FISHERIES RESEARCH BOARD OF CANADA

Translation Series No. 1626

Relationship between physiological state of pelagic fish and peculiarities of their ovogenesis (from "Problems of fish physiology")

By G. E. Shulman, N. I. Revina and T. E. Safyanova

Original title: Svyaz' fiziologicheskOgo sostoyaniya s osobennostyami ovogeneza pelagicheskikh ryb ("Voprosy fiziologii ryb")

From: Trudy Vsesoyuznogo Nauchno-Issledovaterskogo Instituta Morskogo Rybnogo Khozyaistva i Okeanografii (VNIRO) (Proceedings of the All-Union Research Institute of

. marine Fisheries & Oceanography). Publ. by: Pishchevaya Promyshlennost, Moscow, 69: 96-108, 1970

Translated by the Translation Bureau(GNK) Foreign Languages Division

Department of the Secretary of State of Canada

Fisheries Research Board of Canada Biological Station, St. Andrews, N. B.

Arctic Biological Station, Ste. Anne de Bellevue, P. Q. Halifax Laboratory, Halifax, N. S.

Marine Ecology Laboratory, Dartmouth, N. S.

32 pages typescript 1971

• 4...one- •

TRANSLATED FROM - TRADUCTION DE INTO - EN.

' - Russian

PAGE NUMBÉRS IN ORIGINAL NUMÉROS DES PAGES DANS

L'ORIGINAL '

PUBL ISH ER - EDITEUR

Poshchevaya Promyshlen-nost'.

DATE OF PUBLICATION DATE DE PUBLICATION

96 - 108. VOLUME

YEAR ANNÉE

ISSUE NO. NUMeR0 PLACE OF PUBLICATION

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•Moscow. 32. 1970 69

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DATE SOMPLETED ACHEVE LE •

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NoVember 19th, 1970.

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'62.4

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AUTHOR - AUTEUR

G.E.Shulmen, N.I.Revina and T.E.S&fyanove.

TITLE IN ENGLISH - TITRE ANGLAIS '

RelationshiP between the PhySioloical State of Pelegid Fish and. • the Peculiaritie of Their Ovogenesis. .Title in foreign languagd (tranÉliterate fereign. charactera)

Syyaz' fiziologicheskogo sostoyaniya s osobennosty ,mi ovogeneza • pelagicheskikh ryb.

•

RF5.RENCE IN FOREIGN ly-ANGUAGE (NAME OF BOOK OR PUBLICATION) IN FULL. TRANSLITERATE FOREIGN CHAttACTERS. , REFERENCE EN LANGUE ETRANGÉRE (NOM DU LIVRE OU PUBLICATION), AU COMPLET. TRANSCRIRE EN CARACTERES PHONETICWES.

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REFERENCE IN ENGLISH - RÉ. FÉRENCE EN ANGLAIS

Transactions of:thu A11-Uniàn Research Institut e forldarine Fisheries rie, Oceanography..

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0544 • Russian G.N.K. FEB' 1 0 197t ,

Relationship between the Physiological State of Pelagic Fish and the

Peculiarities of'Their'OvogeheSis.

By G. E. Shulman, N. I. Revina and T. E. Safyanova.

(Prim:: "Trudy VNIRO" / Transactions of the All-Union Research Institute

for Marine Fisheries and-Oceanography I. Published by uPishchevays

• PromyShlentost" / Food Industry /, Mosàow; Volume LXIX, 1970.

When studying the pattern of the reproduction and dynamics of

the numbers of organisms it is extremely important to have data on the

peculiarities of the gametogenesis. In particular, • of considerable

importance is the effect of the quality of ancestral organisms upon

the numbers of progeny and upon the latter's condition. It is natural

that this may be traced by analyzing the sexual cells and glands of the

animals. The,determination of criteria of the quality of parents, of

their sexual products and of the progeny may be approached in various

ways. Let us discuss here only some of the aspects of these approaches,

namely: the age and growth rate of fish, the level of energy resources

in their bodies, content of fat and plastic substances in the gonads,

DP,AFT .TRANSLATION Cy for

i\iON RI:VIZ E

•

S - 200 — 1 0 —31

.1■••

These problems are still poorly studied in respect to fish. At

the saine time, the relation betweàn the energy resources accumulated

in the organism of higher animals as reserve fat, between their age,

and between the peculiarities of the ovogenesis are common knowledge.

In livestock production, for example, an entire complex of measures

is developed on this basis; these measures increase the reproductive

capacity of domestic animals and the quality of their offsprings.

Only recently,studies were carried out to determine the

relation character of the quantity and quality of the ripening o f .

sex products in fish and of the viability of their offspring, to the

qualitative indices of the parents constituting the spawning popu4-

tion*. Published materials indicate a complexity of the character

of interrelations, therefore, in individual cases the obtained data

appear to be contradictory. Unfortunately the presented data are

not treated in all the papers with the application of the variation

statistics. 1 further accumulation of data on various ecological

groups of fish is required for a more complete development of the

discussed problems.

The purpose of our research is to study the relations

between the age, the growth rate and the level of the energy reserves

in the body of horsemackerel Trachurus mediterraneus ponticus Aleev,

one of the major pelagic fish in the Black Sea, the chemical composition

of its gonads, the size of the ovocytes and the fertility. Furthermore,

in our research we use data on the intensity , of maturation of the

Atlantic sardine Sardinella aurita Valens. in relation to the level of

fat reserve in its body.

* References to corresponding papers are given later in the text.

-3-

Data and Metteds

The studies were carried out in 1962 and 1963. In 1962,

we analyzed/females from 12 to 25 and front 41 to 48 centimetres long*,

two to seven and eleven to fifteen years old respectively. In 1963,

we analyzed fish from 13 to 15 centimetres long (basically three-year

olds). The data were gathered over short periods of time (two to

six days); for the fish up to 25 centimetres long, in June of 1962

and 1963, from catches in trap nets at the Olginka village (the Tuapse

region), i.e., were taken both years from closely related populations;

for the fish ranging from 41 to 48 centimetres in length, in May, 1962,

from two catches with purse net in the eastern part of the Black Sea.

Specimens of the stage IV maturity (according to Meyen's**

scale) with relatively close maturation coefficients (cv. <20) were

selected for the analyses. Consequently, the analyzed fish were in

an approximattly identical physiological state. For each fish the

length and the weight (mass) of the body were measured, the age was

determined (according to the otoliths), as well as the growth rate

and the weight (mass) of the ovaries. Half of the avary-filled , sac was fixed in a 4% formalin solution for subsequent analyses of

the fertility and size of the ovocytes.

The fertility was determined by the method of direct count

in a 100 milligram batch, the number of ovocytes was recalculated to

the weight (mass) of the ovaries. It was established that a count of

* Fish of these dimensions constituted the.bulk of the spawning population. In recent years, as we know,. fish from 26 to 40 centimetres long is practically absent in commercial catches ,. In the course of analysis the horsemackerels were not divided,into "large" and "small" forms, since it is established now, that both forms ripen for the first time at about the same.age / 40 1.

** Spelling of this scientist's name is unknown.to the translator, phonetic spelling is given here.

fOrt

-4-

ovocytes in a bata of 100 milligrams, 200 or 300 milligrams and more

gives analogous restilts. Because of the intermittent spawning'in the

horsemackerel, we counted in this batch both vitelline, and non- .

vitelline ovocytes, the ratib of which was approximately the same in

-all the analyzed females. To'measure the diameter of the ovocytes,

the latter were taken in each fish from the same part of the ovary-

sac (in the central portion of the latter). We measured 200 Vitelline

ovocytes in each ovary. The diameter of the ovocytes was measured

under a microscope with a 32-fold magnification by means of an ocular

micrometer.

When the biological analysis was completed, the same fish

underwent a biochemical analysis. From each fish batches of muscles,

liver and ovaries were taken to determine their chemical composition.

The batches were placed in beakers and dried in a drying cabinet at

a temperature of 100°C until a constant weight (Mass) was achieved.

From the difference between the weight (mass) of the raw and the drY

batch the content' of dry substance in theaamples and their humidity .

was determined. The fat content in the desicated samples was deter-

mined in the Soxhlet apparatuses according to the Rushkovsky's method

/ 26 I. The content of defatted dry substance in the sample was -

calculated from the difference between the dry weight.(mass) and the

fat-content in the batch. •

• A total of 117 fish were'analyzed (67 in' 1962, and 50 in

.1963). The entirematerial was processed statistically.according to

• 'formulae used for small batches.

Data on the fat-content in and the maturation-degree - of the

Black Sea horsemackerel and of the Atlantic sardine were also analyzed.

These data were collected in 1958 and in 1969, respectively (data on

horsemackerel werè collected in the Inapse region, thosè on sardine,

in the region of Dakar); these data represented the averages of the

sample, each of which contained 20 fish of uniform size and at identical

stage of . maturity (different from fish and from other samples). Horse-

mackerel of .- 9 to 10 centimetres and sardine of 13 to 14 centimetres were

analyzed,. We have 6 samples on horsemackerel, and 13 on sardine. -

- cn •-c e

ri (13 0

(Co -D

n

• •

25 .

-

In each

4

• 14

19

5

25

.?,

Chemical comPosition cf ovaries, dimersion of ovocytes and fertility of h.orsemackerel

of various ages and lengthS in 1962. •

in % of weight - (mass) of the ovaries. -

e .

CI

-f-je

Cri • o in

o 4

ct-1 0

o

Ame in years

2 it 3

4

5

6H7

13 H 15

Leneth in cent,imetres

12-16

16-20

22-25

40-50

*)

27,8 ± 1,96

31,9±1,98

35, 7.± 1,10

32,6 ± 2

31,0±0,84

26,5±1,09

36,3±0,5.1

34,3± 1,43

31,0±0,84

72,2±1,96

68.1±1,98

6±1,12

67,4±2,6a .

69,0±0,84

73, 5 ±-1,00

63,7±0,51

65,7±1,43

69,0±0,84

8 ,.7+ 0,70

10,4±1,33

12,2±2,31

10,6±1,76

10,5±0,92

7,4±0,27:

12,6±0,47

12,5 ±0,76

10,5±0,92

72. 4-D

42, tr Cfl

c r1:1

• ;-1 ;.-4

o e r,-; • 4 0 4-D (1] (1'g). (C

• E

19,07±1,24

21,5±0,85

23,6±0,72

22,0±2,13

21,2 ±0,42

19,1±0,52

23,7±0,30

23,4±Ô,90

21,2 ± 0,42

mu+

'271 'Di -0

• (I)

•-r<

4-) 0 4-) nee- 0

C CH

29,5±1,73 68,8±9,46

30,8±1,94 69;16±1,94

33,5±1.,31. 166,48± 1,59

32,71;71:i 67,27±1,59

:31,8±1.55 68,i91.,66

26,9±1,64 72,9±1,75

34,1±0,85 65,9±0:85

34,8±1,19 :65,3±1,16

31,8 ± 1,55 68,2±1,66

'• • •

73,5±8,29

305,9±77,7

619,8±86,0

641,6±283,9

3517,3±310,6

86,3±9,1

322,5±19,-0

813,6±48,4

3517,3±310,6

I CH •0 0 r-i ,4---

>, (1) cs

.4) -P • tr

1.%

• Ci 0 4-D CO

H 0 CI • 4

4-D b-

3,1 ±0,31

4,0±0,25

4,6±0:65

4,6±0,16

2,8±0,25

3,1 ±0,22

4,4±0,25

4,9±0,28

2,8± 0,24

c *

r-1 (.1 -H

-p C

• 'd C.)

CH C- o

;•1

f-1 .4) o

C O '"C 0

▪ > 0 0

7,8±0,58

9,4±0,50

10,0±0,36

9,1±0,44

8,2±0,42

7,8±0,41

9,8 ±0;46

10,0±0,23

8,2j0,42

›-» ri • CI e • Ci

el : Cei O 7.1

• •

e z z

1.;

CS

-0

›.>

4.

cs Çx

Value of a division on the ocular micrometer is 0.025 millimeters.

ecg• there are approximately 200 vitelline ovocytes. y:› 99

—6—

Results of the Analysis /99

Between the dimensions and age of a horsemackereL, on the one hand, and the cheMical composition of the'ovaries, ovocyte dimensions

and the relative fertility, on the other hand, we have a definite

relationship (Table 1). The content of fat and Of defatted dry sub-

stance in the ovaries is the smallest in the youngest and the oldest

àge groups of horsemackerel. In fish of medium age (and of medium size)

the fat-content in ovaries and the content of defatted dry matter is the

highest. •

9; -cî ka.)

■ .cieom« gealeady

i35

,c>

1.%

• • 30

h • 28

• 26V • • 50 e +7.«

D 4e ft • .4g e " 40

110)4

5-t„ào7in

-% 7 Fe% M '

” 9 10 11 12 - 13. • y5 ' It 12 g :

• Y(uptiocrie .umt4.1 e % i cepery ffesigcnly •. e

(26 t ert 1 ./Y% o

"I. A .S 14b3.5 tc .r

Figure 1. Con• nectiàn between the chemical composition of gonads, the relative fertility and the dimensions of ovocytes in horse-mackerel (each point on the graph is an average of several determinations). •

1)/1 -\-ov,nc..t '‘'Yt

%-rn e-klua

5/ - CI 78 .1 r- #0,57±0)0

' J e

â g

-,e z

?;-.) le* 0 LI 'k ino

S-e .

e - e4 8,0

7.5

.1t .

-7-

The bulk of the fat in the sexual gland in a horsemackerel ;-

is concentrated in the ovocytes. The fat content in the ovaries

characterizes the energy potential of the eggs /25/. Although, the

composition of "fats" determined during the extraction by means of

ether consists not only of true fats, but also of a certain number of

phospholipids, stearines and others participating in the plastic

metabolism, the major part of fat in the ovaries is used as energy

reserve at a subsequent development of embryos. Furthermore, the

degreased dry substance consists basically of albumens and it may

characterize the amount of plastic material localized in sexual cells.

Consequently, the greatest energy reserve.and the amount of plastic

substances in the gonads, as we see in Table 1, is contained in horse-

mackerels of medium size, i.e., in 5 year old fish. The water content

in the sexual glands of these fish is minimal.

The chemical composition of the sexual glands in horsemackerel

is closely related to the dimensions (average diameter) of ovocytes and

of the relative fertility. The greatest dimensions of the ovocytes

and the highest relative fertility are.also observed in fish of medium

size. The mentioned relationship is established in fish of various

age and size, whiCh were analyzed in 1962 (see Table 1). The connection

between the chemical composition of the ovocytes and the.fertility of

fish is traced also in 1963 in a horsemackerel group of same age (three-

year old) and 13 to 15 centimetres long (Table 2).

Thus, all the mentioned components in the chemical composi-

tion of a horsemackerel, when recalculated both to the raw substance,

and to the dry substance, as well as the number and size indices of the

ovocytes are interrelated. This relation is of a linear character and

is expressed by corresponding equation of regression (Figure. 1).

5 15

-13 12

-8- •

Table 2.

The relation between the fat-content of avaries, the fertility and the,

dimensions of ovocytes in three-year old horsemackerels

13 to 15 centimetres long, in 1963.

1 I •

.,

›-

...

›..t H :Z=1 H • e . 4-) 4-D • _ .4) ei) › • .,'- z... e • r-I • r•I 4 ,e N 4 -.) ,-1 0 C.) H H $10 ',

. .1-1 '"C ;--I •1-1 .I.A ni 4.4 H•› o . 1-1 LI -P • 4-3 4-I C.

Z.H 4-) e-i tO e-o CO •

.r- 4 E 4-) 61 111J rC se I:3 es-1 Cl m 0 -- „o .4-1 . CH c o H

4 •t, (ii ce 4-1 li i,..i b) cli • -I—) CO 4) (I) e m 0 e er-1 -I:, › or r.0

4--% -I-) 7:1 0 ›-. :=1 0 -r4 0 • gt e-t *1-1 . C..) • C..) S..-4 r-1 ..C■ A-) .- te. 113 ›

, '0 0-- '0 0 4i , (1 4-) o (le o 4 •4-) In H •H e ro .-4 ..> 4.1 e .11 (-- 1 -0 u Z •H .• gi n o o 4-) -.,__. o . -- •r4 C" •r-I 4 f-I

5,1-6,0 6,1-7,0 7,1-8,0 8,1-9,0

24,0±1,80 25,5±0,80

27,6 ± 0,65 26,8 ±0,66

106,3± 25,5 151,7 + 12,1 155,5 11-17,2

.171,5 + 15;5

3,9 ±0,74 5,5 ± 0,38 5,3± 0,57 6,0±0,58

,

But for one exception the closeness of this connection is great.

Consequently, any one of the enumerated indices may characterize the

quality of a horsemackerel's * s'exual products, and with the help of

any one of these indices the remaining indices may be derived with

sufficient accuracy.

A relation of the fat-content in the ovaries and other

indices of the quality of sexual products, to the growth rate of the

fish can be traced in horsemackerel (Table 3). In fish with a higher

growth rate, we observe a higher fat content in the gonads, an

increased fertility and a greater size of ovocytes.

* The content of fat and Of the defatted dry . substance characterizes them directly, the remaining indices do it indiredtly...

Table 3.

The relation of the fat-content in the gonads, the fertility, the

dimensions of vitelline ovocytes in four and five year old horse-

mackerels in 1962, to their growth rate.

S), m. S)+ • .

. .-..... ›. 15 4) -l. . .I.J e

',,,,E1. e c.) 4-1 • .4-1 N

.,., • 0 • 0 0 4) r..-1 ›.-•

. M e e › v • -1 el

M c. .ri al .0 W 5 4) (g

.1-1 gi , 4) O. Z •

4-4 -? 4--: (1) 4-4 0 ;-4 e cit 0

S) P ..c2 - 0.4--4 0 4-4 m4

4-4 e v • M .4-4 . 4-4 cIJ

0..-1• 4-) W ;,-4 .4-1 5 0 0 Z ,

4D g: C > › 15

.17.4 0 r'>' 4.) • l ;-4 ••f .ri

4) 0 Os-4 0 rd ni 4J 0 >

b: , c) 1J • e 4-1 •ci g5 .4-1• . 4) C. Z i r-1 E 15

v .. (1 4-4 ••-I •i-1 O. e . 4 .r4 r=.4 0 •---* 0 MI › :2.1 ..r-1

Four-yeer-olds;

12,I--16,0 16,I--20,0 22,I--25,0

26 ,6 ± 2,71 33,4 ±I ,00

8, 5±0 , 90 9 ,9 ±0 , 52

10,7±0,20

3,2±0,10 4 ,7 ±0 ,25 4 , 8±0,45

5 4. 5

• Fiveyclim-olds. I2,1--16,0 16,1-20,0 22,I--25,0

23,1+ 3,14 35,4±1,38 34,5± 1 ,48

10,1 ± 0, 74 9 ,9± 0, 36

fe"!

'1

• 0 0 •

• • 0 • •

0 , 0 Q5

18 20 - 22 24 28 31

o Z

_X

-10-

There is practically no connection between the fat content in

muscles and in the liver of a horàémackerel, on the one hand, and between

the fat-content in the gonads and the fertility, on the other hand

(Figures 2 and 3).

An analysis of data on the fat content in horsemackerel at

various stages of maturity collected during the spawning period of 1958

indicates that a more mature fish has a higher fat content (Figure 4).

Analogous data are obtained in 1959 for the Atlantic sardine.•

o

•

4 c.

>44

c4-4 244 o

-4-)

4r.) 8 - 4-)

o o 2

4

o

0,

o

o o

.•, • 0 • 0 00

o él • • • • . o I b oo, •

• • o 00 • . 0‘,.. • .

0 • '. O. .8 . . 0. ,

0 . • • . • le . 0 0 .

o

o

Fat content in p;onads,'In % of dry substance.

Figure 2. Interrelation between the fat content in the muscles, the

liver and the gonads in a horsemackerel:

1 - muscles; 2 - liver.

-11-

e 0•

• e Co

4-) . - m . „ . .... .,.... .. .

•0 .»: .. 32 .. o W

• 30 - o ,

•

›-, 28 - • G o

;-4 % ' e ° 26- .

-a 0.. . 00 . . .24- ..

c.--, a- • • • .,,,,, • •

'GI zu- O. ' . .

e • • - . -

%.. 14- • .0 0 • . j

/6- • e : • o . .

g

erA rii/- 0 g

v). î • • 0 12 1 .. o o • • „, î •

• ' II

4 W- coo o • 9' qf • ,.

✓ 8- 0 :. 0

. % .

-1--) 6 .

e •4- ° • • o •1 '

o • '0• • -• o2,

c..) 2 • - 1 1 I ° °Ill • I •, 1 !

4-.) • ' 3 4 5 6 7 8 9 10 • - — m

... .

Zelative fertility, in thousands.

Figure 3. Inçerrelation between the relative fertility, fat content

In the muscles and in the liver in a horsemackerel:

1 - muscles; 2 - liver,

5>

0

4-1 20 o

•

• m • m

E

5 o 4^

fr,

28/E ' A<

- me ,e/ . •

„ere • •:

28 eye .- •

.02

:e Jer. •if

Stages of 'maturation.

Figure 4. Connection Of the fat content in the body of sardines and

to the maturation stages of their gonadsr.'

1 - • horsemackerel; 2 - sardine.

-12-

Discussion of the Results

As shown in the preceding chapter, fat content in the sexual

products of horsemackerel (as well as the content of defattened dry

substance) may characterize their quality with sufficient accuracy.

This opinion is shared by the majority of authors who have analyzed

the connection between the chemical composition of sexual glands of

fish and the peculiarities of their ovogenesis /23, 25, 35 etc./.

I. N. Petrenkà and A. A. Karasikova /39/, for example, point out that

when the fat content in the eggs and in the muscles is decreased, their

amino-acid content shifts away from the ideal, since the albumen and

the carbohydrate metabolisms are involved. It should, however, be

noted, that in carps according to O. B. Chernyshev's /45/ data a

considérable percent of fat is not contained in the sexual cells,

but in the intercellular tissue and is used as energy substance in

the maturation of gonads. • Therefore, we may characterize the quality

of sexual products in carps according to the fat content in gonads only

with extreme caution.

The horsemackerels have a distinct positive connection

between the content of the plastic and the energy substances in the

ovaries, on the one hand, and the ovocyte dimensions, on the other

hand. M. N. Krivobok and O. I. Tarkovskaya /24/ indicate a positive

connection between the fat content in the sexual products and the

dimensions of the ovocytes in the Baltic herring; Yu. I. Cheprakova

and S. G. Vesetsky /44 1 indicate this for the Caspian roach (vobla);

while M. T. Shatunovsky /47/ does the same for the White Sea flounder.

S. G. Kryzhanovsky /25/ showed that the value di the fat

reserves in fish eggs reflects the ecological specificity of definite

taxonomical groups. The highest fat content in the sexual products

(ten to twenty percent of the raw substance) is found in the sharks,

sturgeons, salmons and mullets /19/. Embryos and larvae of these fish

-13-

are found under the most favourable cbnditions from the euiit of view- • “

of their supply' with'endogenous ntiérients. The minimum content of fat

in sexual products is found in herrings, cods and flounders,.from.one

to three percent of the body weight (mass) and less; these are marine

fish spawning basically pelagic eggs. The low fat content in sexual

produçts of the mentioned fish is explained 'bye high water cotent

of their eggs, thus ensuring them of a satisfactory buoyancy /17, 25/.

As pointed out In the foregoing, the horsemackerel, we may say, has a

rather high fat content in its sexual products, about ten percent of

the raw substance, and it approaches the values obtained for the white-

fishes /19/. An egg of horsemackerel, like in other fishes of the,

perch family, has a relatively high fat drop /25/, which fulfills an

important hydrostatic fundtion and which furthermore Supplies.the

developing embryo' with endogenous food. •

The fat content in the sexual products in fish is determined

not only by the.adaptive peculiarities of the development of embryos

and larvae, but also by the qualitative state and the life conditions

of a parental organism. One of the most important indices of the .

qualitative state of thé organism is its age. .It has been proved not

only on horsemackerel, but on many other fishes, such as herrings /23/,

salmons /57/, carp's /43, 16/, that the highest quality of sexual

products (maximum fat content of egg size). are observed in individuals

of medium age.. An analogous connection is established between the

age of.fish, quality of their offspring . (i.e., of their survival rate),

growth rate etc. /20, 1, 11, 10/: These data are of considerable

importance to the theory and Practice of the fish inchistry, and

simultaneously they indicate the importance of studying the age

structure of the spawning fish population, when analyzing the repro- .

ductive capacities of the populations. Furthermore, these data agree

well with the well-known position determined mainly on mammals on the -

gradual formation of the Metabolic prOcesses in ontogenesis and concern- ,

ing their deterioration'in the old age/à2/. _

.

-14-

The linear growth rate and the intensity of fat accumulation

are important indices of the state of fish in the spawning population

indicating the living conditions. These indices change sharply when

the food supply, length of fattening period etc. change. As noted

earlier, a horsemackerel with an increased growth rate has also the

highest quality of sexual products (the fat content of the gonads and

the dimensions of the ovocytes). Identical data are obtained for the

Caspian vobla (roach) /43, 44, 23, 24/.

Parallel with an improvement in the quality of sexual products,

in the horsemackerel we observe an increase in relative fertility.

N. V. Lebedev and Chêng Chêng-Tun /27/ have demonstrated on the example

of roach and carp that the feeding conditions of the producers not only

determine the amount of yolk, and thus the dimensions of ovocytes, but

also the quantity of produced sexual cells, i.e., the value of the

fertility. When the arrival of the nutrients with blood into the

germinative tissue of the mentioned fish is decreased the resorption

of the ovocytes is observed. The same phenomenon occurs in the pink

salmon /38/ and trout /60/. Thus the fattening conditions of the

producers, the indices of the quality of their sexual products and

the indices of the quantity of the ovocytes are interrelated.

Certain authors, however, find an inverse relation between

the dimensions of ovocytes and the relative fertility of fish 1 24,

47/, particularly in the Baltic herring, in the Baltic flounder and

in the White Sea flounder. On this foundation, M. N. Krivobok /21,

22/ bases his belief that a high fertility contributes to a decrease

in the fat content, in the plastic substances in gonads, and in the

dimensions of ovocytes, in other words, it has a negative effect

upon the quality of sexual products in fish.

If,

-15-

This . contradicts the conclusions resultant from a research

carried out by N. V. Lebedev and diiêng Chêng-Tun /27/. Furthermore,

it is necessary to take into consideration that improvement in the

feeding period conditions of the fish, not only has a favourable effect

upon the quality'of sexual products, but also increases the growth rate

of fish and its absolute fertility, which is connected with the size

of the fish /2, 35/. • We believe, therefore, that the fertility and

the quality of seXual products cannot be contrasted.

We have a considerable amount of data on the connection

between the level of fat reserves in the fish body and the fertility

or fat content of their gonads /2, 7, 8, 39, 36, 23, 47/, particularly

in the Kura salmon, the Baltic and the White Sea flounder, the Baltic

herring, the Onega herring and in the Black Sea anchovy. The high

level of fat content in a fish body reflects good conditions of the

feeding period; these conditions have a positive effect also upon

the ovogenesis processes. Therefore, the indices indicated above

are often intercorrelated. The connection between the feeding

conditions, the amount of fat reserves in the body and the

condition of sex products is easily traced also in insects /42, 30/

and in mammals /18/.

The connection between fat reserves and fish fertility, and

the fat content of gonads can be traced not only when the annual

fluctuations of the mentioned indices within the spawning population

or the individual fluctuations inside the population are compared,

but also when we compare groups of fish different in ecological

respect. Thus, closely related races of European anchovy (Engraulis

encrasicholus), such as the Azov and the Black Sea anchovy, differ

sharply in the intensity of fat accumulation and the level of fat

reserves in the body. The fat-content of the former attains towards

the end of the feeding period 20 to 25% of the raw substance, while

the fat content of the latter does not exceed 10 to 15%.

-16-

Differences in the intensity of fat accumulation in the

Azov and the Black Sea anchovy ara'explained by the fact that the

former occupies during the spawning, and after-spawning feeding the

exceptionally rich feeding base in the Azov Sea, while the second

one lives in the Black Sea, which is relatively poor in respect to

food. We know that the fat reserves in fish are utilized to a consider-

able degree as energy supply for the purpose of ovogenesis /55, 54/.

The fat content in the gonads of the Azov anchovy is twice as high, as

in the gonads of the Black Sea one: Ten and five percent of the dry

substance /50, 36/, and the fertility of the first one is by 25%

higher than of the second one, namely, it is 25,000 to 20,000 thousand

eggs /46/. This is an object example of the effect of living condi-

tions upon the fat metabolism, and through the latter, upon the ovo-

genesis in fish. Similar data are obtained on related subspecies of

the river flounder Pleuronectes flesus, both from the White Sea and

from the Baltic Sea /48/. The first subspecies has a higher fertility

and a higher level of fat reserves, than the second one.

We must point out, however, that the connection between

the fat content in the fish body and the fertility or fat content in

their gonads is not always traceable. Such a connection has not been

found in the White Sea herring /2/ and in certain other fishes. As

indicated earlier, there is no relation between the amount of fat in

the muscles and in the liver of the horsemackerel, the fertility and

the fat content in the ovaries.

Undoubtedly, these data do not indicate the absence of inner

relations between the level of fat reserves in the fish body, the

number of ovocytes and the quality of sexual products, since it is

exactly the fat reserves which to a considerable degree ensure the

processes of ovogenesis. The fact is that in the maturation period of

-17-

sexual products in many species of fish a mobilization of fat reserves

is taking place, consequently, th è fat balance becomes negative for a

long period of time. Therefore, the ripening of sexual products takes

place on the background of a sharp drop in the fat reserves in the body

of fish. The lowest fat content in a fish often,indicates here the

higher energy expenditures on the ensurance of albumen synthesis in

the gonads. As an example we may mention the Far Eastern salmons 137/

and the Azov. Sea anchovy /49/.

In the foregoing chapter we have proved that there is a direct

relation between the' fat content in the body of the Black Sea horse-

mackerel and of the Atlantic sardine, and the ripening degree of their

ovaries, if we analyze fish taken from the same accumulation at a •

certain time. Similar data were obtained by M. N. Krivobok and

O. I. Tarkovskaya /23, 24/ for the spring-spawning subspecies of the

Baltic herring. We believe that'all these data indicate an earlier

and more intefisive maturation of fish with a considerable energy

potential, and, undoubtedly, they demonstrate a relation between the

condition of the producers and the intensity of the ovogenesis processes.

At the same time, the discussed data cannot serve as a basis

for the conclusion arrived at by the mentioned authors /23, 24/, that

when the sexual products ripen in the Baltic herring, its fat content

in the body increases. If we consider the relation between the fat

content in the Baltic herring and the ripening stage of their gonads,

not in a definite moment of time, as the mentioned authors do, but

during the course of the entire pre-spawning period /56, 59, 58, 28/,

then the decrease in the fattiness of fish (we are speaking here about

the spring-spawning Baltic herring) can be traced quite distinctly along

with the progress of the gonad ripening. It should also be noted, that .

a decrease in the fat content in fish, that takes place in the course

of the gonad ripening (if this process is considered in time), is

observed also in the Black Sea horsemackerel and in the Atlantic sardine /53, 53/.

e4.

0

4-1 o

•

W

4J Li

4)

o 4;2

-18-

Fipre 5. Diagram of changes in the fat content in the body of sardine

and horsemackerel during the ripening of their gonads

(A, B and C are the periods of time).

Changes in the fat content in,the body of horsemackerels and of

sardines and of the ripening stages of the gonads are represented on

the diagram following the degree of their maturation (see Figure 5),

where the letters A, B and C symbolize the three consecutive periods

of time, during which the maturity of fish increases and the fat

content decreases.

It is interesting to note that the'sardine in the Dakar area

does almost not feed at all during the pre-spawning period /15/, a

fact stressing the decisive role in the energy supply foi the ovogenesis

processes played by the fat reserves accumulated in the body during the

preceding periods of the annual cycle.

.7

7197

The material discussed in this paper demonstrates the intimate

connection of the reproductive po .Èential of fish to its living conditions

manifested through metabolism. In partiçular, in many fish species the

food supply determines the level of fat reserves in the body, while the

latter, in its turn, affects the quality of their sexual products and

their fertility. Consequently, through shifts in the intensity of

fat metabolism in the fish body, the connection between the change in

the conditions of habitat of the populations of a given species and

the dynamics of their numbers is to a certain degree accomplished. This

problem is elucidated in detail by G. V. Nikolsky /33 to 35/, who placed

the mentioned connection as the basis for the concept of autoregulation

of the fish numbers.

Howevet, there is no doubt that the effect of the living

conditions of the population upon the reproduction of their numbers is

not limited to this connection. The connection between the numbers of

the populatiod and the conditions of their. existence (in other words:

within the system "organism - environment") is carried out not along

one, but,along several channels. Each of these channels plays a

considerable part in the reproduction of animal numbers.

It has been convincingly proved by many authors that the

yield of juvenile fish is determined not only by the quality of sexual

products in the producers and by the amount of the eggs laid, but also

by the biotic and abiotic conditions formed in the water basin during

the pre-spawning and the spawning periods of . the annual cyèle and,

particularly, by the conditions present during the development of the

laid eggs and larvae /6, 12, 13, 14, 29, 36, 41/. Similar conclusions

are also obtained for insects /9/. The value of the replenishment of

many fish species depends both on the hydrometeorological circumstances

under which the development of embryos is taking place, on the supply

of fish larvae with food during the transition to-the active feeding, .

on the pressure from the predators, and on other causes.

-20-

Either of the enumerated factors, as well as the quantity and quality

of the sexual products, may be deèisive for certain species (or even

for the same species in different years), but most often the effect

of these factors upon the reproduction of the population numbers is

a collective one.

The determination of the importance of sone factor in the

reproduction of fish numbers is extremely interesting and must become

one of the most important tasks in the studies of ichtyologists. In

particular, when we continue our studies in the mentioned direction

we should establish how the quantity and quality of the sexual products

produced by the fish,the quality of parental organisms, the food supply

of the larvae, the number of consumers of these larvae, the temperature

conditions of the development of embryos, affect the yield of juveniles.

We must also point out that an important stage in these

studies is the determination of new and more refined criteria for the

quality of the sexual products of parents and their interrelation,

than the criteria used in the present studies. For this purpose, in

particular, we must study in detail the morphophysiology of embryo-

genesis, the exact biochemical composition of fish, their sexual

cells (both male and female) and their eggs, the antigennic properties

of the developing sexual products and embryos, who open the processes

of chemical differentiation. A part of these studies is already

underway for fish in the Azov and the Black Sea basins /4, 5/. It

is obvious, that new data may be obtained as a result, which will

elucidate the complex and contradictory problem of the interrelations

between the parental organisms, the sexual products and the offspring

in fish.

-21-

-

CONCLUSIONS

1. Between the age, dimensions and growth rate of the Black

Sea horsemackerel, on the one hand, and the content of fat and of the

deflattened dry substance in the ovaries, the ovocyte dimensions and

the relative fertility, on the other hand, there is a definite rela-

tion. The best qualitative indices of the sexual products are found

in fish of medium age (and size), as well as in individuals with an

increased growth rate.

2. • The content of fat and of plastic substances in the

ovaries of horsemackerel, the dimensions of ovocytes and the relative

fertility are closely interrelated. The relation between these indices

is linear. When we know one of these indices, we may calculate the

other ones.

3. There is a relation between the fat content in the Black

Sea horsemackerel and the Atlantic sardine, on the one hand, and the

degree of the ripeness of their gonads, on the other hand. This

relation indicates that the intensity of the ripening of sexual

products in the mentioned fish depends on the level of fat reserves

in their body.

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