•rFISHERIES RESEARCH BOARD OF CANADA

Translation Series Noa 2545

Methods and efficiency of breeding Ropshian carp. II•Methods of artificial seleçtion

by V. S.' Kirpichnikov, K. V. ponomarenkoN. V. Tolmacheva; and R. M. Tsoi

Original title:Metody i effektivnôst' selektsii

karpa. Soobshcheniè II. Metody pr

From:Gènetika (Genetics)', 8(9) : 42-53' 1972

Translated by the Translation Bureau(JW,).Foreign Languages Division

Depârtment of the Secretary of.State of Canada

Department of the EnvironmentFisheries Research Board of Canada

Halifax LaboratoryHalifax, N. S.

1973

20 pages typescript

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Methods and efficiency of breeding Ropshian carp. IL Methods of artificial selection

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Metody i effektivnost" selektsii ropshinskogo karpa. Soobshchenie 11. Metody provedeniya otbora.

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Methods and efficiency of breeding Ropshian carp

IL Methods of artificial selection UNEDITZD TRANSLATION

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V.S.Kirpichnikov, et al UDC 639.215.2

All-Union Scientific Research Institute of Pond Fisheries, Dmitrov; State Scientific Research Institute of Pond and River Fisheries, Leningrad

Introduction

In the course of breeding a new variety of Ropshian carp [1],

methods of artificial selection and cross breeding were used and improved

upon. Contemporary genetic methods of selection received considerable at-

tention as well. In this paper we consider questions relating to various

ferms of selection in connection with breeding Ropshian carp.

Our samples were selected from hybrids of the 2nd to 6th genera-

tions produced by cross-breeding carp with "eastern carp."* The European

and "eastern carp" are isolated from one another by geologic time and be-

long to different subspecies [2]. Hybrid populations are sufficiently

*Translator's Note: "eastern carp": see under sazan amurskii C.c.haematopterus (Russian--English Glossary, W. E. Ricker).

SOS-200-1 0-31

by

7 t330-2 1-0:19-6332

2

heterogeneous to ensure the effectiveness of large-scale selection in a

number of generations [3].

The main factors in our selection were the selectional differen-

tial (S), that is to say, the difference between the average value of the

Character of the selected and nonselected specimens, and the tension of

the selection, which is calculated according to the formula

n.100 v

where N and n are the number of specimens before and after the selec-

tion [4]. In a number of cases the selectional differential was expressed

in standard deviations, that is to say, the intensity of selection was

. determined according to Falconer [5]:

Experiment

Mass selection for weight in subsequent hybrid generations. The

selection was begun after the second generation of hybrids. A summary of

the general tension and intensity of the selection is given in Table 1.

In the second generation the tension of selection used in the first /43

and second year of their life was almost identical; the rejection was not •

very strictly applied and about 5% of the raised fish were kept for breed-

ing purposes.

In the 3rd generation we tried to increase the intensity of the

selection, making maximum use of the high fertility of the carp. The

main weight of the selection was shifted to the first year, when the number

of fish can be very large. The selection among 2-year olds had only a cor-

rective significance. This method enabled us to bring the intensity of

selection to 3 and more. Fewer than 1% of the fish were kept for breeding

3

T a 6 :t -,I u a . I

HaupxnemlOCTh (V) It 1111'rLMCIIIIHOCTIt (i) OTfiopzi valmoa no tiecy B iteptibie ;tu .a ro :~a unamt

c I" " :,

Holm-

C

234

6

b

cU:tP ;t(-r, r1 .,*!1,1

0 + , t I I -. , 2

2 27,7

7 1,4

8 2,06 52,85 13,85 89,4

54,3n the

ccc

d gram'

I e

I

f

up t t22,1 5,1 0 ) i - 01I _

2 : P ! L!P -I 1 1; 3~i".)

66,0 0,9 17-97 ()-59 >3,0 276 O'SO79,9 1,6 4-70 0 _- "' ;9,8 5,2 0-12 17-1-57 f 8 127 100

55,2 7,6 5-!)1 1) -16i) 2 , ~3 1 .34,8 31,1 0-3 3- 136 1 1 2 J5,3 2,9 7 17S 1 >3,0 1 11 970

I st and 2nd year of the i r I i fe these wer, .-

ILO J!,kJJ11)~rKCJJKOCTIL oTtio[)a it" nuptsuol it Hlojjwt IUAY itilialut Cupe mill"I 1111 i t

rpy-1-div'ided into two !groups according to tension of,elect-ion-

Table 1 . Tension (v) and intensity (i) of selection of carp byweight in the first two years of life .a-generation ; b-number of crossbreeding ; c-(v)average% ; cc-age, years ;ccc-over two years ; d-selectional differential(S) in grams ; e"(i)aver-age ; f-number of specimens before selection, N .

_J

purposes, and in individual crossbreedings fewer than 0 .1% of the bred fish

were kept .

In the 4th generation the method of selection was changed . Around this

time it was shown that 16] the correlation between the growth rate of the

first and of the second year .of life was not very high. Moreover, it was

established that the inheritability of the distinctions between the record

breakers (fish with thelargest weight in the family) and the remaining spec-

imens was minimal [7--10] . In this connection, almost half of the hybrids

were subjected to a rigorous selection among the 2-year olds, with very

moderate selections in the first year . In the other families, the selec-

tions were made as before--mainly among fingerlings and y6arlings . The

overall intensity (during two years) of selection remained rather high

4

(see Table 1).

In the 5th generation the overall tension and intensity of selection

were considerably lowered. The main reason for-this was the undesirable

correlational changes which frequently accompany overstrained one-sided

selections among animals (appearance of disfiguration, deterioration of

the biochemical indicators, overall reduction in vitality, and so on)

[5, 11]. Signs of a certain reduction in resiliency were also noted in a

detailed study of Ropshian carp fingerlings [12].

With the-6th generation the selection was conducted almost completely

in the second year of their life. More than 50% of the fingerlings and a-

- round 5% of the 2-year olds were retained for breeding purposes. The selec-

tion was made more rigorous again. In selecting the 2-year olds, more at-

tention was paid to their external appearance and to the absence of abnor-

malities.

The selectional differential in many cases was considerable (Table 1). /44

As à result, the intensity of selection sometimes was quite high (more than

3 and even 4). But it should be noted that i is determined on the assump-

tion of a normal distribution of specimens possessing the selectional feature.

With regards to the weight of carp, as a rule we find hère either the larg-

est or smallest deviations from normal distribution. Even when the finger-

lings are caught in the fall they are already characterized by variational

curves that suggest a statistically reliable dissymmetry. The asymmetric

index frequently does not exceed 0.1--0.3 (fig. la), but sometimes it is

quite significant (fig. lb). The analysis of the data from.individual ponds

indicates that the dissymmetry sharply increases during food shortages and /45

when there is intense competition for food. Such a situation obtainèd in

1956 in Pond Bystryanka 4 (fig. lb), in which the coefficient of

,c1 p/0 1?.% 30

I ..... 20

;.0m6e3.e l ec_

i on 10 • • • • •

320 450 4'30 560 6-#0 4.0 k300 <MO \ 250 Bec, c, we gilt

r a g

20

10

2,60 2,70 lg Seca b d

2,80 290 3,00' log , of weight

4 8 12 16 20 2428 3236 8e. 8 We 1 ght

a • ( g ) e% 50

se I ect 10

40

30

20

10

5

30

20 se I ect i on vlic. 1. Pacunegezeaue cero2e-rKoe poruituEr

1 0moop.._

eltdro uapua uo Becy: ,.....

l.t.,.A.

a — ripy 1-imrteFur, 2,'Ponaia, 1958 r.; 37— 15,9 a; (cpeamtil Bec 0T06pafilibIX 111,1 (5 ) -= 29,5 a; S

= 13,6 e: i (11FITelfC1113Hnell> OTtlopit) =- 3,1; v (Ha-riptia<etutocTb 0T6opa)-= 1,0°' : --- 4,4 a; C • V.

27,S?;) ; A., (tioaebnititettr ancummeTptitt)-=±0,22-.

+0,11; 6 ntly Ebtc-rptintia 4, Portura, 1956 r.;

211.11 x, 52.4 a; S = 31,5 a; i 4,0; v = =0,4%; 1=7,9 z; C.V. ==. 37,7%; "Is = + 1,04

--0.14

Puc. 2. Pacupe,qe.-tenue gnyxnerRou poumuu- cRoro apua uo Becy. IlpyA kluanoucRuik,

Pouum, 1959 r. 06oallameinin cm. pttc. 1; u — oilbtmubtii macarra0,.. 6 — -torlpsubmuttocititit = 547 a; ze, = 772 a; S = 225 a; = 2,37; v = 6,6%; G

= 95,0; C. V. = 16,9%; A s +0,39 + 0,21

10 20 30 40 50 60 70 Beg, e we i ght (9') be

Fig. I

Fig. 1. Distribution of Ropshian carp fingerlings by weight:

a-pond Kipen' 2. Ropsha, 1958; xc (average weight of selected fish)=29.5 g; i (intensity of selection)=3.1; v '(tension of selection)=1.074 A

s(coefficient

of disproportion)=+0.22±0.11; b-pond Bystryanka 4. Ropsha, 1956.

Fig. 2. Distribution of 2-year old Ropshian carp by weight. Pond Ivanov. Ropsha, 1959. For designations see Fig. 1.

a-usual scale; b-logarithmic scale.

10

6

disproportionate weight in fingerlings in the fall exceeded +1. Contribu-

ting to the disproportionate weight is a greater variation among the carp

larvae, induced by their introduction into the lake over a period of a few

days, or mixing larvae from different spawning ponds.

Weight disproportions frequently can be observed among 2-year olds

in feeding ponds. We have an example (fig. 2a) of clear weight dispropor-

tion among 2-year olds in one of the Ropshian lakes; there were cases when

the weight disproportion was even greater.

When the disproportionate weight is favorably distributed, the in-

dex of the selection intensity (i) is exaggerated, as a rule, due to the

stretched right arm of the changeable weight 'curve. In the logarithmic

scale the weight distribution curves in most cases are :close to normal

(fig. 2b). This makes it possible to determine the intensity of the selec-

tion more accurately. In very rigorous selections the true intensity of

selection (in the 3rd, 4th, and 6th generations) usually does not exceed 2.5

and rarely 3, but this is less well shown in Table 1. It should be noted

that, given a disproportionate weight, the coefficient of the tension of

selection is a more objective criterion of the intensity of selection.

Optimal time for conducting the selection for weight. After Stegman

showed that between the first and second year the growth rate in Polish carp

has a low correlation, it was time to test our own carp in this regard.

In 1960 and 1961 we conducted two experiments in jointly raising selected

and unselected fish (Table 2).

In the first experiment about 5% of a amall group of fish in the

second year of their life had reached the same weight as the larger fish.

When the fish were divided into three weight categories (2nd experiment),

approximately 20% of the medium weight fingerlings were classified into

7

the large group, but from the small group only individual specimens quali-

fied for this transfer.

In two new series of tests with Ropshian fingerlings of the 5th

selected generation, conducted in 1966 and 1967, a selection of large,

medium, and small fish was made in the fall. The irregular weight distri-

bution of these fish did not permit an exact determination of the index of

the intensity of selection, but the selection, especially on the plus side,

was rather rigorous (Table 3). Subsequently the three groups of fish were

reared jointly, in large feeder ponds. The difference in weight among year-

lings of the various groups was retained in the second and third year of

their life (fig. 3). If we introduce a special correction factor(equaling

3) for the difference in weight whenthey were introduced into the feeder

ponds, then the average increment values for two-year olds, in 1967, will

be: large fish: 612 grams; medium fish: 594 grams; small fish: 457 grams.

There was not much difference between the large, and medium 2-year /46

olds, but the increment in the small fish is much less. Unfortunately, we do

not know to what degree the correction factors determined for subtropical

conditions are applicable in our climate, where the competition for food is

reduced.

In the third year the absolute growth increment in the three groups

of fish were almost identical. The largest growth gain, in terms of per-

centage, was noted in the small carp.

In 1967 we tagged and individually weighed all the 2-year olds we

caught, for the purpose of determining how frequently fish changed from one

group into another. The distribution curves for neighboring groups trans- ,

gressed rather frequently (fig. 4). There is a wide general class (fish

weighing from 320 to 400 grams) which comprises about 18% of fish from the

:

8

1400[_

400

200f-

1966 1967 1968

Pue. 3. KpunMa pocra pourlM:acl;us uap-IIOB, oroôpaBUbls ua uepsou rouy x;uauu:1, 2, 3 - xpyaavie, cpexfine it Dle:'[Fitte ptiu66i,reHepaqnsc 1066 r; •l. J. fi- TO N<i., reFteplmur

^ 1967 r. Oruo;tra B1I. Boama, 1986-106i rr.

Fig. 3. Growth curves of Ropshian carpselected in the first year of their life:

1, 2, 3--large, medium and small fish,genera^ion 1966; 4, 5, 6--same, gene-

ration 1967. Branch VM. Ropsha, 1966-

1968.

"minus" - side, up to 50% of fish from the standard group and more than 13%

of fish from the "plus" - side. About 20% of the 2-year olds from the "plus"

- side.passed into the medium group and vice versa. A similar picture was

obtained in 1968 when we investigated the fish that were crossbred in 1967.

According to Korovin the Ignat'ev's data [14], the

correlation between the first and second-year

0.7. Dense seeding of fish and the resulting

buted to higher coefficients of correlation.

of those of other investigators

considered to be near 0.5--0.6.

growth rate

competition

coefficient of

in carp exceeds

for food contri-

On the basis of our data and

the true coefficients

Although the quickly

lings generally retain their advantages in the second

of correlation can be

growing carp finger-

year as well, not less

than 15--20% of their "heaviest specimens" pass into the medium growth rate

/47

group.

r T a Cia if 1~a 2

CHIMT, pocerom porillililic!'Jo. n,"ImOB -, llefllil~'411 ATOI :CM FVJ~' M11-31111 (f - OTI-OAIZi~

.11, A-e iiaxooemie. r . ; 2 - OTf1O;k',l '11 . 4-e 11"1", w

a lb'f""" bS"'" . C1 I c2 I d

5 tM. ..6 i 0 J1 ISO* Vj 22 :")0 575 OIZO ;11)1 about

2 161 1-116 .6 nj" 6105227 -1 .9 Q 207 560--) 9- 4-42 0,3

A., 0 7 Be :) OTTIOP-1 I : - ~ :111 Iunsele'cte d

b I C L!IYN

i", - ~ (

1 :Z1 :T~ - I. t it

r.N' . rilynuy .

,r:11VNt1,-111(11CCFI lit) ;Xn.-,e TU1C:MACTBeIIHOCTIr

. . . .. .. I PNCIOAX Aff-Tng .

T a 5 ji it q a 3

OT60pa Ili)' I;Pcy epelit cern .leTuOu B Orll,lTn-. (V - R04(~11AITeRT manpnKeH-

imcm . nowazIaTC.lh 111ITCHNIRTIOCTIl OT61111,1, S - ceaeuquoiiHbTjj Aii(D(De_peHqilaji-

P011111a )

Mt~ .wae 3

1!167 1,'pyiviw eClo-almO Z-

CZ' . ~ .t 1 j'No uf-rJe

.Ilt~:Lllllü recI)Llrl ~XO OTr)O- S, c

V, 0 ;1 i

;J0 ~,jj1 aOTI pa, a

C

11 .3 -50 0

,96 700

129150200

1500100.106

-13,8

17,7

+46,9 O'l+2 6-6,0 0,2-L45,1 0,2-~2,3 --8, 6' . 0, -t

>3

>3

01-0-TO 1 . 5~Z'Out

Table 2 . Relation of growth of Ropshian carp in the first and

second year of their life (1--branch VM, 4th generation, 1960 ; 2--branch

M, 4th generation, 1961)* .

a-experiment ; b-yearlings ; bl-number ; b2-average weight (g) ; b3-ten-sion0,ht (g) ;of selection v, % ; c-two-year olds ; cl-number ; c2-average weig

d-specimens that were classified into a higher group (%) .

*VM and M are groups of breeding stock of Ropshian carp which by heredity

differ from the eastern carp (2] .

**Inaccurate count resulting from unclear labels .

Table 3 . Selection indicators for weight among fingerlings4 D(v--coefficient of tension ; i-indicator of intensity of selection ;

S-selectional differential . Ropsha)

a-fish group (1-large ; 2-medium; 3-small) ; b-quantity of fish ; bl-before

selection ; b2-after selection; c-average weight of fish (g) before 5elec-

tion .

10

•—•

80 -160 240 320 400 480 560 540 720 ec, 2

Pue. 4. KpuBble pacupe,galleŒrm uo Becy Rsys..Tie -rizos,

. oToCipanags Ha c-rap,un rogouuKa:

1, 2, 3 — upyrIume, epeutute T meauue ;pbt6b1. Orunusa BM, reuepaum 1967 r. TOHEtIM nyriurupou uniiaualio nepournoe gertcruurenbuoe pacupeue:leaue rpyunax upyruibtx u meauux pm6

Fig. 4. yearling stage; generation 1967 distribution in

Weight distribution curves of 2-year olds selected at the 1, 2, 3--large, medium,. and small fish. Branch VM,

. The broken line indicates the probable actual weight the group of large fish and in the group of small fish.

It may be concluded that it is necessary to conduct the selection

among carp in two stages. Of the fingerlings (or yearlings) not fewer

than 10--20% should be kept for future breeding purposes. The main empha-

sis should fall on the selection from among 2-year olds, which constitute

the marketable commodity in most fishing concerns of the USSR. The tension

of selection of 2-year olds can be quite high (of the order of 5--10%).

These indicators are close to those recommended by Schaeperclaus [15], and

they promise a high efficiency rate of mass selection for weight.

Appearance and other exterior features were only of secondary signi-

ficance in our selection. The only selective criterion with respect to

fingerlings was weight. However, in selecting the large fingerlings, we

couldn't help comparing their external features as well. Most notable were

the "correlated" variations in the value of the conditioning factor and the

height of the back (ratio of the length of the body to its height). The

,

11

data in Table 4 are an example of several carp families of the 4th selected

generation. Selection almost always resulted in a higher conditioning in-

dex and in an increase in relative body height.

In selections for weight, correlative variations in appearance could

also be observed in 2-year olds (Table 4); in large fish both indexes tend

to improve. "However, in this case, in addition to correlative variations,

direct selection by exterior features occurred as well. This kind of selec-

tion played a somewhat more prominent role in raising the 4th and 5th selec-

ted generations of carp. We selected the best fish from among the 2-year

olds in two stages. A rough selection by weight was made at the time when

. we caught them; at this time we kept approximately 1.5--2 tines more than

we needed of the largest fish. Two.days later we examined and measured

them a second time and rejected the specimens with obvious body defects.

In some cases, the large fingerlings and 2-year olds seemed even

more run-down and less conditioned than the unselected ones (Table 4). /48 .

This may have been the result of insnfficient food in the ponds; large carp

are especially sensitive to food shortages.

Further elininations in the 3rd and 4th year of their life involved

only individual specimens with clearly expressed defects. The tension of

selection among the breeders was not very rigorous either. The males and

females that were rejected were those that matured poorly and showed little

suitability for breeding.

On the whole, the mass selection based on external features played

no significant role in the selection of Ropshian carp. External changes

could have resulted only from correlations between weight and external fea-

tures. Our main task in the first stages of our selectiOnal work was to

increase the growth rate in carp under conditions of relatively low summer

12

T a û_:r rrq a 4

. .9 It^7PE;P 1, C?11 bt3

IIiemekrrlrrc*

rnYrrrr,i

a

Al

rI0IM3:1TC:Ill y cCrn:ie'PBnrI it [I)}:C:Iel'lit)II 4-ro ce:ierir(IIOHrInr'o

;(ü it IrUCae orÜ(p:l Uo rSeC;J. l'QIlirra

1 1 't ocr., r-

195919l)019;i1

B\1

V M

li

V

196019601q631064•1964

1961196010'ri21963

r,, ;-Pat-T.ru:rr,i

Cr^ly:I

pond (lake)

0-11- 1 ^)lirnlerrr. 2#3GICT1)AIiita 4

%,ITeM n,tucrn+IiÎ

:3per.ilrrci;nir 1;peICIIIICr:lIli 7

^ 6et;tlricl:rlü i,ri,^^JtI:IITICI;iIiÎ J

^ OeKlrücICIIi! J1

3t)exrlncr;Itii 4A'li a prI0i3-!i a prll.l'r3

3perirlIlcrClTr( J

.31)eKlrücKII12 2

IIUrw:I@HIfA

e ffi,,:np anlucelrr}•urrrLU[HUCriIrio (py.lr,mr(y

Sltrrri<c krr,rco-TU re:ra(l;Fq

•r., ru,r,aa Tto rlocse jo;r,p, ur,inpct oTt)Dj)A or6opa orûoPa

C h-^ h-^_

é,1 ,13-S .S442

41,524,619,4314259

20 ,9474•198354

83,4105 ,0

579

117,065 ,228,3

.'^j42

334

44,5570244364

2,842,b32,71

2,5221652,402,9f2 ,65

2,572,912,962,91

3 ,042:362,84

2,7121672,542,882,76

2,572,962,822,93

2,762,842,92

2,852,983,022,69' ,81^

3,002,842,862,76

IIOc.leor6o )a

k^

2,67

2,84

2,912,912,932,752,80

2,932,822,822,74

Table 4. Exterior indices for fingerlings and 2-year olds ofthe 4th selected generation before and aftér selection for weight. Ropsha.

a"breeding branch; b-generation; c-age (years).; d-average weight (g);

e-Fulton's conditioning factor; f-index of body height (LJH); g-before

selection; h-after selection.

(1)-Kipen' 2; (2)-Bystryanka 4; (3)-Artem`evskii; (4)-Karpov-Karasev;

(5)-Zrekinskii.

water temperature (while at the same time maintaining a high winter resis-

tance), and this task was solved mainly through intensive weight selection

among fingerlings and 2-year olds.

Individual selections in breeding Ropshian carp. In the course of

the selection, but mostly in the 3rd, 4th, and 5th generations, we repeat-

edly evaluated the breeders on the basis of their progeny. Such evaluation

was conducted for two purposes: 1) for the rejection of specimens hetero-

zygous with respect to the s-gene ("scattered" mirror scales) and thereby

eliminate the segmented character of Ropshian carp scales; 2) to identify

the best genotype breeders ("record" types).

13

The results of both efforts have been discussed elsewhere [4, 16--21],

and we shall not return to them here . We should only point out that in

testing the Rops-11iian carp breeders for heterozygote characteristics with

respect to s-genes (over 440), we identifi~_d 236 (53.1%) homozygous (SS)

carp, and this made possible, beginning with the 5th generation, completely

to eliminate undesirable "mirror" fingerlings . Evaluating the breeders by

their offspring made it possible to select a small quantity of elite breed-

ers for raising other'branches of breeders .

Individual selection, not counting selection for homozygosity with /49

respect to the S-gene, did not have an important role in our project .

Comparison of the efficiency of mass and individual selection in carp breed-

ing has shown that individual selection can be more efficient only whe n

the selectional character has a very small value, not exceeding 0 .1--9 .15

[4] . Considerable technical difficulties connected with the evaluation

of breeders on the basis of their offspring or family*selection make wide

use of these methods in carp breeding impossible .

The main method in breeding Ropshian carp has been by mass selection,

--a method that will probably retain its significance in the future as well .

Combined selection . As early as 1966 it was suggested that in th e

interest of efficiency carp should be bred by a composite method, that is

to say, the same generation should be subjected to a mass selection, family

selection, and the males should be evaluated on the basis of thei r

progeny 141 . In practice we managed to combine only the first two methods--

while raising the branch W1 . A mass selection for weight was conducted in

1966-67 among fingerlings and especially among 2-year olds of the 5th selec-

ted generation with a moderate tension of 12 .6 and 13.6% and an intensit y

of selection of 1 .0--l .5 . The family selections involved four crossbreedings 9

14

conducted in 1966, each involving 2--3 females and 3--6 males. In two

of these crossbreedings (crossbreedings 1 and 3) the males and females

came from different backgrounds, in the other two crossbreedings (2 and

4) they came from the same background, one of mass crossbreeding.

The 4th crossbreeding was graded with regards to scale segmenta-

tion in the progeny. The fingerlings of the remaining crossbreedings

were raised in various ponds and their rate of growth could not be com-

pared. We conducted the first evaluation with respect to resistance to

winter conditions; the third crossbreeding turned out to be somewhat more

resistant to winter conditions than the second, despite the lower initial

weight of these fish. The main comparison of the three families was car-

ried out in the second year. Their' weights when first planted were almost

uniform; the evaluation was carried out in three ponds, two of which con-

tained specimens of the three families that had been placed there as year-

lings and one contained specimens of two families. The results indicate

sufficiently clearly that the growth rate of the 2-year olds of the 2nd

crossbreeding lagged behind that of the 1st and 3rd crossbreedings, amount-

ing to an average of 42 and 59 grams, respectively, that is to say, con- /50

stituting more than 10%. They could not be evaluated for vitality owing

to differences in their planting weights and poor preservation of the

labels.

Based on the results obtained from the resiAtance to winter condi-

tions and from the raising of the 2-year olds, the second crossbreeding

was discarded. The two remaining families possessed approximately the

sanie qualities and were kept for further breeding.

The coefficient of tension of family selection can be assumed

to be 66.7% (2 out of 3), since the 4th crossbreeding cannot be taken intO

là

• 'I' a a g a 5

Cemeilumii onlop ua izapttax 5-ru cezettimiottuort) (01130,11M 11M)• coamecruoe ubtpauoutallue

year

age of f ih Itpy

pond ( I ak

Cptutinn nec uNt coi.stnuit r;;• ■ ppeinutp•mmuliblii C;b1:1 ■ ffiC, ib Mml,"cr,a'

C t) ciZ1 1 1 , 114 1 'nlifilirl

a C;b1:1ffitC,

19t;7 2-year o I ds

Ilitttuoncuitii AprmGeitc.uni'l

poucKuii

3 ;)C, KtIFICIarii Tptcx:pyriut 1993 3;)(2 1-zaricurfil I 1203

49$ 357 448

1176

••••;0

526 :359

t;22

327 309 411 595**

Wt3 3-year o l_ç_ls

1 ■■ IppruittrqDplip ,, t - ,a upc‘p.e:o:ult >îa rionpailmi r[..1mm pamilutt, 1 " Paa:itunic c,, c‘;in..1;;;11,,t.iw y m 4.,: -;-,,.:e11:1K) 1;91 yr,,IIHe;

t:Tn

Table 5. Femily selection of the 5th generation in éàïp (6HREh VM). Ropsha, joint breeding.

a-average weight when caught (g); b-average adjusted weight gàih (g5; c-number of crossbreeding; •

*The weight gain adjustment was based on a computed correttiôh of à eâffià for each gram of difference in planting weight.

**The difference with crossbreeding 2 is reliable at a signifiéâhêé of 0.05.

account. The selectional differential (according to the weight Êgilh 8É

the 2-year olds) amounted to 52 grains, and the intensity of âétéét18h ih

this case was 1.5. Given a high inheritability of differénêéà ih

growth rates (in the order of 0.8-0.9), one might have expeétéà iâfâé

shifts in the growth rate of the following generation, aceettdihg tb the

formula of selective efficiency R = S.h2

[5]. However, - thé àêêêlefâted.

growth of carp from the 1st and 3rd crossbreedings may be linked tà Saiite

extent to the greater heterogeneity of the fish from these éteêàbf-éêàihgs,

owing to the more distant relationships among the breeders. Ih this êàse,

weight variations may be due partly to a non-additive propétty, in hih

the inheritability of different growth rates would be signifiéànti

I

and the efficiency of family selection would also be lowered.

The best two crossbreedings (1 and 3) turned out to be distinct

with regards to conditioning and appearance. The average indices for the

3-year olds, which over a period of two years were raised together as

follows:

Coefficient of Body Height, Body thickness,

conditioning, K LfH B/L, %

Crossbreeding 1 2.66 2.86 17.81

Crossbreeding 3 2.89 2.78 18.35

Significance ofdifferential, p 0.001 0.001 0.05

Discussion

The main emphasis in our project fell on mass selection. Carp

breeding, with mass selection, and aided by the fertility of the female

carp, can produce good results. The.production of tens and even of

hundreds.of thousands of offspring by one female makes possible such

abundant fish harvests--yields unimaginable with regards to the majority

of other animals and plants.

But survival of only one specimen in a hundred, or sometimes in

a thousand, can be linked to rapid undesirable changes which would lower

the vital and productive qualities of the fish. Practical experience in

selection has taught us to limit the tension of selection and to conserve

not less than 5, or in an extreme case, 1--2% of all fish. Toorigorous a

selection is a danger testified to by other carp breeders as well. Thus,

Moav and Wohlfahrth [10] showed that a selection of only high-back carp

results in offspring whose distinction is a slower growth rate. Such

correlative changes in selected fish are obviously very widespread and

17

probably in all other animals as well.

Composite selection is seen as a real possibility of increasing

efficiency in carp breeding. This kind of selection makes it possible,

without losing time, to combine mass selection with family selection

(with an evaluation on the basis of the sibs) and to evaluate the breeders

on the basis of their progeny. The advisability whether or not a composite

program should be adopted varies with the circumstances of each individual /51

case, depending mainly on the availability of technical facilities such as

ponds and basins for raising and evaluating a sufficient quantity of young

fish.

We have enumerated elsewhere the conditions that must be observed

in mass selection [4, 16, 19]. Here we shall recall only the two most

important ones: 1) the entire breeding stock must be raised, before the

selection, in the same pond and under identical conditions, including time

limits for reproduction and transplantation into other ponds. Mixing fish

from various ponds before selection (without labelling) is not to be per-

mitted; 2) the selection should be carried out preferably at an age when

the fish would normally be marketable. In the USSR, where carp are market-

ed at the end of their second year, the selection should be carried out

primarily among 2-year olds.

Finally, it should be noted that Ropshian carp of the 6th selected

generation are still rather varied with regards to growth rate and appear-

ance and that this variability, to a great extent, is genetic in nature.

Hopefully, the selection in the 6th and in subsequent generations will re-

tain its efficiency, which should result in still further improvement in

the productive qualities of carp. Contributing to such inprovement will

also be heterozygous commercial crossbreeding, which will be discussed in

our next report.

18

Conclusions

We conducted a mass selection for weight in the 2nd to 6th hybrid

generations resulting from crossbreeding European carp with "eastern

carp". At first the weight of the selection fell on the fingerlings

(age 0+), but beginning with the 4th, and especially the 6th, generation

the selection was conducted mostly among 2-year olds (1+). This shift

became necessary when it was shown that the correlation between the growth

rate of fingerlings and that of 2-year olds in carp was too low.

The tension of selection (the portion of selected specimens ex-

pressed as a percentage) and the intensity of selection (selectional dif-

ferential expressed in terms of standard deviations) reached very high

values in the first generations. Excessively rigorous selections result

in undesirable correlative changes; hence in the 4th to 6th generations

the coefficient of tension was kept at 1-5%.

The uneven weight distribution in carp, which increases with food

shortages, precludes an exact computation of the intensity of selection.

Hence the tension of selection must be considered the main criterion of

the strength of selection in carp breeding.

A high fertility rate in carp, on the one hand, and technical dif-

ficulties in raising and evaluating large numbers of descendants on the

other, combine to make mass rather than individual selection more efficient

in carp breeding. In order to raise the selection efficiency, we used a

composite selection, that is to say, we evaluated several families of the

sanie generation and selected the best specimens from the best families.

The composite selection can be improved upon by including in its method an

evaluation of the breeders (the males) on the basis of the progeny.

This paper contains 5 Tables, 4 figures and 21 bibliographic

19

entries.

Submitted for publication,21 October 1971.

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I •• 20

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