Aquaculture 243 (2005) 83 – 92

www.elsevier.com/locate/aqua-online

Genetic (co)variation in harvest body weight and survival in

Penaeus (Litopenaeus) vannamei under standard
commercial conditions
Thomas Gitterlea, Morten Ryeb, Ragnar Saltec,d, James Cocka, Harry Johansenb,
Carlos Lozanoa, Jorge Arturo Suáreza, Bjarne Gjerded,*
a
CENIACUA (Centro de Investigaciones para la Acuicultura en Colombia), carrera 8A número 96-60, Bogotá, Colombia
b
AFGC (Akvaforsk Genetic Center), 6600 SundalsØra, Norway
c
Departament of Animal and Aquacultural Sciences, Agricultural University of Norway, N-1432 Ås, Norway
d
AKVAFORSK (Institute of Aquaculture Research), 5010 N-1432 Ås, Norway
Received 13 July 2004; received in revised form 30 September 2004; accepted 13 October 2004

Abstract

In 1997, CENIACUA (Centro de Investigaciones para la Acuicultura en Colombia) in collaboration with AKVAFORSK
(Institute of Aquaculture Research, Norway) initiated a family-based selection scheme in Penaeus (Litopenaeus) vannamei to
improve growth rate and survival in ponds and tanks. This paper reports results from a series of tests in which a total of 430 full-
sib families (representing 204 paternal half-sib families) were tested for harvest weight and survival in standard commercial and
intensive growing environments. The families originated from two selected lines. The estimates of heritability (h 2FS.E.) for
harvest weight for the two lines were 0.24F0.05 and 0.17F0.04, and the corresponding estimates for survival in pond/tanks
were 0.04F0.02 and 0.10F0.02. Results showed a favorable correlation between the estimated mean full-sib family breeding
values for harvest weight and pond/tank survival (0.42 in line 1 and 0.40 in line 2) indicating that selecting for growth will cause
a positive correlated response in overall survival. The genetic correlations between body weights and pond/tank survival in
different grow-out environments were high, demonstrating low genotype by farm interaction (GXE) for both traits.
D 2004 Elsevier B.V. All rights reserved.

Keywords: Shrimps; Heritability; Genetic correlations; Harvest weight; Survival

1. Introduction

The global shrimp farming industry grew contin-

* Corresponding author. Tel.: +47 6494 9514; fax: +47 6494
9502.
E-mail address: bjarne.gjerde@akvaforsk.nlh.no (B. Gjerde).
0044-8486/$ - see front matter D 2004 Elsevier B.V. All rights reserved.
doi:10.1016/j.aquaculture.2004.10.015
uously until the early 1990s when production became
highly unstable and total output declined. It was only
at the turn of the century that production again began
84 T. Gitterle et al. / Aquaculture 243 (2005) 83–92

to grow consistently (FAO, 2001). Infectious diseases rearing seasons and years and over consecutive
represented a major cause for the variation in generations and thus reduce the efficiency of selecting
production, and the rapid spread of diseases has been for this trait (Gjedrem, 1983).
ascribed to environmental degradation and regular use Nevertheless, pond survival and growth rate
of larvae or broodstocks captured from wild popula- determine the total harvest yield (kg/ha/year) in
tions for aquaculture purposes (Gjedrem and Fimland, shrimps. Therefore, estimation of the genetic param-
1995). The latter substantially increases the risk of eters for these two traits including their genetic
introducing diseases to the commercial farms, and correlation under standard commercial rearing con-
also reduces the potential benefits of domestication ditions is required for evaluating the potential to select
and genetic improvement. for and thus maximize the response in total yield.
Selective breeding of farmed populations can adapt This paper presents results from the selective
the animals to artificial conditions and also improve breeding program for P. vannamei in Colombia in
traits of economic interest such as survival and growth terms of genetic variation for harvest body weight
rate (Gjedrem and Fimland, 1995). Although the and survival in ponds and tank and their genetic
benefits of selective breeding have been amply correlations.
demonstrated in livestock and several fish species
(Gjedrem, 2000), systematic selection has rarely been
applied in shrimps. 2. Materials and methods
On the Colombian Atlantic coast the major disease
problem from 1994 to 1999 was due to the Taura 2.1. Genetic material
Syndrome Virus (TSV), which caused high mortalities
during the first years after its appearance. However, The data were obtained from animals of the on-
after two consecutive generations, the survival rates of going breeding programme initiated by CENIACUA
the selected animals were on average 20% higher than in 1997. Three base populations (batches 1, 2 and 3)
those of the unselected ones (Gitterle, 1999). By 1999, were established from a total of 87 sires and 154 dams
imported seed was no longer stocked in Colombia, originating from eight countries in Latin America
TSV was considered to be under control. Presently, (Table 1). Batches 4, 5, 6 and 7 were the corresponding
TSV occurs only sporadically and is not reported to descendant generations from the three base popula-
cause mortalities exceeding 40%. tions (Table 2). Two genetic lines were founded from
Due to the good results obtained with the indi- the three base populations. Line 1 (L1) is made up of
vidual selected seed, CENIACUA in collaboration
with AKVAFORSK initiated a breeding program for
Table 1
Penaeus (Litopenaeus) vannamei in 1997 based on a
Number of P. vannamei from different geographical areas used to
combined family and within family selection scheme produce the three base populations
to improve growth rate and pond survival. Growth
Country Batch 01 Batch 02 Batch 03
rate is one of the most desirable economic traits in
Males Females Males Females Males Females
aquaculture and it has been shown to have moderate
to high heritabilities in Penaeus species (Benzie et al., Colombiaa 4 28 5 52 6 35
ColxVenea,* 8 12
1997; Hetzel et al., 2000; Argue et al., 2002; Goyard Costa Ricab 5 1
et al., 2002; Perez-Rostro and Ibarra, 2003a,b) as well Ecuadorb 4
as in many fish species (Gjedrem, 2000). Pond Hawaiia 5 3
survival is another desirable economic trait in Panamab 3 18 12 9 2
shrimps. However, less attention has been paid to Perub 5
Salvadorb 4
selecting for this trait since pond survival is influ- Venezuelaa 4 6 9 21
enced by a number of environmental factors including Total 26 52 26 52 35 70
accidents or management problems (Fjalestad et al., a
Cultured.
1993). Furthermore, the causes of pond mortalities b
Wild stocks of P. vannamei.
may differ across rearing environments, between * Cross between Colombian and Venezuelan stocks.
 T. Gitterle et al. / Aquaculture 243 (2005) 83–92 85

Table 2 after 8 h and a random sample of about 30,000 nauplii

Association between P. vannamei batches and generations for Line
1 and Line 2
Line Batches Generations
1 1 F0
4 F1
6 F2
2 2+3 F0
5 F1
7 F2
batch 1 and their selected generations (batches 4 and 6)
and Line 2 (L2) was based on batch 2 and 3 and their
selected generations (batches 5 and 7) (Table 2). Each
batch was produced over a period of 1–3 weeks (Table
3) mainly using a nested mating design with females
nested within males (full-sibs and paternal half-sib)
and on a few occasions with males nested within
females (full-sibs and maternal half sibs), depending
on the availability of mature breeders (Table 3). In total
448 full-sib families representing offspring from 204
sires and 442 dams were produced (Table 3).
2.2. Production of families
To produce the full and half-sib families, two
mature female breeders were each artificially insemi-
nated by covering their thelycum with the spermatic
mass stripped out from one of the two spermatophores
of a male breeder. The inseminated females were then
placed in individual spawning tanks. Spawning took
place after 1 to 4 h, and the eggs from each female
were collected in 9-l tanks and washed. Eggs hatched
from each full-sib family were then stocked in
separate 250-l larvae-culture tanks. Larvae were fed
a mixed diet of Chaetoceros sp., Artemia sp. and
micro pellets. During the larvae-culture period 11, 1, 4
and 2 families were lost in batches 1, 3, 4 and 5,
respectively.
At the post-larvae 10 stage (PL10) random samples
of 600 postlarvae per family were transferred to
separate net cages (2.5 m3, 1000 Am mesh size) in an
outdoor concrete tank for on-growing until they
reached a minimum tagging size of 1–2 g. In batch
7, 280 animals per family were stocked in two
different nets in order to reduce the density within
net and thus the time until tagging. In the nets, the
postlarvae were fed ad libitum four times a day.
Prior to tagging, seven animals per family were
sampled for the following diagnostic analyses: In situ
hybridization and histopathology analyses were run
for Taura Syndrome Virus (TSV) and Necrotizing
Hepatopancreatitis (NHP) and from batch 3 onwards
the same diagnostic analyses were run for White Spot
Syndrome Virus (WSSV) and additional nested PCR
tests were done (Lo et al., 1996). All samples were
negative for all diseases in both Lines, except for
batch 1 where a TSV outbreak occurred during the
growing period in the net cages and most of the
families were infected. However, only 20% of the
animals died during this period.
Animals were tagged after spending from 5 to
20 weeks in the nets at an average weight of 1.1 to
6.3 g (Table 4). The families of batch 3 and 5

Table 3 Table 4
Production date and period, number of full-sib and paternal and
Number of weeks P. vannamei spent in the nets until tagging,
maternal half-sib families of P. vannamei produced for each batch
average tagging size (g), number of families tested for harvest body
Batch Production Number of families weight and pond/tank survival for each batch
Date Period Full-sib Paternal Maternal Batch Number of Average Number of full-sib familiesa
(weeks) half-sib half-sib weeks in tagging
Ponds Tanks
1 Apr-1998 2 52 26 0 the nets size (g)
2 Oct-1998 1 52 26 0 1 12 2.8 41 (2) –
3 Feb-1999 2 70 35 0 2 10 1.8 52 (2) –
4 Oct-1999 3 68 31 2 3 20 2.8 69 (1) 69 (1)
5 May-2000 2a 68 32 0 4 9 1.1 64 (3) 64 (1)
6 Feb-2001 3 68 28 6 5 15 6.3 66 (2) 66 (4)
7 Aug-2001 3 70 26 4 6 5 2.1 68 (1) 68 (5)
Total 448 204 12 7 8 2.2 70 (1) 70 (3)
a a
Two full-sib families were replaced in week 4 after the first Number in parenthesis indicate no. of rearing environments
mate. (farms or tanks).
86 T. Gitterle et al. / Aquaculture 243 (2005) 83–92

remained in the cages for longer period than the
other batches: batch 3 as a biosecurity measure
until the selected rearing farms where screened for
the presence of WSSV and families of batch 5
because of problems with the importation of the
colored fluorescent elastomers for tagging the
families.
A family color code was given to a random
sample of individuals from each full-sib family by
injecting differently colored fluorescent elastomers
into the 6th abdominal segment of each animal
(Godin et al., 1996). The tagged individuals from
each full-sib family were transferred to commercial
grow-out ponds at different farms and to bio-secure
tanks within the breeding nucleus in the CENIA-
CUA research facility on the Atlantic coast (Tables
4 and 5). Between 28 to 115 and 20 to 164 animals
per family and batch were stocked at the commer-
cial grow-out ponds and tanks, respectively. The
number of families evaluated per batch is given in
Table 4.
2.3. Growth and survival test
Grow out tests were conducted in four commer-
cial farms and in tanks in the CENIACUA research
facility. The farms were located in different areas of
the Atlantic coast of Colombia and represented a
range of commercial production environments, in
particular with respect to salinity (range 0–35 ppt)
and natural pond productivity. The animals in the
farms were raised under standard commercial
conditions during the grow-out phase. Standard
rearing densities of 18 to 22 animals per square
meter were obtained by stocking ponds or tanks
with both tagged individuals and untagged animals
of similar size from commercial hatcheries. The
ponds had a water exchange rate of 15–30% of the
total water volume per day. From batch three
onwards, a random sample of tagged individuals
from all families were kept at CENIACUA’s
research facilities as a biosecurity measure. For this
purpose, on average 117 (batch 3) and 164 (batch

Table 5
Number of P. vannamei tested (total and per family), mean and coefficient of variation (CV) for harvest body weight, survival from tagging to
harvest for each farm and batch, and mean sex differences
Line Gen Batch Test Number tested No. of Body weight Pond
environment
weeks survival
farm Total Per family Mean (g) CV (%) Sex differenceF Sexb difference (%)
S.E. (g)a (%)
1 0 1 P1 2388 58 14 13.8 24.6 1.00F0.16* 7.0 77.7
P2 1887 46 18 17.8 19.1 0.57F0.15* 3.2 53.9
1 4 P1 1267 20 21 21.1 22.3 0.99F0.28* 4.7 42.8
P2 1207 19 18 22.8 17.1 1.10F0.22** 4.8 66.2
P5 1688 27 16 18.2 18.1 0.75F0.16* 4.1 54.4
T1c 5601 89 20 27.2 20.2 3.50F0.14** 12.9 74.2
2 6 P2 2366 35 17 28.2 14.9 1.49F0.17** 5.3 69.6
T1c 6743 20 18 21.7 19.4 1.13F0.10** 5.2 56.9
2 0 2 P2 2245 43 33 23.9 16.7 1.54F0.17** 6.4 37.4
P3 937 18 31 17.4 14.9 0.27F0.21 1.6 18.6
0 3 P2 2893 42 23 20.8 16.3 0.29F0.13* 1.4 51.8
T1c 7042 102 21 22.6 21.2 1.31F0.11** 5.8 62.1
1 5 P1 1678 25 19 13.8 29.0 0.25F0.20* 1.8 53.3
P5 1033 16 15 16.3 18.4 0.15F0.19 0.9 32.5
T1c 4080 15 16 22.2 24.3 0.91F0.17** 4.1 77.2
2 7 P3 1519 22 17 18.6 17.7 0.96F0.17** 5.2 43.4
T1c 8186 39 15 18.0 18.9 0.46F0.08* 2.6 76.4
a
Females minus males.
b
(Sex difference divided with respective pond or tank mean)
100.
c
Based on tanks means within the breeding nucleus at Ceniacua’s facilities.
* Pb0.05.
** Pb0.001.
 T. Gitterle et al. / Aquaculture 243 (2005) 83–92
4) tagged individuals from each full-sib family were
stocked in a concrete tank of 125 t capacity, while
20, 34 and 50 individuals from each full-sib family
from batches 5, 6 and 7 were stocked in four, five
and three fiberglass tanks of 50 t capacities,
respectively. The families in the tanks were main-
tained with low water exchange (less then 5% of
the total volume daily) and strong aeration. The
stocking densities in the tanks ranged from 26 to 90
animals per square meter. A total of 173 full-sib
families from Line 1 and 257 from Line 2 were
tested for harvest body weight and pond and tank
survival (Table 4).
The families grew in ponds and tanks for a
period between 14 and 23 weeks in all batches
except batch 2, which was maintained in ponds for
31–33 weeks due to low growth rate. The harvest-
ing and recording procedures were similar in all
batches. In the ponds tagged animals were sepa-
rated from the untagged ones. The family code and
sex of each tagged animal was determined, the
weight recorded and a unique numeric eye tag was
placed on each individual. At the commercial farms
all surviving animals were transferred to new ponds
for rearing at lower density until sexual maturity,
except for one farm from which the animals from
batch 2 were transferred to a tank at CENIACUA’s
facilities as a safeguard due to reported diagnosis of
White Spot Syndrome Virus (WSSV) on the
Colombian Pacific coast. At CENIACUA’s facili-
ties, surviving animals were transferred to different
concrete and fiberglass tanks and reared until sexual
maturation.
2.4. Statistical analyses
The appropriate variance components for the
random effects in each batch for the two studied traits
harvest body weight and pond and tank survival were
estimated using a univariate mixed linear animal
model (Madsen and Jensen, 2002). In matrix notation
the model can be written:
y ¼ Xb þ Za þ Wc þ e ð1Þ
where y is the vector of observations (for pond/tank
survival the vector is the scores 0 and 1 assigned to
dead and surviving animals in each batch, respec-
tively; for harvest body weight the vector is the
87
individual body weights), b is the vector of fixed
effects (farm by pond/tank by sex effect for harvest
body weight and farm by pond/tank effect for
survival in ponds and tanks), a~(0, Ar a2) is the
vector of additive genetic values, c~(0, Ir c2) is the
vector of random effects common to full-sibs caused
by factors other than additive genetics (i.e., environ-
mental effect caused by the separate rearing of each
full-sib family until tagging, non-additive genetic
effects and maternal effects) and e~(0, Ir e2) is a
vector of random errors. X, Z and W are known
design matrices assigning the observations to levels
of b, a, and c, respectively. A is the additive
genetic relationship matrix, and I is an identity
matrix. Additionally, an overall estimate for the
variance components for both traits harvest body
weight and pond and tank survival were obtained
for each line using model 1 including batch as
additional fix effect.
The heritability was calculated as h 2 =r a2 /
(r a +r c2+r e2) and the effect common to full-sibs other
2
than additive genetics as c 2=r c2/(r a2+r c2+r e2).
For each test environment, full-sib family breed-
ing values for harvest body weight and pond/tank
survival were obtained as 1/2a s+1/2a d, where a s
and a d is the estimated breeding values of the dam
and the sire, respectively, obtained from the above-
mentioned animal model and with the estimated
variance component as inputs (Madsen and Jensen,
2002).
2.5. Genetic correlation
Only animals alive at harvest provided data on
harvest body weight. Therefore, the genetic correla-
tion between harvest body weight and survival could
not be calculated. Instead, product moment correla-
tions between the full-sib family breeding values for
harvest body weight and pond/tank survival were
calculated. When the number of full-sibs per family is
high this correlation is close to the true genetic
correlation between the traits.
Estimates of the genetic correlations between
harvest body weight recorded in different tests
environments (ponds and tanks), and similarly
between survival recorded in different test environ-
ments, were calculated for each batch using a multi-
trait analysis (model 1). In these analyses records
88 T. Gitterle et al. / Aquaculture 243 (2005) 83–92

obtained for the same trait in different environments Table 7

were treated as different traits (Falconer, 1952). Correlations between P. vannamei full-sib family breeding values
for harvest body weight and survival in ponds/tanks for each batch
and line
Line Batch Correlation
3. Results
L1 1 0.48*
4 0.24**
Descriptive statistics for harvest body weight and 6 0.16
survival from tagging to harvest for each batch and All 0.42***
farm is given in Table 5. Mean harvest weight varied
substantially between batches. These large differences L2 2 0.28**
3 0.18
could be due to the differences in the grow-out period
5 0.10
and differences in the growing conditions with respect 7 0.11
to temperature, salinity and pond productivity. In all All 0.40***
batches except for batch 6 the growth rate in tanks * Pb0.01.
were greater than those in ponds (Table 5). The ** Pb0.05.
coefficient of variation for harvest body weight *** Pb0.001.
averaged 19.6% over all batches and farms, and
showed large variation between farms. 3.1. Genetic parameters
With the exception of one farm in batch 5 and one
farm in batch 2, females were consistently heavier The estimated heritabilities for harvest body weight
than males at the end of the grow-out period (Table 5). varied substantially between the different batches
The mean sex difference expressed as % of the pond (Table 6). The overall estimate for Line 1 (0.24F
mean at harvest weight was 4.5%, but varied 0.05) was slightly higher than for Line 2 (0.17F0.04)
substantially among test environments (Table 5). In (Table 6). Estimated heritabilities for pond/tank sur-
ponds, the product-moment correlation between the vival were in general low, and lower for Line 1
mean harvest weight and the sex difference was 0.72 (0.04F0.02) than for Line 2 (0.10F0.02) (Table 6).
( Pb0.001) showing that the sex difference increases For both studied traits the estimates of the effect
with increasing mean body weight at harvest. common to full-sibs other than additive genetics (c 2)
Survival varied substantially between batches and varied substantially between the different batches. For
between ponds and tanks within batches. However, in body weight these estimates were in general lower
spite of this there was no verification of the presence than the heritability estimates, while the estimates for
of specific pathogens, either in ponds or tanks. pond/tank survival were equal to or higher than the
heritability estimates. For body weight the estimate of
Table 6 Line 1 (0.07F0.01) was slightly lower than for Line 2
Heritability (h 2FS.E.) and the effect common to full-sibs other than (0.09F0.02), while for pond/tank survival the esti-
additive genetics (c 2FS.E.) for P. vannamei harvest body weight mate was slightly higher for Line 1 (0.05F0.01) than
and survival in ponds/tanks for each batch and line for Line 2 (0.02F0.01).
Line Batch Body weight Pond survival The correlations between the full-sib family
2
h FS.E. 2
c FS.E. h 2FS.E. c 2FS.E. breeding values for harvest body weight and pond/
1 1 0.54F0.22 0.00F0.05 0.12F0.06 0.00F0.02 tank survival were all positive (Table 7). The overall
4 0.21F0.11 0.08F0.04 0.04F0.04 0.06F0.02 estimates for each line were very similar, 0.42 for Line
6 0.15F0.08 0.08F0.03 0.04F0.03 0.04F0.01 1 and 0.40 for Line 2, and both were significantly
All 0.24F0.05 0.07F0.01 0.04F0.02 0.05F0.01 different from zero.
2 2 0.23F0.11 0.02F0.04 0.11F0.05 0.01F0.02
The genetic correlations between harvest body
3 0.11F0.15 0.17F0.08 0.03F0.09 0.11F0.05 weights recorded in the different commercial farms
5 0.01F0.09 0.13F0.04 0.03F0.03 0.03F0.01 were very high and the genetic correlations between
7 0.40F0.11 0.01F0.02 0.02F0.02 0.02F0.01 harvest body weight in ponds and tanks were
All 0.17F0.04 0.09F0.02 0.10F0.02 0.02F0.01 generally lower but also high (Table 8).
 T. Gitterle et al. / Aquaculture 243 (2005) 83–92 89

Table 8
Genetic correlations (FS.E.) for harvest body weight (BW) and pond/tank survival (S) of P. vannamei recorded at different test environments;
farm vs. farm and farm vs. tanks
Line Batch Farm vs. farm BW S Farms vs. tank BW S
L1 1 1 vs. 2 0.98F0.08 0.97F0.66
4 1 vs. 2 0.99F0.09 ne 1 vs. t 0.71F0.22 0.99F5.13
4 1 vs. 5 0.99F0.16 0.99F3.77 2 vs. t 0.73+0.21 0.19F1.3
4 2 vs. 5 0.98F0.20 0.33F0.68 5 vs. t 0.65+0.29 0.99F0.52
6 2 vs. t 0.81F0.24 0.99F3.3
L2 2 2 vs. 3 0.97F0.05a 0.99F0.11
3 2 vs. t 0.94F0.02a 0.99F2.6
5 1 vs. 5 0.97F0.04a 0.04F0.7 1 vs. t 0.82F0.07 0.29F2.4
5 5 vs. t 0.81F0.07 0.68F1.0
7 3 vs. t 0.83F0.13 0.85F2.2
ne=not estimable.
a
Estimate obtained after excluding the random effect common to full-sibs.

The genetic correlations between survival in the
different ponds and between ponds and tanks were
generally positive, although a couple of negative
estimates were obtained (Table 8). However, most of
the estimates had very high standard errors, probably
caused by the low genetic variance for pond survival
in most of the farms (some close to zero). The two
estimates that were significantly different from zero
were positive and very high (batch 2 and 4 in Table 8).
4. Discussion
An important point in a commercial breeding
program is to emulate commercial conditions so that
animals evaluated and selected for superior traits
during the selection process manifest the same traits
under commercial conditions. Therefore, in all
batches the performance tests were done in a standard
Colombian farming system. On the Colombian
Atlantic coast the normal farming practice is semi-
intensive. Earthen ponds of average size of 6.5 ha are
stocked at an average density of about 20 animals per
square meter. The standard period between stocking
and harvesting is about 16 weeks, with a final
survival commonly between 60% and 80% and
harvest weights of 12–14 g (Anonymous, 2003). In
this study the period from stocking to harvest was in
general longer than in the industry, mainly since
families had to be reared separately until they reached
a size suitable for tagging. In order to ensure a longer
rearing period in ponds after tagging than in net-
cages before tagging, the former period was
extended. This decision probably favored the trend
of higher harvest weights and lower survival than in
the industry. Nevertheless, the harvest weight and
pond survival obtained in all batches are within the
ranges obtained under the commercial conditions in
Colombia.
4.1. Body weight
The heritability estimates for body weight in Line 1
(0.24F0.05) and Line 2 (0.17F0.04) are in accord-
ance with previous findings in shrimp. Additive
genetic variance for body weight has been demon-
strated in several shrimp species. In P. vannamei,
Argue et al. (2002) reported very high realized
heritability estimates (outside the parameter space)
for harvest weight. However, Carr et al. (1997),
reported lower heritability estimates for the same trait
(0.42F0.15). And more recently, Perez-Rostro and
Ibarra (2003a,b) reported heritability estimates of
0.17F0.06 and 0.34F0.18, respectively, based on
full-sib information. In Penaeus japonicus, Hetzel et
al. (2000) reported a realized heritability ranging from
0.16 to 0.31 and in Penaeus monodon Benzie et al.
(1997) reported a paternal half-sib heritability of 0.10
and a maternal half-sib heritability of 0.39. In Penaeus
stylirostris, Goyard et al. (2002) reported a realized
heritability of 0.47.
The large number of full and paternal half-sib
families involved and the application of an animal
model that accounts for effects common to full-sib
other than additive genetics, makes the current
heritability estimates highly accurate.
90 T. Gitterle et al. / Aquaculture 243 (2005) 83–92
The lower heritability estimates obtained in
Batches 3 and 5 compared with the other batches
may be due to the longer period of separate rearing
before tagging in these batches (Table 3). This view is
supported by the higher effect common to full-sibs
(c 2), which could explain the lower heritability
estimates for harvest body weight in these batches.
The effect common to full-sibs is caused by the
separate rearing of the families in the net-cages until
tagging and by possible maternal and non-additive
genetic effects, which are confounded in the present
study. For harvest body weight this effect accounted
for from 7% to 9% of the total phenotypic variance.
The magnitude of this effect in the present study
shows that body weight records should take account
of this effect when estimating heritability and breed-
ing values for harvest body weight. Efforts should be
made to reduce this effect as much as possible by
standardizing the environmental conditions for all
families and by keeping the period until tagging size
as short as possible.
The very high genetic correlations between
harvest body weight in the different test environ-
ments (between ponds and between ponds and tanks)
suggest that the genotype by test environment
interaction (GXE) is low. Recently Perez-Rostro
and Ibarra (2003b) reported low GXE in several
growth estimates, but the authors indicated that these
low estimates might be due the short period that the
families spent in a common environment as com-
pared with that in separate environments. In our
study, the full-sib families spent from 22% (batch 6)
to 48% (batch 3) of their total production time, from
hatching to harvest, in the net-cages. Despite the
large differences in common rearing between
batches, the genetic correlations between harvest
body weights in different test environments were
high and highly consistent (Table 8).
These results strongly indicate a low genotype
by test environment (i.e., ponds and tanks) inter-
action for harvest body weight in P. vannamei
under the range of test environments investigated.
Consequently, the testing of families in the breeding
program and the commercial rearing of the off-
spring (i.e., the grow-out animals) could take place
in ponds as well as in tanks. However, to reduce
risk, testing should take place in the most prevailing
grow-out environment. In Colombia this is earthen
ponds, but production from intensive farming may
increase in the future.
The observed sexual dimorphism, with females
heavier than males, agrees with observation by Chow
and Sandifer (1991), Argue et al. (2002) and Perez-
Rostro and Ibarra (1999, 2003b). This sexual
dimorphism has also been observed in Penaeus
aztecus (Parrack, 1979), Penaeus indicus (Devi,
1986), Penaeus longistylus (Dredge, 1990), and P.
monodon (Hansford and Hewitt, 1994). The positive
correlation coefficient between mean harvest weight
and sex difference in the ponds implies that sexual
dimorphism is more pronounced at high than at low
body weights. Perez-Rostro et al. (1999) reported
sexual dimorphism only in animals heavier then 10 g
and Chow and Sandifer (1991) found that females
were only heavier than males when they were heavier
than 13.2 g. In P. monodon the differences in growth
between females and males appear to result from a
greater increase in weight per molt cycle, rather than
a higher molting rate (Hansford and Hewitt, 1994).
For commercial purposes it would be desirable to
select for a sex ratio that favors females, however
Argue et al. (2002) found no genetic variance for this
trait in P. vannamei.
4.2. Survival
The heritability estimates for survival in both tanks
and ponds were low but significant in both lines. Low
heritability estimates for survival (0.11 or lower) have
also been reported in abalone (Jonasson et al., 1999);
Atlantic salmon (Standal and Gjerde, 1987; Rye et al.,
1990; Jonasson, 1993), chinook salmon (Withler et
al., 1987) and rainbow trout (Rye et al., 1990).
Somewhat higher estimates (0.2–0.3) were reported
for coho salmon (Swift et al., 1991; Murray et al.,
1993). There are few reported estimates of genetic
parameters for survival in shrimps. Wong and
McAndrew (1990) reported a realized heritability of
h 2=0.24 for freshwater tolerance in larvae of Machro-
brachium nipponense, and Argue et al. (2001)
obtained a half-sib heritability between 0.11 and
0.16, when families of P. vannamei were exposed to
high concentration of ammonia. Argue et al. (2002)
reported a low heritability estimate for survival
(0.10F0.07) in a recirculating raceway and a negative
estimate for the same trait in a round pond.
 T. Gitterle et al. / Aquaculture 243 (2005) 83–92
As already mentioned the causes of mortalities
may differ among test environments within a batch,
as well as among batches representing different
generations. In spite of TSV being endemic on the
Colombian Atlantic coast, there was no confirmed
TSV or other specific disease outbreak in the ponds
or tanks during the grow-out period. However, the
high values of the genetic correlations between the
test environments where the estimates were signifi-
cantly different from zero indicate that the causes of
mortalities could be similar in these test environ-
ments. Survival in the test environments could thus
be due to an innate resistance to diseases or to
environmental stress that could be important to select
for even though the heritability of this trait was found
to be low.
The correlations between harvest body weight and
survival were all positive. This positive correlation is
a further indication that the main source of the
mortalities in the ponds was not TSV since the
genetic correlation between TSV resistance and
growth in P. vannamei has been found to be negative
(Argue et al., 2002). The sign and the magnitude of
the genetic correlation between harvest body weight
and survival found in this study would indicate that
selecting for growth will cause a positive correlated
response in overall survival.
A susceptible population under constant infection
or stress pressure may reduce its production capacity
(yield, growth, etc.) and therefore the observed
production is less than its production potential (Van
der Waaij et al., 2000). By deterministic modeling,
Van der Waaij et al. (2000) showed that by selecting
for production under constant infection or stress
pressure, the levels of resistance increases as well,
and the increased production during the first gen-
erations are mainly caused by improving the level of
specific resistance. Therefore, a positive genetic
correlation between harvest body weight and survival
is to be expected, at least in the first generations of
selection of any of the two traits.
5. Conclusions
The findings in this work suggest that harvest
body weight and survival in ponds and tanks are
under additive genetic control and are expected to
91
respond to selection. Since body weight and survival
both have a major influence on total yield in shrimps,
genetic improvement of both traits is economically
very important. Furthermore, the favorable correla-
tion between mean full-sib breeding value for these
two important traits indicates that both can be
improved simultaneously and result in increased
yields (kg/ha/year).
Acknowledgements
The authors wish to thank Andres Suarez, Jaime
Faillace and Rafael Vergara for the production of the
families.
This work was financially supported by a joint
grant from Colciencias, Proexport, and the Colombian
Ministry of Agriculture.
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