Quantitative genetics of body

weight, fillet weight and fillet

yield in Nile tilapia
(Oreochromis niloticus)
BJARNE GJERDE A,B, SAMUEL BEKELE
M E N G I S T U B , E , J Ø R G E N Ø D E G Å R D A , B , . H A R RY
J O H A N S E N C , D A N I L O S O Z A A LTA M I R A N O D
Abstract
With the objective of how to best improve fillet weight in Nile tilapia (Oreochromis niloticus)
through selection round and gutted body weight and fillet weight were recorded at harvest
over four generations (G0, G1, G2 and G3) and from which the following traits were derived:
Fillet yield (fillet weight×100/round weight), non-edible part of the fish (NEP=round
weight−fillet weight), and non-meat part of the gutted weight (NMP=gutted weight−fillet
weight). The round weight was recorded on 42,795 offspring of 327 sires and 422 dams, and
the other traits on a random sample of 3361 of the offspring of 277 sires and 359 dams. The
fish were reared in net-cages in a lake or in earthen ponds. Selection was performed for
harvest weight and fillet yield with an equal relative economic weight (50%) on each trait.
The heritabilities for round, gutted and fillet weight, NEP and NMP were all moderate (0.16–
0.19) and lower for fillet yield (0.06). The effect common to full-sibs (c2) accounted for a
large proportion of the phenotypic variance for all traits (0.11– 0.19).
The genetic correlation between round weight of fish reared in cages and in ponds was unity
(1.00), indicating no genotype by environment interaction for this trait. The genetic correlations
between round, gutted and fillet weights were all close to unity, as were the genetic correlations of
round and fillet weight with the calculated traits NEP and NMP. The close to unity genetic
correlation between round body weight and fillet weight indicates that genetic improvement of one
of the traits without achieving a proportional genetic change in the other is difficult or even
impossible. This implies that improvement of fillet yield through direct selection is difficult to
achieve. Still, based on the current results, fillet yield is expected to increase with increasing body
weight, both genetically and phenotypically. However, if increased growth leads to harvest at a
younger age rather than higher weight, response in fillet yield may not be observed. Improvement
of fillet weight can be obtained through selection for increased round body weight, a more efficient
strategy as the trait can be recorded on live breeding candidates and as higher economic weight
can be put on this trait instead of on fillet yield
Introduction
Carps, tilapia and salmonids dominate the farmed finfish production in the given order
(El-Sayed, 2006; Fessehaye, 2006). Tilapia is geographically the most widely grown
farmed fish (FAO, 2005- 2009), with farming in more than 100 countries in 2002 (El-
Sayed, 2006). In the market, tilapias are sold as either whole fish or as fresh or frozen
fillets. The US and European markets prefer fillet. The desirable fish size to produce
depends on the type of product; for whole fish preferable 400–500 g, which can be
attained at the age of 8 to 10 months, while for fillet 700 to 1000 g is preferred (11 to 14
months) (FAO, 2005–2009; Tilapia Profile, 2008).
• Among the few less desirable characters of tilapias is their low fillet yield (fillet
weight/body weight) which has become more of an issue as processors implement
additional trims and deeper skinning at the request of customers (Fitzsimmons et al.,
2011).
•In this context it should be noted that the trait fillet yield has no economic value in itself,
only through its relation to total fillet output (i.e., high fillet yield in a small fish has
smaller value than the same fillet yield in a bigger fish)
•In a recent study on Nile Tilapia, it was estimated that multi-trait selection for growth until
a fixed body weight and fillet yield (at this weight) reduced genetic gain in growth by
20% as compared to single-trait selection for growth only (Thodesen et al., 2011).
• In fish, the most commonly used selection methods are individual and/or sib selection
(Gjedrem, 1992). Sib selection has to be performed for traits that cannot be recorded on
the live breeding candidates, but on their sibs; e.g., fillet weight and fillet yield.
• Both fillet weight and yield are laborious and costly to measure, and research on
improvements of these traits has therefore centred on selection for different correlated
body measurement and body shape traits
•The main objective of this study was to estimate the genetic and phenotypic parameters
for round body weight and for the carcass traits; gutted body weight, fillet weight and
fillet yield recorded at harvest size in Nile tilapia. Furthermore, based on these
parameters, recommend how to improve fillet weight through selection.
Materials and Methods
Base population
• The studied traits were recorded on animals representing the base population (G0) and
the first (G1), second (G2) and third (G3) selected generation produced by Nicanor,
Nicaragua. The selection was conducted mainly for increased growth through selection
for harvest body weight, but also for increased fillet yield when producing G1 and G3.
• The parents of the base population (G0) were obtained in 2004 from a Vietnamese
family-based breeding program started in 1999. This nucleus population was mainly
offspring of the fifth generation of the GIFT breeding project conducted in the Philippines
(Eknath and Acosta, 1998) in which growth already had been improved over five
generations.
Mating design, rearing and grow-out system

• The mating design used was nested with one sire mated to two different dams and each dam to one sire.
However, mortalities until tagging size resulted in many pair matings, in particular G1 and G2. There was
no reuse of parents across generations. The families were produced through natural mating in separate
breeding hapas (1 m×1 m×1 m mosquito net in generations G0 and G1 and 2 m×2 m×1 m in generations G2
and G3)
• Inspection of the presence of swim-up fry was done two times per week and the period from
detection of fry in the first and the last produced family was 46 (G0, April 30 to June 14, 2004),
109 (G1, June 17 to October 3, 2005), 140 (G2, July 10 to November 27, 2006) and 70 (G3,
September 24 to December 22007) days resulting in some variation in the age at harvest within
each generation
• When swim-up fry were detected, the sire was removed and placed together with
another dam in a different hapa. After the yolk sack was absorbed also the dam was
removed while the fry were kept in the hapa until tagging. The number of fry of each full-
sib family (hapa) was counted at an average age of A50 days after swim-up and
reduced to a maximum of 500 fry per family.
•After this reduction the average number of fry per family was 437 (G1), 413 (G2) and
438 (G3). In G2 the number of fry per family was further reduced to a maximum of 300
fry at an average age of 76 days.
•The fish were individually tagged with PIT tags. All available fingerlings in the base
generation (G0) were tagged
• After tagging those to be grown in ponds or raceway were sexed based on external
appearance and stocked into two separate 0.05 ha ponds (one for males and one for
females, in G1, G2 and G3) or a 35 m×2.5 m×1 m concrete raceway (one half for males
and the other half for females, in G0). The remaining tagged fish (in G0, G2, and G3)
were stocked into one cage (12 m×12 m×5 m deep, mesh size 10 mm)
• Individuals of G0 were sexed and stocked in two separate parts of the cage, while
individuals in G2 and G3 were reared in mixed sex culture. In cages stocking densities
in ponds varied from 12 to 14 fish/m2, in cages from 5 to 6 fish/m3, and 6.6/m2 in the
raceway.
Traits recorded at harvest
• At harvest size the round body weight (RW1) was recorded on all the fish, while the
carcass traits gutted body weight (GW2) and fillet weight of the two untrimmed fillets
with skin (FW2) were recorded some days or weeks later when slaughtering a random
sample of the fish, at which also the round body weight (RW2) of these fish were
recorded
• Within a generation the traits recorded at slaughter (RW2, GW2 and FW2) were only
recorded on fish reared in cages (G0 and G3) or in pond (G2); on males only in G0 and
G2, while on both males and females in G3. Thus, within generation G2 and G3 the trait
Rw1 was recorded on fish reared in both cages and ponds, while the traits RW2, GW2
and FW2 were recorded on fish reared in either pond (G2) or cage (G3)
Selection of parents
• For the recorded animals of each generation single-trait BLUP (Best Linear Unbiased
Prediction) breeding values were obtained for round body weight and fillet yield based
on all available information on the candidates and their full- and half-sibs and by taking
into account all additive genetic relationships among all animals back to the base
population (G0)
• In G1 selection was performed for round body weight only as fillet yield was not
recorded. To keep rate of inbreeding at acceptable levels, a restricted number of sires
and dams were selected from each of the highest ranking families, and the production of
a new generation of families was performed by omitting mating of full- and half-sibs.
Statistical model
Genetic parameters
• Given a significant genotype by test environment (cages vs. ponds) interaction (G×E)
for the studied carcass traits it may not be appropriate to pool the data from these
different environments and thus assume that e.g. fillet weight or fillet yield is the same
trait when recorded in ponds and cages
• Gutted weight and fillet weight within a generation, and the traits derived from these
traits (fillet yield, NEP and NMP), were not recorded on fish reared in both ponds and
cages the magnitude of the G×E cannot be estimated. However, for round body weight
an estimate of the magnitude of this G×E can be obtained by defining round body weight
in ponds and cages as separate traits.
• First the additive genetic, full-sib family and random error variances for round body
weight of fish reared in ponds and for fish reared in cages (excluding the G0 data
from the two raceway at Tipitapa) were obtained by restricted maximum likelihood
(REML) using a bivariate animal model in ASReml (Gilmour et al., 2006). In a matrix
notation the model (Model 1) can be written as:
• Secondly, the data from all generations and test environments were used to obtain
(co)variance components for the random effects in Model 1 for all traits (RW1, GW2,
FW2, FY, NEP, NMP) across all generations, sites, ponds or cages or raceways and
sexes. A model with maximum three traits at a time had to be used to obtain
convergence, in which round body weight recorded on all fish (RW1) was always
included to account for possible non-random sampling of fish when GW2 and FW2 were
recorded (Model 2).
Results
• Descriptive statistics for the studied traits at the two test sites within each of the three generations.
The few females with gutted weight and fillet weight in G0 (n=17) and G3 (n=30) were misclassified
with respect to sex just prior to slaughter (G0) or just prior to stocking in ponds (G2). For round weight
the males were from 32% to 58% heavier than the females (Pb0.001). The coefficients of variation
(CV) for round body weight varied from 18.4 to 27.6 for the different generation by site combinations
and of similar magnitude in males (23) and females (24). For gutted weight and fillet weight the sex
differences and CV were very similar to those for round weight
• Average fillet yield varied from 41.3% to 45.1%

• In G3 the fillet yield of males was 0.8%-points higher than that of females (Pb0.05).

•In G0 and G2 the number of females was too low to obtain reliable estimates of the difference in fillet
yield between males and females. The coefficients of variation for fillet yield were low and varied from
3.4 to 6.8%.
Parameter estimates within cages and ponds
• For fish reared in cages the heritability for round body weight was lower than for fish
reared in ponds. The genetic correlation between round body weight in ponds and cages
was unity (1.00±ne; i.e., the correlation was restricted at the upper boundary of the
parameter space, consequently ASRemle did not provide standard errors).
Parameter estimates across cages and ponds
• Estimates of heritability and the effect common to full-sibs (c2) for each trait are given.
The heritability's of the recorded traits round weight, gutted weight and fillet weight and
of the two calculated traits NEP and NMP were very similar and of medium magnitude,
while the heritability of fillet yield was lower and not significantly different from zero
(P>0.05). The effect common to fullsibs (c2) was highly significant (Pb0.001; based on a
likelihood-ratio test, Lynch and Walsh, 1998) and accounted for a large proportion of the
phenotypic variance for all traits
Prediction of fillet yield

The first-order polynomial for corrected round weights (RW2) on corrected fillet weights
(FW2) was highly significant (Pb0.0001) and explained a large proportion of the variation
in the corrected fillet weights (R2 ranging from 92.5 to 97.4% for the four studied levels
of the combined generation, site, cage (or pond) and sex effects). The second-order
polynomial for corrected round weights (RW22) was not significantly different from zero.
The regression equations for the four levels of this fixed effect were:
FW=−6.669+0.457RW for G0-Om-Ca-M, FW=−12.451+0.434 RW for G2-Ti-Po-M, FW=
−9.231+0.433RW for G3-Om-Ca-M and FW=−6.669+0.457RW for G3-Om-Ca-F.
Conclusion
The close to unity genetic correlation between round body weight and fillet weight
indicates that genetic improvement of one of the traits without achieving a proportional
genetic change in the other is difficult or even impossible. This implies that improvement of
fillet yield through direct selection is difficult to achieve, also indicated by the negligible
predicted selection response for this trait despite being strongly selected for over several
generations. Still, based on the current results, fillet yield is expected to increase with
increasing body weight, both genetically and phenotypically. However, if increased growth
leads to harvest at a younger age rather than higher weight, response in fillet yield may
not be observed. Improvement of fillet weight can be obtained through direct selection for
increased round body weight, a strategy that is both more efficient and less costly, due to
the fact that round body weight can be recorded on the livebreeding candidates .
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