Iranian Journal of Fisheries Sciences 11(1) 214- 222 2012 Genetic parameters estimation of growth in trutta caspius as a function of body weight and Length

Yousefian M.1*, Hosseinzadeh-Sahafi H.2, Golshahi, H.3, Laloei, F.1, Tagavi, M.3;Taheri A.4, Seidanloo Y.4

Received: March 2011 Accepted: June 2011

Abstract There is no available adequate information on the biology and genetics of the Caspian salmon (salmo truta Caspiu) living in the Caspian Sea with other species such as common carp families, sturgeons and etc., therefore we are interested in knowing whether selection for rapid growth rate in salmo trutta Caspius will potentially lead to correlated genetic response in body weight and length. The potential for correlated responses was evaluated by the heritability estimation, estimating heritabilities of body weight and length. The genetic parameter estimations were obtained for different stages of growth from 5400 fish consisting 27 unrelated families that reached maturity at 36 months of age. The parent which used to produce separate family groups had been caught from the Caspian Sea and spawned during the autumn season. The estimated heritability of body weight and length was 0.42±0.13 and 0.49±0.01, respectively based on differential and selection response between parent pairs and their offsprings. Several factors may have affected the creation of this high level of estimation, dominant effect, inherited from different breeders to offspring.

Keywords: Salmo trutta caspius, Genetic parameters, Heritability, Caspian Sea

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______1-Caspian Sea Ecology Research Center, P. O. Box: 961, Sari, Iran. 2-Iranian Fisheries Research Organization, Tehran-Karaj Highway, Sarve-Azad, P.O.Box: 14155-6116 Tehran, Iran. 3- Shahid Rajaei Propagation center for Fresh water fish, Sari, Iran. 4-Fish Disease Dept., Faculty of veterinary, University of Tehran, Tehran, Iran. *Corresponding author's email: [email protected]

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Introduction The genetic management of breeding analysis of developmental stage of this stocks is very important in order to fish. ensure proper and long-term There are several technical harvesting of the breeders. Increasing studies in genetic analysis of the production and growth rate are Salmonids. In a study done by usually the main traits to be improved Ferguson et al., 1985 the effect on by individual selection of breeders, hatching time rainbow was without having any information of found. Also, Smoker in 1986 observed genetic basses of fish which usually a small maternal effect and a the farmers have special interest in, significant sire and dam effect in this since it is simple and cheaper to set up fish. In , the heritability of in practical condition while family early survival is about zero (Kanis et selection seems to be more effective al., 1976). No such data for Salmo (Gjedrem 1983; Hershberger et al., trutta caspius are reported in literature 1990). at this time. In the present study, in The Salmo trutta Caspius is a order to better define a set up of the very important fish in the Caspian Sea response to selection of Salmo trouta and due to its high market price and as Caspius, an experiment was designed an alternative of , it is of to determine the genetic parameters of interest to be propagate and rearing. growth-related traits at different stages The Salmo trutta Caspius is a very of development. slow fish at early development. We Materials and methods designed a program to enhance The prepared fish for the present study individual selection. Growth rate is the were reared in fresh water during the main breeding goal of fish breeders, whole experiment at Propagation and but individual selection has often Culture Station S.B. Kelardasht, Iran. In shown poor responses in fish species. December 2006, 27 mature 3-6 year old The technique introduced by female fish were artificially striped and Chevassus and his teamwork in 2004, mated with 27 males with the same ages, was developed Archive to overcome possible in whicofh three SIDmales were mated with one factors that may contribute to this low female. A total of 27 unrelated families success using a high number of were produced as the base population of breeders and repeated growth gene bank. Fertilized eggs were incubated challenge. This technique was used by separately for 30 days, after which, they some modification to establish the were transferred into indoor 150-L tanks lines as gene bank for managing a (2000 larvae in each tank). The order of cross breeding design for restocking egg tray and larval tanks were randomized the fish in the Caspian Sea. Yet, there in incubation and rearing facilities. In are not any references on the genetic spring 2007, in each family tank, family

www.SID.ir Iranian Journal of Fisheries Sciences, 11(1), 2012 216 size was standardized to 200 individuals. SS♀♀ = ab Σ( X ♀- X )2; During rearing in tanks, the fish were fed SS♂♂ = n Σ ( X w- X ♀)2; with commercial dry pelted feed (Chine product, Tehran, Iran), composed of 45% SSw =Σ (X- X w)2; protein, 21% fat, 1% fiber and 7% ash. SSph=Σ (X- X )2, Water oxygen concentration was where X is a common mean for all fishes, maintained above 95% over the duration X of the experiments daily. The temperature w mean values for different offspring. of the experimental station was not X ♀ mean values for different offspring of constant from 2C in winter to 15C in one and the same female X is individual summer. Therefore the amount of feed was measurements, n is the number of adjusted for temperature and fish biomass individuals in a given offspring. a is the in the tanks and in the pond. Further, the number of females. b= the number of condition of the fish was monitored at least males crossed with one female. twice. The fish were weighted and The so-called observed variances (MS or measured at different stages of growing. Y) may then be calculated. They are The hierarchic complex was employed. obtained after the division of the sum total The first experiments with hierarchical of the squared values by the number of complexes were performed by Nenashev degrees of freedom: a-1 for SS♀♀.a (b-1) (1966, 1969) with the common carp.The for SS♂♂. ab(n-1) for SSw and abn-1 for external fertilization of eggs in this fish SSph. After this division we obtain: and the high fertility of females facilitate V♀♀ or VD- the variance of mean values the simultaneous conduct of a large for females (dam component); number of crosses. In the hierarchical V♂♂ or Vs - the variance of mean values complex one female from several different for males crossed with one female (sire males are half sibs with respect to each component); other, while the individuals within each V random or VW -the variance of given offspring represent full sibs. An differences within separate offspring: analysis, of the variation of different V Ph - the variance for all descendants. groups of offspring makes it possible to Each of these observed components of calculate the heritabilityArchive of the trait under 1111: of variance SID is non- uniform and study. In the treatment of lengths or weight contains a number of components of measurements for a limited number of different origin (Falconer, 1960): individuals originated from each cross the V   2  n 2  bn 2 ; D W S D first task involves calculation of the sum V   2  n 2 ; V   2 total of square deviations from the mean S W S W W Then it is easy to determine the values for (SS) separately for females. For males it  2 2 crossed with one and the same female s and  D within different broods and for all the 2 VD Vs 2 VS VW  D   S  fishes measured. The calculations are bn ; n made using the following formulae:

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 2 2 additive genetic variation of the patents. Each of the variances s and  D Heritability can be obtained from the contains. as calculated, one quarter of the following formulae:

Results weights at the four time of sampling The mean weight and length of were 0.8, 4.5, 21.3 and 98.6 g. Also, offspring, based on family crossing, at the average length s of fish during four different time of sampling are shown times were 4.2, 7.7, 12.5 and 21.2 cm, in table 1. The average over all fish respectively.

Table1: Mean and standard deviation of weight and length at different age of growing (n=30 sample number) th th th th Time of 1 spring 1 summer 1 winter 2 spring sampling

Mean± SD Mean± SD Mean± SD Mean± SD Weight(g) 0.8±0.4 4.5±1.1 21.3±6.7 98.6±21.8 Range(g) 0.3-1.8 2.4-6.3 7.9-33.8 67.1-150.4 Length(cm) 4.2±0.6 7.7±0.9 12.5±1.6 21.3±3 Range (cm) 3.4-5.6 6.4-10.8 9-16 18.2- 33.7 No of offspring 5400 5082 5011 3968

The survival rate was 92% over year stages. It gave the same results for rearing out of 5400 fingerlings. Table length of fish at different sampling 1 shows the high variation between stages. The heritability estimation for weight and length at different families growth parameter at four interval at the first stage of growing. A periods are presented in table 2. comparison was done on mean weight Heritability of growth was quite high of the offspring in different families in Salmo trutta Caspius of the Caspian (Fig. 1). The difference was highly Sea. significant (P<0.00) at all sampling

Table2. Heritability estimation and additive genetic correlations TimeArchive of Sampling for length of SID th 1 spring 0.5021* th 1 summer 0.0229 0.4909 th 1 winter 0.0498 0.1112 0.5087 th 2 spring 0.1333 0.0229 0.0642 0.5070

Time of Sampling for weight th 1 spring 0.4421 th 1 summer 0.0715 0.4674 th 1 winter 0.0025 0.0159 0.5024 th 2 spring 0.0022 0.0854 0.0551 0.5043 *The italic number shows the heritability estimation

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Weig ht at firs t s pring Weig ht at firs t s ummer 2.50 8.00

2.00 6.00 1.50

4.00 1.00 weig ht (g ) weig ht (g ) 0.50 2.00

0.00 0.00 1 2 3 4 5 6 7 8 9 101112131415161718192021222324252627 1 2 3 4 5 6 7 8 9 101112131415161718192021222324252627 S almo trutta cus pius families S almo trutta cus pius families Weight at first winter Weig ht at s ec ond s pring 200.00 40.00

150.00 30.00

100.00 20.00 weig ht (g )

weight (g) weight 50.00 10.00

0.00 1 2 3 4 5 6 7 8 9101112131415161718192021222324252627 0.00 1 2 3 4 5 6 7 8 9 101112131415161718192021222324252627 S almo trutta cus pius families Salmo trutta cuspius families Figure 1: The variation in fish weight at different stage of sampling between 27 families of Salmo trutta Caspius

Discussion Genetic studies have revealed According to McKay et al., (1986), moderate levels of genetic variation the heritability estimated in rainbow for growth rateArchives of salmonids, trout of for weightSID and length ranged allowing its genetic improvement from 0.13±0.17 to 0.38±0.22, (Gjerde and Schaeffer, 1989; respectively. Heritability based on Crandell and Gall, 1993b; Elvingson the sire component of variance for and Johansson, 1993; Winkelman rainbow trout traits were 0.21 for and Peterson, 1994). The estimation body weight and 0.18 for body of heritability at different stages of length (Gjerde and Schaeffer, 1989) rearing and condition were varied (Crandell and Gall, 1993a). The based on the several experimental heritability at swim-up in brown works done on Salmonids. trout (salmo truta fario L.) for initial

www.SID.ir 219 Yousefian et al., Genetic parameters estimation of growth in Salmo trutta caspius… weights was h2= 0.23±0.13 (Wallace and Aasjord, 1984; (Vandeputte et al., 2002). In spite of Vandeputte et al., 2002). the high variation in genetic The breeders used in the components of Salmonids, in the present experiment had a different present work the heritability at age and weight, therefore the eggs of different times of sampling was some females were smaller than the approximately in the same range. others, consequently such effect There was high family persisted at swim-up stage. Analysis variation on first stages of growth in variance also revealed significant salmo trutta Caspius (0.3-1.8 g). differences between different Several factors may have affected families. In addition, the basic the creation of this high variation. protein energy requirements of The variable between different Salmonids showed that a significant family individuals may be growth improvement can be obtained considered as dominance effects, when using specific diets with inherited from different breeders to protein corresponding to the offspring. Dominance effects for all requirement of the species. In this individual variables were moderate experiment we used the trout diets, to large (0.14-0.39) for brown trout which may be far from the optimum (salmo trutta fario L.) which is need of brown trout. Up to now, concluded in the work of Vandeputte precise monitoring of the food et al., 2002. conversion index for Brown trout of In spite of having the the Caspian Sea was not obtained experiment in a common precisely by experimentation. environment and randomizing the Therefore, the experiment may not experiment to eliminate have been performed under optimum environmental effectArchives, yet the eggs environment of SID conditions to estimate of each family were incubated the correct heritability for this fish. separately, which may have The effect of breeders may permitted the onset of a common also be explained by the effect of environment different for each inbreeding depression which family. The large dominance effect occurred in offspring, that is in is based on the fact that accelerated accordance with Quillet, 1994. He larval growth allows an earlier onset stated the fact that growth rate is of exogenous feeding, and improves especially affected by inbreeding the survival during this phase depression during early stages which

www.SID.ir Iranian Journal of Fisheries Sciences, 11(1), 2012 220 is linked to the existence of oriented Selection Evolution. 36, 643– dominance variation. 661. The lower variability and Crandell, P. A., Gall and G. A. E., heritability of domesticated strain in 1993a. The effect of sex on many cases of Salmonids indicates heritability estimates of body that they were propagated with small weight determined from data on effective numbers of breeders, individually tagged rainbow resulting in some inbreeding. This trout ( mykiss). initial inbreeding in the domesticated Aquaculture, 113, 47– 55. strains could somehow hamper the Crandell, P. A., Gall and G. A. E., possibilities of genetic gain from 1993b. The genetics of body selective breeding (Vandeputte, weight and its effect on early 2002). maturity based on individually The Brown trout of the tagged rainbow trout Caspian Sea is quite distant from (Oncorhynchus mykiss). other species of brown trout in Aquaculture, 117, 77– 93. Europe and may then have Elvingson, P. and Johansson, K., conserved different alleles 1993. Genetic and environmental influencing the species with high components in body traits of genetic variability or/and heritability rainbow trout (Oncorhynchus which in this species may be a good mykiss) in relation to age. material to start selective breeding Aquaculture 118, 191–204. programs with. Ferguson, M. M., Danzmann, R. G. and Allendorf, F. W. 1985. References Developmental divergence Chevassus, B., Quillet, E., Krieg, among hatchery strains of F., Hollebeq,Archive M., Mambrini, ofrainbow SID trout (Salmo gairdneri) M., Faure, A., Labbe, 1; Pure strains. Can. J. Genet. Laurent., Hiseux, J. and Cytol. 27, 289-297. Vandeputte, M., 2004. Gjerde, B. and Schaeffer, L. R., Enhanced individual selection 1989. Body traits in rainbow for selecting fast growing fish: trout. II. Estimates of the “PROSPER” method, with heritabilities and of phenotypic application on brown trout and genetic correlations. (Salmo trutta fario) Genetic. Aquaculture 80, 25–44.

www.SID.ir 221 Yousefian et al., Genetic parameters estimation of growth in Salmo trutta caspius…

Gjedrem T., 1983. Genetic variation different characters in fishes. in quantitative traits and Genetika,11, 100–108. selective breeding in fish and Nenashev, G. A., 1969. Heritability shellfish. Aquaculture, 33, 51– of some selective characters in 72. Ropsha carp. Izvestija Gosud. Herbinger, C. M., Doyle, R. W., Nauchno-issled. Inst. Ozern. Pitman, E. R., Paquet, D., Recn. Rybn. Hos (GosNIORH) Mesa, K. A., Morris, D. 65, 185–195. B.,Wright, J. M. and Cook, D., Quillet, E., 1994. Survival, growth 1995. DNA fingerprint based and reproductive traits of mitotic analysis of paternal and maternal gynogenetic rainbow trout effects on offspring growth and females. Aquaculture, 123, 223- survival in communally reared 226. rainbow trout. Aquaculture, 137, Vandeputte, M., Peignon, E., 245–256. Vallod, D., Haffray, P., Kanis, E., Refstie and T. and Komen, J. and Chevassus, B., Gjedrem, T., 1976. A genetic 2002. Comparison of growth analysis of egg, alevin and fry performances of three French mortality in salmon, strains of common carp and rainbow trout. Aquaculturem (Cyprinus carpio) using hemi- 8, 259-268. isogenic scaly carp as internal Kirpichnikov, V. S., 1981. Genetic control. Aquaculture, 205, 19– Bases of Fish Selection. ISBN. 36. 3-350-10911-0-Springer-Verlag. Wallace, J. C. and Aasjord, D., New York. P. 410. 1984. An investigation of the McKay, L. R., Ihssen, P. E. and Friars, consequences of egg size for the G. W., 1986.Archive Genetic parameters of ofculture SID of Arctic charr, growth in rainbow trout Salmo alpinus (L.). J. Fish gairdneri, as a function of age and Biol. 24, 427– 435. maturity. Aquaculture, 58, 241-254. Winkelman, A. M. and Peterson, Moav R. and Wohlfarth G. W., 1976. R. G., 1994a. Heritabilities, Two way selection for growth rate dominance variation, common in the common carp (Cyprinus environmental effects and carpio L.), Genetics. 82, 83-101. genotype by environment Nenashev, G. A., 1966. The interactions for weight and determination of heritability of

www.SID.ir Iranian Journal of Fisheries Sciences, 11(1), 2012 222 length in Chinook salmon. Aquaculture of Cyprinids. Aquaculture, 125, 17–30. INRA, Paris, pp. 195–208. Winkelman, A. M. and Peterson, Wohlfarth, G. W., Moav, R. and R. G., 1994b. Genetic Hulata, G., 1987. Breeding parameters (heritabilities, programs in Israeli aquaculture. dominance ratios, and genetic In: Tiews, K. (Ed.), Proceedings correlations) for body weight of World Symposium on and length of Chinook salmon Selection, Hybridization, and after 9 months of saltwater Genetic Engineering rearing. Aquaculture, 125, 30– inAquaculture, Bordeaux 27–30 36. May 1986, vol. 2. Heenemann Wohlfarth, G. W., 1986. Selective Verlagsgesellschaft mbH, Berlin, breeding of the common carp. pp. 393–405. In: Billard, R., Marcel, J. (Eds.),

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