Development of Breeding Programs

for Aquatic Species Should be

Given High Priority
Trygve Gjedrem 1 and Kari Kolstad

W orld aquaculture production has grown 7.1 percent slight improvement since previous estimates: 1 percent
in 1993 (Gjedrem 1997) and 5 percent in 2002 (Gjedrem
per year during 2000 - 2009 and reached 55.7 million t of fish
and shellfish and 17.3 t of aquatic plants in 2009 (FAO 2011). 2004). In addition, some commercial farms may undertake
If expansion of fish and shellfish continues, production will some individual selection for growth rate but the majority of
reach 68.4 million t in 2012, which is greater than the amount aquaculture production is based on wild individuals or stocks
projected from capture fisheries (FAO 2009). The prognosis is that are not genetically improved.
that fisheries production will stabilize at around 90 million t in
the coming years (FAO 2008). The potential for further growth What Has Been Accomplished?
in aquaculture production seems to be very good, considering Over the last 30 years, a number of selection experiments
coastal and land areas suitable for aquaculture in different have been carried out (summarized by Gjedrem and Thodesen
countries. However, competing activities in these areas and a 2005). The average genetic gain from 12 estimates of growth
lack of availability of feed may limit expansion. rate in different fish species was 15 percent per generation, two
Production of shellfish and to some extent herbivorous fish estimates for shrimp report 8 percent, four estimates for oysters
species opens possibilities for harvesting some of the extensive report 14 percent, one estimate for clams reports 9 percent and
primary production that occurs naturally in the sea because two estimates for scallops report 17 percent. The average of 21
these species feed on algae and other plankton. Combining fish, estimates was 14 percent genetic gain per generation. Similar
shellfish and seaweed (IMTA; integrated levels of genetic gain are reported for coho
multi-trophic aquaculture) is an interesting It should no longer salmon (Neira et al. 2006), channel catfish
concept for sustainable development be acceptable that 90 (Rezk et al. 2003), and European sea bass
(Chopin 2011). percent of aquaculture (Vandeputte et al. 2009). This indicates
Production of diadromous and marine production is based on that growth rates can be more than doubled
species is low compared to freshwater wild and genetically in less than six generations of selection.
species (Table 1). This is somewhat unimproved animals. Two examples have shown that this has
surprising because the potential for been accomplished.
aquaculture production is much larger in
the marine environment compared to freshwater. Differences in Example 1: Nile tilapia (Oreochromis niloticus), Philippines
value among groups of species should be noted. Highly prized The base population consisted of hybrids among eight
species—diadromous and marine fishes and crustaceans—are strains that were used to make the GIFT strain. Selection was
usually produced under intensive farming conditions compared for growth rate only. The response during five generations of
to freshwater fishes, where lower-priced cyprinids represented selection was 85 percent (17 percent per generation) (Bentsen
76 percent in 2005 (FAO 2007). The main purpose of this paper et al. 2003). Fish of the GIFT strain have been distributed to
is to discuss the role selective breeding can have on the potential several countries. According to Neira (2010), 10 of 20 tilapia
expansion and efficiency of future aquaculture production. family breeding programs are a continuation of GIFT. One
of these breeding programs, mainly based on the fifth GIFT
Status of Breeding Programs for Aquatic generation, has resulted in a further genetic gain of 59 percent
Species after 6 generations of a multi-trait selection program (Thodesen
The majority of fish culturists raise animals derived from et al. 2011).
wild stocks or from those that have been selected for only a few
generations. In contrast, genetically improved stocks and seeds Example 2: Atlantic salmon (Salmo salar), Norway
are widely used in terrestrial farm animal and plant production. Four base populations based on 41 wild strains were
Thus, fish culture lags far behind terrestrial animal husbandry established (Gjedrem 2010). The first round of selection started
and plant production (Tave 1986). in 1975. Thodesen et al. (1999) studied the response over five
In surveys, Neira (2010) and Rye et al. (2010) recorded 101 generations of selection in one of the populations by comparing
family-based breeding programs in aquaculture, emphasizing selected and wild animals. Selection was mainly for growth rate,
that there may be more. Based on these, 8.2 percent of world with some emphasis on age at sexual maturity. Improvement
aquaculture production is estimated to be based on family in growth rate was 113 percent or 22 percent per generation. In
breeding programs (Gjedrem et al. 2012). There has been addition, the correlated effects of protein and energy retention

10 S E P T E M B E R 2 012
 TABLE 1. World aquaculture production of various species groups in 2009 (FAO 2011).

Species group Production (million t) Value (US$ 1000) Value (US$ per kg)
Freshwater fishes 30.64 44.19 1.44
Diadromous fishes 3.53 14.00 3.96
Marine fishes 1.95 7.10 3.64
Crustaceans 5.30 24.13 4.55
Molluscs 13.52 13.13 0.97
Miscellaneous aquatic animals 0.73 2.75 3.77
Aquatic plants 17.34 4.82 0.28
Total aquaculture production 73.02 110.12 1.51

were improved by 9 percent and 14 percent, respectively. Further, performed over multiple generations using few broodstock
feed conversion ratio (FCR) was improved by 20 percent after without control of the relationship between mating animals.
five generations of selection. This has resulted in inbreeding, leading to high mortality and
low performance. This leads to a loss of confidence in selective
In terrestrial livestock, genetic gain in growth rate obtained breeding among farmers.
in selection programs is typically 5 percent per generation or 1-2 3. Fry and fingerlings are easily available from wild stocks
percent per year, which is 5-6 times less than in aquatic species at a low price for a number of species.
(Gjedrem and Baranski 2009). 4. Lack of knowledge in quantitative genetics and selective
It is also possible to obtain high genetic gain in fish by breeding among farmers and researchers.
selecting for disease resistance. This has been documented for 5. Lack of knowledge of the high benefit/cost ratio of
resistance against IPN virus in Atlantic salmon (Storset et al. investing in breeding programs.
2007, Housten et al. 2008, Moen et al. 2009), pancreatic necrosis 6. Lack of farmers’ co-operatives and private companies
in rainbow trout (Okamoto et al. 1993), Areomonas hydrophila willing to develop breeding programs.
in rohu carp (Sahoo et al. 2011), and bacterial cold water disease 7. High investment required to begin family-based breeding
in rainbow trout (Leeds et al. 2010). Applying challenge tests programs.
under standardized environmental conditions opens possibilities 8. Farmers are not willing to pay more for improved stocks.
for developing resistant strains for specific diseases. 9. National and international organizations have not
It is only possible to obtain high genetic gains when advocated and stimulated development of breeding programs
breeding programs are well planned and carried out. The genetic for aquatic species, with the World Fish Center (formerly
gain (ΔG) per generation depends on three parameters: ΔG = i ICLARM) a notable exception.
• h2 • σp where i is selection intensity, h2 is heritability and σp is
phenotypic standard deviation. Initiating Breeding Programs for Aquatic
The primary reasons for the high genetic gain obtained for Species
aquatic species are: It is essential to start a breeding program with a broad
1. Fecundity is very high when females contribute genetic base. For species with ongoing breeding programs, such
thousands or even millions of eggs to each spawn. Selection as those for the GIFT strain of Nile tilapia and Atlantic salmon,
intensity can, therefore, be very high. it is possible to get a ‘flying start’ by importing stock that has
2. Phenotypic and genetic variation is large for growth rate already been genetically improved. Most governments will
and most other traits of economic importance. require detailed health certificates for translocated animals and
3. Heritability for economically important traits is of that quarantine and other conditions are met. This procedure
medium magnitude. can be advocated because genotype by environment interaction
Because the generation interval is rather low (compared to is low and insignificant for tilapia (Eknath et al. 1993), Atlantic
terrestrial livestock), the yearly genetic gain can be very high. salmon (Gunnes and Gjedrem 1978), rohu carp Labeo rohita
(Reddy et al. 2002) and white shrimp Litopenaeus vannamei
Reasons for Few Breeding Programs in (Fjalestad et al. 1997). The alternative is to start from scratch by
Aquaculture collecting animals of different origin to form a synthetic base
There may be many reasons for the infrequent use of population.
breeding programs for aquatic species improvement. In general, There are two recommended breeding strategies: individual
aquaculture farmers have not adopted the genetic improvements selection and family-based breeding programs. Individual
obtained. Some of the reasons are: selection may be efficient in improving growth rate. If mass
1. We deal with small animals, each individual with a low spawning is used without knowledge of the relationship
economic value. between mating animals, inbreeding will increase rapidly and
2. There are several examples of individual selection being (CONTINUED ON PAGE 10)

W O R L D A Q UA C U LT U R E 11
destroy improvements obtained in the previous generations.
DNA tagging should be used to keep track of relationships and
control inbreeding. Still, individual selection is limited to those
traits that can be measured. Family-based breeding programs
have the advantage that economically important traits, such as
product quality and disease resistance, can be selected for based
on information from siblings of breeding candidates. Adding
genomic information to these schemes increases accuracy.
Controlled single-pair mating between unrelated individuals
should be practiced in individual selection and family-based
breeding programs. The number of progeny per mating should
be standardized. In family-based breeding programs, it is
advantageous to rear families separately until tagging and then
use communal rearing thereafter. This means that infrastructure
must be established to produce a high number of families (at
least 100) per generation. With genetic markers for identification,
communal rearing is possible, reducing the need for
infrastructure. However, this approach may result in a reduced
number of families and large variation in family size, thus
introducing genotyping costs. For all approaches, to keep the
generation interval as short as possible, estimation of breeding
values should take place as soon as data is available.
Consequences of Using Selective Breeding in
Aquaculture
Assuming a genetic gain in growth rate of 12.5 percent
per generation, estimated global aquaculture production
increases with greater use of selective breeding (Table 2). The
impressive effect of selection is that genetic gain is cumulative
over generations. Global production would be doubled by 2020,
representing 5-6 generations of selection, if all of it was based
on genetically improved stocks (Table 2). Breeding programs for
Nile tilapia and Atlantic salmon have already shown that this is
possible. In addition, commercial Atlantic salmon aquaculture
is an example where nearly all production is currently based on
improved stocks (Gjedrem 2004).
It is not realistic to expect that all of global aquaculture
production will be based on improved stocks because of the high
number of species in aquaculture, with a resulting need for a high
number of breeding programs. According to FAO (2007) there
are 241 species used in global aquaculture and, among them, 124
species with production above 1000 t.
From a resource use efficiency point of view, there is much
to be gained by applying selective breeding. Doubling growth
rate with 5-6 generations of selection reduces the production
time by almost 50 percent. Food production will increase and
there will be more efficient utilization of production facilities
(i.e., ponds and cages), labor, water and feed (Thodesen et al.
1999), making aquaculture an even more cost-efficient and
sustainable proposition.
Investments in efficient breeding programs are cost
effective. For fish, the estimated benefit-cost ratios for breeding
programs vary from 8-1 to 60-1 (Gjerde et al. 2007, Gjedrem
1997, Ponzoni et al. 2007, Ponzoni et al. 2008, Robinson et al.
2010). The economic value of better feed efficiency in Atlantic
salmon after six generations of selection is estimated to be US$
0.50 per kg fish produced (Gjerde et al. 2007).
12 S E P T E M B E R 2 012
Aquaculture for the Future
The paramount objective for aquaculture in the future is
the sustainable production of animal protein. This can only be
obtained if we use animals that have been selectively bred for
improved performance in aquaculture production environments,
not by using wild animals that do not thrive under these
conditions. By increasing growth rate, more foodfish will be
produced in a shorter time, production will become more cost-
efficient, feed will be more efficiently converted into animal
protein (Thodesen et al. 1999), and mortality will be reduced
because less time will be required to reach market size. The
potential to improve disease resistance is well documented
(Leeds et al. 2010, Sahoo et al. 2011, Storset et al. 2007).
Selective breeding is crucial for cost-efficient production,
animal welfare, and minimizing the impact of aquaculture on
wild populations.
These aims can be obtained only by efficient breeding
programs. It should no longer be acceptable that 90 percent
of aquaculture production is based on wild and genetically
unimproved animals. The status quo is an inefficient way of
using natural resources, particularly feed-related resources, and
is a misuse of valuable land, water, and labor. Selective breeding
increases the rate of domestication and improves animal welfare
(Doyle 1983) and results in more predictable and reliable
production. We should learn from terrestrial farm animal and
plant agro-industries that have been using selectively bred stock
for many decades.
The time has come for aquaculture producers and other
stakeholders to cooperate with government authorities and
institutions to plan and start breeding programs for each of the
most important species in aquaculture. There is a foundation
of available literature, knowledge and experience on which to
build.
Notes
1
Nofima, PO Box 210, 1431 Ås, Norway
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