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The origin of Fulton's condition factor - Setting the record straight

Article  in  Fisheries · May 2006

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ESSAY:
FISHERIES HISTORY

The Origin of Fulton’s Condition Factor—

Setting the Record Straight Richard D. M. Nash
Antonio H. Valencia
Audrey J. Geffen
In many instances in fisheries science, the understanding fundamental principles about Nash is a principal scientist at the
origin of an idea seems obvious and there fish growth. Institute of Marine Research, Bergen,
appears to be a complete trail of references Norway. Nash can be contacted at
Fulton’s condition factor is most often
which record the development of the idea. In Richard.Nash@imr.no. Valencia is an
attributed with the citation Fulton (1911)
reality, references to older literature are some- associate professor at Facultad de
The Sovereignty of the Sea. This citation is
times misquoted or misused and there may Estudios Suporiores, Zaragoza,
incorrect as it has no reference to “condition” Universadad Nacional Autonoma de
even be misinterpretations of the original arti- in fish; rather, it is a history of how parts of Mexico, Biología. Geffen is a professor
cle. As a case in point, the current attribution maritime law came in to being, including the in the Department of Biology,
of the condition factor to Fulton is misleading, 3-mile zones around each country’s shore line. University of Bergen, Norway.
and obscures the contributions of early fishery If authors over the years had included the full
scientists who first developed the methods for title of the book (The Sovereignty of the Sea:
studying natural populations and methods for An Historical Account of the Claims of England reference was cited in Ricker (1975) but the
management. This essay is intended to rein- to the Dominion of the British Seas and of the words “cited in” were lost during the conver-
state the history of the development of this Evolution of the Territorial Waters, with Special sion from a thesis chapter to a paper.
widely used tool, and to acknowledge the Reference to the Rights of Fishing and the Naval The other widely used reference for condi-
insight of the scientists who first linked weight tion factor, Fulton 1904, is more relevant
Salute) in their reference lists then the error
and length measurements to condition. It is since here Fulton referred to the increase in
would have been obvious (Valencia
not the intention of this essay to discuss the weight conforming to the cube of the length
Hernández 2003). RDMN will admit to being
merits or disadvantages of using length and (see argument in Valencia Hernández 2003).
one of these authors (Nash 1982), and this is
weight measures as an indicator of the “condi- Fulton stated here (1904) that the cubic rela-
a good example of how attribution errors
tion,” “well-being,” “plumpness,” etc., of a tionship is “broadly” true but it does not
occur. To be fair, Nash (1980) stated that the
fish. Errors and incorrect attributions can
become incorporated into subsequent publi-
cations and textbooks over time. Whether it
is to correct a fundamental flaw or simply to
set the record straight, these matters should
be brought to the fore, as they become
known.
Fulton’s condition factor is widely used in
fisheries and general fish biology studies. This
factor is calculated from the relationship
between the weight of a fish and its
length, with the intention of describing
the “condition” of that individual. The
formula is of the form:
K = W3
L
where K = Fulton’s condition factor, W =
the weight of the fish, and L is the length
(usually total length). A scaling factor is
usually applied to bring the factor close to
1. The history of the search for a rela-
tionship to describe fish condition, and
the development of a mathematical tool
for quantifying variations in fish popula-
tions, is an example of how rapidly the Fulton’s condition factor is most often attributed with the citation Fulton (1911) The Sovereignty
exchange of ideas can lead to progress in of the Sea.

236 Fisheries • VOL 31 NO 5 • MAY 2006 • WWW. FISHERIES . ORG

Thomas Wemyss Fulton William Edwin Ricker James “Jas” Johnstone Alexander Meek D’Arcy Wentworth Thompson

accurately reflect the true relationship noted that the ratio varied with species, loca- Fulton (1904) was unaware of these activities
between length and weight in fish. tion, and season. He suggested that the ratio and these results. Johnstone (1912) noted
To trace the origin of “Fulton’s condition varied due to reproductive state and the gen- that Henking and Heincke (1907) adopted
factor,” citations to Fulton’s condition factor eral “conditions of nutrition” brought about the formula given by D’Arcy Thompson. This
and the use of condition factors in general in by seasonality or “other circumstances” in the exchange of ideas and applications was also
fish biology and fisheries were located using environment. The reference to “conditions of recorded separately by Hecht (1916) when he
electronic bibliographic search tools. The pur- …” could be construed as the first reference reported that Heincke and co-workers
pose of this listing was to determine the source to a measure of “condition” of a fish but adopted the cubic relationship, based on sug-
of the earliest mention of Fulton’s condition Fulton’s meaning here is not clear. gestions by D’Arcy Thompson, and used it for
factor—who first attributed K and its equa- Johnstone (1912) reviewed the develop- all their work on plaice.
tion to Fulton. Given that neither Fulton ment of the weight to length cubed formula in Heincke (1908) also published the
1904 nor Fulton 1911 specifically mention a fish studies. He pointed out that Meek (1903) length-weight formula but additionally
condition factor, a further search was con- was the first to show a cubic relationship described the variability in the length-weight
ducted directly through the writings and between length and weight. According to relationship over seasonal cycles and in rela-
bibliographic citations of Fulton and his con- Johnstone (1912), the formula: tion to the spawning cycle. Significantly, he
temporaries. Original published documents (Length(cm))3 goes on to define K as a measure of the nutri-
Weight(g) = K •( )
from the period 1890–1925 were consulted, 100 tional condition of a group of plaice caught at
and the focus of the research was the use of was first proposed by D’Arcy Thompson for the same time and location. Heincke was
the weight-length relationship as an index of fisheries investigations. During the previous probably the first to link K with fish condi-
condition in fish. decade other fishery biologists were busy tion, and stated that the better the condition
Fulton (1902, 1904) explored the rela- studying the seasonality of fish growth, and of the fish, the higher the value of K. Thus, it
tionship between length and weight of a exploring the relationships between increases was probably Heincke (1908) who first intro-
number of different fish species. In the 1904 in length and increases in weight. They prob- duced K as an index of “condition” of fish
paper he studied fish growth through mea- ably were able to measure fish length with into the literature.
sures of length, volume, and weight but opted greater precision and accuracy than fish Johnstone (1911) commented on the
in the end to primarily use the weight of the weight, especially at sea. An understanding of dependence of the value of a (normally
fish, because it was easier (more “satisfac- weight-length relationships would allow fish referred to as K) on season and that this was
tory”). He went on to say on page 142: length to be converted to weight for the study the only “convenient index of condition” of
“According to the well-known law, that the of growth. However, the relationship between plaice inhabiting a particular fishing ground
volume of similarly-shaped bodies of the same weight and length was not stable. For exam- (in this paper he wrote the word condition in
specific gravity vary directly as the cube of ple, Johnstone (1905) reported on weights quotation marks). However, Johnstone
corresponding dimensions—a law which was and lengths of Irish Sea European plaice (1912) gave a fully developed description of
brought prominently forward by Herbert (Pleuronectes platessa), and referred to Fulton’s fish condition. This was probably the first
Spencer in his Principles of Biology—a fish (1904) comment that the weight tended to comprehensive definition of the “condition
which has doubled its length should have increase more rapidly with length than the index.” In this paper the equation
increased its weight eight times.” Spencer cubic law would suggest. This inconsistency K = W3
L
(1884:123) wrote that doubling of size results probably provided the basis for the concept of
was called the Meek-D’Arcy Thompson for-
in an organism being eight times more heavy; condition.
mula.
however, he did not specifically refer to fish Hecht (1916) also reported on the devel-
Thompson (1917) utilized the formula
here. Fulton considered this “law” conve- opment of the condition factor idea. He stated
nient for considering growth rates of fishes as that Hensen (1899), at the suggestion of K = W3
L
all that is required is that one calculates the Reibisch (1899), “divided the weight of a and illustrated the change in K of plaice with
relationship between weight and one of the series of plaice by the cube of the length, in an length (in this case K decreases after sexual
dimensions (linear) at a particular size and attempt to get some weight-length relation- maturity) and seasonal changes in K with
then calculates the ratio between that dimen- ship.” He also pointed out that Meek (1903) spawning season. Thompson (1942) showed
sion and the weight at different sizes. Fulton found a relationship between weight and the that the relationship between K and length
(1904) made observations on a number of cubic function of length and expanded on this varied seasonally due to spawning, e.g., prior
species at different times of the year, and in Meek (1905). Hecht (1916) suggested that to the spawning season the value of K

Fisheries • VOL 31 NO 5 • MAY 2006 • WWW. FISHERIES . ORG 237

increased with length. condition factor to any one Laboratory of the Liverpool Biological
Thompson (1942) called K source. In fact many authors Society, Liverpool, further clarified the use of
the ponderal index and sug- still do not provide a refer- K as an index of condition in fish. The litera-
gested that variations in K ence for the equation. ture is often confusing since there are many
reflected not only the spawn- However, the origin of the arguments surrounding the use of condition
ing state but also changes in name “Fulton’s condition fac- factors and what they mean. Many of the
appetite and general condi- tor” is undoubtedly Ricker authors mentioned above (including
tion. Thompson (1942) (1975). This appears to be Johnstone and others in the early 1900s)
attributed the ponderal index the first time that the idea of
to Livi (1897, cited in pointed out the problems with using K as a
the “condition factor,” and
Thompson 1942) where it was simple measure of condition. This essay pro-
the equation
used in the form W vides the correct historical background as a
3 Friedrich Heincke K= 3 platform for those arguments concerning
rW L
L x100 was attributed to Fulton. Whilst we do not appropriate measures of condition.
After the early 1900s the use of the advocate that the condition factor’s name be ACKNOWLEDGEMENTS
weight-length relationship as a tool for mea- changed and Fulton should retain credit for
suring fish condition became accepted and wrestling with the problem of using the The authors are very grateful to the librar-
the formula assumption of the cubic “law,” the present ians of both the University of Liverpool
W generation of fisheries scientists should be (PEML and main campus), UK, and Institute
K= 3 aware of how this “condition factor” came
L of Marine Research, Bergen, for being so help-
(where K is condition) became commonplace attributed to a specific fishery biologist, with a ful in tracking down some of the older
in the literature. Hile (1936) discussed the use citation referring to maritime law. The correct
references and the Dove Marine Laboratory
of the cubic relationship for describing condi- citation for Fulton’s condition factor should
tion of fish and these ideas were developed be Ricker (1975). for the portrait of A. Meek. The authors are
further by Weatherley (1972). Carlander In summary, it was F. Heincke in the also indebted to Chris Zimmerman (Institut
(1950) presented the cubic relationship as a Helgoland Laboratory who first utilised K fuer Seefischerei, Bundesforschungsanstalt
standard method of assessing condition, and it (what is now known as Fulton’s condition fac- fuer Fischerei, Germany) for translating the
was also discussed in Beverton and Holt tor) as a measure of the condition of fish relevant parts of Heincke’s (1908) paper into
(1957). None of these sources attributed the (Heincke 1908). J. Johnstone of the Fisheries modern English.

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238 Fisheries • VOL 31 NO 5 • MAY 2006 • WWW. FISHERIES . ORG

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