current.

and the head is towards the kathode

making, with a perfectly fresh specimen the
effect is to cause a cessation of the gliding movement and a
change to a very slow crawling. The direction of the movement is not changed. There is a well-marked contraction of
the anode (posterior) end of the body. The reaction of the
direction of the current,

at the

moment

of

animal in this position

currents have either no
or else

may

is

shown

effect

in Fig. 45,

on a specimen

J).

Yerj weak

in this position

cause a very slight contraction of the ventral

longitudinal fibres mentioned above.

When

the long axis of the body

of the current,

and the head

is

is

parallel to the direction

towards the anode at the

Diagram showing tlie elcctrotactic reaction of Plan aria when

Fig. 45.
the long axis of the body is in line with tiie current direction, and
Contracted portions are indicated
the head is towards tiie katliode.
by heavy lines.

moment
is

of making, the effect of a current of

to cause the gliding

there

is

movement

a very definite contraction of

of the body.

medium

intensit}-

At the same time

the anode (head) end

to stop.

As the current continues

to act the

specimen

begins to squirm about, and very soon gets out of line Avith
the current. Then the anterior end is turned toAvards the

MOVEMENTS, ETC, OF FRESH-WATER PLANARIAKS.

689

may be continued until combrought

about
and
the animal comes to lie
plete reversal
again in line Avith the current, but with the anterior end now
directed towards the kathode. This reversal into the usual
orientation is the typical reaction for fresh specimens at the
kathode slowly,

unci this process

is

the current

first trials of

it is

shown

in Fig. 46.

In case the

specimens have been under the action of the current for some
The specimen
time, there is no reversal of the position.

J +
I'lG.

46.
l)iai,Miim bliowiiig the elect rolactic reaction of Plaiiaiia wlien
the long axis of the body is in line with the current direction, and
the head is towards the anode.
Contracted portions are indicated

by iieavy

lines.

simply renuiins in the same position and contracts strongly at

the anode (head) end of the body.

Strong currents have the same effect as described in the

preceding experiment. Very weak currents cither have no
definite effort, or else cause a slight jerking back of the head,

and turning a

little to

one side at the

moment

of

making.

After the animals have become partially paralysed by the

liAYMOND riOAUL.

600
iU'tion

ol'

till'

iMiiTi'iit

iKiliin'

llu'

tlu-

ol'

contract ions and

relaxations of dilTiTi-nt pai-ts of the body can

and since

\>c

very fU-arly

most signiiicant features of thi>

animal's reactions to the electric current, they niay be iK'scribed a little more fully.
These reactions for tlie three
scHMi,

tlicsc arc the

shown in I'ig. 47. The essential features

anode eml of the body Avlien in line
with the cui'rent, and convexity on the anode side when at
right angles.
IJesides this there seems to be some slight
exj)ansion at the kathode end of s]X'cimens in line with the
positions are

cliii'f

art"

contraction of

tlu'

curi'ent,

but this a])])earance

current

in

is ne)t

constant.

Iveversal of the

these paralysed specimens causes contraction at

/^^-

Yic.

4-7.

Diagram

contractions caused by tlie current with

three principal pot-itions.

bhowiii;;; llic

tlic l)ody in tiic

new anode imuI or bending towards the new kathode.

breaking the current the contracted portions relax.

the

c.

Mechanism

of the Iveactions.

It will

be seen from

the figures, and the accemnt which has been given

n>sponses to the electric current, that there

is

On

ol'

tlu'

an apjiarent

anomaly in the lu'liaviour. The specimen contracts always

at the anode end of the body, 1)ut a]i])ai'ently not on the
anoiU' side of the body.
believe that the explanation foi1

behaviour

this ap]iarent dilferenci' in

structure of

tlu-

animal, and

in a

the constant current which has binm

When

the animal

is

in line

is

to

be found

pi'culiarity in

noted

in

in

the

the action oi

another case.

with the current the contraction

MOVKMKNTS,
(>l>.sei'VC(l

KTC.,

at Llio uiiodu

ol'

uud

KliKSII-WATEIf.

shown

is, tis

fi])rcs_,

wliilc

fibres

lie (il)rc's

of tlic circiihir

Jn other words, the

which hear

has been shown

an

in

which

produced

a contraction

l>y

dorso-ventral

fibres

oi'

luli

taken

nal

\>y

niiisclc-

;ui<l

transverse Kystem

only alTects those

oi'

tnrnin<^-

(})rincipa]ly

001

A NS.

ciii-i'ent

ai-e ])ar!iHcl

eai-lier ])ai-t

ordinary ne^^ative reaction the

is

(^-i

definite orientation with

a,

jhnv, vi/. those

ti(jn oi' its

l.'l

])y tlic loiiii

the part reacting-, a contraction oftlio Ion

are relaxed.

l'I-AX.\

the

this

r(rl:i,li(jn

with

])a|)(M'

to direc-

Now

it.

that

in

it

the

awjiy i'rointhe stimulus

cii-culai-,

transverse,

the cii'cular)

on

the

and
side

stimulated.
I'^videntiy when the ;iiiiiii;il is at right angles to
the direction of the flow of the current the only muscle-fibres
in

tiie

body

\vli(;se

longitudimil axes are in line with the

current are the fibres of the circular and transverse systems.

assumed that the current acts differently in one

theic is no ajtparent i-eiison why, when
the animal is in the ti"insvei-se ])osition, the; fibi-es which are
in line with the direction of the curi-ent (jii th(! anode side of
the body should iK.t conti-act.
if the filn-es fullilbng tliese
Unless

is

it

case from

in iinothei'

conditions as to location and o]-ientati(ni (the circular system)

do contract, they will cause the anterior end to be turned

towards the kathode and the anode side to become convex,
in other words,

produce the actually observed result. The

system sliould not be affected, and
there is no evidence; that they are. This explanation assumes
that the current produces its effect by directly causing the
fibres of the longitudinal

contraction

of

prf)perly oriented rnuscle-fibres, possibly, or

even probably, witlujut relation to the stimulation of any

sense-organs of the aniniiil. Or, to ]nit it in another way, the
responses according tfj this view might not necessarily be
reactions of the oi-ganism at

all, in tlie sense of being something that the animal does after receiving and transfoi-ming

a stimulus, but are direct effects of the stimulus acting on the

motor organs.

It

has doubtless occurred to the reader

another explanation
that they are in no

be produced

if

is

way

the

tliat

possible for these reactions, namely,

essentially different

from what would

animal were given strong mechanical

KAYWOND PEARL.

692

stimuli on tliosc parts of the

in the several positions.

body wliich arc nearest the anode

In other words, the constant current,

from the standpoint of the planarian, produces the same

effect on the anode side or end of the body that a strongmechanical stimulus applied in the same place would.
Which of these two views is the correct one the planarian
does not show cleai'ly. Yet there is some inferential evidence
which makes it seem probable that the first view as to the

cause of the reaction

the correct one, viz. that the current

is

produces direct contractions of muscle-fibres oriented in line

with its direction. The evidence for this view is as follows
:

(1) In the case of specimens which have been for some time
under the action of the current, and are, as has been

mentioned, almost completely paralysed, the essential features

anode side or end can still be produced by a fairly weak current. At the same time it takes a
very strong mechanical stimulus to get any reaction from
of contraction on the

these

pieces, indicating that their

sense-organs are almost

completely paralysed, and their general sensitivity gone.

merely as a stimulus
others which produce the same reactions,

the

why

current

it

acts

should be effective in

weak

If

qualitatively like
is

not apparent

intensities

when another

it

If it acts directly on the

would be capable of producing an effect after the general sensory functions had been
lost.
current are
(2) The contractions produced by the

stimulus fails in strong intensity.

muscles we should expect that

sharply localised,

i.

it

they involve only a certain definite

e.

part of the body whether the current

is

strong

or

weak

whereas mechanical stimuli applied

to the same places with an intensity sufficient to cause the
same definitive reaction will also cause a marked general
response of the whole organism. This is just what would ])c
expected if the current affects only the muscles oriented in
line with it and lying at the anode pole of the worm,
(o)
By analogy with other forms for example, the Protozoa it
Avould be expected that the curreiit would |)roduce some
other effect than that of an ordinary stimulus applied at the
(within certain limits)

693

M0VE.MENT8, ErC, OK FRKSll-WATKR ri,ANARIANS.

same

In

point.

tlie

case of the Ini'ui^oria the current causes

an entirely different reaction from

other

known

pi'oduced by any

tliat

stimulus.

For these reasons, then,

am

inclined to think that in the

case of the flat-woinn the current affects certain definitely

oriented nniscle-fihres directly, and by this means ])roduces in

the main the characteristic reactions.

That the current does

not also stimulate the sense-organs, and so act like other
stimuli applied to the same places, I am not })repared to say,
but it seems probable that the phenomena observed are not
primarily caused by such action.
It has been brought out l)y inference that the cilia play
no part in the electrntactic reaction of planarians. This is
the true state of the case. The current in any intensity
sufficient to cause the definite reactions stops

far as I

have been able to observe,

evidence for this

is

twofold.

ciliary

First, all

immediately, so

movement. The

gliding

movement

and second, by direct observation

specimens crawling ventral side up on the surface film no

stops in effective currents

of

all

ciliary currents
acts.

This

reactions

can be observed while the electric current

result

of

the

interest

of

is

rhabdocoele

connection

in

with

the

Steno stoma leu cops,

O.

Schm. This form, which normally moves freely through the

water by the activity of its cilia, reacts to the electric current
in essentially the same way as do the Infusoria (cf. Pearl, 00),
That is to say, the cilia on the kathode half of the body take
a reverse position when the current is made, and their effective
The different relations of
stroke is towards the anterior end.
the cilia in different positions of the body are shown in
:

This relation of the ciliary beat, coupled with the

Fig. 48.

form of the body, causes, as a uiechanical necessity

Ludlolf,

^95),

the

animal

towards the kathode.

I

to

orient with the anterior

This method of reaction of S t e n o s t o m

worked out by precisely the same methods

])revious study of the

electrotaxis

This reversal of the position of the

action

of

the

(cf.

end

current

of

the

cilia

as I used in a

Infusoria

(:0()).

as a result of the

has hitherto been observed only in

EAYMOND PKAHL.

694
the Infusoria, and to

the samo tiling in a multicollular

fiiul

It is outmatter of considerable interest.

side the scope of the present paper to discuss the relation
of this result to current theories of electrotaxis, as I hope

organism

to be

is

able to do in a later paper, but

it

may be

said that

this furnishes another strong piece of evidence that in the

case of these loAver organisms the current does not cause the

observed reactions in any way comparable to that in which a

mechanical stimulus causes a reaction,

certain "sensation."

On

i.

e.

by furnishing a

the contrary, the current acts as

a physical force on a structure organised in a certain way.

Experiments on the electrotactic reaction

of cut pieces of

lfy^^yyy^^--y^^A~,-~^.^^^^^y^ ^^,

Fig. 4S.

Diagram

sliowiii^ (he electrotactic reaction of the rhabdocele,

St en OS to ma leucops, O. Schm.

planarians have been tried in considerable numbers, but Avith,

on the whole, unsatisfactory results. Anterior pieces result-

ing from transvei'se cuts are the only ones from which I have
been able to obtain any constant results. Such pieces react
Posterior pieces from
like the normal animal in every way.
transverse cuts show the contractions on the anode side and

ends in a slight degree, but there

of orientation.
line

from

the

tions of

end

is

no constant production

longitudinally in the middle

nearly

to

the

head

react

like a normal specimen, although much more

have observed in one case fairly precise orientasuch a specimen. From specimens slit longitudinally

essentially

weakly.

Specimens
posterior

slit

KTC, OF FRI'JSH-WATPIIJ

:\[OVE.MI'JNTS,

PLANAlv'IAN.S.

()95

the iniddlo line from in front backwards 1 have never been

ill

any definite results.

Tliey simply s(|uirm
an aimless way for a mcjnu'ut when ihc eunent is
made and then become quiets and remain so while the current
able to obtain

about

in

passes.

no

'J'he

direction in which

diifereiice

in

tlie

current

their behaviour.

is

flowing makes

All operated specimens

become very quickly paralysed by the cun-ent.

d. Summary.
The constant current very quickly para-

lyses planarians.
of the

anode side

Its specific effect

end

is

to cause a contraction

of the body.

This produces in the

case of fresh specimens a well defined orientation, with the
anterior end towards the kathode.
All prog-ressive move-

ment

after the

making

ing method, the

in their beat.

ment

oi-

cilia

The

of orientation

of an effective current

is by the crawlbeing stopped or very greatly slowed

electrotactic reaction, so far as the attainis

concerned,

is

essentially the

the negative reaction to mechanical stimuli.

same as

In the rhab-

docccle Stenostoma leucops there is found to occur a

reversal of the cilia on the kathode half of the body, such as
occurs in the case of the Infusoria.

V. Keaction to Desiccation.

series of

experiments Avas performed to determine the

reactions of the aninud on drying.

This is an environmental
condition Avhicli planarians ])robably have had to meet with
relative fre(|uency in the course of their history as a species,

and it is a matter of interest to determine whether they have

any method of reacting which protects them from it.
Experiments were flrst performed in the following-

manner

Specimens were taken from the aquarium dish on

the point of a scalpel or a spatula, and lightly touched to a
:

moment to remove any adherent water, and

then laid upon a dry surface either glass or ])aper. The
behaviour was usually as follows
The worm would curl up
filter-paper for a

and thrust the head under the body, as shown in

Fig. 49.
The purj)()se of their behaviour seems to be to get
the body into as small space as [jossible, and especially to keep
closely

RAYMOND PEARL.

696
the head from

At

drying.

fairly

intervals the

frequent

animal straightens out and extends the head in front as far

It is then
as possible^ and makes "feeling'^ movements.
After the drying

withdrawn, and the animal curls up again.

has proceeded
of the

for

some time the

This

whole reaction appears.

posterior part of the

end then attaches

body

inost characteristic feature

is

a lengthening of the

The

to its fullest extent.

itself to

posterior

the surface, and strong waves of

contraction, like those in the crawling

movement, pass over

the body from the posterior end forward.

No

progressive

made, but backward crawling is evidently

attempted, and is only prevented by the dry surface Avhich
There may be considerable variation in the
the animal is on.
this
first part of the reaction with regard to the curling up
may appear or may not, but the attempted backward crawl-

movement

is

iG.

-t'J.

Diagram showing

the reaction of

Planaria

to desiccation.

have found
I have
performed. When the dorsal surface of the worm becomes
dry all movement ceases. If quickly put back into the water
the worm will usually recover completely, even though all

ing
to

movement

of the posterior part of the

body

be a constant feature in the experiments which

movement has ceased

in the air.

put on a slide in the centre of a small area

which has been wet, but on which there is no standing water,
it will squirm about and extend the head frequently, as in
If the head goes outside the wet area
the last experiment.
If the

Avorm

is

very quickly jerked back, and the specimen gives the

negative reaction, i. e. turns away from the side stimulated.
it

is

The attempted backward crawling occurs

in this case just as

in the others, a short time before the dorsal surface dries

It is to

be noted that there

is

off.

never any actual progressive

MOVEMENTS,

movement

of a

on very Avet

water
is

at

KTO.,

specimen

in tlie nir.

filter-])a])er

it

is

any time snrronnded by a

of this lack of ability to

If a

s])ecinien is

placed

not ahle to jtroo-ress nnless

kept constantly clroppino- on

is

G97

OF TUKsn-WATKl^ rLANATHANS.

it

from above, so that it

On account

laj^er of water.

move when out of water, there is no

movement towards Avater.

true hydrotaxis in the sense of

As has been mentioned before, specimens placed on a dry

surface dorsal side down do not show the rightino' reaction.
To sum up, it is found that planarians, when removed
from the water and subjected to a ])rocess
unable to make progressive movements. At
in the drying* process they

form

of

of tlryiny, are
a

certain stage

attempt to crawl backwards

movement which, under

get the animal back into water.

On meeting

might

certain circumstances,

a dry surface

with the anterior end the animals give a well-marked negative

The animal does not give the righting reaction on

reaction.

being inverted on a dry surface.

On the whole, the general behaviour when subjected to
that is, it would tend to prevent the
drying is purposeful
;

animal ever becoming dried up under natural conditions.

There is nothing in the behaviour of planarians to indicate

how the change from aquatic to terrestrial life could be

brought about. T^he fresh-Avater Triclads, so far as I have
observed them, never leave the water and crawl u}) into the
air above the surface film as some other forms do.
VI. Rheotaxis.

large

number

of

experiments were performed early in

the course of the work with various sorts of devices to deter-

mine whether the animal shoAved any

distinct reaction

currents in the Avater, but without success.

from a

pipette, currents

made by

filling

to

Streams of Avater
the tube

of

the

above (pp. 661, 662) with water

and other methods gave no results. If the

diffusion apparatus described

and bloAving

into

currents Avere

ment

it,

made

of the animal

Avith sufficient force to

from

its

threaten dislods"-

hold on the bottom

it

Avould stop

movino- and contract longitudinal) v, and thus attach

itself

698

more

I^AYMOND rEARL.

Weaker

firmly to the substrate.

currents caused no

was inclined to believe that the lono-jtudinal contraction and the gripping" of the bottom Avere tlie
only rheotactic reactions which the organism exhibited. It
was found later, however, that there was a very precise
effect wliatever.

rheotactic reaction of a different character.

In the course

of the experiments on reactions to localised chemical stimuli

by the

capillary tube method,

was discovered that by using

it

a tube with a relatively large opening (from j to 4 mm. in

diameter) and letting the ordinarj^ tap-water in which the

animals were flow out of

it,

by

its

own

weight, a current of

just the right intensity to cause a positive reaction could

produced.

be

The animals would turn very sharply towards the

source of such a current, the reaction being evidently the

same as that given

mechanical).

to

other

This reaction

the usual positive reaction.

is

directed against the head

is

weak

stimuli

It is

and
same way as

(chemical

localised in the

given only when the current

or anterior part of the body.

thus seen that the planarian is positively rheotactic to

very weak currents, the form of the reaction being precisely
the same as that given to other weak stimuli. It seems very
doubtful if this reaction is of any impoi'tance in the normal
It is

activity of the animal.

G.

General Summary and Discussion of Results.

As was
this

stated earlier in the paper, the problem witli which

study deals

common

is

the

analysis of

fresh-water planarian.

the beliaviour of

The movements and

the

reac-

more important stimuli, with the exception of

and heat, have been described and analysed into their
component factors in the body of the paper. It is believed
that it is of the greatest importance to have as complete and
detailed an account of the various activities as possible, and
as a consequence full details have been given in the case of
Since this method of treatment
each subject treated.
necessarily makes the account of considerable length, it has
a tendency to obscure the general and significant results in a
tions to all the
lio-ht

MOVEMENTS,

ETC., OF

FEESn-WATKK PLANAKTANS.

G99

mass of detail. It is desirable, then, to state clearly at the

end the impoi'tant general facts which have been bronoht ont
by this study, and to discuss to some extent their significance.
In this place I shall state the results in a categorical manner,
making no attempt to indicate the evidence on which the
This will avoid needless repetition.

conclusions are based.

The locomotor movements of PI an aria are of two sorts,

gliding and crawling. The gliding movement is produced
by the beating of the cilia on the ventral surface of the
organism. It is by far the most usual method of locomotion.
For its production it is necessary that thei'e be a layer of
sticky, mucous slime between the ventral surface of the body
and the substrate. In this slimy secretion the cilia beat and
so propel the animal (cf. pp. 544 and 545).
The organism
never moves freely through the water without some sort of
mechanical support. The rate of the gliding is changed by
1.

the action of various agents, such as light, chemicals, electricity, etc.

Its direction is

always forward.

The crawling movement is produced by strong longitudinal

waves of muscular contraction passing over the body from
the anterior to the posterior end.
It is more rapid in rate
than the gliding.

It

the organism, and

of

appears only after strong stimulation

its purpose is evidently to get the

animal quickly away from harmful

stinuili.

Its

direction

may be either forward or backward.

Periods of movement alternate with periods of rest in the
When at rest the flatcourse of the animaPs daily activity.

worm

is

in a condition of relaxation

and generally lowered

tonus, corresponding to the condition of a higher organism

in sleep.
(a)

The causes which induce the coming

the primary cause

is

tive.
tlie
its

to rest are

a more or less fatigued condition of the organism.

{!))

without

it

This

the other causes are ineffec-

relatively low intensity of light,

(c)

Koughness

of

This brings the body into a position such that

different parts form angles with one another, and causes
substrate.

the animal to couu' to rest as the result of a reaction wiiich

I

have called goniutaxis

(p.

5G2).

(d)

Certain chemical con-

BAYMONP

700
ditions.

As

riOAKL.

a result of the action of

some one

oi" all

of these

above-meutioiied factors^ collections or groups of ])lanarians

are frequently formed.

Planarians which have been injured by operative procedure

move comparatively little during the course of regeneration,
thus showing a sort of regulation or correlation between
behaviour and morphogenetic processes (pp. 573, 574).
2.

There are two principal qualitatively different reactions

and negative reactions.

to stimuli, the positive

The negative reaction

is

given

in

response to strong

unilateral stimulation of the anterior portion of the body.

essentially in a turning of the

It consists

head away from

the side stimulated. It is brought about by the extension of

the body on the side stimulated. This extension is produced

by a contraction
systems
reaction

of
is

of the circular, dorso-ventral,

muscle-fibres.

The purpose

of

and transverse
the

negative

evidently to get the organism aAvay from harmful

stimuli.

The

positive reaction

is

given only in response to weak uni-

lateral stimulation of the anterior portion of the bod}-.

It is

essentially a turning of the head towards tiie source of the

This reaction is one of considerable precision,
stimulus.
bringing the anterior end into such a position that it points in

most cases exactly towards the source of the stimulus. The

turning is brought about by the contraction of the longi-

The evident
tudinal muscle-fibres of the side stimulated.
animal into
the
is
get
to
reaction
positive
purpose of the
regions of beneficial stimuli.
3.

Whether

the negative or the positive reaction shall be

given in response to a particular stinndus depends primarily

on the intensity of the stimulus, and secondarily on its locaNeither reaction is given unless some part of the body
tion.
in front of the pharyngeal region is stimulated. The negative
reaction is given only in response to stimuli above a
certain intensity (strong stimuli). This relation between
intensity of stimulus and form of reaction holds for both
mechanical and chemical stimuli.

MOVKMI'INTS, ETC., OF
4.

The reactions

of

FRESH-WATER PLANARFANS.

Planaria

cluding- representatives

of

701

to a variety of chemicals, in-

several of the

chemical groups, were studied.

It

most important

was found that

to a

weak

any substance, regardless of its chemical composithe organism gave a positive reaction identical with the

solution of
tion,

To strong

positive reaction to mechanical stimuli.

same

of the

solutions

substances (with a single exception, see p. 657)

the organisms responded by a negative reaction identical with

that caused by strong mechanical stinmli.

Planaria does not

orient itself to a diffusing chemical in

such a way that the longitudinal axis of the body

to the lines of diffusing ions.

is

parallel

chemicals are

Its reactions to

motor reflexes identical with those to mechanical stimuli. The

is an orienting reaction in the sense that it
directs the anterior end of the body towards the source of the
stimulus with considerable precision, but it does not bring
about an orientation of the sort defined above.
5. Several important features in the normal behaviour of
the flat-worm are found upon analysis to have their explanation in the positive and negative reactions to mechanical and
positive reaction

chemical stimuli.

The method by which the organism gets

a special case of the positive reaction.

its

food

From

is

simply

substances

which serve as food for the planarians, various juices diffuse

When the planarian meets any
into the surrounding water.
of these diffusing substances

that

gives the positive reaction,

it

turns in the direction from which the stimulus comes.

is,

The food substance

acts

which the animal reacts

as a

in the

weak chemical

same way

stimulus, to

as to all other

weak

chemicals.

The

direction of the planarian's

with reference to obstacles in

movement, and its behaviour

its

path, are usually deter-

mined by its reactions to mechanical stinuili.

The behaviour of the organism with reference to the
surface film is determined by its reactions to mechanical
stimuli.
0.

either
NKW SERIES.

Strong stinmlation

VOL. 46, PAKT 4.

mechanical or chemical
Z Z

of

RAYMOND PEARL.

702

the posterior portions of the body induces the crawling movement. This is to be regarded as the specific reaction of this
portion of the body. Weak stimidation of the same region
causes local contraction at the point stimulated in the case of
mechanical stimuli, while weak chemical stimuli applied to
this res'ion are ineffective.

The ventral surface of the body of PI an aria is .strongly

positively thigniotactic, and the dorsal surface is negatively
7.

thigmotactic.
8.

When

the organism

is

placed in an inverted position

it

This reaction consists in a

performs the righting reaction.

turning of successive parts of the body about the longitudinal

During the process the animal takes the
axis through 180.
form of a spiral. The anterior end is brought into the upright position

first.

On

analysis

the righting

reaction

is

be a special case of the reaction to strong stimuli

It is brought about by an extension
(the negative reaction)
of one side of the body^ while the other side maintains its
The reaction is given when679).
original length (pp. 676

found

to

ever the ventral surface

is

removed from a

solid or the surface

film of the water.

9.

To the constant

electric

current

Planaria

turning the anterior end towards the kathode.

orientation and movement towards the kathode

reacts

by

Complete

may

occur.

towards the kathode is brought about by an

extension of the anode side of the body. The current causes
a contraction of muscular elements whose long axes are
The
693).
parallel to the direction of the current (pp. 690
Tlie turning

current very quickly paralyses planarians on which

The rhabdocoole Steno stoma leucops

current with the anterior

it

orients

acts.

to

the

end towards the kathode, and

moves towards this pole. This orientation is brought about

by changes in the positions and consequent effective beat of
the cilia, exactly like those which occur in the case of the
Cilia, on the portions of the body directed
ciliate Infusoria.
towards the kathode pole, take on reversed positions.
10. All the normal reactions to stimuli are of the nature of

MOVEMENTS, KTC, OF FKESH-WATER PLANAKIANS.

more or

reflexes,

complex.

less

What

the animal

703

do

will

after a given stimulus, or in a given situation, can be predicted

There is, however, some variation

in the behaviour, depending on the physiological or tonic
condition of the individual at the time of stimulation. Thus
with reasonable certainty.

a stinuilus sufficiently

one specimen

may

weak

to induce the positive reaction in

cause the negative reaction in another

or

same individual may show different reactions

either the positive or negative' to the same stimulus.
The objec11. Psychological Position of Planaria.
tive psychological position of any organism is evidently determined by the relative simplicity or complexity of what it
does.
With a view of determining what the position of

at different times the

Planaria in the psychological scale

make a catalogue of the things which

is, it
it

may be

well to

does in the course

of its ordinary existence.

The animal performs the following acts

It moves progressively by two methods, a
and a muscular motion.
a.

It turns,

h.

weak
c.

of simple reflex acts,

motion

towards

all

stimuli investigated.

It turns,

all strons:

d.

by a complex

ciliary

It

by another

set of simple reflex acts,

away from

stimuli investisj'ated.

comes

to

rest

in

certain

definite

environmental

situations.
e.

and

When

stimulated in a certain

way

it

extends the pharynx

feeds.

Wlien its ventral surface is removed from contact with

a solid body (or the surface film), a reflex of essentially the
same character as that of c brings this surface again into
/.

contact with the solid.

From

these

essential

factors

whose complexity one has only

is

composed a behaviour

to study to realise.

The behaviour is thus seen to be, in the main, what may

be characterised as reflex. It is very simple to say that an
animal's activity is composed of a series of invariable reflex
acts in response to stimuli, but I doubt whether the full

EAYMOND PKAUL.

704

significance of such a condition is always realised.

It implies
that the animal as an individual " does " nothing in the sense

that a

man

place by

" does " things.

its

It is

locomotor organs;

and invariable

relations

to

it

its

moved about from

is

place to

put into certain definite

surroundings by

its

reflex

mechanisms.

Considered as a wholes such an organism is a

sort of shell to hold a series of mechanisms, each of which is
independently capable of doing a certain thing, and in the
doing produces some effect on the shell as a whole. We may

perhaps got a clearer picture of what such a reflex existence

means by considering
if all

for a

moment what would be

the effect

a man^s activities were composed of invariable reflexes,

off by the appropriate stimuli.

Under such circumwhenever a man saw or smelled food he would have
to go to it and eat it.
Whenever anything touched him he
would have to move in a new direction very closely related
to the position of the object which touched him.
Whenever
he touched water he would have to take a bath^ or perhaps
drink till he could hold no more. During the day he would
have to move always in a definite direction with reference to
the sun, and so on ad infinitum. All he did would be
definitely fixed and, in a sense, predetermined by the things

to

be set

stances,

about him.
apparent that the behaviour of Planaria is not thus
entirely and purely reflex, because there is a certain amount
It is

of variation in

in the

body

As has been brought out

it.

in several places

and in paragraph 10
variation in the behaviour is the

of the paper,

conclusions, this

the physiological condition of the individual.

a more concrete form,

we may

of

these

result of

To put

this in

say that a fatigued animal

or an animal in a state of great excitation does not always

react to a certain stimulus by the same set of reflexes as that

by which a normal animal would

is

variation

in

the

intensity

Furthermore, there
the negative reaction

react.

of

dependent upon the intensity of the stimulus producing it.

Another point in which the reactions of Planaria differ
from what would obtain in the case of an organism whose

MOVKMRNTS,

ETC.,

OF FRKSII-WATKIJ I'LANAUFANS.

705

behaviour was composed of invavial)le reflexes is foniicl in tlie

behaviour foHowing repeated stroug stimuli applied to tlie
anterior end (vide pp. 580, 581).
In this case the organism
shows an evident modihability in reaction, for after giving
for some time the ordinary negative reaction, and not thereby
getting

away from the

stimulus,

towards the source of the stimulus.

body we

reactions of pieces of the

680

turns directly

finally

it

Again, in the righting

see entirely

new forms

of

reaction appearing (pp.

683).
In order to give a concrete idea of the psychological

Planaria it may be well to present in parallel

columns the principal factors which make for simplicity in
the behaviour on the one hand, and for complexity on the
position of

other hand.

r actors which tend to make

tlie Behaviour Simple.
A. Essential reflex character at the
basis of

all

Factors which tend to make

the Behaviour Complex.
A'.

Comparatively

the reactions.

large

qualitatively

number

different

of

general

reactions,

B. General lack of modifiability of

Marked

B'.

qualitatively

different

re-

actions to differing intensities of

reactions,

stimulus.
C. Comparatively

small

qualitatively

composing

number

different

C.

Definite

reflexes

general

the

of

relations of

reactions

to

location of stimulus,

reac-

tions.

D'.

Rather close dependence of reactions on the physiological condition of

tlie

individual. This brings

about variation

The behaviour

of

Planaria

is

evidently

in the reactions.

much more com-

plex than that of the Infusoria, as described by Jennings

(loc. cit.).

In the case of the Infusoria,

which make the behaviour

are nearly or quite absent and

C,

D',

isms are at a

much lower

of

all

the factors A', B',

Planaria

in respect to

stage

than

so complicated,

these organ-

Planaria.

The

Infusoria have practically but one purely reflex reaction to

nearly all stimuli, and this reaction is not localised with

RAYMOND PEARL.

706

reference to the location of the stimuhis.

soria do not

show

Again, the Infu-

qualitatively different reactions to differing

PI an aria to a marked degree.

Planaria stands considerably higher in

intensities of stimuli, as does

We

thus see that

scale than the Infusoria, and that the

development is taking place along two main lines (a) the
higher organism reacts differentially with reference to the
and (b) the physiolocation and intensity of the stimulus
logical balance in the higher organism is much more deli-

the psychological

cately adjusted than in the lower,

see

much more

These variations

variation

in

and

as a consequence

the physiological

in the ]3hysiological condition

we

condition.

bring about

variability in the reactions.

In the case of the ctenophore

Mnemiopsis Leidyi we

have an intermediate stage between the Infusoria and

Planaria, Here the animal reacts with reference to the
position, but not the intensity of the stimulus.

This condi-

which an organism reacts with relation to the position

and not to its intensity, must be for the individual a precarious one, because the animal must either go
towards or away from all stimuli alike, whether good or
harmful. Chances are theoretically equal that after each
stimulus it may get a toothsome morsel of food, or, on the
Further development
contrary, serve in that capacity itself.
where Planaria
series
in
the
behaviour
point
beyond the
tion, in

of a stimidus,

stands must be in the line of further differential reactions

with reference to quality of stimulus. A beginning along
this line is made by the planarian, and the process is carried

a step farther in the case of

described by Yerkes
12,

llelation of

G onion em us,

as

recently

(loc. cit.).

Behaviour and Structure.

tions of organisms are evidently, in

any

case,

The reac-

very closely de-

pendent on the structural relations of the given organism, and

on the conditions under which it lives, i, e, its environment
in the broadest sense.
Thus we find the asymmetrical Infusoria, which live freely in the water and move about by means
of cilia, all reacting in the

same way, and the determinative

MOVEMENTS, KTC, OF FRESH-WATER PLANAIflANS.

factor in the reaction

707

the asymmetry of the body (cf.

further founds that
certain rotifers, Avhich live freely in the water and
move
about by the activity of cilia in a similar way, and further-

Jennings,

:00).

Now

is

Jennings has

more are asymmetrical

in fundamentally the same way that

the Infusoria are, react in essentially the same manner
as do
the Infusoria.
Similarly, I believe that the general reactions
method of the planarians may be found to be in the main the

method by which

organisms presenting the same general

and mode of life react.
Only one
example on which this conviction is based may be given
here.
In the case of such fresh-water molluscs as Physa it
is apparent that the actual locomotor and
sensory organisastructural

tion

all

relations

symmetrical in form, and furthermore these forms live

on the surface of solid bodies just as do

is

in fresh Avater

planarians.
Now I have found, in a series of observations
not yet published, that in the case of several of these molluscs
the fundamental scheme of reaction is like that in
the
planarian.
They react in the same way with reference to

the location and intensity of the stimulus, and these are the
fundamental things. In fact, the general behaviour is
strikingly alike in the two widely separated groups.
13.

Purposive Character of Reflexes.A

fact

which

strongly imjiressed on one working on the behaviour of an

organism whose activities are largely reflex is the purposive
character of these reflexes. They are so adjusted that in the
is

long run they keep the animal out of danger, and get it into
favourable conditions. In the flat-worm these two things are
very well done in general by the negative and positive
reactions.

positive
fact that

is
it

Of these two reactions it is easy to see tliat the

the more highly developed, in particular in the
is much more precisely localised with reference
to

We

the position of the stimulus.

can see a reason for this in
the fact that under the conditions of the planarian's life the
Complete observations not yet published. For preliminary account see
N. S., vol. xv, pp. 521 and 525 and Jennings, 01, in bibliography
the end of this paper.

'

'

Science,'

at

RAYMOND PEARL.

708
getting of food
existence than

is

of far

more importance

the avoidance of

in tlie strnggle for

danger.

This point has,

however, been discussed earlier in the paper, and need not

detain us here. The real problem is presented in the attempt
to discover

how any

of

the

purposive reflex acts in the

organisms arose. I see no reason for denying that man}- of

them such as, for example, the positive reaction which gets
the animal its food were developed by natural selection.

There are other evidently purposeful reactions, however, with

whose development it hardly seems as if natural selection

could have had anything to do, since they cannot themselves
be of selective value. This point has been well brought out
in a recent paper by Morgan (: 02, p. 281).
I think a possible explanation of some of these may be found in their
analysis into component factors, when it may appear that
only a very few simple reflexes had to be formed by natural
selection, and then all the reactions are built up from these.
An example will make my meaning clearer. In the righting
reaction of the planarian we have a fairly complex reaction
which is evidently immediately jDurposeful. Yet we find on
analysis that this reaction is at bottom nothing but a slight
modification of the ordinary negative reaction, which might
very well have been developed by natural selection. And
thus it is with other reactions and pieces of behaviour. They
are for the most part built up from a very few simple
purposive reflexes. If we can get them subdivided and
spread out, as it were, so that we can see what goes to
compose them, we may find that our problem has diminished
very much, and we shall have to deal with only a few factors
where before there appeared to be many.
A difficult problem in purposeful behaviour presents itself
when we find that new methods of reaction appear at once if
the usual reaction is prevented. The best examples of this
are found in the righting reaction of cut pieces of planarians.

Here we find pieces of the body, in which the noannal

mechanism of the reaction has been destroyed, immediately
reaching a certain end (the righting) by a method differing

MOVRMKNTS,
entirely from

any

ETC., OF
tliat

FKESH-WATER I'LANARIANS.

709

planarians ever nsed before to attain

we have evidence. Tliese ]ihenomena

have a eonsi(U'ral)le reseml)liince to sucli phciKHiiena as tlie
well-known reo'eneration of the lens from the iris in some

the same end, so far as

It is

iiaturiil

belono's

snch behavionr comes

seh'ction helps ns very

apparently to the same

What

same explanation.

little.

The matter

of

])henomcnii

and probably

h:is

ultimately

this explanation

is

morpliological regulations,

14.

how

not easy to see

Amphiljia.
about, and

class

;i's

the

we do not know.

Functions of the Nervous System. The most

portant function of the brain

of the organism.

diminishes,

After

and on

its

is

removal the general tonus rapidly

which

account the positive reaction

this

im-

the preservation of the tonus

depends rather closely on the physiological condition c:in

be obtained only with great ditticulty in such deca])itated
specimens. There is no evidence of the presence of special
centres in the brain.
The nervous system, as a Avhole, has
its main function in the rapid conduction of impulses.
15. Subjective
Psychic Attributes. One of the
principal questions which forever recurs with regard to work
on animal behaviour is, does the animal possess consciousness ? Now although it has been shown what the component
parts of the activities of the planarian are, yet it cannot be
said, as it seems to me, that the planarian does not, or, on the
other hand, that it does, possess consciousness. All that any
such an organism ever has done in the past, or ever will do
in the future, cannot tell us whether it was conscious in the
doing or not. Any " objective criterion " of consciousness
does not exist. Furthermore, whether consciousness is or is

not present in any given case

concern of the physiologist,

tive explanation of

of

what the animal

subject Claparede

how
is

(:

not, in

is

who

any event, the greatest

rests content with the objec-

results are

brought about, regardless

thinking about the matter.

On

this

01, p. 24), in concluding an interesting

and valuable discussion, has

animaux sont-ils conscients ?

said,
la

"

la

physiologic

question

et

psychologic en tant que cette science est explicative

meme

les

la

doivent

KAYMOND PEARL.

710

done repoudi'e
'

seulementj

iion

Pen m'iniporte

"

'
!

With

Je I'ignore/ mais encore,

'

am

this standpoint I

thorough

in

accord.

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