NON-VASCULAR PLANTS • One species of completely colorless

• DIVISION BRYOPHYTA (mosses) liverworts that live underneath mosses is

• DIVISION HEPATICOPHYTA (liverworts) totally dependent nutritionally on its fungal
• DIVISION ANTHOCEROPHYTA associate.
(hornworts) • The gametophytes of more advance plants
are completely dependent on their
Introduction sporophytes for their nutrition. The
• The material nurses used was a species of widespread peat mosses are ecologically
Sphagnum moss (bog or peat moss), which very important in bogs and in the
has since been experimentally demonstrated transformation of bogs to dry land.
to have antiseptic properties. This moss has
specialized water absorbing ‘’leaves” and has Peat mosses sometimes form floating mats over
been used as a packing material in the past. water and keep conditions acid enough to inhibit
• It is still widely used as a soil conditioner. the growth of bacteria and fungi
• The ‘’bandage’’ Sphagnum is one of about
The luminous mosses are found in caves near the
23,000 species of bryophytes that include
entrance and in other dark damp place. They are
mosses, liverworts, and hornworts, many of
called luminous because they glow an eerie
which frequently makes a soft and cool-
golden-green in reflected light. The upper
looking green covering on damp banks, trees,
surfaces of their cells are slightly curved, each cell
and logs that are shaded for at least a part of
functions as a magnifying glass, concentrating
the day.
light on the chlorophasts.
• In contrast, some bryophytes can withstand
long periods of desiccation and are also found None of the bryophytes have true xylem and
on bare rocks in the scorching, while others phloem, and be able to sexually reproduce, all
occur on frozen alpine slopes. bryophytes must have external water, usually in
• The habitats of bryophytes range in elevation the form of dew in rain.
from sea level near ocean beaches up to 5,500
or more meters (18,000 or more feet) in Many mosses do have special water-conducting
mountains. cells called hydroids in the centers of their stems
• Few species are found only on the antlers and and a few have food conducting cells called
bones of dead reindeer. leptoids surrounding the hydroids. Neither type
of cells, however, conducting as efficiently as
• Others are confined to the dung of
herbivorous animals, while others still grow vessel elements or tracheids of xylem and sieve
only on the dung of carnivores. tube members of phloem and most water is
absorbed directly through the surface.
• A few tropical bryophytes thrive only on large
insect wing covers. • The absence of xylem and phloem makes most
The pygmy mosses, which appear annually on bryophytes soft and pliable and it is not
surprising therefore, that birds often use them
bare soil after rains, are only 1 to 2 millimeters
(0.04 to 0.08 inch) tall and can complete their to line their nests.
whole life cycle in a few weeks.

Bryophytes of all phyla often have mycorrhizal

fungi associated with their rhizoids. In some
instances the fungi apparently are at least
partially parasitic
ALTERNATION OF GENERATION The midrib (absent in some genera)
• More conspicuous in bryophytes and ferns occasionally projects beyond the tip in the
than in most other organisms. form of a hair or spine.
• In mosses, the ‘’leafy’’ plant is a major part of • The leaf cells usually contain numerous lens-
the gametophyte generation that produces shaped chloroplasts, except at the midrib. The
the gametes. leaves of peat mosses, however have large
• The sporophyte generation which grows from transparent cells (without chloroplasts) that
a ‘’leafy’’ gametophyte, produce the spore. It absorb and store water. Small, green
usually resembles a tiny can with a rimmed lid photosynthetic cells are sandwiched between
at the tip of a slender upright stalk. the large cells.
• All bryophytes have similar life cycles and • The axis is somewhat stemlike but has no
habitats. However, based on their structure xylem or phloem, although there is often a
and reproduction they are separated into distinctive central strand of hydroids.
three distinct phyla. • At the base, there are rootlike rhizoids
consisting of several rows of colorless cells
None of the bryophytes appear closely related to that anchor the plant. Some water absorbed
other living plants. Little evidence is provided by by rhizoids rises up the central strand, but
fossils about the relationship of each phyla to most water used by the plant apparently
other phylum. travels up the outside of the plant by means of
Botanists speculate that the three lines of capillarity.
bryophytes may have arisen independently from
ancestral green algae. Asexual Reproduction
• Some reproduction in mosses does not
DIVISION BRYOPHYTA
depend on such a sexual cycle and its
• Mosses – almost any greenish covering or Alternation of Generations.
growth on tree trunks and forest floors has
• It has been demonstrated under laboratory
probably been called ‘’moss’’ at one time or
conditions that cells of archegonia and
another. antheridia, paraphyses, ‘’leaves’’, stem, and
• Ex.: lichens (reinder moss), red algae (irish rhizoids can develop protonemata.
moss), flowering plants (Spanish moss) and
club mosses Human and Ecological Relevance of Bryophytes
• Club mosses look somewhat like larger true • Mosses in particular retain moisture slowly
mosses but are vascular plants with xylem and releasing it into the soil. They reduce flooding
phloem. About 15,000 species of mosses are and erosion and contribute to humus
currently known. These are divided into three formation.
different classes • Some mosses grow only in soils that are rich in calcium; the
presence of others indicates higher than usual soil salinity or
• Commonly called peat mosses, true mosses, acidity. When certain mosses are present in dry area, it is a good
and rock mosses indication that running water occurs there sometime during the
year. A few mosses are occasionally a problem in water
• Mosses are distinct.
reservoirs, where they may plug entrances to pipes. A few
• The ‘’leaves’’ of moss gametophytes have no bryophytes are reported to be grazed, along with lichens, by
mesophyll tissue, stomata, or veins such as foraging, mammals in arctic regions, but bryophytes are not
generally edible. Some mosses have been used for packing
those of the leaves of more complex pants. dishes and stuffing furniture, and Native Americans are
• The blades are nearly always one cell thick reported to have used mosses for diapers and under splints
when setting broken limbs.
except at the midrib, which runs lengthwise
• Most important bryophytes to humans are
down the middle and they are never lobed or
mosses. Its extraordinary absorptive capacity
divided nor do they have a petiole (leaf stalk).
 made it very useful as a soil conditioner in and the rhizoids, which look like tiny roots,
nurse and as a component of potting mixtures. anchor the plants.
Live shellfish and other organisms are shipped
in it ‘’Leafy’’ Liverworts are often abundant in tropical
forests and in fog belts. Always have two rows of
• The natural acidity produced inhibits bacterial
partially overlapping “leaves.” The “leaves” have
fungal growth and gives it antiseptic
no midribs.
properties.
• The absorbency, which is greater than that of • In the tropics the lobes form little water pockets
cotton, combined with the antiseptic in which tiny animals are nearly always present.
properties made it a useful poultice material It has been suggested that these water pockets
for application to wounds. Peat, like the may function like the pitcher of pitcher plants.
decomposed peat mosses, is used around the • A third row of ‘’underleaves’’ are smaller than
world as a soil conditioner and as a fuel. the other leaves and not visible from the top. A
• In the manufacture of Scotch whiskey, few rhizoids that anchor the plants develop
sproute barley is dried on a screen over a peat from the stemlike axis in the base of the
fire. The smoke permeates the barley and ‘’underleaves’’.
imparts a smoke flavor to the beverage. • The archegonia and antheridia of the ‘’leafy’’
DIVISION HEPATOPHYTA (LIVERWORTS) liverworts are produced in cuplike structures
The word wort simply means plant or herb. In composed of a few modified leaves, either in
the axils of leaves or on separate branches. At
medieval times when the Doctrine of Signatures
maturity, the sporophyte capsule may be
held sway the herbalists of the day through some
of the bryophytes. pushed out from among the “leaves” as the seta
elongates. When a spore germinates, it
Structure and Form produces a protonema consisting of a short
• There are about 8,000 known species of filament of photosynthetic cells. The
liverworts. The most common and widespread protonema soon develops into a mature
liverworts have flattened lobed, somewhat gametophyte plant.
leaflike bodies called thalli (singular: thallus)
Asexual Reproduction
• The thalloid liverworts however constitute only
• Marchantia reproduces asexually by means of
about 20% of the species.
gemmae (singular: gemma).
• The other 80% are ‘’leafy’’ and superficially
• Gemmae are tiny lens-shaped pieces of tissue
resemble mosses.
that become detached from the thallus.
• Liverworts differ from mosses in several details
• Lunularic acid – inhibits development of of
and are considered complex.
gemmae while in the cup.
• Their thalli or leafy stages (gametophytes)
develop from spore. When the spore Sexual Reproduction
germinates, they may produce a protonema, Gametangia
which is an immature gametophyte consisting • Produced on separate male and female
of a short filament cells. gametophyte more specialized than those of
• Thalloid liverworts have smooth upper surface other liverworts.
as well as various markings and pore, and the • Formed on gametophores (umbrella-like
corners of cell walls of most liverworts are structures borne on slender stalks rising from
specially thickened. the central growth of thallus).
• The lower surfaces have many one-celled
rhizoids. Growth is prostrate instead of upright,
• Antheridiophore: male gametophyte, disclike Sexual Reproduction
with a scalloped margin • Archegonia and antheridia are produced in
• Antheridia: male gametangia produced in rows just beneath the upper surfaces of the
rows just beneath the upper surface of the gametophytes.
antheridiophore. • Like both mosses and liverworts some species
• Archegoniophore: female gametophyte, like of hornworts have unisexual plants, whereas
the hub and spokes of a wagon wheel other species are bisexual.
• Achegonia: contains a single egg. • The distinctive sporophytes of hornworts have
• Fertilization occurs before the stalk of the numerous stomata. They have no setae (stalk)
archegoniophore have finished growing after and look like tiny green broom handles or
which, the zygote develops into a multicellular horns rising through a basal sheath from a foot
embryo. beneath the surface of the thallus.
• Capsule: main part of the sporophyte in which • A meristem above the foot continually
different types of tissue develop. increases the length of the sporophyte from
• Sporocytes undergo meiosis producing the base when conditions are favorable.
haploid spores. • Sporophytes undergo meiosis, producing
• The capsule splits at maturity, and air currents spores.
carry spores away. • Diploid elaters that function similarly to those
of liverworts are intermingled with the spores.
DIVISION ANTHOCEROPHYTA (HORNWORTS)
Structure and Form
• Mature sporophytes look like miniature SEEDLESS VASCULAR PLANTS
greenish to black rods that may curve slightly, ❖ DIVISION PSILOPHYTA (Whisk Ferns)
have gametophytes that resemble filmy ❖ DIVISION LYCOPHYTA (Ground Pines, Spike
versions of thalloid liverworts. They are Mosses, And Quillworts)
usually less than 2 centimeters (0.8 inch) in ❖ DIVISION ARTHROPHYTA (Equisetum)
diameter and thrive mostly on moist earth in ❖ DIVISION PTERIDOPHYTA (Cryptograms)
shaded areas, although some occur on trees.
DIVISION PTERIDOPHYTA (CRYPTOGRAMS)
There are only about 100 species worldwide
• These plants are the ones that do not have
they are uncommon in arctic regions.
any flowers or seeds.
• They differ from liverworts and mosses in
• They include ferns and horsetails. In fact, they
several aspects and appear to be only distantly
can be considered as the first terrestrial
related to them.
vascular plants, showing the presence of the
• Hornworts usually have only one large
vascular tissue, xylem, and phloem.
chloroplast in each cell.
• They can be found mostly in damp and shady
• Each chloroplast has pyrenoids similar to
places. Most ferns are grown as ornamental
those of green algae.
plants.
• Nitrogen-fixing cyanobacteria often grow in
the mucilage. Features of Pteridophytes
• Rhizoids anchor the plants. • Pteridophytes display differentiation – true
root, stem, and leaves.
Asexual Reproduction
• A saprophyte is the main plant body.
• Hornworts reproduce asexually primarily by
• Some of the species belonging to this division
fragmentation or as lobes separate from the
have small leaves called as the microphylls.
main part of the thallus.
 For example, Selaginella. Megaphylls are the • Loosely resemble small green whisk brooms.
large leaves that some pteridophytes have. Exactly where these plants fit in the Plant
• For example: Fern plants. The main plant Kingdom is not clear.
bears the sporangia. These bear some leaf- • They have been associated with a number of
like appendages called the sporophylls. In few extinct plants called psilophytes that
species such as Selaginella and Equisetum, flourished perhaps 400 million years ago, but
the sporophylls form compact structures there is not much fossil evidence to
called cones or strobili. substantiate the link. because whisk fern and
true fern gametophytes share several
Reproduction
features, some botanists classify them with
• True alternation of generation. Here, the the true ferns.
dominant sporophyte produces spores
through meiosis. The gametophyte Structure and Form
generation forms gametes by mitosis. The • Psilotum sporophytes consists almost entirely
spores are produced by the sporangia in the of dichotomously forking aerial stems and are
spore mother cells. These spores germinate unique among living vascular plants in having
and give rise to gametophytes. neither leaves nor roots.
• These gametophytes are free-living • The above-ground stems arise from short,
multicellular and photosynthetic. They are branching rhizomes just beneath the surface
called as the prothallus. Generally, the of the ground.
gametophytes require damp and cool places • Enations, which are tiny, green, superficially
to grow, due to their dependence on water. leaflike, veinless, photosynthetic flaps of
For this very reason, the growth of tissue are spirally arranged along the stems.
pteridophytes is confined to certain • Photosynthesis takes place in the outer cells of
geographical areas. the stem (epidermis and cortex). A central
• The male sex organs are called the antheridia. cylinder of xylem is surrounded by phloem.
• The female sex organs are called archegonia. The xylem is star-shaped in cross section.
• The male gametes are called antherozoids, Short roots, aided by mycorrhizal fungi, are
which release by the antheridia. scattered along the surfaces of the rhizomes.
• When the spores of the plants are similar then 1. Immature sporangium
these plants are called homosporous plants. 2. Sporangia that are fused together in threes
• Heterosporous plants are the ones that have and resemble miniature yellow pumpkins
two different kinds of spores. are produced of the tips of very short,
• In these heterosporous plants, the stubby braches in the upper parts of the
megaspores and microspores germinate and angular stems.
give female and male gametophytes 3. Spore released from the sporangia
respectively. germinate slowly in the soil, in tree bark or
tree fern “bark” crevices, or in other similar
DIVISION PSILOPHYTA habitats.
The Whisk Ferns 4. The gametophytes, which lack
• There is nothing very fernlike in the pigmentation, develop from the spores
appearance of whisk ferns, commonly beneath the soil surface and are easily
referred to by their scientific name of overlooked. Resemble tiny, transparent
Psilotum. dog bone. Cylindrical and may branch
dichotomously. Have no chlorophyll,
 absorbs nutrients via one-celled, rootlike species, and Selaginella, with over 700
rhizoids aided by mycorrhizal fungi. species. They are distributed throughout the
5. Archegonia and antheridia develop world, but they are more abundant in the
randomly over the surface of the same tropics and wetter temperate areas.
gametophyte.
6. After a sperm unites with an egg in an LYCOPODIUM – GROUND PINES
archegonium, the zygote develops a foot Structure and form
and a rhizome. • Lycopodium plants often grow on forest
7. As soon as the rhizomes becomes floors. They are sometimes called ground
established, upright stem are produced pines, partly because they resemble little
and the rhizome separates from the foot. Christmas trees, complete with ‘’cones’’ that
are upright or in a few species hang down.
One group of these fossil plants, of which • The stems of ground sporophytes are either
Cooksonia and Rhynia are examples had simple or branched. The plants are mostly less
naked stems (no photosynthetic flaps) and than 30 centimeters (1 foot) tall, although
terminal sporangia. some tropical species grow to heights of 1.5
Cooksonia is the oldest plant known to meters (5 feet) or more. The upright or
have had xylem. A second group of fossils, sometimes pendent stems develop from
represented by Zosterophyllum, had branching rhizomes. The leaves may be
somewhat rounded sporangia produced whorled or in a tight spiral and are rarely more
along the upper parts of naked stems than 1 centimeter (0,4 inch) long. Adventitious
Whisk ferns are of little economic roots, whose epidermal cells often produce
importance. Their spores have slightly oily root hairs, develop along the rhizomes.
feel and were once nudes by Hawaiian men
to reduce loincloth irritation of the skin. Reproduction
Hawaiians also made a laxative liquid by 1. At maturity, some species of ground pines
boiling whisk ferns in water. produce kidneybean-shaped sporangia on
short stalk in the axils of specialized leaves.
DIVISION LYCOPHYTA Such sporangium-bearing leaves are called
The ground pines, spike mosses, and quillworts sporophylls. In other species, the sporophylls
have no chlorophyll are smaller than the other
• The 950 or so known species of club mosses
leaves and are in terminal conelike clusters
(ground pines and spike mosses) mostly look
called strobili (singular: strobilus).
enough like large true mosses that the
2. In the sporangia, sporocytes undergo meiosis,
Swedish naturalist Linnaeus lumped both
producing spores that are released and
together in a single class.
carried away by air currents.
• Once details of the structure and the form of 3. After germination, independent
club mosses were known, however, it became gametophytes develop from the spores. The
obvious that they are quite unrelated to true
gametophytes vary in shape. Gametophyte
mosses. Today, there are living body usually develops in the ground in
representatives of two major genera and two association with mycorrhizal fungi. Some
minor genera of club mosses. gametophytes develop primarily on the
• The sporophytes of all species have surface.
microphylls that are usually quite small; they 4. All types produce both antherida and
also have true stems and true roots. The two archegonia on the same gametophyte that in
major genera: Lycopodium, with about 50 some species may live for several years. Since
 the sperm are flagellated water is essential for Reproduction
fertilization to occur. Reproduction is similar to that of spike
5. Zygotes first become embryos, each with a mosses, except that no strobili are formed.
foot, stem, and leaves which then develop Both types of sporangia are produced at the
into mature sporophytes. Underground bases of the leaves. Up to 1 million
gametophytes’ chlorophyll doesn’t develop in microspores may occur in a single
young sporophyte unless it emerges into the microsporangium.
light. A number of ground pines also
reproduce asexually by means of small bulbils Human and Ecological Relevance of Club Mosses
(bulbs produced in the axils of leaves) each of and Quillworts
which is capable of developing into a new • Manufacture of theatrical explosives and
sporophyte. photographic flashlight powders.
• In the past, druggists mixed spore powder
SELAGINELLA-SPIKE MOSSES with pills and tablets to prevent them from
Structure and form sticking to one another. The spore powder
• The sporophytes of Selaginella, the larger of itself has been used for centuries in folk
the two major genera of living club mosses are medicine, particularly for the treatment of
sometimes called spike mosses. The urinary disorders and stomach upsets.
approximately 700 species are widely • Some native Americans used it as a talcum
scattered around the world in wetter area, but powder for babies, snuffed it to arrest
they are especially abundant in the tropics. A nosebleeds, and applied it following childbirth
few are common weeds in greenhouses. They to stanch hemorrhaging. It was also
tend to branch more freely than ground pines, sometimes used to stop bleeding from
from which they differ in several respects. wounds.
• Reduce fevers, but partly because of
The two most obvious differences are these:
undesirable side effect, medicinal use of club
1. Their leaves have a tiny extra appendage or
mosses has now largely been abandoned.
tongue called a ligule on the upper surface
Native American of Washington, Oregon, and
near the base and
British Columbia becomes mildly intoxicated
2. They produce two different kinds of spores
after chewing parts of one local species of club
and gametophytes on advance feature
moss. It is reported that they become
referred to as heterospory.
unconscious.
ISOETES-QUILWORTS • Ground pines and spike mosses have been
Structure and Form used ornamentally indoors as ground covers.
• there are about 60 species of quillworts. Some ground pines are spray painted and
• Most are found in areas where they are at used as Christmas ornaments or in floral
least partially submerged in water for part of wreaths.
the year. Their leaves are slightly spoon- • Several species of lycopodium have been
shaped of the base and look like green exploited to extent that they are now on rare
porcupine quills, although they are not stiff and endangered or threatened species lists;
and rigid. Arranged in tight spiral on a stubby they should no longer be collected.
stem resembling corn of gladiola or crocus. • Quillwort corms have been eaten by domestic
Ligules occur toward the leaf bases. and wild animals, waterfowl, and humans.
• The corms have a vascular cambium and may
live for many years.
DIVISION ARTHROPHYTA umbrella-shaped sporangiophore. This has a
• The division Arthrophyta (also called short stalk and a flat, shield-shaped head form
Sphenophyta) contains several genera of which the sporangia project parallel to the
extinct plants and one genus, Equisetum, with sporangiophore stalk.
15 extant species known as horsetails or • Equisetum is homosporous, so all plants have
scouring rushes. only one type of sporangiophore and
• The living plants are all herbs without any strobilus, but is some species the strobilus
secondary growth, and although certain occurs at the tip of a green photosynthetic
species may attain a height of up to 10 m, they shoot, whereas in other it is borne on a
are usually less than 1 m tall. Their aerial special, colorless reproductive shoot.
stems have a characteristics joined structure, • After spores are released they germinate on
with a whorl of fused leaves as the nodes. moist soil and develop into small (1 mm
• The leaves are small and have just a single across) green gametophytes. They have no
trace to vascular tissue, but they are small vascular tissue at all and no epidermis or
megaphylls, not microphylls. stomata; the body is just a small mass of
• The vertical aerial stems from deep parenchyma. Whereas the gametophytes of
subterranean rhizomes, in harsh habitats Psilotum are still somewhat complex, those of
aerial shoots die during winter, but rhizomes Equisetum, ferns, and seed plants are all
persist and plants can spread vigorously. simple. The gametophytes of Equisetum are
• True roots are present being at the rhizome’s either male or bisexual. Antheridia release
nodes. The evolutionary origin of roots is still numerous multiflagellate sperms that swim to
uncertain. the archegonia and eggs. The megagamete
• Equisetum plants are small, having vertical and zygote are never released, but rather are
shoots that arise from subterranean, highly retained and nourished by the gametophyte.
branched rhizomes. In some species such as E.
debile, each shoot is both photosynthetic and
reproductive; in other species such as E.
telmateia two distinct types of shoots are
produced. (R. and L. Mitchell; W.E Ferguson)

Internal structure is a distinctive as external

morphology.

• Stems have pith, so these are siphonosteles,

not protosteles.
• Vessels are rare outside the flowering plants,
occurring only in Selaginella, four ferns, three
genera of gymnosperms, and Equisetum. Stem
elongation causes some cells to be stretched
and torn, forming canals.
• The epidermis contains stomatal pores and
guard cells as well as such large amount of
silica that the stems are tough and rigid.
• Reproductive structures in Equisetum are
specialized sporangia never occur singly, but
always in groups of five to ten located on an