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




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2. Genetic variability of the species

Wild genetic resources available

Today, it seems doubtful that real wild resources are available, and all “wild” carp in nature are thought to be of feral origin, at least in Europe [6]. In East-Asia, carp from the Amur river are thought to be wild carp. They are known for their cold tolerance. The original Western limit of common carp natural distribution was the Danube basin [6]. Population genetics studies show that there are two subspecies of common carp, C. carpio carpio from Europe and C. carpio haematopterus from Asia [3, 6-8]. The Central Asian “subspecies” (sometimes called C. carpio aralensis) is very close to the European one [3]. Within each subspecies, the genetic distance between populations is very low [8, 9].


Differences between wild and/or domesticated populations

Genetic variation within domesticated stocks is lower than within feral populations (4.4 alleles/microsatellite vs 8.2 in “wild” populations [3]. Phenotypically, wild/feral carp are usually long, torpedo shaped, fully-scaled fish. European domesticated strains are more humpbacked, which in some cases (Aischgrunder carp) is caused by a fusion of the proximal vertebrae [10]. Scale patterns are highly variable. Most European carps are of the mirror carp type with much only few large scales. The trend is to breed for strains with as few scales as possible (the “framed mirror” type). Body shape and number of vertebrae show phenotypic plasticity, and are dependent on water flow (pond or river) and temperature.



Interaction between wild and domesticated stocks

There are no studies on such interactions. However, the existence of wild common carp stocks in Europe is questionable (see above)



Inbreeding effects

Inbreeding is suspected to lead to decreased growth rate and viability (reviewed in [11]). High levels of inbreeding have been achieved through gynogenesis (e.g. [12, 13]), and affected reproductive traits (lower fertility and fecundity). It is not clear whether the rate of performance loss would be the same under slow cumulative inbreeding.



3. Reproduction

Fecundity and main reproductive features

Gonad weight in mature females is between 15 and 30 % of total body mass. Males have relatively large testes (5-10% of BW). Egg size averages 1000 eggs /gram (depending on age). Carps reproduce in spring time, and spawn only once in temperate climates. Under laboratory conditions, (see earlier) carp females can be stripped every 8 weeks. Males can be stripped every 10 days. Ovulation is induced with carp pituitary, either as crude preparation, or as extract, in two injections. Stripped eggs have limited storage time (2-4 hours at room temperature). Cooling of eggs is not recommended. Milt can be diluted in extenders, and stored at 0-4ºC for several days. Eggs are degummed prior to fertilization and incubated in conical upflow jars. Hatching is temperature dependent and takes place after 3-5 days of incubation (range for 18 - 25ºC).


Cryopreservation

Adequate methodologies for sperm cryopreservation are available [14]. Live gene banks of common carp exist in Szarvas (Hungary), Vodnany (Czech Republic) and Golysz (Poland).


Genetic and environmental sex determination, sexual dimorphism

Male and female carp have similar phenotypes. Sex determination is male dominant, with females being “XX” and males “XY” [15, 16]. YY males, produced by androgenesis are fertile and sire all male offspring. Sex determination is labile, at least in females, and hermaphrodites are frequently observed. Sex can be changed by exposure to endocrine agents [17] but probably not by temperature [15]. A recessive gene, causing male sex reversal in females, was identified in clonal carp [15] and is thought to be related to low cortisol production [18].



4. Selection

Genetic variability per trait

Scale cover variation is governed by a 2 locus system S/s - N/n with epistatic interaction. Homozygous recessive ss animals have a mirror type scale pattern. The N gene reduces scale cover even more but has pleiotropic effects on viability, development and fin shape [11]. Homozygous NN fish die at hatching, which is why the N gene has been removed from most commercial carp strains, except Koi. Some colour variants in Koi have been studied and also have a Mendelian basis [11, 19].

Different cultured strains of common carp have been tested for their productivity traits. In some cases, very high differences can be seen between the strains (1:1,3 to 1:2,0 for growth - [2, 20-22] ; 1:1,3 to 1:2,1 for survival - [23-25]). Occasionally, differences between strains were reported for other traits: resistance to Koi Herpes Virus [26], natural antibodies level [27], and seine catchability [28].

Many estimates of heritability for growth rate have been published, but are not fully reliable due to confounding of environmental effects and small designs. A mean value of 0.3-0.4 for heritability of weight seems reasonable [29], and is confirmed by a study using a marker-based pedigree [30]. Genetic variability of quality traits has been studied recently and shows good heritability values [31]: h²=0.58 for fat-meter value, 0.28 for gutted weight, 0.38 for fillet with skin. Other traits for which genetic parameters are available are stress response (h²=0.60, [32]), condition coefficient (h²=0.3-0.4, [30, 32]), female gonad weight (h²=0.75, [33]) and percent deformed larvae (h²=0.31, [33]). Heritability of mouth and fin deformities seems to be very limited [34].


Genetic correlations and undesirable side effects

Very few genetic correlations have been documented in common carp. The correlation between growth and shape seems to be unstable, sometimes absent [35], sometimes negative (-0.17 between K and weight, -0.38 between K and length [30]). A positive correlation exists between growth and muscle fat content (+0.6-0.7 [31]), and a strong negative genetic correlation was found between relative head length and skinned fillet yield (-0.86 [31]), opening the possibility to select for improved fillet yield on individuals without killing the candidates.


G*E interactions

G*E interactions have been documented essentially in comparisons of Chinese and European carp strains, and their hybrids. Strains show GXE, were environment is defined in terms of pond management, and measured by the mean growth of different genotypes in a given pond. These studies show that growth differences between carp strains can be modulated by the environment, to the point that rankings may be modified [2, 36, 37].


Genetic responses documented, progresses and control lines

While many breeding programs have been operated in the past on this species all over Europe and in Russia, most of them were not reported in scientific journals with all necessary details on the protocols and results. Until now, no well documented breeding project has shown any significant improvement of growth rate (reviewed in [29], see also [38]). Response to selection has been obtained for Height/Length ratio (0.33-0.47 realized heritability, [35]) and resistance to dropsy (0.15-0.20 realized heritability, [39, 40]).


Dominance and intraspecific crossing

As most lines have been kept probably with low effective sizes, and selective breeding has not proven efficient till now, crossbreeding and search for heterosis have been widely applied in common carp (reviewed in [41]). Heterosis has been demonstrated on growth rate [20, 24, 25, 42] and survival [24, 25] mainly, with one result on flesh yield [43]. In these experiments, heterosis can account to 20-40% of the trait mean, except on flesh yield (2.7%). Recent results suggest that the relative weight of additive and dominance variance could largely vary over development time in carp [44].



5. Polyploidisation and monosexing and hybrids

Gynogenesis, androgenesis and mitotic clone performances

Many uniparental reproduction experiments have been performed in common carp, including meiotic and mitotic gynogenesis and androgenesis (see reviews by [45] and [46]). Doubled haploid (DH) carp can be produced by gynogenesis and androgenesis but with low yields (1-15% of treated eggs). Fertility of DH females is often problematic, but males have good fertility.([47, 48]. Some clones have been produced and are maintained by Wageningen Univeristy, The Netherlands. They are used for immunological and stress physiological research and genetic studies (reviewed in [46]).


Triploid induction and performances

Induction of triploidy has been optimised, using cold and heat shocks, in the 1980s-1990s (see review by [45]), and reliable methods (80-100% triploids) have been developed, and made applicable to hatcheries [49]. However, triploids are clearly outperformed by diploids for growth and survival [50], and have therefore never been used in practice.


Tetraploid induction and performances

Tetraploidy has been achieved [51, 52], but with very poor survival, and hence tetraploids have never been used in practice.


Interspecific hybridisation

Interspecific hybridization has been tested with many other cyprinid species (reviewed in [41]) . Tench x Carp hybrids are viable but have not been used in practice. The only hybrid which has shown sufficiently high performance to be cultured in significant quantities in China is the crucian carp x common carp hybrid.



6. Genomics

The carp genome typically consist of 96-100 chromosomes and is believed to originate from a relatively recent genome duplication event. Some traits still show tetraploid inheritance (e.g. blond, in [47]). There is no genome map of common carp. A preliminary linkage map of 272 markers is available, including 105 gene markers, 110 microsatellites, and 57 RAPD [53]. A comparative carp- zebrafish EST database has been published [54]. Some duplicated genes show high levels of redundancy and neofunctionalization (e.g. CYP17, Komen et al., in preparation).


Tools to evaluate population genetic variability

Many types of markers have been used to estimate genetic variability of carp populations, including allozymes [7-9, 55], mt-DNA [56], AFLPs [57] and microsatellites [3, 9, 57-59]. There are no reports on the use of SNP’s in carp.




Genetic markers for genealogical traceability

Scale cover genes may be used to identify genetic groups and to produce internal control lines in strain testing protocols [36;60]. Many microsatellite markers are available (e.g. [57;61]), and have been used with variable success (75-100% unique assignment) for parentage identification [30;31;62]. Problems are occasional tetraploid inheritance, null alleles and limited numbers of alleles.


QTL and Marker Assisted Selection

There are no systematic studies on QTL detection for commercially interesting traits in carp. Gynogenesis and androgenesis can be used for linkage analysis and QTL were detected for growth, length, and glucose levels after stress in progeny from domesticated X feral carp [63]. A QTL for cold tolerance has been mapped in a cross with Amur carp [53].


Transgenesis

Growth hormone transgenesis has been done in several instances in common carp [64-66], and their evaluation for growth rate gave very variable results. When large growth gains are announced, either there is no valid control line and a short-term growth experiment (8 weeks) in aquaria [67], or the sample size is very small (10 transgenics, 10 non transgenics), the growth period very short (3 weeks), and the rearing system very specific (1 fish per aquarium), as in [68]. On longer periods in ponds, the growth gains are moderate (+17% on average, [69], same growth in summer, higher growth in winter, [70]). Therefore, we can consider to date that proven growth gains with transgenic carps remain moderate. To date and to our knowledge, no transgenic carp is produced for human consumption anywhere in the world, including China where quite restrictive regulations have been taken for the evaluation of this possibility [71] .



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[71] C. Fu, W. Hu, Y. Wang, and Z. Zhu, "Developments in transgenic fish in the People's Republic of China," Rev. sci. tech. Off. int. Epiz., vol. 24, pp. 299-307, 2005.





Review on Breeding and Reproduction of European aquaculture species

Charrs (Salvelinus alpinus and Salvelinus fontinalis)



Salvelinus alpinus Salvelinus fontinalis

Pierrick Haffray (SYSAAF, France)


January 2008

1. General information on production and breeding

History of the domestication

Charrs are salmonid species originating from cold regions of the northern hemisphere. Charrs live in freshwater, though some populations migrate to brackish water in coastal areas. The only charr species originating from Europe is the Arctic charr Salvelinus alpinus, found in northern Europe and in the Alpine and Pyrenean mountain areas.

The charrs were domesticated in different regions of their distribution area at different dates. Some broodstocks have probably been domesticated since the middle of the 20th century. The first reported domestication trials on the Arctic charr were made in France in 1860. New domestication processes have been described over the last 20 years in Canada, Sweden, Iceland and Norway.
Production


  • Annual production by country in EU, in the World (reviewed by FEAP)

The process of Arctic charr farming has been reviewed by [1]. Three different species of charr are farmed in Europe: the Arctic charr (Salvelinus alpinus), the brook trout (Salvelinus fontinalis) and to a lesser extent the lake trout (Salvelinus namaykush). Arctic charr and brook trout are the more important of these species farmed in Europe.

As with all cold water species, charrs are mainly reared in the north of Europe (Finland, Sweden, Norway, Iceland, and Ireland) and also in mountain areas: as in Scotland, or in the Alps or Pyrenees (France, Switzerland, Germany, and Austria). Their adaptation to sea water is poor and most production is done in fresh or brackish water.

Production is limited as these species are highly susceptible to furonculosis and cannot stand rearing temperatures higher than 12 to 14°C, even if some brook trout broodstock are raised in rivers in the south of France that may reach 18 to 20°C. They are farmed mainly for table consumption but also for restocking lakes. European Arctic charr production was estimated at 3,130 T in 2004 and brook trout production at 300 T (source FEAP, 2004). Canada is the main producer outside of Europe and some importations are made for the fillet market.


  • EU consumption

No precise data but at least equal to the total European production.
Biological features of interest for breeding practices

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

Charrs are slaughtered at pan size (250-350 g) as their growth is limited. Today, this weight is reached between 12 to 18 months old in fresh water in the France, Italy and Germany.

  • Feeding practices during life cycle

Charrs are reared with same feed and management as other salmonids.


  • Generation interval for males and females

Farmed charrs can have a generation interval from 2 to 3 years, depending on water temperature and salinity. Females and males can produce ova and sperm at 1 year of age in the South of Europe and in fresh water. When reared in brackish water, Arctic charr become sexually mature at 2 years of age. Under some conditions a high rate of precociously mature males and females is observed. The use of specific photoperiods can limit precocious maturation [2].
Main diseases

Charrs are susceptible to most diseases recorded in Salmonids, but Arctic charr, brook trout and lake trout are also resistant and healthy carriers of two major virus: Viral Hemorrhagic Syndrome (VHS) and Infectious Haematopoietic Necrosis (IHN) [3]. However, the charrs, as species living in cold water, are highly susceptible to furonculosis Aeromonas salmonicida. They are often vaccinated against these bacteria, although repeated vaccinations are necessary to provide a long term protection.



2. Genetic variability of the species

Wild genetic resources available (Figure 1)

For lake trout and brook trout, wild genetic resources are only present in North America. Wild genetic resources exist for Arctic charr in Europe and North America [4]. Genetic differences based on allozymes, microsatellites or mtDNA support the hypothesis of a high level of subspecies differentiation. Large differences in morphology and performance have been reported between populations. Arctic charr populations are highly segmented geographically and significant genetic differences can exist over as little as 6 km. All over Europe, small genetic units exist in different lakes, river drainage systems or amongst coastal rivers [5, 6]. Population genetic studies have been made all over Europe on the Arctic charr but no single study has made a complete survey of genetic variability in the species.

In Arctic charr and brook trout, migrating and non migrating populations are described with differing physiological ability to smoltify. Anadromous populations in Norway have mainly been reported to the north of latitude 65°N, while populations in the south are considered as non-anadromous [7]. The Hammerfest strain is northern and anadromous, and also shows significantly higher growth. Genetic differences in growth and salinity tolerance have been demonstrated between 2 anadromous strains of Arctic charr in Canada [8]. This suggests that adapted broodstock could be chosen according to the water salinity of the farming environment. Large differences were also reported in growth and age at sexual maturation in both males and females between Svalbaard and North of Norway wild populations [9].





Figure 1: Distribution area of wild populations Arctic charr. Source: FAO Fishbase.

Differences between wild and/or domesticated populations

Data on evaluation of performances of wild and domesticated populations are limited even if domestication of some broodstock has been performed over at least 10 and probably 20 generations.


Interaction between wild and domesticated stocks

There are few data on genetic interaction between wild and domesticated stocks of Arctic charr in Europe [10]. For the other species, the lack of wild populations in Europe means that this topic is beyond the scope of the present review. Arctic charr was introduced to most of the Alpine lakes since the end of the 19th century, using broodstock propagated from the pioneering hatcheries. The influence of stocking on the genetic integrity of the Königssee population has been negligible for example [11]. Diploid or triploid hybrids are also used for restocking in Alpine lakes, but the genetic impact of such practices is unknown and should be evaluated.


Inbreeding effects

High inbreeding of aquacultural stock supposedly occurs due to commercial practices [12] and the low level of technical support and advice. However, in many cases commercial broodstock can present higher genetic variation than its wild counterparts [13]. No study gives an up-to-date view of genetic variability in commercial stocks.

Expected negative effects of inbreeding include a high rate of malformation and poor survival, both seen to be linked with low genetic variability [10].

3. Reproduction

Fecundity and main reproductive features

Fecundity is in the same range as other salmonids. Artificial reproduction is performed as in rainbow trout. When Arctic charr are grown in brackish water, males and females need to be transferred to fresh water a few weeks before the initiation of ovulation. Water temperature is the key factor to success in charr reproduction. The optimal temperature for egg quality, fertilisation and incubation is 6-7°C [14]. At 10°C, egg over-maturation occurs very quickly and a low rate of success is seen at the eyed stage. A transfer to adapted temperature (7°C) one month before the beginning of normal spawning provides high egg quality. Hormonal spawning induction with GnRH solves most of the problems encountered when rearing temperature is above 7°C and below 10°C [15]. The only GnRH registered in EU (Gonazon) is efficient for this means in charr [16]. The charrs reproduce under short day length, and protocols for manipulation of the spawning season are the same as for rainbow trout [17].


Cryopreservation

Sperm cryopreservation can be performed with same protocols, extenders and cryoprotectants (glycerol, DMSO) used in rainbow trout, using TBS or CBS straws in nitrogen vapour prior to immersion in liquid nitrogen [18]. Commercial extenders are supplied by companies, such as IMV, specialised in sperm freezing.


Genetic and environmental sex determination, sexual dimorphism

The genetic sex determination of the brook trout [19] is based on female homogamety and male heterogamety (XX/XY) as in all salmonids, but no data has demonstrated this in Arctic charr. In brook trout, females grow faster than males [20]. Protocols for sex reversion of breeders using methyltestosterone have been proposed for brook trout [19] and Arctic charr [21].


Cryobanks and reproductive biotechnologies (cloning, cell transfer, germ cell transfer)

No cryobank has been reported.



4. Selection

Genetic variability per trait

The first genetic basis of an heritable trait reported in the charrs concerned disease resistance to furonculosis in brook trout. By the systematic selection of surviving fish, the survival rate at 6 months of age was increased from 2 % to 69 % in 3 generations (Embody and Hayford (1925) cited by [22]). Heritability for growth and production traits is well documented in the Arctic charr, mainly from experiments performed in Sweden and Canada. An important tank effect was observed in the evaluation of genetic parameters for growth [23]. The first evaluation of heritability for growth reported realised heritabilities in the range 0.22-0.30 [24]. In the lake Hornovan strain, maternal effect on growth at 2 years old, age at maturity and muscle fat lipid content were not significant. Growth presented a high heritability in terms of weight and length (h2 > 0.35). This heritability increased between the ages of 2 and 3 years, from 0.35 to 0.45 [25], and these two traits were highly positively correlated (close to unity). Depending on site, heritability of condition factor or age at maturity varied widely from 0.45 to 0.19. Heritability for carcass component traits [26], e.g. fat, protein, dry matter and astaxanthin contents were low to medium (0.0 to 0.28). Genetic basis of heart development was also estimated [27]. Estimated heritabilities for weight were 0.54 ± 0.07, 0.45 ± 0.07, 0.36 ± 0.06, and 0.31 ± 0.05 at days 342, 458, 591 and 695 respectively. Estimates at slaughtering weight (day 695) for gill and heart weight heritabilities were in the medium range (0.30 ± 0.03 and 0.20 ± 0.03). Recently, heritability of body weight in brook trout was estimated at 0.57, genetic resistance to furonculosis at 0.51 [28].


Genetic correlations and undesirable side effects

In Arctic charr, genetic correlations between weight, length and condition factor were positive and high. Genetic correlation between sexual maturation and growth can vary between sites from low positive to highly negative. Correlation between weight and pigmentation is high but genetic correlation between weight and fat content is low even if fat variation is closely associated with variation in body weight [26]. Four generations of selection for growth have not been seen to modify aggressiveness [29].

In brook trout, the genetic correlation between growth and survival in a furonculosis challenge test was low (0.15) indicating the relative independence between these two traits [28].
G*E interactions

Not documented.


Genetic responses, progress and control lines

Very few experiments report genetic response to selection in charrs compared with control lines. Only [24] report a realised heritability of 0.25-0.30 for growth.


Dominance and intraspecific crossing

Not documented.



5. Polyploidisation and monosexing and hybrids

Triploid induction and performances

The sterility of spontaneous triploids was observed in 1978 [30]. Triploidy was initially induced by heat shock, but variable results for survival and triploidisation success were reported [31, 32, 33]. Triploid brook trout and Arctic charr are now produced using pressure treatment. Triploid brook trout perform as well diploids for growth and food consumption [35]. The same observation was also made in Arctic charr for growth up until 3 years of age [34]. There are now triploids are on the market from at least Canadian, Icelandic and French hatcheries.


Tetraploid induction and performances

No success of tetraploid induction has been reported in Arctic charr or brook trout.


Gynogenesis, androgenesis and mitotic clone performances

There are no reports of gynogenetic, androgenetic or mitotic clones in the Arctic charr. All-female gynogenetics were reported in brook trout [19].


Interspecific hybridisation

Abundant literature exists on interspecific hybridisation [36] between the Salvelinus genus [37] and Salmo trutta or Onchorhynchus sp (coho and rainbow trout). Although hybridisations are sometimes lethal at the diploid level, triploidisation can restore viability in certain hybrids. Triploid hybrids between female rainbow trout and male brook trout or Arctic charr have the VHS and the IHN resistance of these species [3]. However, these hybrids show intermediate growth rates and have a higher sensitivity to furonculosis at temperatures over 10°C. The most common hybrid farmed in Europe for fishing is the “Tiger” between brown trout and brook trout, and potential of selection to improve its survival in the triploid hybrid form has already been demonstrated [38].



6. Genomics

Tools to evaluate population genetic variability

Allozymes were routinely used until the end of the last century. Mitochondrial DNA markers were highly efficient in determining sub-structure. Now a large number of microsatellites (> 50) have been published for the brook trout [39, 40], some of these also have a high level of cross species utility for the Arctic charr.


Genetic markers for genealogical traceability

No publications report the use of microsatellites for parentage assignment in the Salvelinus genus. No European lab proposes such a service to the industry at present, probably due to the lack of demand.


QTL and Marker Assisted Selection

A linkage map has been developed for the Arctic charr [41] based on 301 microsatellites, AFLP and SNP markers. The total map lengths were estimated at 390 cM for the male map and 992 cM for the female map, and 46 linkage groups were identified. Potential QTLs for growth, identified in Atlantic salmon, are conserved in Arctic charr [42]. Tao and Boulding [43] evaluated associations between SNP in candidate genes and growth rate in this species. QTLs for temperature tolerance were identified [44] and there was also an unsuccessful trial to identify a QTL associated with sex in the Arctic charr [45].


Transgenesis

No transgenesis has been reported, except in the Arctic charr by the company Blue Revolution in Canada.


References

[1] M. Jobling, H. Tveiten, and B. Hatlen,. ”Cultivation of Arctic charr: an update,” Aquaculture International, vol. 6, pp. 181–196, 1998.

[2] J. Duston, T. Astatkie, and P. F. MacIsaac, “Long-to-short photoperiod in winter halves the incidence of sexual maturity among Arctic charr,” Aquaculture, vol. 221, pp. 567–580, 2003.

[3] M. Dorson, B. Chevassus and C. Torhy, “Comparative susceptibility of three species of char and of rainbow trout * char triploid hybrids to several pathogenic salmonid virus,” Dis. Aquat. Org., vol. 11, pp. 217-224, 1991.

[4] P.C. Brunner, M.R. Douglas, and L. Bernatchez, “Microsatellite and mitochondrial DNA assessment of population structure and stocking effects in Arctic charr Salvelinus alpinus (Teleostei: Salmonidae) from central alpine lakes,” Mol. Ecol., vol. 7, pp. 209-223, 1998.

[5] K. Hindar, N. Ryman and G. Ståhl, “Genetic differentiation among local populations and morphotypes of Arctic charr, Salvelinus alpinus,” Biological Journal of the Linnean Society, vol. 27, pp. 269-285, 1986.

[6] A. J. Wilson, D. Gislason,, S. Skulason, S. Snorrason, C. E. Adams, G. Alexander, R. G. Danzmann, and M. M. Ferguson, “Population genetic structure of Arctic charr, Salvelinus alpinus from northwest Europe on large and small spatial scales,” Mol. Ecol., vol. 13, pp. 1129– 1142, 2004.

[7] Norberg

[8] J. L. Delabbio, B. D. Glebe, and Sreedharan, “Variation in growth and survival between two anadromous strains of Canadian Arctic charr (Salvelinus alpinus) during long-term saltwater rearing,” Aquaculture, vol. 85, pp. 259-270, 1990.

[9] B. Damsgar, A. M. Arnesen., M. Jobling, “Seasonal patterns of feed intake and growth of Hammerfest and Svalbard Arctic charr maturing at different ages,” Aquaculture, vol. 171, pp. 149–160, 1999.

[10] C. R. Primmer, T. Aho, J. Piironen, A. Estoup, J. M. Cornuet and E. Ranta, “Microsatellite analysis of hatchery stocks and natural populations of Arctic charr, Salvelinus alpinus, from the Nordic region: implications for conservation,” Hereditas, vol. 130, pp. 277-289, 1999.

[11] C. C. Englbrecht, U. Schliewen, D. Tautz, “The impact of stocking on the genetic integrity of Arctic charr (Salvelinus alpinus) populations from the Alpine region,” Mol. Ecol., vol. 11, pp. 1017–1027, 2002.

[12] T. A. Lundrigan, J. D. Reist, and M. M. Ferguson, “Microsatellite genetic variation within and among Arctic charr (Salvelinus alpinus) from aquaculture and natural populations in North America,” Aquaculture, vol. 244, pp. 63–75, 2005.

[13] D. Ditlecadet, F. Dufresne, N. R. Le François, and P. U. Blier, “Applying microsatellites in two commercial strains of Arctic charr (Salvelinus alpinus): Potential for a selective breeding program,” Aquaculture, vol. 257, pp. 37–43, 2006.

[14] C. Gillet, “Egg production in Arctic charr (Salvelinus alpinus) brood stock: effect of temperature on the timing of spawning and the quality of eggs,” Aquat. Living Resourc., vol. 4, pp. 109-116, 1991.

[15] C. Gillet, B. Breton and T., Mikolajczyk, ”Effect of GnRHa and pimozide treatments on the timing of ovulation and on egg quality in Arctic charr (Salvelinus alpinus) at 5 and 10°C,” Aquat. Living Resourc., vol. 9, pp. 257-263, 1996.

[16] P. Haffray, W. J. Enright, M. A.. Driancourt, T. Mikolajczyk, P. Rault, and B. Breton, “Optimisation of breeding of Salmonids: Gonazon®, the first officially approved inducer of ovulation in the EU,” World Aquaculture Magazine, pp. 52-56, 2005.

[17] C. Gillet, “Egg production in Arctic charr (Salvelinus alpinus L.) broodstock: effects of photoperiod on the timing of ovulation and egg quality,” Can. J. Zool., vol. 72, pp. 334–338, 1994.

[18] J. Piironen, “Cryopreservation of sperm from brown trout (Salmo trutta m. lacustris L.) and Arctic charr (Salvelinus alpinus L.),” Aquaculture, vol. 116, pp. 275-285, 1993.

[19] P. F. Galbreath, N. D. Adams, and L. W. Sherrill, “Successful sex reversal of brook trout with 17α-methyldihydrotestosterone treatments,” North American Journal of Aquaculture, vol. 65, pp. 235-239, 2003.

[20] Y. Boulanger, “Performance comparaison of all-female diploid and triploid brook trout,” Can. Rep. Fish. Aquat. Sci., vol. 1789, pp. 111-121, 1991.

[21] M. Chiasson and T. J. Benfey, “Gonadal differentiation and hormonal reversal in Arctic charr (Salvelinus alpinus),” J. Exp. Zool., vol. 307A, pp. 527-534, 2007.

[22] B. Chevassus and M. Dorson, “Genetic of resistance to disease in Fishes,” Aquaculture, vol. 85, pp. 83-107, 1990.

[23] B. G. E. de March, “Tank effect as confounding factors in genetic experiments: experiences with Arctic charr (Salvelinu alpinus),” Can. Tech. Rep. Fish. Aquat. Sci., vol. 1761, pp. 61-68, 1990.

[24] B. G. E. de March, “An evaluation of the importance of within-family selection for juvenile growth characteristics in Arctic charr,” The Progressive Fish-Culturist, vol. 57, pp. 192-198, 1995.

[25] J. Nilsson, “Genetic parameters of growth and sexual maturity in Arctic charr (Salvelinus alpinus),” Aquaculture, vol. 106, pp. 9-19, 1992.

[26] P. Elvingson, and J. Nilsson, “Phenotypic and genetic parameters of body and compositional traits in Arctic charr, Salvelinus alpinus (L.),” Aquaculture and Fisheries Management, vol. 25, pp. 677-685, 1994.

[27] M. A. Montañez, R. Ginés, A. Navarro, J. M. Afonso, and H. Thorarensen, “Heritability estimations for growth characters and size of the cardiorespiratory organs in Arctic charr (Salvelinus alpinus),” Aquaculture (in press), 2007.

[28] G. M. L. Perry, P. Tarte, S. Croisetière, P. Belhumeur and L. Bernatchez, “Genetic variance and covariance for brook charr (Salvelinus fontinalis) weight and survival time of furonculosis (Aeromonas salmonicida) exposure,” Aquaculture, vol. 235, pp. 263-271, 2004.

[29] E. Brännas, T. Chaix, J. Nilsson and L. O. Eriksson, “Has a 4-generation selection programme affected the social behaviour and growth pattern of Arctic charr (Salvelinus alpinus) ?,” Applied Animal Behaviour Science, vol. 94, pp. 165-178, 2005.

[30] S. Allen and J. G. Stanley, “Reproductive sterility in polyploid brook trout, Salvelinus fontinalis,” Trans. Am. Fish. Soc., vol. 107, pp. 473-478, 1978.

[31] B. D. Glebe, P. Delabbio, R. L. Lyon, R. L. Saunders and S. McCormick, “Chromosome engineering and hybridization of Arctic charr (Salvelinus alpinus) and Atlantic salmon (Salmo salar) for aquaculture,” EIFAC/FAO Symposium on Selection, Hybridization and Genetic Engineering in Aquaculture and Fish and Shellfish for consumption and stocking. Bordeaux (France), 27-30 May 1986.

[32] P. Dube, J. M. Blanc, M. Chouinard, and J. de la Nouë, “Triploidy induced by heat shock in brook trout (Salvenilus fontinalis),” Aquaculture, vol. 92, pp. 305-311, 1991.

[33] P. F. Galbreath and B. L. Samples, “Optimization of thermal shock protocols for induction of triploidy in brooktrout” North. Am. J. Aquac., vol. 62, pp. 249-259, 2000.

[34] C. Gillet., C. Vauchez and P. Haffray, “Triploidy induced by pressure shock in Arctic charr (Salvelinus alpinus): growth, survival and maturation until the third year,” Aquat. Living Resour., vol. 14, pp. 327-334, 2001.

[35] R. A. O’Keefe and T. J. Benfey, “Comparative growth and food consumption of diploid and triploid brook trout (Salvelinus fontinalis) monitored by radiography,” Aquaculture, vol. 175, pp. 111-120, 1999.

[36] B. Chevassus, “Hybridization in Fishes,” Aquaculture, vol. 33, pp. 245-262, 1983.

[37] S. Dumas, J. M. Blanc, C. Audet and J. de la Noüe, "The early development of hybrids between brook charr (Salvelinus fontinalis) and Arctic charr (Salvelinus alpinus),” Aquaculture, vol. 108, pp. 21-28, 1992.

[38] J. M. Blanc, H. Poisson and F. Vallée, “Survival, growth and sexual maturation of the triploid hybrid between rainbow trout and arctic charr,” Aquat. Living Resourc., vol. 5, pp. 5-21, 1992.

[39] B. Angers, L. Bernatchez, A. Angers and L. Desgroseillers, “Specific microsatellite locci for brook charr (Salvelinus fontinalis Mitchill) reveal strong population subdivision on a micro-geographic scale,” Journal of Fish Biology, vol. 47, pp. 177-185, 1995.

[40] G. L. Perry, T. L. King, J. St Cyr, M. Valcourt and L. Bernatchez, “Isolation and cross-familial amplification of 41 microsatellites for the brook charr (Salvelinus fontinalis),” Molecular Ecology, vol. 5, pp. 346-351., 2005.

[41] R. A. Woram, C. McGowan, J. A. Stout, K. Gharbi, N. M. Ferguson and B. Hoyheim, “A genetic linkage map for Arctic charr (Salvelinus alpinus): evidence of higher recombination rate and segregation distorsion in hybrid versus pure strain mapping parents,” Genome, vol. 47, pp. 304-315, 2005.

[42] D. P. Reid, A. Szanto, B. Glebe, R. G. Danzmann., and M. M. Ferguson, “QTL for body weight and condition factor in Atlantic salmon (Salmo salar): comparative analysis with rainbow trout (Oncorhynchus mykiss) and Arctic charr (Salvelinus alpinus),” Heredity, vol. 94, pp. 166-172, 2005.

[43] W. J. Tao and E. G. Boulding, “Associations between single nucleotide polymorphisms in candidate genes and growth rate in Arctic charr (Salvelinus alpinus L.),” Heredity, vol. 91, pp. 60-69, 2003.

[44] I. M. L. Somorjai, R. G. Danzmann, and N. M. Ferguson, “Distribution of temperature quantitative trait loci in Arctic charr (Salvelinus alpinus) and inferred homologies in rainbow trout (Oncorhynchus myskiss),” Genetics, vol. 165, pp. 1443-1456, 2003.

[45] H. K. Moghadam, M. M. Ferguson, and R. G. Danzmann, “Linkage variation of the sex determining locus in Arctic charr (Salvelinus alpinus),” Aquaculture (in press), 2007.




    1. Industrial Survey

A questionnaire focusing on aquaculture breeding practices was prepared and sent in 28 countries through national contacts. The answers received from single breeding organisations were aggregated into a final report. An updated document has been prepared lately to include breeding programs set up in Central and East European countries. The results give an extended view, even if not exhaustive, of current commercial breeding activities in Europe (37 selective breeding programmes listed, 14 selected species) with a description of practices split by strategy, country and species.
This survey brings a clearer view of the European aquaculture breeding sector, today still small and fragmented, showing a trend of development across species and countries. It highlights the diversity of strategies implemented and brings to light specificities linked to the species biology and markets structure. It also shows strong disparities among countries, suggesting measures of readjustment where few or nothing is done.
The survey will contribute to shape the image of the European aquaculture breeding sector, diverse among countries and till now lacking of contour. It is expected, with the mergence of the proposed Network, that such a survey be updated every second year in order to monitor the evolution and needs of the sector.
All the project’s partner brought support to the build up of the survey.
The work performed is presented in the Deliverable 2.



DELIVERABLE 2

Questionnaire and Survey



Questionnaire on the breeding practices

in the European aquaculture industry

Company form






AquaBreeding project (2006 – 2008) – www.aquabreeding.eu

“Towards enhanced and sustainable use of genetics and breeding

in the European aquaculture industry”








  1. Breeding strategy




 

 

    1. Latin name of the species

      

    1. Number of lines1 or closed populations2

Points 1.3 to 1.10 should be completed separately for each line

 precise nbre of age class if there are

    1. Number of selected generations performed

      

    1. Selected traits

growth

processing yield

product quality

disease resistance

reproduction (maturity, fecundity)

morphology

others


   
   

   

   

   

   

   










If others, please specify:

      

    1. Reproductive technologies

artificial fertilisation

monosexing

gynogenesis


   

   

   

1.5 Reproductive technologies (continue)

mass spawning

cryopreservation

hormonal spawning induction

hormonal spermiation induction

environment manipulations

T°C


pH

photoperiods



   

   

   

   
   

   

   


    1. Mean number of parents per line

< 100

100 to 200

200 to 500

500 to 800

> 800


   

   

   

   

   

    1. Type of selection

crossbreeding3

individual selection

within family selection4

between family selection5

combined selection6


   

   

   

   

   

    1. Molecular tools

fingerprint for parentage assignment

marker assisted selection

others


   

   

   










If others, please specify:

      

    1. Alternative genetic improvement technologies

triploidisation

tetraploidisation

hybridisation (between species) 7

others


   

   

   
   










If others, please specify:

      

    1. Protection strategies

Sterilisation

genetic traceability

hybridisation (within species) 8


   

   

   

1   ...   6   7   8   9   10   11   12   13   ...   16


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