AquaBreeding Project title: “Towards enhanced and sustainable use of genetics and breeding in the European aquaculture industry”




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Review on Breeding and Reproduction of European aquaculture species

Gilthead seabream (Sparus aurata L.)



Hervé Chavanne (Istituto Spallanzani, Italy), Béatrice Chatain (Ifremer, France), Pierrick Haffray (SYSAAF, France), Kostas Batargias (Technological Educational Institute of Messolongi, Greece)


January 20008

1. General information on production and breeding

History of the domestication

Before seabream aquaculture began, the species was reared traditionally in lagoons all along the Mediterranean coast. As with seabass, seabream domestication is recent and started concomitantly with the development of reproduction and husbandry methods in the 1980s. The first breeding programmes appeared in France and Israel in the 90s, followed by a few initiatives based on family selection in Greece and Cyprus [1-3]. A high proportion of seabream fingerlings produced today is derived from unselected broodstocks.


Production (Figure 1)

Seabream production reached 104,000 tonnes in 2006, with major contributions from Greece (47%), Spain (20%) and Italy (17%). The number of fingerlings produced in 2006 was 520 million [4].



Figure 1. Distribution area of wild populations (red) and production countries (green).

Source: FAO Fishbase, FEAP.


Biological features of interest for breeding practices

  • Main commercial data on weight and age at harvest and slaughtering performances

Rearing temperature plays a fundamental role in seabream growth. There is a 5-6°C difference between the extremes of yearly average culture temperatures in Europe (from the Atlantic coast to the eastern Mediterranean or even the Red Sea), and culture duration for a given weight is highly variable among rearing sites. Low winter temperatures (below 12°C) encountered in some rearing sites drastically slows down growth for three to four months during the year. The production time to slaughter may vary from 18-24 months post-hatching, given the above conditions and depending on the time of birth.

More than 40% of the European market volume is represented by portion size fish of 300/400 g, followed by 33% of 400/600 g fish and 14% of 200/300 g fish [4]. Fish above one kilo represent a niche market. As in seabass, Turkish production is characterized by lower fish size while Spanish fish tend to represent larger portion sizes (no 200/300 g fish and 1% of production based on fish above 1 kg).

Seabream is a hermaphrodite protandric species. Maturing males are observed under farm conditions after the first year and sexual inversion usually occurs between the second and third years, but can be later [5, 6].


  • Generation interval for males and females

The maturation pattern is quite clear for males, which all become spermiating within the second year of life. The timescale of the sexual inversion into females is less clear. The evolution of any individual is difficult to predict, with sex inversion occurring generally after 1 to 2 years of male function [5], or from the second to the fourth year of life [7]. Males exceeding 4 kg are commonly encountered in cultured broodstocks. The underlying mechanisms of inversion are still unknown, even if social interactions are though to play a role [1, 8, 9] and can be used to optimize broodfish sex ratios. The breeding scheme proposed by Brown [1] leads to a generation interval of 4 years with a rate of replacement that gives any broodfish three consecutive spawning seasons, though none are kept beyond five years old.
Main diseases

Lymphocystis disease (lymphocystis iridovirus) is a viral infection associated with hypertrophy of connective tissue cells; it induces external clinical signs that make affected fish unmarketable.

High losses are reported on cultured seabream affected by bacterial infections like: Pasteurella piscicida, with typical extensive, acute multifocal necroses in the spleen and kidney; the opportunistic pathogen Flexibacter maritimus, which provokes hyperplasia and erosion of the gills; or vibriosis caused by Vibrio sp., mainly represented by Vibrio anguillarum, characterized by external ulcers and internal haemorrhages. Winter disease (Pseudomonas anguilliseptica) outbreaks, affects both juvenile and adult seabream when the temperature decreases below 12-13°C and can induce high mortalities by hemorrhagic septicaemia.

2. Genetic variability of the species

Wild genetic resources available

The structure of wild seabream populations remains unclear. The first studies reported conflicting data on the presence of panmictic or subdivided populations, as referred to in [10]. From the latest results, a weak but significant structuring pattern is suggested. A fragmentation into subpopulations appears in the western Mediterranean, revealing groups that are genetically different from Atlantic Ocean and Adriatic Sea samples [11]. A genetic differentiation between northern and southern west Mediterranean populations and a trend for lagoon populations to be differentiated from their marine counterparts were suggested by [12] using RAPD and microsatellite markers. Analysing six wild populations from the Atlantic Ocean and Mediterranean Sea, [13] also observed also a slight degree of differentiation but without association with geographic factors. A small scale study revealed a genetic structure between the Adriatic Sea, the Tyrrhenian Sea and the Sardinian channel [10]. The species subdivision still needs to be clarified, in particular through further analysis covering the whole species distribution and also seasonal or year to year variation.


Differences between wild and/or domesticated populations

Comparing six wild and five cultured populations using molecular markers, [13] and [14] reported higher heterozygosity in wild individuals and a slight loss of genetic variation in cultured stocks. Microsatellite and AFLP markers were presented as useful tools for tracking the geographical origin of individual breeders [15, 16]. Further efforts have been made to set up analytical methods based on chemical and isotopic fingerprinting to differentiate between wild and farmed individuals [17].

Strain comparison studies lack for seabream, as only one experiment has reported performance differences (based on weight) between Atlantic and Mediterranean strains [18].
Interaction between wild and domesticated stocks

The introduction of non autochthonous populations occurs through the exchange of commercial fry all over Europe and escapes during on-growing phases (no monitoring of escapees is made for seabream), a phenomenon that may be further enhanced when farmers rely on non-local broodfish [3]. Moreover, females can spawn a large quantity of eggs in the cages and increase genetic escape by this means. The success of survival and reproduction of escapees is unknown, and many factors could limit this (size at escape, prey abundance in the season of escape, adequate environment, etc). However, the use of local stock was not supported by experts (see conclusions of GENIMPACT: http://genimpact.imr.no/compendium) as the changes in performance associated with the selection are expected to be similar between local and non local stocks. Sterilisation by triploidisation or hybridization is recommended as a way of limiting the risks associated with escapement.


Inbreeding effects

Few data are available to assess the effects of inbreeding on seabream. Only [9] have produced progenies by self-fertilization of eggs with the frozen sperm of the same individual, and subsequent survival at hatching was very low.



3. Reproduction

Fecundity and main reproductive features

Seabream is a protandric hermaphrodite species with asynchronous ovarian development, spawning at 24 hour intervals over a period of up to three months in conditions of captivity [19].

Brown [1] noted that in the adult stages, faster-growing fish normally undergo sex reversal earlier than the slower-growing remainder of a population: a maturation process that could lead to large sex ratio imbalances in selected broodstocks. [30] calculated a genetic correlation of 0.90 between body weight and sex reversal while the phenotypic correlation was 0.31, supporting the hypothesis that fast-growing fish have the intrinsic tendency to reverse sex earlier (high genetic correlation) but this is influenced by non-genetic factors (medium-low phenotypic correlation).
Cryopreservation

The milt characteristics of seabream contrast with those of seabass by a long duration of motility (22 to 50 min) and a relatively low sperm concentration (0.72 to 26 x 109 spz.ml-1) [20, 21]. Different cryopreservation protocols have been proposed [20, 22-25], with similar fertilisation results after freezing [22].

Research is being conducted on embryo storage through microinjection of different cryoprotectants into the yolk sac of seabream embryos, as are the relevant toxicity studies [26].
Genetic and environmental sex determination, sexual dimorphism

Studies on the physiological effects triggered in females by the removal of males from a group of spawning fish, [8, 9] concluded that an endocrine response to socio-sexual stimuli exists during the reproductive process. Brown [1] supported the view that the sex ratio is socially determined in this species and noted a tendency of the broodstock to approach an equilibrated sex ratio regardless of the number of fish replaced.

Three forms of gonadotropin-releasing hormone (GnRH) controlling the final oocyte maturation, ovulation and spawning have been isolated in seabream [27]. Long acting synthetic GnRH agonist (GnRHa) treatments have proved to be efficient in increasing egg production, with daily peaks of 105 eggs kg-1 female body weight. However, such treatments perturb the 24 h ovulatory cycle displayed by untreated females. This unstable duration of ovulatory cycles, combined with the daily fluctuation in volumes of spawned eggs and the short time during which oocytes remain fertile (1-2 hours [28]), make artificial fertilization difficult to perform in this species [19].

4. Selection

Genetic variability per trait

In spite of the important commercial value of seabream, little attention has been paid so far to experiments aiming to estimate genetic parameters in seabream. The first results, published by [29], give a realized heritability for weight of 0.34 ± 0.02 at 2 years of age using replicated selected lines. Using an animal model, [30] calculated a heritability for weight 0.40 ± 0.20, 0.52 ± 0.22, 0.55 ± 0.18 at 6, 10 and 22 months old, respectively. The heritability of the gutted body weight was high (0.51) whilst the estimate of dressing was lower (0.31) [30]. The genetic correlations with body weight at 22 months old were very high (>0.90) [30]. More recently, [3] reported high heritability for body weight (0.61 ± 0.06) and moderate to low hertiabilities for external colour and spinal deformities (0.21 ± 0.02 and 0.12 ± 0.02 respectively).

The heritability estimates of the morphometric characters displayed a wide range [30]. The estimates for the lengths ranged from 0.11 to 0.36. The heritabilities of body heights and width were higher and ranged from 0.32 to 0.54. The genetic correlations among them and with the body weight at 22 months were very high (>0.90).

A genetic component was suggested as an explanation of the family dependant structure of spinal column deformity [31].


Genetic responses, progresses and control lines

Industry records report a selection response ranging from 5 to 10% per generation [32]. Thorland et al. (2006) [3] provided the first predictor of response to selection for weight, based on an estimate of the selection differential, that reached 22% for the first generation. Similar results (23%) were obtained in a field test comparing the selected F1 progeny with a control line.

A phenotypic deviant (ebony), controlled by a simple recessive Mendelian factor, has been isolated by the IOLR [32].
No information is available regarding G*E interactions and dominance effects in seabream.

5. Polyploidisation and monosexing and hybrids

Triploid induction and performances

Triploidy has been successfully induced in seabream by cold shock treatment, which produces a high percentage of triploids (90-100 %) as confirmed by karyological analysis [33] or flow cytometry [34]. Hatching rate of triploids was significantly lower than diploid controls [33]. Triploids develop as males but do not sex reverse to females as diploids do. Their spermatocytes are blocked at meiosis II and do not develop any further [34]. Until the beginning of the natural sex inversion of diploid males to females (480 g, which is also the most frequent commercial weight), triploids do not exhibit a difference in growth, survival, processing yields or quality traits compared with diploids [34]. For larger sizes, triploids have a lower growth performance but are leaner and have a better gutted yield. Comparative spawning behaviour now needs to be evaluated in sea cages [34].


Gynogenesis, androgenesis and mitotic clone performances

Progeny have been obtained by self fertilisation, which are genetically equivalent to meiogynogenetics or androgenetics [9]. Meiogynogenetic seabream were produced by cold shock treatment [33, 35]. Similar fertilization rates were obtained using homologous and heterologous semen (from seabass), while the hatching rate of meiogynogens was lower than that of diploid controls.


Interspecific hybridisation

Diploid or triploid hybrids between Sparus aurata and the other Sparids: Diplodus puntazzo, Diplodus vulgaris, Pagrus major or Dentex dentex are sterile and have limited survival rates or poor growth [36-39]. Viable hybrids were also produced by an artificial hormonal-induced cross involving seabream and red porgy (Pagrus pagrus). The best hybrids (crosses between female seabream and male red porgy) had too low a performance for aquacultural use [40], although these fish had normal gonad development either as immature females or maturing males [41]. Hybridization between gilthead seabream females and white seabream males (Diplodus sargus), undertaken in a commercial hatchery trial, resulted in high rates of fertilization and survival. Enhanced aggressiveness was observed in post-larvae.

The conditions necessary to obtain triploid hybrids with gilthead seabream have been evaluated [36-38, 42], though none of these experiments produced data on early survival or growth performances.
Tetraploid induction and performances

Not documented.



6. Genomics

The haploid genome of the seabream consists of 24 pair of chromosomes [43], that weigh 0.95 pg [44] and contains approximately 930 Mb. A first-generation genetic linkage map based on 204 microsatellite markers distributed over 26 linkage groups and covering 1242 cM (Kosambi function) is available [45]. A radiation hybrid map made of 440 markers (288 microsatellites, 80 gene based markers and 70 STS) counting 28 RH groups with a combined size of 5680 centirays was published the same year [46]. Numerous evolutionarily conserved regions were found with the Tetraodon nigroviridis genome on both maps. Integration of additional genomic data will contribute to building more robust genomic maps.

Clone sequences obtained from embryonic/larval stages and liver of seabream produced two distinct cDNA libraries which led to a collection of both 5’ and 3’ ESTs [47]. Today, the total number of available ESTs is estimated at up to 5.000 [48].

Tools to evaluate population genetic variability

First studies based on mtDNA variation [49] were followed by work using allozymes, RAPDs and microsatellite markers [10-14, 50]. With the rapid development of genomics resources [46] range of available markers is increasing.

Solutions have been proposed to construct new broodstocks for commercial hatcheries, based on the genetic variation shown by sets of microsatellites [51].
Genetic markers for genealogical traceability

More than 30 primers of polymorphic microsatellites have been published by different authors for the seabream [52-56, 56]. The recent development of a radiation hybrid map added another 288 microsatellite markers, among which 134 have been shown to be polymorphic [46].

The large number, variability, stability and low genotype error associated with SNPs make these markers potentially useful for fingerprinting, but their relative higher cost is still slowing down their diffusion (4 to 5 more SNPs than microsatellite markers are needed for similar discriminatory power). The screening of 76 new SNP markers is reported by [57].
QTL and Marker Assisted Selection

The development of genetic maps is recent [45, 46], and has not yet permitted the identification of QTLs in the seabream.


Transgenesis

Trials have been made in Israel.


References

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