[ARCHIVES
FISHERIES RESEARCH BOARD OF CANADA
Translation Series No. 907
Meth,ods of studying the food supply of fish, in
relation to the problem of their abundance
(Paper No. 4 in "Collection of papers on methods
of estimating stocks and making
catch predictions of commercial fishes")
By M.V. Zheltenkova
Original title: Metodika izucheniya obespechennosti ryb
pishchei v svyazi s problemoi ikh chislennosti.
(Iz sbornika statei po metodike otsenki zapasa i
sostavleniyu prognoza promyslovy4 ryb).
From: Trudy Vsesoyuznogo Nauchno-Issledovatelskogo Instituta
Morskogo Rybnogo Khozyaistva i Okeanografii (VNIRO),
Vol. 50, pp. 89-107. 1964.
Translated by the Translation Bureau (MJK)
Foreign Languages DiVision
Department of the Secretary of State of Canada
Fisheries Research Board of Canada
Atlantic Oceanographic Group
Dartmouth, N. S.
1967
,
SECRÉTARIAT D'ÉTAT
BUREAU DES TRADUCTIONS
'DEPARTMENT .OF THE SECRETARY OF STATE
BUREAU FOR TRANSLATIONS
FOREIGN LANGUAGES
DIVISION
CANADA
TRANSLATED FRoM - TRADUCTION DE
INTO
DIVISION DES LANGUES
ÉTRANGÈRES
-X
English
Rus sian
SUBJECT - SuJET
Fisheries
AUTHoR - AUTEUR
M.V. Zheltenkova
TITLE IN ENGLISH - TITRE ANGLAIB
m
•
(?,
u
Methods of studying the avallehili-ty of-ealecipear fish in conneakeimik-wIththe problem of their abundance
• .•;^
■
(Paper N ° 4 in ' Collection of papers on uastimat} Imethods ofIstocks and emk-eetettlefIfirang
,/
.
TITLE IN FoREIGN L A
' NGUAGE - TITRE EN LANGUE IÉTRANGèRB /
eoreeestr-of-eitteWai
)
Metodika izucheniya dbespechennosti ryb pishchei v sviazi s problemoi ikh chislennosti
(Iz sbornika statei po metodike otsentki zapasa j sostavleniyu prognoza promysloyykh ryb)
REFERENCE - RdFÉRENcE INAmE or NOOK OR PUBLICATION - NOM Du LIVRE OU PUBLICATION,
Tom L
Trudy VNIRO (V.sesoyuznyi Nauchno-Issledovatelskii Institut Morskogo Rybnogo Khozyaistva
i Okeanografii)
Re-gear
611-tet—e
VUJ...ps irel,epcUJUJ CITOff—C1
PUBLUHER-EWTEUR
and Oceanography)
Izdatelstvo "Pishchevaya Promyshlennost" ( Publishing House st Food Industry ")
CITY - VILLE
DATE
Moscow
REQuEST RECEIVED FROM
REQUIS PAR
'
Fisheries Research Board
of Fisheries
YOUR NUMBER
VOTRE DOSSIER No
DATEREcEivED
REÇU LE
1964
Dartmouth Research Station
DEpARTMENT
MINISTRE
769-16-14
20 February 1967
PAGES
89 - 107
56 typed pages
OURNUMBER
NOTRE DOSSIER NO
7123 - 4
TRANSLATOR
TRADUCTEUR
sKruynski
DATE COMPLETED
REMPLIE LE
26 June 1967
›
..,
, Tom L. Trudy Vsesoyuznogo nauchno-issledovatelskogo institute.
morskogo rybnogo khozyaistva i okeanografii (VNIRO) 1964
(Vol. L.Transactions of the All-Union Scientific Research Institute
of Marine Fiaheries and Oceanography 1964)
597.0 577.472
yA;‘Lain ede ;5
Methods of studying the 4.veri-letketi-1-Meof elegelemier fish
in-reaneebeen--wIth.the Droblem of their abundance
M.V. Zheltenkova
The availability of food for fish is considered to be one
of the essential factprs determining their abundance.
An in-
dication of the(actualftylof this problem is the last conference
on the dynemics of fish abundance during which many lectures
were devoted to the influence of food availability on the formation of fish productivity in seas (69).
m4-1
The availability of food permits in many cases to understand the(caUSed)condltioning the abundance of a species end
the peculiarities of growth of fish, the abundance of the population and the variations of abundance of individual generations
offeme or another species in the same water body (27, 30, 51).
The essence of the problem, according to Rikker (64), is
in the clarification of the degree of influence of various
factors of the environment on the abundance of the fiSh population.
In this connection, the solution of the problem of food
availability for fish dhould help in compràhending the laws
governingthe formation of fiàh resources in water bodies,
create the possibility ,of forecasting the state of stocks of
commercial flail and point out the means of the most rational
exploitation of the natural resources of water bodies.
•••2
-
. 2 e
The expression "food availability" introduced into the Soviet
ichthyological literature by G.V. Nikoiskil (.55, 5 6) ha s attained
wide popularity in recent time.
Nevertheless this problem has been attracting for a long
time the interest of research workers. Particular attention was
paid to it in lake and pond fisheries.
Investigations of the
Danish school (Blevece Petersen et al.) represent an elaborate
treatment of the problem connected with food evailability for
fish.
Assuming that in a water reservoir only such a quantity
of fish is able to exist "as can find nourishment for themselves",
Ber connected the resources of water reservoirs with
food availability for fish. Particularly, analysing the problem
of alleged dying-out of mollusks in the Caspian Sea,Ber (5)
believed that the mollusks of the Caspian Sea do not die out
but on the contrary are represented by very abundant, presently
existing species and that there is no reason " to deprive our
children and grand children of the hope of eating caviar".
Assuming that mollusks are the main food of sturgeons, Ber
referred directly (using contemporary terminology) the abundance
of species of sturgeonsto the food availability for them.
According to G.V. Nikolskii (55, 56) the food availability
is determined not only by the quantity and quality of the nouridhment available for fish, but also by the possibility of its
assimilation. G.S. Karzinkin (39) complements this attitude, ,
Translator's note:
The correct spelling is probably "Blegvad" pronounced Blyvad.
•••3
.3•
connecting the notion of "food availability" with the notion
of "satisfying the needs of the feeding organism" • Thus, the
expression "availability of food for fish" represents a short
formulation of the notion "availability of the needs of fish
with regard to nutrition".
It appears that going out from such a definition, the
measure of "food availability" should be the difference between
.
the nourishment required by the fish and that of leich it partakes actually.
The availability of food for fish is judged by the state
of the fish population:
abundance, rate of growth, condition
of fatness end fullness, variability of Characteristics etc.
according to the character of the nourishment resources of the
water body* and conditions of foragineA
The food availability for fish varies considerably, because
the conditions of foraging by fish, on which essentially their
food availability depends, are extremely unstable and subjected
to space and time Changes depending on hydrological conditions
(temperature, salinity), feeding relations of the fishes etc.
(76, 19, 26, 71).
The notion "availability of food for fish" is complex,
representing in extreme cases a function of three values:
the
The nouriehment resources of the water body is the totality
of all animal end vegetation organisms inhabiting the water
reservoir and the products of their decay; the nourishment base
for fish are the organisàs consumed directly by the fish as food.
Only part of the nourishment resources of the water reservoir
forms the nourishment base for fish (65).
x*
The conditions of foraging or the conditions of fattening
are the complex of conditions ensuring feeding of fish, i.e. the
presence of a nourishment base of a determined composition and
magnitude, mutual relations of fish with regard to feeding and
hydrological conditions (24).
•••14
'
(,)
.4.
nouriehment resources of the water body, the needs of the fish
with regard to food, - and conditions ansuring satisfaction of
these needs. Therefore it is not sufficient to judge the extent
of food availability for fish in any water body on the basis of.
only one of those factors.
The Methods of obtaining the indices of food availability
for fish are manifold.
In 'the essence, all the basic indices
obtained during investigations of the biological conditions of
the fish population and nourishment resources of the water
reservoirs are used in drawing conclusions as to the availability
of food for fish.
It would be impossible and47e-v7n7(Pexpedient to recite
all those methods, since this has been done in special investigations and instructions.
Therefore we shall restrict our-
selves to indices of fôod availability connected with peculiar-
ities of feeding by fish and A exploitationjeeemm of the nourishment base, limiting ourselves in other cases only to references
to corresponding instructions.
We shall first of all attenpt
to esttmate what is the contribution of this or some other index
for the solution of questions connected with the problem of the
abundance dynamics of fish end to what extent are reliable the
conclusions drawn on the basis of indices obtained by various
methods.
National literature has been made use of in this paper.
As has been mentioned before (24) foreign literature does not
•••5
•%. •
• 5 •
pay particular attention to the problems of the influence of
feeding on the abundance of fish and of the availability of food
for them in natural conditions, assuming that feeding plays an
established role.
Foreign papers are either devoted to prdblems
of feeding byfiah, or the food resources of water bodies (65).
Feeding,rate of growth and condition of fish as indices of food
availability for them
The study of feeding, rate of growth and other biological
indices of the conditions of the fish population is not only
important as such, but also in connection with the problem of
abundance of fish.
In these works e ecological methods are applied in combination with the results of experimental investigations.
Feeding of fish in natural conditions is studied on the
basis of the content of their intestinal tractse using data on
the ecology of fish and the organisms forming their food, and
also experimental data on the effective action of the foods
and the physiology of feeding.
In the analysis of the contents,
the weight and quantity of specimens of individual organisms
and their frequency are established, the degree of fullness of
stomachs is determined visually em degrees of fatness), the
quantity of empty stomachs is counted, individual and general
indices of fullness and other indices are calculated (65).
During investigation of the contents of the intestinal tracts
•••6
t
6
it is very Important to determine the dimensicins of the organisms
consumed. This makes it later possible to select from the food
resources of the water body the organisms forming the nutrition
base of the fish and to analyse more profoundly the ability to
select, the competition in feeding, the food morphology and the
activity of the fiàh.
All these points, as shown in the investigations by
A.A. Shorygin (76), influence directly the conditions of feeding
and at the same time the food availability for fish. The composition of the food and magnitude of the index of fullness
enable us,'Isihen studying feeding, to make a judgment as to the
availability of food for fiàh.
However, this must be done very cautiously, because we do
not yet have enough data with regard to the standards of food
requirements by fish in-natural conditions. Therefore, apart
from data on the character . of feeding, in order to judge the
food availability for fiàh, it is necessary to utilize also
other indices. Thus, G.V. Nikolskii (55, 56) proposed to use
for that purpose the number of food components.
A narrow range
of foods is considered as an index of good food availability,
a wide range is an index of bad food availability. However,
in some cases, e.g. with "vobla" (19, 21) a narrow range of
foods is not a result of a rich nutrition base and high availability of food, but on the . contrary a result of poor and monotonous fauna leading to a reduction of feeding intensity by the
* Translatorts note:
"vobla" Rutilus rutilus
•••7
'
‘‘.
. 7.
li
vobla".
Therefore, to be able to judge the availability of
food from the number of components, it is necessary to apply
also other data, particularly indices of fullness of stomachs
and characteristics of the nutrition base.(!1M.I. Shatunovskil
used A..t
in conjunction with data on variations in the quality
of the population.
Mostly, the food availability for fish is judged by the
rate of their growth. Indeed, the rate of growth of fiSh
appears to be a resultant of the availability of food for
them. It depends on all the biotic and abiotic conditions
favorable for the fiSh in exploiting the nutrition base. A
special textbook is devoted to the methods of studying the
• rate of fiàh growth (74).
However, it is not sufficient to draw conclusion as to
the availability of food solely on thelDasis of standard data
on the rate of growth.
The rate of growth permits only to state
that there was availability of food in the preceding years, the
very mechaniem however of food availability remains undetected.
It is not known what in the final account has determined the
change of the rate of growth of fish - abundance of food,
hydrological conditions or change in the numbers of consumers,
i.e. their competitive relations.
Promising are in this respect
the detailed investigations of size or weight increments of fish
in intervals of determined periods of time.
Thus, N.I. Chugunova
(73) relying on data of the nutrition base, has shown various
Passage omitted by typographical error:
•••8
M.I. ShatunovskiiMetermining the food availability for Baltio herring (Clupea harengus
membras) used the 4 content of the essentàal oomponent in the food. The higner its oontent,
the better the food availability. However, also thin index alone has little applioation.
Therefore...•
•to.
.8.
availabilities of food for the n voble *, dwelling in various
parts of the Northern Caspian Sea and discovered the influence
of this - availability on the speed of maturing of the fish.
K.A. Zemskaya (29, 30) herself succeeded te provVa con-
nection between the rate of growth of bream of the Northern
Caspian Sea and the availability of its food. As an index of
food availability, the author used in one case the quantity
of the whole nutritious base for bream (29), in another the
quantity of food for one individual (30).
Yearly variation
of the rate of growth of bream and roach dwelling in the
Azov Sea also correspond to the conditions of their foraging
(22, 26).
Cseeftels indices of the conditions of foraginOsorvod tile
In
r'
indices of feeding (the magnitude of indices of fullness of
intestines, composition of food, presence in it of organisms
of high caloric value) and indices of the nutrition base (total
,
L
organisms and the quantity of food organisms biomasefd
per single specimen)41/4.
The food availability for fish is judged also on the basis
of their condition and fatness. However the works by G.E. Shulman
(78) and D.A. Shubnikov (77) show how cautiously one must
approadh the estimation of food availability on the basis of
those indices.
Increase of weight, accumulation and distri-
bution of fat in various organs
of the fish depend not only
Translator's notes:
* Rutilus rutilus caspicus
" Rutilus rutilus heckeli
•••9
• 9 •
'on the environment i.e. the supply of food substances, but also
on interna], regularities created in the process of evolutionary
development. When food availability is not sufficient, the
growth in length is reduced, but the accumulation of fatty
substances necessary for formation of sexual products continues.
Methods of studyinF the consumption of the nutrition base by
fie in reference to abundance , distribution and migration of
the fish
A representation of the nutrition base for fish must not
be construed only on the basis of studying the nutrition resources
of the water body. It is also necessary to know the particular
features of consumption of those nutrition resources by the fish.
Actually, the presence of same or other organisms in the
water body is (:eetr-et-s4D-steletfor a judgment of possible but
not necessarily existing conditions of food availability. for
the fish populating this water body.
Rich food resources of
the water body maY be distributed in places having hydrological
conditions unfavorable for fie: inappropriate salinity, temper-
ature or depth. Thus, in the Azov Sea itself, the major part of
mollusks and worms is inaccessible for young bream because of
too high salinity in locations of their habitat (22, 40, 68).
In the Bering Sea (53), substantial masses of food organisms
cannot be consumed by the flounder because too low a temperature
A
of the water.
Towards the end of the fifties a mass of mollusks
in the Caspian Sea was found to be not available for the "vobla"
because of an increase of salinity (67).
Rutilus rutilus caspious
..1 0
• 10
On the other hand, organisms which would appear fully suitable
as fish food actually are not available for those fish because
of their constitutional features (size, crust or shell, poisonousness) and behaviour (speed of movement, use of shelters) (4).
Milia—celmmbe—ef suah inaccessibility of food organisms e c
regular in natural conditions. This phenomenon has extremely
A
variegated forms, whiCh fact has been illustrated by a multitude of examples (4, 50, 76).
The mutual relationship of food
organisms and the fish consuming them i.e. the relationship
between the prey and the Predator lead in the food organisms
to creation of defensive features rendering them inaccessible
or accessible in 0 limited measure for the fish.
Expressly
because of that, apart from an account of food resources of
the water body, it is necessary to know that is actually con11,Jtsumed by fish as food, and this can be judged ini first place
by the results of autopsy of their stomachs.
Ueing generally accepted methods, the nutrition base of
fish is determined.
For that purpose the food resources of the
water reservoir are investigated accurately, i.e. its plankton,
benthos, fish population and detritus, (3). Plankton and
benthonic organisms are subdivided into organisms forming the
nutrition base for various kinds of fish in different stages of
their development, and into non-food organisms. Having studied
the feeding of fish and-determined the role of some or other
...1 1
• 11 •
organism in the nutrition, the nutritional organisms are
segregated. Analysing the samples for that purpose it is very
Important to determine the stages of development and the sizes
of the organisms composing the plankton and benthonic population of the water body. The arrangement of such work gives
eg
1,1
the possibility ±1, form an idea on the dynamics of the nutrition
base for fish. A. P. Kusmorskaya (45) showed the dynamics of the
nutrition base for fish feeding on plankton in the Black Sea,
for a period of many years;
I.N. Stark (68) presented the
dynamics of the nutrition base of bottom-feeding fish in the
Azov Sea.
Ya.A. Birshtein and N.N. Spasskii (2), L.G. Vinogradov
(8), A.K. Saenkova (66) determined the yearly changes of the
nutrition base of bottom-feeding fish in the Caspian Sea.
Sim.-
ilar works were conducted by many research workers for commercial
fishes of our landlocked seas and open parts of the ocean (9,
12, 38, 46, 53,
54,
70).
The determination of the capacity of
the nutrition base for commercial fishes in combination with
data on the food requirements by fish makes it possible to cornpute orientation-wise the abundance of fish stocks.
The nutrition base and its exploitation by • fish in connection
with the problem of food availability is also investigated when
studying the distribution and migration of fish.
It is generally known that the distribution and the migrations
of fish are determined by conditions favorable either for spawning
...12
. 12 •
and development of the young fish, or for foraging.
Of course
during the period of foraging the decisive factor is the distribution and abundance of accumulations of food organisms.
When exploring new commercial fishing regions, the food
resources of the water body are investigated, i.e. its plankton
and benthos.
The general representation of feeding by some or other
fish, being a reassumption of the current ichthyological literature, makes it possible to segregate organisms composing the
nutrition base of fish, or at least to eliminate the non-food
organisms.
Informations on the distribution of the nutrition base for
hyd(kes
fish 43;4--the possibilbe to concentrate the search for fish
in locations of foraging l in the most promising parts of the
explored region. There is without doubt a connection between
the distribution of fish during the period of their fattening
and the food organisms in our landlocked seas i.e. Black Sea
(47), Azov Sea (11), Caspian Sea (16, 19, 41, 76) and in open
seas and oceans, the Barents and Norwegian Seas (28, 49, 51)
in the Pacific Ocean, the waters of the South Atlantic (70);
however this problem requires detailed investigations.
All these investigations are based on accurate data concerning the nutrition base and the feeding by fish.TrYarticularlf,
N.V. Lebedev (47) and V.P. Vorobev (11) conducted the exploration
...13
. 13 .
e fish on the basis of determination of the regions in which
fish feed intensively on food organisms, and later N.V. Lebedev
(48) and N.S. Novikova (59, 60) proposed te judWthe stability
of accumulations on the basis of the sizes of the daily food
rations of the fish.
Investigation by B.F. Manteifel (50) and S.G. Zusser
•
(33, 34) show that not only the horizontal but also the vertical
distribution of fish can be explained by the distribution of
their food organisms.
The exploitation of the nutrition base and the magnitude
of the foraging area not only have an influence on the distribution of fishes during the foraging period, but also are one
of the essential factors of evolution, determining the-abundance
of the population of certain species.
IU.Yù. Marti (Si) connects
the magnitude of the stock of herring in various seas with the
extent of their migrations.
G.V. Nikolskii (57) considers the
magnitude of the foraging area mastered by them as one of the
points determining the abundance of fish.
G.N. Monastyrskii
(52) also assumes that the area of foraging, together with
the spawning area determine the abundance of the population.
er.
T.F. Dementeva (69) differanààuting , in this situation, notes
that the abundance of the population in a water body during
the process of formation of its population depends on the
total quantity of food resources.
But the yearly variation
«
c.
. 14 .
In the abundance of generations are determined in the majority
of cases by changes in the biomass of plankton ensuring the
survival of fish in the early stages of development.
The most vigorous,ase abundance and rate of growth of the
population are concerned, are roaches, which multiply in areas
preceding the estuary and- sete foragte In certain sections of
lluted
the sea retreehee by siZe4 water, most. abundant in feed objects,
In—f-laUe41U'e in mollusks (18).
Thus, the nutritioni base not
only determines the total strength of the population, but its
changes cause Variations in their abundance and the stock by
weight.
All this indicates that investigations of the nutrition
base and its exploitation by flail are indispensable. The
results of those investigations have both a practical (establishing the optimum magnitude of catches and survey of fish distribution) and a theoretical importance (understanding the laws
governing the evolution process and formation of the abundance
of fish).
The gm litude of changeability of features as
availabili y for fiàh
an index of food
km
A series of works were publiàhed in recent time, discussing
Imter ,
the connection between the amplitude of changeability of
individual features of fish and the charaCter of feeding and
the degree of food availability for them.
...15
■
e
• 15 •
The authors of those works sul›—eut fram the position taken
by G.V. Nikolskiie who assumes this changeability to be the
result of adaptation to a ahange of the nutrition base (56).
Investigation of the amplitude of variations of length,
weight and rate of development of carp foraging in ponds with
various nutrition bases (62, 63) has shown that in carp well
provided with food the coefficient of variation of dharacter-
istics is considerably lower, than in those badly provided.
In the first case it equals 14-25%, and in the second it sometimes exceeds 60%.
The same is observed with regard to body
length and rate of development. With a lowering of the food
availability the development of individuals occurs unequally.
According to G.V. Nikolskii and V.A. Pikuleva (58) three
kinds of gudgeons live in the river Moskva: short-whiskered,
medium-whiskered and long-whiskered. Gudgeons of different
kind may dwell in the same sector and be found in the same
sample taking. When the food availability is low or medium,
the dharacter of feeding of gudgeons of different kinds is
species-wise
different both in/x±xxxi composition and the width of range;
when it is high the dharacter of feeding is the samet
The degree to which the fish was fed satisfactorily,equal
i.e. the relation between the weight of the food lump (P) and
the length (1),served as an estimate of food availability.
The index P/1 is identical with the index of fullness of the
intestines P/Q, where P is the weight of food and Q is the
weight of the fish.
It is technically simpler to obtain,since in this case
there is no need to weigh the fish, but in contrast with the
usual index expressing a concrete value , the quantity of food
per unit of fish weight (lg, 100 g or lo kg of fish, depending
on the method of calculating the index) this index is but a
conventional value.
• . .16
9
16 •
According to the opinion of A.V. Chepurnov (72) the anplitude
of the changeability of the number of whisker appendages in
the Norwegian herring * and the Baltic herrine* is connected
with food availability for them during the period of formation
of the whisker appendages (the rate of growth serves as the
index of food availability).
In individuals growing slowly
the amplitude of changeability of the number of whiskers appendages was the largest, in fast growing individuals, the
emallest. M.I. Shatunovskii (75) assuminp food availability
as the possible cause of formation of the spring and autumn
forms of the Baltic herring, separated anong yearlings (on the
basis of scale structure) individuals belonging to the one or
the other form. It transpired that areas of foraging of the
different forms are located at different depths.
This permits
the species to embrace larger foraging areas. Applying as
index of food availability the percent content (by weight)
of the essential component, M.I. Shatunovskii makes the deduction that with better food availability the coefficient of
length variation of Baltic herring is reduced.
All those investigations show that with a lowering of food
availability for fish, the amplitude of changeability of their
characteristics increases. Peculiarities of feeding characterized by different indices serve here usually as the criterion.
Only G.D, Polyakov uses the nutrition base of the water body
as criterion of food availability for fish (62, 63).
Translatorls . notes:
e "serd" Clupea harengus
* u salaka n Clupea harengus membras
...1 7
• 17 •
The study of the amplitudes of dhangeability of the population in various conditions, supplying a complementary criterion permitting„I to judge the food availability for fish, shows
how conditions of foraging influence the formation of biological
and morphological characteristics and how the intensity of
mutual relationships between species are moderated. .
Studv of the requirements of food by fish
A judgment of the food availability for fish in natural
conditions can be made from comparative standards.
These
standards themselves can only be obtained on the basis of the
food requirements by fish. In this context the requirements
-bo
of food by fish give an idea on the degree el which the food
resources of the water body are exploited and permits, as was
tve--
mentioned before, to estimate at least in general outlines il the
magnitude of the fish population inhabiting the water body.
Experimental works give the most accurate representation
of the food requirements by fish; they make it possible to
establish the quantities of nutritious substances indispensable
for the basic metabolism and growth of the organism. The essential aspects are focussed in this case on the fact that the
food requirement for various species and various age groups of
the saine species varies and depends on the features of the food
organisms, the water temperature and a series of other factors
(6, 7, 39).
...18
. 18
e
According to G.G. Winberg (6), the food consumed by fiàh
is used up on the essential metabolism and growth following the
formula:
P = 1,25 (g4-T )
where P is the ration for a determined period
If is the increment in weight (g)
T is the loss for essential metaboliem
,
The expense of nutritious substances for metabolism is the
same for all species and all age groups within the species and
can be expressed, according to G.G. Vinberg (6) and V.S. Ivlev
(35) by the formula:
r, 0.3 W°
.8
where W is the weight of the fish.
,
Thus, in order that the fiàh does not perish from hunger,
the food dhould ensure it of a supply of energy at least equal .1(1
Q.
Anything that the fiàh consumes in excess of this quantity
is used for growth. Consequently the rate of grawth depends
on the quantity of food. However this relationship is valid
up to a known limit. G.S. Karzinkin (39) established e.g. that
when carp is fed in excess with Chironomus , the latter passes
through the intestinal tract of carp completely undigested end
remains alive.
Translator ' s note:
*Bloodworm
•••19
• 19 .
V.S. Ivlev (36) proposed a method and worked out a model of
estimating the food availability for fish feeding on plankton,
gâti
4.4-t from the balance n of input and output" of energy ob-
tained from the food. The estimation of the food availability
is based on the functional relation between the density of
food, the daily ration, expenditure on metabolism and the
growth of the fish.
Low concentrations of food, in the opinion
of V.S. Ivlev lead to a reduction of rations, starvation and
perishing of organisms.
One of the most accurate methods of
determining the food requirements by fish is the investigation
of nitrcemetabolism 001).
In pond fish establishments and fisheries the problem of
an optimum ration and desired rate of growth of fish is the
subject of profound and thorough considerations; the density
of stocking with fish, the quantity of food and fertilization
of the ponds are determined.
There are very few data, which would make it possible to
draw conclusions on the food requirements of marine commercial
fish. The problem of standards concerning normal feeding and
growth of sea fish has been worked upon to a very small extent.
Usually the research workers, without mentioning the food requirements of the fish, embark directly on the problem of good
or bed- food availability in concrete conditions (in this or
some other year, in this or some other water body, or part of
it).
Although standards characterizing normal food availability
are not cited, they are actually implied. Each investigator
...20
+di
20 •
has a general idea what should be this or some other population
In normal or rather in optimum conditions, i.e. when all its
requirements are satisfied, in thsgl4mulbes
ments.
food require-
The researcher compares the data obtained from actual
material (although this is not mentioned directly) with indices
characteristic in his opinion tle the optimum state of the
population.
When e.g. the author wishes to give an estimate of food
availability in various years, he can do that by only establishing (even if for himself) what indices within the range of the
order considered, express a good, medium or poor food
availability.
Only in recent time works began to be publishede eving
directly quantitative indices,which can be used for frcharacterm* ,
illtir-dee
the requiraments or the degree of satisfying the food
requirements of sea fish itexxiteme in natural conditions.
'Mils,
R.M. Pavlovskaya (61) has shown that in the North-Western part
of the Black Sea, with a plankton density of less than 1,000
specimens per m
empty intestinal tracts were found in fry of
anchovy; when the density was more than 5-6,000 specimens per
m[3 the quantity of food rose sharply and finally with 10-14,000
specimens per m3 it reached its maximum. A further increase of
the density of plankton (up to 28,000 specimens per m 3 did not
)
change the conditions.
According to K.I. Vonokova (10) in the
. • •21
,t.
• 21 •
water bodies of the Volga river delta the maximum intensity
of feeding by the fry of n vobla e is observed at a concentration
of food organisms above 200-250,000 specimens per m3 ; of bream
fry at a concentration above 225-275,000 specimens per m3 ; of
perch-pike fry at 25-35,000 specimens per m 3 . i.i. Nikolaev
(54) assumes that at a density of Limnocallianus of less than
50-70,000 specimens per m3 the Baltic Sea herring stops -tme feed1 ,.1
on it and changes to the consumption of Mysidaeand Amphipoda.
As direct indices of the requirements by fish, on the
basis of which is construed a judgment of food availability
for them in natural conditions, can be considered the quantity
of nutritive substances necessary for fish, according to the
formula by G.G. Vinberg, at the maximum of growtb ald also the
ration and indices established in aquarium conditions with
excessive feeding. Thus, the quality of bream fingerlings
3-4 cm long in the Taganrog Bay in 1955 and in the Don river
in 1951 and 1953 was estimated by us on the basis of the values
of indices of fullness of the intestinal tracts. The latter
were compared with indices of fullness of intestinal tracts of
fingerlings in the ponds of the Aksai experimental base, for
which it has been established experimentally than an index
70-150%er,
characterizes satisfactory conditions of foraging (23).
Usually, however, to make a judgment of the food avail14,41
ability for fish it is necessary to teeke recourse to indirect
4 "voblail Rutilus rutilus caspicus in this case.
• 22 .
-SMs
indices determining these requirements and on thoir basis to
find the standards of good and poor food availability.
When judging the food availability by the character of
feeding, the character of feeding of the population in good
conditions of foraging is used as standard.
When comparable
material from the sanie water body is not availabl; data from
other water bodies are used.
Thus, in order to solve the question if the "rechneya"
flounder in the Baltic Sea was eufficiently provided with food
in 1948 and 1949, the character of its feeding was compared
with the character of feeding of the "rechnaya" flounder from
the Barents Sea (20).
To solve the question of the degree of
food availability for the roach, >` in the Azov Sea in 1950, the
character of its feeding was compared with the character of
feeding of the "vobla
* of the Northern Caspian Sea in 1 935$
when the state of the "vobla" population (abundance and rate
of growth) was fairly satisfactory (22).
The starting position in determination of food availability
for fish by the character of their feeding is the aspect of the
typical food of the fish (18, 65).
As typical is understood
the food ensuring the existence of the species or subspecies
In water reservoirs of a determined type with a typical composition of the nutrition base and the fish fauna.
Translator's notes:
* rechnaya kambala
Pleuronectes flesus trachurus
Rutilus rutilus heckeli
"* Rutilus rutilus caspicus
...2 3
• 23 •
The typical aharacter of feeding is determined, going out
from the anatomical and physiological peculiarities or the'fish
and the composition of the fish population in various conditions
of existence. The typical character of feeding by the fl yable
was e.g. established on the basis of the character of feeding,
rate of growth.and . abundance of representatives of the species
Rutilus rutilus in water bodies of various types.
It was accepted
as one of the standards of godd condition of the roach population when roach attained a length of 21 am in five years (18).
Knowing the mbde of feeding typical for a given species
and taking into account deviations from it in some concrete
conditions it'is possible to presume a probable rate of growth
of the fish and their abundance in a certain body of water..
However euah an assumption cannot serve yet as a final conclusion, because data are scarce in this domain;
they are now
only in the stage of being collected. When investigating a
series of.problems connected with estimating the magnitude of
possible catches of fish and acclimatization'wnrk, it is
often necessary to establish the food reeirement of fish inhabiting the body of water.
It is determined on the basis of
yearly rations, or feed coefficients of the stock or catches of
om
fish and of data ef their distribution.
The actual rations are known for a small number of fish
and usually it is necessary to make use of calculated rations.
24
A.A. Szorygin (76), studying the feeding and food relationships of fish, eaeteeed on the basis of. experimental data icï a
number of research workers, that the yearly rationper hm:GAF-ef
fish coneuming molluskS exceeds 23 times the weight of the
fish; the ration of fiàh consuming worms, crustaceans and fish
exceeds its own weight 9 times.
The ration of fidh with a mixed
mode of feeding, is determined by the relation of the first and
second components. Going out from the food requirement of the
fiàhes and the biomass of food'organisms, A.A. Shorygin determined the intensity of the competition for food and presented
a conclusion as to the intensity of this competition. Comparing the biomass of fish and their food, he was able to Make
a judgement of food availability for fish in the Caspian Sea.
An analysis'of the rate of growth, - condition, feeding . and
nutrition base of Acepenseridae of various seas made it possible for A.A. Shoryginto form a conclusion as to a ccmparatively
poor food availability for the sturgeons of the Caspian Sea.
This conclusion confirmed the expediency of acclimatization
of Nereis'and Corbulale proposed by L.A. Zenkevidh and Ya.A.
eirshtein (32).
•
In 1951 the quantity of food and.different food organisms
necessary for the fish of the Azov Sea was establidhed according
to their yearly ration (22, 25).
On the basis of production of
Translatorrs note:
"Korbulomina" in Russian. - Not found in dictionaries. Assume
that molluskà of the corbula species are meant here.
...25
• 25
•
benthonic organisms (68) the coefficient of exploitation of food
was calculated. It was found that in 1951 this coefficient was
equal .0.27, i.e. was lower than in 1935, when it was O.L. (11).
When determining the food requirements of fish of the Azov
Sea,E.A. Yablonskaya (79) made use of food coefficients O (for
• plankton-eaters) and 30 (for bottomrfeeding fish). Ge-ing-out
âikeut ›,
' 1 and bOttom organisms and the feeding
from plankton 1.;t."eïzà,44coefficients of fish,E.A. Yablonskaya obtained the probable fish
production. Comparing the probable and the actual production
s..he showed that during the period 1950-51 the fieh of the Azov
Sea consumed plankton very intensively, the exploitation of
benthos being considerably lower.
To determine the quantity of food required by the Salmonidae
of the North-West part of the Pacific Ocean,B.M. Mednikov
applied the feeme-le-ce-th.e balanceàequation introduced by G.G.
Vinberg (6) i.e. P = 1.25
consume less than
5%
T). It was found that Salmonidae
of the whole yeal” productiàn of zoo-
plankton.
B.M. Mednikov came to the conclusion that irrational fishveset e rve
ing of the Salmonidae may cause a ealekemeef the nutrition base
LitU
and development of fish of little valuee competitors of the
-
Salmonidae.
The problem of the food'requiraments of sea fish is of
extraordinary importance:
it presents the possibility -tT-o makEil , c1
a correct judgment of the food availability for them. Nevertheless,
...26
,
26
é
this damain requires further experimental investigations, because
the data obtained in this area are very scarce.
Some indices of food availability in an example of bottom feeding
Trsh of the Azov Sea
When studying the feeding and the exploitation of the nutrition
base by bottam-feeding fish of the Azov Sea, it was necessary
to analyse in detail the problem of food availability for them.
This was caused by the fact, that the conclusion by E.A. Yablonskaya
(79) concerning poor exploitation of the benthos of the Azov Sea
by bottom feeding fish in 1950-1951 contradicted the indices of
poor food availability for bream.
In 1950-1951 the index of
intestinetract fullness of bream, particularly of older age
groups was found to be considerably lower than in 1935 , when
feeding of bream was studied by V.P. Vorobev. Also Une importance
of mollusks in the food of bream increased. This supplied a
basis for the assumption that the conditions of fattening by
bream in 1950-1951 were poor. Later this assumption was confirmed by a decrease of the benthos biomass on the Zhelezinsk
bank, where bream of older age groups usually fattens, and by
a lowering of the average weight of bream (22, 25). It transpired that in spite of the good food availability for bottom
feeding fish in 1950-1951, the food availability for bream, at
that time the most important of the benthophagi anong the commercial fish, was low.
...27
• 27 .
In order to determine the food availability for bream and
other bottom-feeding fish, the difference was found between the
quantity of food required by fish at normal or speaking more
accurately, good conditions, and the quantity of food consumed
by fish in the conditions of 1950-1951.
The quantity of food required by fish in normal conditions
was determined on the basis of yearly rations calculated according to A.A. Shorygin (76).
The consumed food was computed with
the help of indices of the fullness of intestinal tracts. In
the application to bream, the indices were used with a correction
in relation to the food required;
yeCevt,re
In-elerIi.eate to other fish
In the form of initial values.
et
' For bream of . various age groups the absolute value ofp food
deficit.was determined, Which in 1951, totally for the WhIole
bream.population, proved to*. equal 895,500 centners.
These
(44u_e_e(1
data were deduete4=by calculating the deficit of the biomass of
tohe
the result of ,& reductionAm 195i)of its rate
bream (25
of growth (in comparison with 1935).
The deficit of the biomass
%-nof
o
of bream in 1951 amounted to 60,000 centners. The qie.e41-epet
ketie
e4 _tbe-445414,4tee of the food deficit ever the biomass deficit
to
of bream was equal 14.9. The commensurability of this value
A
•
with the food coefficients determined for other fish confirmed
tle
the reasonableness of the value of food deficit. Detailed methods
of computation were presented in special papers (22, 25) therete
fore here ate- resUlt oallete' is shown.
...28
. 28 .
Data for obtaining absolute values of food availability
for àther fish could not be obtained; however, an analysis of
the composition of their food gave some notion g the relative
degree of this availability.
In Table .l is shown the exploitation of the nutrition base
by different species of fish according to individual requirements
and yearly rations.
Below are presented the Initial data, the yearly rations
not included in the table and catches of fish in 1951.
•
•
Catch
thousands of
centners
Bream
' '
Azov roach
Vimba vimba .
Acipenseridae
Gobildae
Yearly
rations*
77.0
5.0
15
23
16.
13.9
16
194.1
21
80.7
j( The yearly ration is-the:quantity of food ransomed by a fiàh
.during the year. It represents the ratio of food consumed by
the fie to the weight of the fish.
Working out individual indices for catches of fie produces
weighed individual indices, conventional units characterizing
an individual requirement of food by fish i.e. the quantity of
some or other organisms preàent at some defined moment in the
.
stamaChs of all the fish caught. BY thiS computation, bream
had consumed 5.1% of all the food consumed by the bottom feeding
fiàh, the Azov roach 11,4%, the gobies 79.1%
•••29
29
•
A computation of the exploitation of the nutrition base
on the basis of yearly rations produced a different relationship: at normal, or rather optimum conditions, bream should
have consumed 15.6% of all the food, and the gobies 55.8%.
A
comparison of the value of exploitation of food organisms by
indices and by yearly rations shows that in the conditions of
1951,bream and Azov roach made less intensive use of the nutrition base, and the gobies fed more.intensivelyethan eke>.
would be the case in normal conditions.
B"rom the divergence between the intensities
.04f
exploitation
of .the nutrition base by individual species of fish computed
from indices and yearly rations, a preliminary conclusion was
drawn that in 1951 the food aVallability for gobies waà
relatively higher than for bream.
.
As has been said before', in 1951 the food availability for
bream, particularly for older age groups was low, Which reflected
in the large deficit of food end reduction of tle average weight
of bream. Completely different was the situation as far as gobies
are concerned. ,The conditions of foraging for gobies, particularly
Neogobius melastomus (43), were good in 1951: the index of
stomach fullness was high, the round goby grew well and was well
fattened.
A conclusion can also be drawn from the table that in 1951•
availability for sturgeons was good.
The Azov roach
.had a somewhat worse food availability than the Acinpenseridae .,
but still better tàan bream.
...30
thefod
• 30 •
This conclusion finds se confirmation also in other indices.
V.A. Kostyuchenko (42) noted, that fattening of sturgeon increased
in 1952, apparently because of an Intensified consumption by
it of Corbulae (in 1951).
The conditions of foraging for the
Azov roach in 1951 were also fairly advantageous, although
Worse than in 1950 (22).
* See note on page 24.
...31
TABLE
EXPLOITATiON OF THE NUTRITION BASE IN THE AM 5EA BY _BOTTOM FEEDING ,FISH
Lcno.11,30Baline 6eHroco5ummmil pbalamil KopmoBon 61131,1
cingle
Préquener indiees ,
Food
opra:1113MEI
organism's
4
U•
=
d
0.52, .ot
d
O.
0
N . to
I*
e--Cardium
Monsds.kne
gydrobla
3,6
9,2 .
6,3
0,3 61,6
Fn2poümq
Sindtsmia
'too
,11
••Az.,
oonvIentional units
m
a leg
o':-
d
s.
!..4 0 Zet
te
.0
=0
rf:
w_ 01
-.0,
,E.c ;
o,
.4'p-I -.., _a'
,..7: Xi ri :::.:, 77,14 i
- -,
34,6 290,5 708,4 9,0
2,2
.r..0
-4 '11. =el
.7:0 ;o :=,.d
C.
-,.: fen
•,'Q
14,8
--
--
1,8
--
0,5
19,9
508,4 --
6712,4 7750,9; 3,7
30,6
--
74,0
582,11 87,4
.97,0 4904,2' 0,5
24,2 4 774,0 9,0
: ra
•j
,.., . 5..
«.
ee
7e,
1> .
0, 1
9,1
.4- -1.:
-a-
0,4
12,6 1 --
--
97,4 ; 0,1
--
0,2
-
0,3 41,3
66,9
145,3 261,8: 1,5
569,9 12978,6 13957,1 1,0
1,9
001 1 4,1
2,2
3,6
14,8
80,7 15,4 11,0
50,0 2871,2 3028,3 2,6
0,5
--
18,5
0,2
32,2 --
--
2,0
--
--
161,4 .--
:-
gammari da e
. I aMMaplubi
0,01 0,001
5,8
0,82
0,8
0,1 --
80,6
I\OpOcplllUibl
0,07 0,06
2,5
--
0,07
5,6
4,6 . 12,5
--
13,6;
0,02'
0,3
0,03
0,02 72,6
0,4
3,9
1,7
8,5
KopCiynomiisi
Norois
Hep eue
Nophtbys
neTTnc
Hipania
--
1,8
3,4
1,0
0,2
0,4
rilrra rim
elm,91,e.
0,4
•
Ostrasoda
0,9
OCTIl 'ali02b1
12,5
Piii6a ?tell
0,2
Sce opra1-1133MEI
All • rganis a s
0,7
33,3 77,0
1,5
1,5
0,041 968,4 130,9 42,5
29,05 13,0
16,0
--
38,8
264;1
1
1,6
0;4
--
335,1 9,5
1 79,0
•-
.
0,3
3,5,3 15,6
79,9. 91,0
1
7,8 1149,6 84,2
3,5, . 403,1 2916,0 3332,6
35,6 123,2 213,2 2663,1 5929,0 183:0 2112,8 41322,0 52509,9
-
-o
d
-• 0.-zo
t•
=11.
.0 <
12,8 t 34,6
1,8 ;
11,3 ,
5,1 ! 11,4
1,8
87,4 -- 12,6 -2,0 0,898.2 0,3 '--
5.6
-0.7
2,3 0 4,0
92,99; 4,4 5,0 0,01, 4,8o 8 5,7
!
'
94,9 10,4 1,6 1,4 1, 9 8 1.7
65,4
11,5 26,2 .
8 ,7
_
38,3
-
11,4
--
37.0 22,913,352,4
0,5
4,9 9 5,2 1,5 1,8
0.3 . 1,2
0,7 . 88,8 7,3 3,7
0,2
3,8 '
; . 0,1
7er4
0
33,4 65,27 1,9 0,1 --
0,03
0,5
o
_d
- .0 =0--••••• 0.z
86,7 12,621,6 0,4 ' 0,1
:100
-- 100,0
159,1 . 210,6'
•
1,8 . 97,3 3,1 0,6, --
278,0 14472:4 14886,1' 0,7
1,3
yearly rations
a
0 .4
se .d •o".z:
..d
.' .zii
' .:
74,6 ; 104,9 30,8 --
Corbulomia
7:
*
O
1,8
1
a 5 :ill
ces Fee4yelluir,e,aent,1;iy
1
I;(-roma, ye.i.
Kopmonie
IN 1951
!Food
ime,teMl-ec.! ,rood
mu.e,cm.
luot r.
A3OBCK 0r0 M 0p2
T
12,1
0,3 • 4,1
I
Transliterated. The corresponding English or Latin names could not b. located
87,3 :2,4 --
0,4' 11,9
79,1 15,624,3 1,1
s2.3
3:2 55.8
32
of
A comparison of the indices/fullness of the intestines
101
of Azov roaéh in 1950-1951 and the "vobla tt*. of the Caspian
Sea in 1935 when the condition of the n vobla p -population was •
good, served as the basis of such an estimate.
Thus, the conclusion as to the.availability of food for
fish, which was made on the basis of the difference in the
intensity of consumption of the nutrition base by various
fish species determined from indices and yearly rations, was
growth
confirmed by the rate of/fteur and a comparison estimate of
feeding conditions (Table 2).
Table 2
Index of food availability for fish in 1951
Exploitation of Exploitation of. Index of Condition of
population
food
food according
food according
availto rations
to indices
Fish
Bream
Azov roach
Vimba vimba
Acipenseridae
Gobiidae
5.1
11.4
0.3
4.1
79.1
15.6
24.3
1.1
3.2
55.8
0.33
0.47
0.27
1.3
1.4
poor
medium
lack of data
good
good
Note: The ratio of food exploitation according to indices
to the food exploitation according to yearly rations
is considered as the index of food availability for
various species of fish.
Good food availability is dharacterized by indices higher
than one, poor by indices lower than one.
As can be seen from the table, the food availability is in
conformiti with the condition of the fish population.
'1"
Rutilus rutilus caspicus
...33
I
'
•
• 33 .
Table _3
Indices of food availability for breaxn and percarina
Locality and time Daily ration in % of
of observation .
weipht of the fish
Species
Index of
food availability
In natural in aquarium
conditions or computed
.Pream
young
adult
'Pond of Aksai base
1June 1956
eaganrog Bay
(June 1956
Percarina. Taganrog Bay
(June 1956
Author
19.50
38.4
0.51
5.00
9.0
0.55
7.04
6.5
1.03
M.B.
Zeltenkova
t
I.P.
Kanaeva(37)
Note: the daily ration for'young bream was determined in
the aquarium, for adult bream and percarina it was
computed by the method of A.A. Shorygin on the
basis of the yearly ration and the seasonal course
•
of feeding.
The ratio of the value of the daily ration obtained in natural
conditions (59, 60) to the ration determined in aquarium conditions, or computed by the method of A.A. Shorygin (76) can
also serve as an index of food availability for fish with regard
to the degree of their foraging.
As can be seen from table 3, the index of food availability
of young immature bream in the pond of the Aksai fishery station
and adult bream in the Taganrog Bay was low, the index of food
availability for percarina was high.
This is oonfirmed also by the condition of the fish population.
In 1953 bream was growing better, than in 1951, but
worse than in 1935.
The condition of the percarina population
(according to I.P. Kanaeva) was satisfactory.
...34
102‘.
. 34 •
Influence of food availability on abundence, rate of_prowth
end catches of fish
It was mentioned in the opening paper (15) that "limitation
of food availability for the population causes a reduction of
the abundance predominantly in the early stages of development.
In youngimmature and in adult fishes insufficiency of food may
ceuseA strain in relationships within the species and have an
influence on the growth of the fish".
Numerous exemples of this are presented in a series of
papers including works of the author (24, 27).
It seems now to be expedient to dwell once again on some
investigations which are the most characteristic in this aspect.
The influence of food availability on the abundance of
anchovy was studied in detail.
T.F. Dementeva (14) observed ttel
the survival of fry of Azov anchovy -tgi dependLee on various
As indices of food availability , were used the cmposition of the contents of the intestinal tract of anchovy
factors.
fry, indices of fullness, the rate of growth of the fry and
the character of their nutrition base. The euthor maintains
that the anchovy fry possess a high adaptability to food and
that in 1952 and 1953 their food availability was satisfactory
and the abundance of the recruitment was determined mainly by
the survival of embryos ledependge on oxygen supply and
temperature conditions. But in 1954-1955 the food availability
for young Immature anchovy deteriorated as the result of a
.. 35
.
,1
•
35 •
sharp decline of productivity in the Azov Sea. This brought
erNelm(ect/
about the re•ri-sh-tne of a considerable part of the fry and a
sharp diminleh;Ing of the total stock of anchovy.
T.V. Dekhnik (17) conducted in 1957-1958 detailed investigations of the causes of mortality of the Black Sea anchovy in
early stages of development and arrived at the conclusion, that
the largest mortality is observed during the embryonal period
•
and during the period of feeding on yolk.
The change to active
feeding does not cause any increase in mortality, since there
is no lack of food in the sea. P.I. Grudinin (13) studied the
survival of anchovy fry in the Azov Sea in 1958-1959. He judged
the food availability by the magnitude of the nutrition base
(the quantity of the nauplius and other stages of development
of Copepoda) and from the quantity of feeding anchovy fry.
P.I. Grudinin came to the conclusion that the basic factor
determining the abundance of young anchovy is the condition
of the nutrition base during the period of spawning.
Thus, one can see, en the example of anchovy fry,that the
',nay 1)e
in
determining the abundance of anchovy ee-, different
factors
different years. Nevertheless, food availability for the fry
appears to be the decisive factor in a number of cases.
The food availability has an influence on the rate of
growth of adult fish and consequently on their catches. In*
sufficient food availability for the n vobla n of the Northern
Caspia.n roach
00.36
-
e
4
• 36 •
.
Caspian Sea and for bream in the Azov Sea (25, 27) caused by
a decrease of the nutrition base and reduction of the area
occupied by the fish themselves in connection with hydrological
conditions brought about in some cases a decrease in catches
amounting to 25-35%.
Food availability influences not only the character of
the growth of individual generations, but also the(species)
Cate_
T■N
of the flounder
Qm-the-**10;*1e
of
their
growth.
peculiarities
in the Baltic Sealcan be shown very easilyithe connection between
the peculiarities of its growth in the process of ontogenesis
and the food availability for individuals of various age groups,,
The study of feeding by the flounder has shown (20) that
the composition of its food Changes with growth.; small flounder
feeds on crustageans e -and when it reaches the length of 22 am,
it .changes to consuMing màllusks.
...37
m
4
• 37.
ill
e
C
p
,'?, g
lu
•
•
----''''-•\
-',.
,
.,'''./'
E
cl,
r,
a
7,,.
/
/
5.
\ .„ 2
N, N7----,. A
•
<"--; o
200 tz
Y.:4..
:. •
10H- 5 , I,
'`
6
/
c
A
o•
27
1 2.
Qnuna
27
32
KUH(IUJlbI. CN
of flounder
en.
Indices of food availability of the Baltic (rechnaya
flounder in the Baltic Sea in 1948-1949.
1 - general index of fullness of the intestinal tract of flounder
in the region Ventspils-Liepaya 1948;
2 - quantity of components in the food of flounder in the region
Ventspils-Liepaya 1948;
3 - General index of fullness of the intestinal tract of flounder
in the region of Klaipeda 1949;
4 - the same with correction for protein;
5 - quantity of components in the food of the flounder in the
region of Klaipeda 1949.
Fig. 1
,
The food of large (more than 25 cm) flounder consists of
mollusks almost exclusively.
The character of feeding by the
flounder, its competitive relationship in various age groups
with other representatives of the fish fauna of the Baltic Sea,
and an analysis of food availability for flounders of various
sizes, made possible the conclusion that the flounder 20-2)1 cm
long is worst provided with food. In Fig. 1 is shown the
characteristic of food availability for flounder according to
three indices:
indices of fullness of the intestinal tract,
the calori value of the food, and the quantity of components.
1,11-réchnaya Kambala" - Pleuronectes flesus
...38
• 38 •
All indices confirm the poor food availability for flounder 202 cm long.
(.2 ')
Fig. 2
147(
years of gr trot h
Curve of the rate of growth of flounder from the middle
part of the Baltic pea;
(Fig. 1 from the paper by K.A. Zemskaya (31)).
1 - generation 1952; 2 - generation 1953.
This is reflected in a decrease of the values of indices of stomach
fullness and the calorie value of the food lump and an increase
of the components of food. The flounder up to 20 cm long and
longer than 24 cm was found to be better provided with food.
This served as basis for the assumption that the rate of
growth of the flounder longer than 25 am is faster than the rate
of growth of the flounder 20-24 cm long, which was confirmed by
investigations by K.A. Zemskaya (31).
A study by the method of
regression analysis of the rate of growth of flounders of the
generations 1952 and 1953 has shown that the largest linear
increment of flounder is observed in its second year, than the
increment is reduced, to become larger again at the age of
5
years (Fig. 2.).
—.39
•
•
• 39 •
An analysis of the increments of various age groups with
regard to their average dimensions conducted by K.A. Zemskaya
for several years on large quantities of material confirmed this
regularity.
The author mentions that the flounder reaches a
length of 20-22 am in four years and that the characteristic
curve of growth of the flounder can be explained by its change
to feeding on mollusks, after having reached a length of 22-24 cm.
In Fig. 3 are compared the indices of fullness of the
intestinal tract of flounder in 1948, which can serve as indications of food availability for its various size groups and the
linear increments of the fiah in
paper by K.A. Zemskaya.
194e, taken from Fig. 2 of the
As can be seen from Fig. 3,the curve
of growth of various age groups of flounder in 1948 reflects
exactly the curve of indices of fullness of the intestinal tract,
i.e. the food availability for flounder.
In connection with the peculiarities of feeding by flounders
of various age groups and the eundance of mollusks in the
middle Baltic Sea it was proposed (20) to increase the commercial
size of flounder, since the established size removed from the
population individuals before their change to feeding on mollusks.
...40
I.
• 14.0
3110( 6r
z
-•
Mill 4;
°
•
1-1
ItiP
I
2;
,
:II
\
E
•
- o
12
1%
22
27
.fm,,,,,a02, 4 cm
,
e
Length of flounder on
Fig. 3.
11 The
Relationship between food availability and linear
increments of the Baltic flounder in the region of
Ventspils in 1948:
1 - linear increments of flounder (data by K.A. Zemskaya)
2 - indices of fullness of the intestinal tracts of
flounder.
methods of running a fish farm and, in first place, of
fishing" should ensure "the possible maximum of exploitation by
commercial fish of the nutrition resources of the body of water.
(1, 15) - in this manner can be formulated one of the essential
tasks of the science of fisheries.
An analysis of the fish
population, their conditions mad their foraging makes it possible to draw conclusions as to food availability for fish and
this consequently permitvot only to understand the reasons of
variations of the abundance of fish, but also to arrive at a
solution of the problem of rational exploitation of the natural
resources of the water reservoir.
Conclusions
Food.evailability for fish serves as the starting point for
determination of natural mortality, rational fishing, strains
...41
11
à
le
• 11--1— •
in relationships within the species and between them, rational
composition of the commercial fish fauna by age and species,'
acclimatization work etc. In forecasting the survival of young
Immature fish, the abundance of generations, the rate of growth
and possible catches, determination of food availability for
fish appears as one of the essential components of the work.
In the solution of complex problems,like determination of
natural mortality, estimation of stocks and the manner of their
exploitation etc., it is necessary to achieve an absolutely
reliable conclusion as to the magnitude of food availability
-tt'e
for fish. It should be based on all four indices ; food
A
requirements, nutrition base, mode of feeding and the condition
of the fish population, including the indices of the amplitude
of population changeability.
In arder to forecast the magnitude
of stocks, it is first of all necessary to know the changes in
the nutrition base of the fry and the mode of their feeling.
Then, after the connections between the nutrition base of adult
fish, its exploitation and the yield of fish production (abundance,
rate of growth, catches) are determined, it is possible to make
prognoses on the conditions of foraging (nutrition base and feeding)
the conditions of fish in the population with regard to quality,
and their importance in the formation of coinaercial stock.
The problem of food availability for fish became particularly
crucial in recent times, not only because of variations in the
•••42
lO
■
11
. Li2
p.
stocks of fish, but also in connection with the necessity of
deciding to what extent in each separate case the natural
(cosmic, oceanographic and biological) and sociological factors
influence the fish productivity in bodies of water.
Only a
profound understanding of the causes determining in concrete
cases
a certain level of fish stock and consequently also of
catches, will make it possible to work out rational bases of
the economy of fisheries.
\\\--._ ...43
• 43
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•
•
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...45
■
• 45.
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o
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•
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‘.1
•
'
• 48 •
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•••5
1;
.
50
.
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...51
• 51.
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Lebedev N.V.
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v severo-zapadnoi chasti Chernogo morya". "Rybnoe
khozyaistvo' No. 9, 1936
("Method of finding concentrations of Acipenseridae
in the North-West part of the Black Sea". 'Fisheries"
No. 9 1936)
48. Lebedev N.V.
"0 vozmozhnosti opredeleniya stepeni ustoichivosti
promyslovykh skoplenii vobly v period ee otkorma"
Vestnik MGU (Moskovskoi Gusudarstvennyi Universitet)
No. 2 1950.
("On the possibility of determination of the degree of
stability of roach accumulations during the period of
its fattening". Moscow State University Herald No. 2
1950).
49. Manteifel B.P.
"Plankton i sold v Barentsevom more". Trudy PINRO.
Tom VII. Moskva-Leningrad Pishchepromizdat 1941.
("Plankton and herring in the Barents Sea". Transactions of PINRO (see No. 28) Vol. VII MoscowLeningrad. Food Industry Publications 19)4.)
Manteifel B.P.
"Adaptivnoe znachenie periodicheskikh migratsii vodnykh
organizmov" Voprosy ikhtiologii. Vypusk 13 1959.
("The adaptivity aspect of periodical migrations or
organisms dwelling in water". Problems of Ichthyology.
Issue 13, 1959)
51.
Marti Yu.Yu.
"Migratsia i problema obespechennosti pishchei morskikh
ryb". Trudy soveshchaniya po dinamike chislennosti ryb.
Moskva. Tzdate1stvo AN SSSR 1961
("Migration and the problem of food availability for
sea fish". Transactions of the conference on the dynamics
of fish abundance. Moscow. Publications of the Academy
of Science USSR, 1961.)
...52
e
11).
• 52 •
52.
Monastyrskii G.N.
"Dinamika chislennosti promyslovykh ryb" Trudy VNIRO.
Tom XXI Moskva 1952.
("Dynamics of abundance of commercial fish" Transactions "VNIRO" (see No. 8) vol. xxI, moscow 1952.)
53. Neiman A.A.
"Nekotorye zakonomernosti kolichestvennogo raspredeleniya
bentosa v Beringovom more". "Okeanologiya" Tom 1.
Vypusk 2, 1961
("àome regularities of quantitative distribution of
benthos in the Bering Sea" "Oceanology" Vol. 1
Issue 2, 1961.
5 )4 .
Nikolaev
"Plankton kak faktor raspredelenia I produktivnosti
planktonoyadnykh ryb Baltiiskogo morya". Trudy
Soveshchaniya po dinamike chislennosti ryb. Moskva.
Izdatelstvo AN SSSR. 1961.
("Plankton as a factor of distribution and productivity
of plankton eating fish of the Baltic Sea". Transactions of the Conference on dynamics of fish abundance.
Moscow. Publications of the Academy of Science USSR.1961.
55.
Nikolskii G.V.
"0 zakonomernostyakh vnutrividovykh pishchevykh otnoshenii
u presnovodnykh ryb". Byulleten MOIP. Otdelenie
biologii. Tom IV, Vypusk 1. Moskva 1 9 )49
("On regularities of feed relationships within a species
with regard to freshwater fish" Bulletin MOIP (see No. 2)
Biological Division. Vol. IV.
56, Nikolskii G.V.
"0 zakonomernostyakh pishchevykh otnoshenii u presnovodnykh
ryb" Ocherki po obshchim voprosam ikhtiologii. Moskva.
Izdatelstvo AN SSSR 1953.
("On regularities of feed relationships of freshwater
fish". Sketches on general problems of ichthyology.
Moscow. Publications of the Academy of Science USSR.1953.)
57.
Nikolskii
"0 teoreticheskich osnovakh rabot po dinamike chislennosti
ryb". Trudy. Konferentspii po voprosam rybnogo khozyaistva.
Moskva. Izdatelstvo AN SSSR 1953
("On the theoretical bases of work on the dynamics of
fish abundance". Transactions of the Conference on
problems of fisheries. Moscow. Publications of the
Academy of Science USSR 1953.
...53
Dc
• 53
58. Nikolskii G.V. i Pikuleva V.A.
"0 prisposobitelnom znachenii amplitudy izmenchivosti
vidovykh priznakov i svoistv organismov"."Zool. Zhurnal"
Tom XXXVII. Vypusk 7, 195 8
"On the adaptivity aspect of the changeability of the
features of a species and characteristics of.organisms".
"Zool. Journal" Vol. XXXVII. Issue 7. 1958.
59. Novikova N.S.
"Opredelenie sutochnogo ratsiona voble Severnogo
Kaspiya neposredstvenno v more" Vestnik Moskovskogo
Gosudarstvennogo Universiteta No. 5, 1951
("Determination of the daily ration of roachl` in the
North Caspian Sea directly at sea". Moscow State
University Herald No. 5 1951
60. Novikova N.S.
povedenie vobly Severnogo Kaspia na mestakh otkorma"
"Voprosy ikhtiologii" vypusk 7,1956
("Behaviour of roach of the North Caspian Sea in
places of fattening". Problems of Ichthyology, issue
7, 1956.
61. Pavlovskaya R.M.
"Nekotorye voprosy biologii razmnozheniya i razvitia
dhernomorskol khamsy v sviazi s problemoi dynamiki
chislennosti". Trudy AzcherNIRO, vypusk 17, Kerch
1958.
("Some problems of the biology of reproduction and
development of the Black Sea anchovy in connection with
the problem of the dynamics of abundance"
Transactions AzcherNIRO (see No.11)issue 17, Kerch 1958
62. Polyakov
I O prisposobitelnom znachenii izmenchivosti vesa
segoletkov karpa" Zool. zhurnal. Vypusk 3. 1958
("On the adaptation aspect of the changeability of
weight by carp underyearlings" Zool. Journal. Issue 3,
1958
63. Polyakov
"Prisposobitelnoe znachenie izmenchivosti priznakov i
svoistv populatsii ryb" Trudy Soveshchaniya po dinemike
chislennosti ryb. Moskva. Izdatelstvo AN SSSR 1961.
("The adaptation aspect of the changeability of features
and characteristics of the fish population". Transactions of the Conference on the dynamics of fish
abundance. Moscow. Publications of the Academy of
Science USSR 1961.
Translator's note:
* Rutilus rutilus caspicus
...54
.5 4.
64. Rikker
"Kratkaya istoria razvitiya teorii dinamiki chislennosti
populatsii ryb v stranakh Zapadnol Evropy I Severnoi
Ameriki." Trudy Soveshchaniya po dinamike chislennosti
ryb. Moskva, Izdatelstvo AN SSSR 1961
"Short history of development of the theory of dynamics
. of abundance of the fish population in the countries
of Western Europe end North America" Transactions of
. the Conference on the dynamics of fish abundance.
Moscow Publications of the Academy of Science USSR 1961
65.
"Rukovodstvo po izucheniyu pitania ryb v estestvennykh
uslovyakh". Moskva, Izdatelstvo AN SSSR 1961.
("Textbook for the study of feeding by fish in naturalconditions" Moscow, Publications by the Academy of Science
USSR 1961.
66. Saenkova A.K.
"Sezonnaya dinamika bentosa Severnogo.Kaspiya" Trudy
KaspNIRO. Tom 15. Moskva 1959.
"Seasonal dynamics of the benthos in the North Caspian
Sea" Transactions of KaspNIRO (see No. 10) Vol. 15
Moscow 1959.
67 , Sergeeva A.I.
Trudy
"Ob izmelchanii severokaspiiskoi vobly"
Soveschaniya po dinamike chislennosti ryb. Moskva,
Izdatelstvo AN SSSR, 1961
("On the degeneration of the North Caspian roach"
Transactions of the Conference on the dynamics of fish
abundance. Moscow. Publications of the Academy of
Science USSR, 1961.)
68. Stark I.N.
"Godovaya i sezonnaya dinamika bentosa v Azovskom
more" Trudy AzNIIRKh e''' . Tom 1. Vypusk 1. Rostovskoe
•
knizhnoe izdatelstvo.
("The yearly and•seasonal dynamics of the benthos in
.
the Azov Sea" Transactions of the AzNIIRKh.
Vol. 1 Issue 1. The Rostov Book Publidhing House,
69. Trudy Soveshchaniya po dinamike chislennosti ryb
Moskva. Izdatelstvo AN SSSR 1961
(Transactions of the Conference on the dynamics of fish
abundance. Moscow. Publications of the Academy of •
Science USSR 1961).
Translator , s notes:
Rutilus rutilus caspicus
"AzNIIRKh = Azovskii nauchno-issledovatelskii institut rybnogo
khozyaistva (Azov Scientific Research Institute of Fisheries)
...55
1
4
1
. 55 .
70. Khromov N.S.
"Rasprostranenie planktona I pitanie sardinelly v
raione Dakara (Vostochnaya Atlantika)". Rybnoe
Khozyaistvo No. 1, 1960
"Spreading of plankton and feeding of Sardinella in
the region of Dakkar (Eastern Atlantic)". "Fisheries"
No. 1, 1960.
71. Tseeb R. Ya.
"K voprosu o ezhegodnykh kolebaniyakh pitania pikshi
Murmanskogo poberezha" Trudy Murmanskogo Biologicheskogo
Instituta 2 (6). Moskva-Leningrad. Izdatelstvo AN SSSR,
1960
("On the problem of yearly variations of feeding by
haddock of the Murmansk coastal zone". Transactions
of the Murmansk Biological Institute 2 (6). MoscowLeningrad. Publications of the Academy of Science
USSR, 1960.
72. 0hepurnov A.V.
"0 sviazi izmenchivosti kolichestva piloricheskikh
pridatkov u raznykh form atlanticheskoi seldi s
obespechennostyu ikh pishchei" Vestnik Moskovskogo
Gosudarstvennogo Universiteta No. 6, 1960
("On the connection between .the changes in the
number of whisker appendages in various forms of
Atlantic herring and the food availability for them"
Moscow State University Herald No. 6, 1960
73. Chugunova N.I.
"Rost i sozreVanie vobly Severnogo Kaspiya v zavisimosti
. ot us1ovii otkorma". Trudy VNIRO. Tom XVIII. Moskva
1951.
("Growth and maturing of the roach of the North
Caspian Sea in dependence on conditions of fattening"
Transactions "VNIRO" (see No. 8) Vol. XVIII. Moscow
1951.
74. Chugunova N.I.
"Rukovodstvo po izucheniyu vozrasta i rosta ryb".
Moskva. Izdatelstvo AN SSSR 1958
"Textbook for the study of the age and growth of fish"
Moscow. Publications of the Academy of Science USSR,1958
75. Shatunovskii M.I.
e
"Nekotorye osobennosti pitania molodi salaki Rizhskogo
zaliva." Nauchnye doklady Vysshoi Shkoly Biologicheskoi
Nauki No. 1 Gosudarstvennoe Izdatelstvo "Vysshaya
Shkola" 1961
("Some peculiarities of feeding by the immature young of
the Baltic herring in the Riga Bay" Scientific reports
of the Academy of biological Science No. 1. State Publication "Vysshaya Shkola" (Academy or University)
* Translator's note :
salaka Clupea harengus membras
56
4'er *
•
• 56
76. Shorygin A.A.
"Pitanie i pishchevye vzaimootnoSheniya ryb Kaspiiskogo
=rya" Moskva. Pishchepromizdat. 1952
("Feeding and mutual relationship of fish in the Caspian
Sea with regard to nutrition". Moscow. Food Industry
Publications 1952.
77.
•
Shubnikov D.A.
"Nekotorye osobennosti lineinogo rosta atlantikoskandinavskoi seldi v letnii period". Trudy Soveshchaniya
po dinamike chislennosti ryb. Moskva, Izdatelstvo
AN SSSR, 1961
("Sake peculiarities of the linear growth of the
Atlantic-Scandinavian herring during the summer season".
Transactions of thé Conference on the dynamics of fish
abundance. Moscow. Publications of the Academy of
Science USSR. 1961.
78 , Shulman G.E.
. "Dinamika soderZhaniya zhira v tale ryb" Uspekhy
sovremennoi biologii. Tom 49. Vypusk 2, 1960.
("Dynamics of fat contents in the body of .fishes."
Progress of contemporary biology. Vol. 11:9. Issue 2, 1960.
79.
Yablonskaya E.A.
"Vozmozhnye izmeneniya kormovoi bazy ryb Azovskogo
morya pri zaregulirovanii stoka rek" Trudy VNIRO,
Tom XXXI. Vypusk 1. Moskva 1955.
("Possible changes of the nutrition base for fiàh of
the Azov Sea after regulation of the flow of rivers."
Transactions "VNIRO" (See No. 8) Vol. XXXI. Issue 1.
Moscow 1955.