Research Article

Growth and Survival Variations of Penaeus monodon from Six Breeding Families  

Y.D. Li1,2,3 , S.G. Jiang1,2 , J.H. Huang1,2 , Z.H. Ma1,2 , F.L. Zhou1,2
1 South China Sea Fisheries Research Institute, Chinese Academy of Fishery Sciences, Guangzhou, 510300, China
2 Key Laboratory of South China Sea Fishery Resources Exploitation and Utilization, Ministry of Agriculture, Guangzhou, 510300, China
3 College of Fisheries and Life Science of Shanghai Ocean University, Shanghai, 201306, China
Author    Correspondence author
International Journal of Aquaculture, 2015, Vol. 5, No. 39   doi: 10.5376/ija.2015.05.0039
Received: 15 Oct., 2015    Accepted: 12 Nov., 2015    Published: 08 Apr., 2016
© 2015 BioPublisher Publishing Platform
This is an open access article published under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
Preferred citation for this article:

Li Y.D., Jiang S.G., Huang J.H., Ma Z.H., and Zhou F.L., 2015, Growth and Survival Variations of Penaeus monodon from Six Breeding Families International Journal of Aquaculture, 5(39): 1-9

Abstract

The survival and growth of P. monodon from six breeding families were evaluated in this study. The morphometric traits, including the carapace length, carapace width, body length, telson length and body weight were measured throughout 108-days growth experiment. Ratios of the telson length and carapace width to body length, the length-weight relationship (LWR), condition factor (K), weight gained rate (WGR), specific weight rate (SWR), and survival of P. monodon were calculated. The testing traits were varied in selected breeding family. Length-weight relationships of family1 can be expressed as W = 0.0126BL3.064 (R² = 0.9473) which had the biggest slope value (b). The equation W = 0.0563BL2.417 (R² = 0.8318) belong to family2, while the slope value less than 3 indicates that the animal becomes slender as it increases in length. The order of the survival rate was family4 > family1 > family2 > family3 > family6 > family5.The order of the specific weight rate and weight gain rate was family6 > family4 > family2 > family5 > family3 > family1. In comparison of the condition factor between different families, the highest condition factor (1.57) was observed in the family5, and the lowest condition factor (1.40) was the family6. According to above indexes, rearing performance of family4 was the best in the tested breeding families in this study.

Keywords
Penaeus monodon; Survival; Growth; Condition factor; Breeding family

1 Introduction
The giant tiger shrimp Penaeus monodon (Fabricius, 1798) is one of the most important species of Penaeus currently being cultured commercially in many countries, due to its rapid growth, low food cost and ability to reach a larger size than other commonly farmed species (Kenway et al., 2006). P. monodon is widely distributed in the Indo-West Pacific Ocean. In South of China, P. monodon is an important indigenous shrimp resource in aquaculture. With existing culture systems and level of technology, P. monodon has by far the fastest growth rate among all penaeids in aquaculture. Nevertheless, for P. monodon breeding seeds, except a few from wild-caught on the sea, the vast majority of shrimp seeds by artificial breeding. Long-term overfishing and reliance on wild broodstock may lead to degradation of germplasm and impact on the overall prawn farming business due to unavailability of broodstock, introduction of disease, decline of disease resistance and an inability to conduct selective breeding. These problems have contributed to the recent decline in P. monodon production in Asia (Macbeth et al., 2007; Sun et al., 2015). For the protection of wild shrimp resources and prevent the losing of genetic diversity, to form stable genetic breeding strains is particularly urgent.
 
As for most cultured aquatic species, P. monodon was anticipated substantial production benefits from a selective improvement program. Selective breeding programs have been initiated for a several aquaculture species, such as Nile tilapia Oreochromis niloticus (Karisa et al., 2006), Pacific white shrimp Penaeus (Litopenaeus) vannamei (Ibarra et al., 2009), Atlantic cod Gadus morhua(Kettunen et al., 2007) and Giant fresh-water prawn Macrobrachium rosenbergii (Kitcharoen et al., 2012). Selective breeding for traits such as growth, survival, and disease resistance has increased efficiency and profitability of production. To date, for P. monodon there has been some progress towards establishing selective breeding programs in Thailand (Damrongchai, 2002), Australia (Kenway et al., 2006; Preston et al., 2009), China (Huang et al., 2009; Sun et al., 2011), and India (Krishna et al., 2011). In shrimp, growth and survival are among the key traits that drive profitability. Increasing the rate of growth and enhancing the ability of disease resistance were critical goals in breeding works of shrimp. Many indexes can be measured to describe the shrimp growth, such as length-weight relationship, condition factor, weight gain rate (WGR), specific weight rate (SWR) and other index, and were widely used in the practice and research in species such as Macrobrachium rosenbergii (Lalrinsanga et al., 2012), P. monodon, and P. esculentus (Benzie et al., 1997; Gopalakrishnan et al., 2014).
 
In the present study, the morphometric traits including the carapace length, carapace width, body length, telson length and body weight were measured, the length-weight relationship (LWR), condition factor, weight gain rate (WGR), specific weight rate (SWR) and survival of P. monodon in six breeding families were evaluated, aiming to distinguish the growth advantage among different breeding families. Results from the present study will provide valuable information on selective breeding in P. Monodon.
 
2 Materials and Methods
2.1 Broodstocks cultivation and mating
The broodstocks used in this study were wild shrimp collected from southern section of the South China Sea in South-east Asia. Females and males were separately cultured in cement tanks (7 × 3 × 2 m). The broodstocks were fed with a conditioning diet of fresh frozen squid, and a clam worm (Nereis succinea). Female broodstocks were artificially inseminated 2-days post-moult using spermatophores extracted from male broodstocks. Each female was unilaterally eyestalk ablated using heated wire snips. The eye was tagged for individual identification and then returned to the tank immediately after artificial insemination. Each batch of eggs was collected, washed and transferred to a tray and nauplii were hatched in the following morning. Then, the nauplii were separated into family batches.
 
2.2 Larval rearing and fluorescent marked
The nauplii from different families were separately cultured in rearing tanks at a density of 30,000 nauplii per tank. The water quality and culture environment in all tanks during this process were same. At 15 days post metamorphosis, offspring of each family was relocated into plankton mesh cages (3 × 3 × 2 m) suspended in a communal pond (1,333 m3).When shrimp at an average body length of 3cm, each shrimp was individually tagged using visible implant fluorescent elastomers. Visible implant elastomer tags (NMTTM, Shaw Island, Washington, USA) were injected intramuscularly into the dorsal, ventral left and ventral right portions of the last abdominal segment in each shrimp. All families reared in a communal grow-out pond (4,000 m3) to ensure that the rearing environment and condition were identical.
 
2.3 Water quality maintenance
Dissolved oxygen and water temperatures were measured by a portable water quality meter (HACH, America) and salinity and pH were measured using a hand refractometer (ATAGO, Japan), and a portable pH meter (SANXIN, Shanghai), respectively. According to the farm records, the study period was from June 25 to October 13, 2014. The water quality recorded at the sampling sites was within the optimum range for P. monodon growth. The environmental parameters were kept at 25 - 32 ℃, 4.8–8.9 mg dissolved oxygen L-1, 25-35‰ salinity and pH 7.8 - 8.6 In the rearing pond, water was daily exchanged and maintained at 120-150 cm deep. All shrimps were fed with a commercial pellet feed (CP Shrimp Feed, Thailand) three times a day at 00:07, 12:00, and 17:30 h.
 
2.4 Data measurement
At the beginning of rearing trial (June 25, 2014), 50 shrimps from each breeding family were measured in triplicates. After 108 days cultured, shrimp were sampled on October 13, 2014. A total of 705 specimens were harvested and measured in this study. Body length (BL) measured was the distance from postorbital edge to tip of telson, with the shrimp fully stretched, carapace length (CL) was from postorbital edge to posterior margin of the carapace, carapace width (CW) (maximum width), telson length (TL) those measurement was taken with a ruler to the nearest 0.1 cm. Body weights (W) were determined with an electronic balance to the nearest 0.01 g.
 
2.5 Statistical analysis
To remove body length related variation, carapace length (CL), carapace width (CW), and carapace height (CH) were recalculated individually by using the following formulas: CL = CL’ × BL / BL’, CW = CW’ × BL / BL’, TL = TL’ × BL / BL’ (CL: calculated carapace length, CW: calculated carapace width, TL: calculated telson length, CL’: measured carapace length, CW’: measured carapace width, TL’: measured telson length, BL: measured body length, BL’: mean body length).
The Survival rate, rations of telson length (TLR), rations of carapace length (CLR), weight gain rate (WGR) and the specific growth rate (SGR) were calculated as follows: 
Survival rate (%) = 100 × Ns/N0,
TLR (%) = 100 × TL/BL,
CLR (%) = 100 × CL/BL,
WGR (%) = 100 × (Wt – W0)/W0,
SGR (%day-1) = 100 × (lnWt - lnW0)/T 
where Ns was the final number of shrimp following the 108 days experimental period; N0, number of larvae at the beginning of the experiment; Wt was the final weight; W0, the initial weight; T, the duration of the culture experiment.
 
The relationship between body length (BL) and wet weight (W) were calculated by the power regression W = a × BLb. The association degree between BL and W was calculated by the determination coefficient (R2). The value of exponent b provides information on fish growth. When b = 3, the increase in weight is isometric. Otherwise it is positively allometric if b > 3, and negatively allometric if b < 3 (Morey et al., 2003).Fulton’s condition factor (K) was estimated from the mean length and mean weight in the sample estimated from this equation, K = 100 W/L3, where K = condition factor, W = mean weight (g), and L = mean body length (cm).
 
Statistical analyses were performed using Excel and PASW Statistics 20.0 statistical software. All data were subjected to one-way ANOVA after confirmation of normality and homogeneity of variance. If significant differences were indicated at the 0.05 level, then Duncan’s multiple range tests were used to test the differences between treatments.
 
3 Results
A total of 96 to 132 individuals were harvested from each family when sampled after 108 days rearing experiment (Table 1). The survival rate was conservatively estimated based on initial 150 shrimps. Family4 showed the highest survival rate (88.7%, Figure 1), and the lowest survival rate (64.0%) appeared in family5. Family1, family2, family3 and family4 had the indistinctive differences in survival (Figure 1).

 

 

Table 1 Descriptive statistics and estimated parameters of length-weight relationships for Penaeus monodon

 

 

Figure 1 The survival rate of P. monodon from six families (mean ± SD, n = 3)

 

The values for elevation (a) and slope (b) together with their corresponding regression coefficient (r2) for the length–weight relationships in P. monodon of six families are presented in Table 1. The relationships vary with different equations according to each family, indicating that the weight increase will be proportional to the cube of length increment. While in family2, family4, family5 and family6, the slope value less than 3 indicates that the animal becomes slender as it increases in length whereas slope greater than 3 denotes stoutness indicating allometric growth in family1 and family3 (Figure 2).

 

 

Figure 2 Length-weight relationship curves of six P. monodon families

 

Ratios of telson length (TLR) and carapace length (CLR) were calculated and showed in Table 2. Highly significant differences in the telson length ratio and carapace length ratio, with family1 had the maximum ratio both telson length and carapace length.

 

 

Table 2 Carapace width, carapace length ratio and telson length ratio for Penaeus monodon

 

Significant differences existed in weight gain rate of six families. Weight gain rate 2311.9%、2894.2%、2443.9%、3619.2%、2649.3% and 3808.2% for family1-6, respectively (Figure 3). Similar trends were observed on the specific growth rate (SGR) among the six families, with the orders were family6 (3.63%) > family4 (3.56%) > family2 (3.34%) > family5 (3.26%) > family3 (3.21%) > family1 (3.01%) (Figure 4).

 

 

Figure 3 The weight gain rate (%) of P. monodon from six families (mean ± SD, n = 3)

 

 

Figure 4 The specific growth rate (SGR) of P. monodon from six families (mean ± SD, n = 3)

 

Upon completing the experiment, condition factor of shrimps was significantly different in selected breeding families (Figure 4, P < 0.05). Family6 exhibited the smallest average condition factor, and family5 exhibited a significantly larger condition factor than other families (Figure 5, P < 0.05).

 

 

Figure 5 The condition factor (K) of P. monodon from six families (mean ± SD, n = 3)

 

4 Discussions
In current aquatic seedling status, pedigree breeding has become one of the most important methods to get good quality varieties. Family selective breeding was using the whole family as select units, whether the family selected or remained according to the phenotypic characters of families. Compared with group breeding, pedigree breeding has many advantages, such as short period, obvious efficiency, flexible operation and prominent excellent characters (Chen et al., 2008; Huang et al., 2009).
 
In recent years, taking pedigree as the research object to carry out the breeding work was rising up gradually, the breeding object including Pinctada facata (Tang et al., 2011), Argopecten irradians (Zheng et al., 2004), Paralichths olivaceus (Chen et al., 2008), Litopenaeus vannamei (Ruan et al., 2013) and so on. There are already a number of examples of successful selection in penaeid species. For example, one generation of selection produced a 4.4% weight growth increase in P. vannamei (Fjalestad et al., 1997), a 21% growth increase in P. vannamei (Argue et al., 2002), an average 11% growth increase in P. japonicus (Hetzelet al., 2000), and the selected five generation increased a 21% growth rate in P. stylirostris (Goyard et al., 2002). Therefore, taking pedigree as object to carry out the breeding work plays an important role in selective breeding.
 
In a commercial selection program, the genetic traits of survival and growth are monitored to ensure selection effort is directed to improving profitability. In this study, results of the survival rate showed significant differences among families. The highest survival rate (88.6 ± 4.0%) was observed in family4, and the lowest survival (64.0 ± 4.0%) was recorded in family5. The survival observed in this study is lower than that reported on five P. monodon semi-sib families by Huang et al. (2014), but higher than that estimated by Kenway et al. (2006). An additional reason may be that our survival rate was estimated for shrimp at a longer experimental period (108 days) or due to environmental variance between our tank environment compared with theirs. A third possibility is differences in genetic variation (Krishna et al., 2011). Nevertheless, both results indicate substantial survival variation of P. monodon from different breeding families, and selective breeding of this species may improve the rearing performance.
 
Length–weight relationship is a useful index to measure variation in the growth of an individual prawn or a group of prawns (Jayachandran and Joseph, 1988), determine possible differences between stocks (King, 2007), and select genetic traits (Daud and Ang, 1995). The growth parameters estimated from the length–weight relationships in the present study are well fall into the ranges in other penaeid species (Primavera et al., 1998; Abohweyere et al., 2008; Gopalakrishnan et al., 2014). However, the regression parameters in varied between families, family 1(b = 3.064) and family 3(b =3.004) followed the positive allometric growth, while other families showed negatively allometric growth. The value of b parameter presented here is different from studies on wild, cultured, and loose-shell-affected P. monodon by Gopalakrishnan et al. (2014), but in line with studies on Triportheus guentheri by Godindo (1997) and studies on Farfantepenaeus paulensis by Peixoto et al. (2004).
 
Weight of shrimps is another trait of economic importance. That family with different Length–weight relationships in growth tended to be different in weight is of biological interest. However, it should not affect the validity of estimating body weight of market shrimp on the basis of body length as the phenomenon is inconspicuous when the shrimp approach harvest size. In many species, particularly livestock, selective breeding programs often start by increasing weight for age (Henryon et al., 2002). This trait is highly correlated with economic returns, is easy to measure, and commonly has a medium heritability. For shrimps, there have been a number of recent workers who have either selected for weight at age, or estimated heritability for weight at age. These include P. vannamei (Gitterle et al., 2005), Marsupenaeus japonicus (Preston et al., 2009), and Penaeus stylirostris (Goyard et al., 2002). The heritability of size in juvenile P. monodon prawns from half-sib mating were reported by Benzie et al. (1997). It is obvious from the present study that significant differences existed in weight gain rate (WGR) and the specific growth rate (SGR) among families. In addition, similar difference and trend were found in weight gain rate and specific growth rate among families. Family6 always exhibited the largest weight gain rate and the largest specific growth rate, and family4 followed. The specific growth rate of six families ranged from 3.06 to 3.63%, this result was in line with the previous study (2.97-4.00%) on 13 families of P. monodon by Sun et al. (2011).
 
Fulton’s condition factor (K) is often used to quantify an animal’s physical wellbeing, and considered to be a useful complement for growth estimate in crustaceans (Rochet, 2000). Heincke (1908) suggests that condition factor of plaice varies with growth, size, sex, season and degree of gonad development. These observations have been confirmed in many species including newfoundland capelin Mallotus villosus (Carscadden and Frank, 2002), eurasian perch Perca fluviatilis (Kurkilahtia et al., 2002), Lahontan cutthroat trout Oncorhynchus clarki henshawi (Robinson et al., 2008), Rasbora tawarensis and Poropuntius tawarensis (Muchlisin et al., 2010). Currently, the genetic correlations for growth-related traits in P. monodon are very limited (Sun et al., 2015). Since Fulton’s condition factor can be used to estimate the growth of crustaceans, it is likely to be applied in selective breeding on shrimp to compare growth performance between different breeding families. In the present study, in comparison of condition factor between different families, the highest condition factor (1.57) was observed in the family5, and lowest condition factor (1.40) was observed in the family6. This results was significantly higher than the condition factor (0.727-1.092) of wild, grow-out and ‘loose-shell affected’ P. monodon studied by Gopalakrishnan et al (2014). An additional reason maybe that our sampling shrimps had smaller size at a younger age and or due to large environmental variance in their wild environment compared with our tank environment. To the best of our knowledge, there is the first time using the condition factor as an index for selective breeding in P. monodon. However, lack of sufficient sample number of each family and not distinguish gender were likely prohibitive to identifying significant associations of the condition factor with sex and growth period. Future studies should examine larger data sets to better determine if the condition factor has significant differences in sex or growth period of P. Monodon.
 
In summary, we evaluated the survival, morphometric traits and growth of P. monodon from six breeding families. Significant differences in survival rate, length-weight relationship, telson length ratio, carapace length ratio, weight gain rate, specific growth rate and condition factor (K) of P. monodon were found among families. We can draw a conclusion that family5 had an advantage in condition factor and family6 had a better growth rate. Comprehensive comparison all indexes, Family4 was the best in the tested breeding families, was thought to be a relative balanced advantage both in survival and growth. Results from the present study will provide valuable information for the following selective breeding works in P. monodon. Methodology used in the present study can also be applied in other similar species.
 
Acknowledgement 
This study was funded by the National 863 Program (2012AA10A409). China agriculture research system (CARS-47), GuangDong Province project of China (2013B020201001), HaiNan Province project of China (ZDXM2014057) and Guangdong oceanic and fisheries project of China (A201300B03).
 
References
Abohweyere P.O., and Williams A.B., 2008, Length-weight relationship and condition factor of Macrobrachium macrobrachion in the Lagos-Lekki Lagoon system, Nigeria, Research Journal of Biological Sciences, 3(11):1333-1336
 
Benzie J.A.H., Kenway M., and Trott, L., 1997, Estimates for the heritability of size in juvenile Penaeus monodon prawns from half-sib matings, Aquaculture, 152: 49-53
http://dx.doi.org/10.1016/S0044-8486(96)01528-1
 
Carscadden J.E., and Frank K.T., 2002, Temporal variability in the condition factors of Newfoundland capelin (Mallotus villosus) during the past two decades, Journal of Marine Science,59:950-958
http://dx.doi.org/10.1006/jmsc.2002.1234
 
Chen S.L., Ting Y.S., Xu T.J., Deng H., Liu S.T., Liu B.W., Ji X.S., and Yu G.C., 2008, Development and characterization for growth rate and disease resistanceof disease-resistance population and family in Japanese flounder (Paralichthys olivaceus), Journal of Fisheries of China, 32(5):665 -673
 
Damrongchai N., 2002, Agricultural biotechnology in Thailand, Asian Biotechnology & Development Review, 5:23–38
 
Daud S.K., and Ang K.J., 1995, Selection of broodstock of tiger prawn, Penaeus monodon Fabricus, on the basis of morphometric traits, Pertanika Journal Tropical Agricultural Science, 18(1):15-20
 
Gitterle T., Rye M., Salte D., Cock J., Johansen H., Lozano C., Suarez J.A., and Qjerde B., 2005, Genetic (co)variation in harvest body weight and survival in Penaeus (Litopenaeus) vannamei under standard commercial conditions, Aquaculture, 243: 83-92
http://dx.doi.org/10.1016/j.aquaculture.2004.10.015
 
Godinho A.L., 1997, Weight-length relationship and condition of the characiform Triportheus guentheri, Environmental Biology of Fishes, 50: 319-330
http://dx.doi.org/10.1023/A:1007316028288
 
Gopalakrishnan A., Rajkumar M., Rahman M.M., Sun J., Antony P.J., Maran B.A.V., and Trilles J.P., 2014, Length-weight relationship and condition factor of wild, grow-out and 'loose-shell affected' giant tiger shrimp, Penaeus monodon (Fabricius, 1798) (Decapoda: Penaeidae), Journal of Applied Ichthyology, 30(1): 251
http://dx.doi.org/10.1111/jai.12269
 
Goyard E., Patrois J., Peignon J.M., Vanaa V., Dufour R., Viallon J., and Bedier E., 2002, Selection for better growth of Penaeus stylirostris in Tahiti and New Caledonia, Aquaculture, 204: 461-468
http://dx.doi.org/10.1016/S0044-8486(01)00831-6
 
Huang Z., Lin H.Z., Huang J.H., Yang Q.B., Wen W. G., Chen X., Zhou F.L., and Jiang S.G., 2009, Growth, feed utilization and whole-body composition of six Penaeus monodon families, South China Fisheries Science, 5: 42–47
 
Heincke F., 1908, Bericht über die Untersuchungen der Biologischen Anstalt auf Helgoland zur Naturgeschichte der Nutzfische. In: IV/V. Bericht über die Beteiligung Deutschlands an der Internationalen Meeresforschung in den Jahren 1905/6–1906/7, Berlin, pp:67–156
 
Henryon M., Jokumsen A., Berg P., Lund I., Pedersen P.B., Olesen N.J., and Slierendrecht W.J., 2002, Genetic variation for growth rate, feed conversion efficiency, and disease resistance exists within a farmed population of rainbow trout, Aquaculture, 209: 59-76
http://dx.doi.org/10.1016/S0044-8486(01)00729-3
 
Hetzel D.J.S., Crocos P.J., Davis G.P., Moore S.S., and Preston N.C., 2000, Response to selection and heritability for growth in the Kuruma prawn, Penaeus japonicus, Aquaculture, 181: 215–223
http://dx.doi.org/10.1016/S0044-8486(99)00237-9
 
Huang Z., Jiang S.G., Lin H Z., Yang Q.B., Zhou F.L., and Huang, J.H., 2014, Comparison on growth,digestive enzymes and non-specific immunity of five Penaeus monodon semi-sib families. Marine Fisheries, 32(2):163-169
 
Ibarra A.M., Famula T.R., Arcos F.G., 2009, Heritability of vitellogenin in hemolymph, a pre-spawning selectable trait in Penaeus (Litopenaeus) vannamei, has a large genetic correlation with ovary maturity measured as oocytes mean diameter, Aquaculture, 297: 64-69
http://dx.doi.org/10.1016/j.aquaculture.2009.09.015
 
Jayachandran K.V., Joseph N. I., 1988, Growth pattern in the slender river prawn Macrobrachium idella (Hilgendorf), Mahasagar, 21(3):189-195
 
Karisa H.C., Komen H., Rezk M.A., Ponzoni R.W., Arendonk J.A.M., and Bovenhuis, H., 2006, Heritability estimates and response to selection for growth of Niletilapia (Oreochromis niloticus L.) in low-input earthen ponds, Aquaculture, 261(2): 479–486
http://dx.doi.org/10.1016/j.aquaculture.2006.07.007
 
Kenway M., Macbeth M., Salmon M., McPhee C., Benzie J., Wilson K., and Knibb W., 2006, Heritability and genetic correlations of growth and survival in black tiger prawn Penaeus monodon reared in tanks, Aquaculture, 259:138-145
http://dx.doi.org/10.1016/j.aquaculture.2006.05.042
 
Kettunen A., Serenius T., and Fjalestad K.T., 2007, Three statistical approaches for genetic analysis of disease resistance to vibriosis in Atlantic cod (Gadus morhua L.). Journal of Animal Science, 85(2): 305-313
http://dx.doi.org/10.2527/jas.2006-112
 
Kitcharoen N., Rungsin W., Koonawootrittriron, S., and Na-Nakorn U., 2012, Heritability for growth traits in giant freshwater prawn, Macrobrachium rosenbergii (de Mann 1879) based on best linear unbiased prediction methodology, Aquaculture Research, 43(1): 19-25
http://dx.doi.org/10.1111/j.1365-2109.2011.02796.x
 
King M., 2007, Fisheries biology, assessment and management, 2nd edn, Blackwell Scientific Publications, Oxford, pp:1-381
http://dx.doi.org/10.1002/9781118688038
 
Krishna G., Gopikrishna G., Gopal C., Jahageerdar S., Ravichandran P., Kannappan S., Pillai S.M., Paulpandi S., Kiran R.P., Saraswati R., Venugopal G., Kumar D., Gitterle T., Lozano C., Rye M.,  and Hayes B., 2011, Genetic parameters for growth and survival in Penaeus monodon cultured in India, Aquaculture, 318: 74-78
http://dx.doi.org/10.1016/j.aquaculture.2011.04.028
 
Kurkilahtia M., Appelbergb M., Hesthagenc T., and Rask M., 2002, Effect of fish shape on gillnet selectivity: a study with Fulton's condition factor, Fisheries Research, 54(2):153-170
http://dx.doi.org/10.1016/S0165-7836(00)00301-5
 
Lalrinsanga P.L., Pillai B.R., Patra G., Mohanty S., Naik N.K., and Sahu S., 2012, Length Weight Relationship and Condition Factor of Giant Freshwater Prawn Macrobrachium rosenbergii (De Man, 1879) Based on Developmental Stages, Culture Stages and Sex, Turkish Journal of Fisheries and Aquatic Sciences,12: 917-924
 
Macbeth M., Kenway M., Salmon M., Benzie J., Knibb W., and Wilson K., 2007, Heritability of reproductive traits and genetic correlations with growth in the black tiger prawn Penaeus monodon reared in tanks, Aquaculture, 27: 51–56 
http://dx.doi.org/10.1016/j.aquaculture.2007.03.018
 
Muchlisin Z.A., Musmanand M., and Azizah M. 2010, Length-weight relationships and condition factors of two threatened fishes, Rasbora tawarensis and Poropuntius tawarensis, endemic to Lake Laut Tawar, Aceh Province, Indonesia, Journal of Applied Ichthyology,26:949-953
http://dx.doi.org/10.1111/j.1439-0426.2010.01524.x
 
Preston N.J., Coman G.J., Sellars M.J., Cowley J.A., Dixon T., Li Y., and Murphy B.S., 2009, Advances in Penaeus monodon breeding and genetics. In: The Rising Tide, Proceedings of the Special Session on Sustainable Shrimp Farming, Aquaculture, (ed. by Browdy, C.L. & Dory, E.J.),  World Aquaculture Society, Baton Rouge, Louisiana, pp. 1–6.
 
Primavera J.H., Parado Estepa F.D., and Lebata J.L., 1998, Morphometric relationship of length and weight of giant tiger prawn Penaeus monodon according to life stage, sex and source, Aquaculture, 164:67-75
http://dx.doi.org/10.1016/S0044-8486(98)00177-X
 
Oujifard A., Amiri R., Shahhosseini G., Davoodi R., and Moghaddam J.A., 2015, Effect of gamma radiation on the growth, survival, hematology and histological parameters of rainbow trout (Oncorhynchus mykiss) larvae, Aquatic Toxicology, 165: 259-265
http://dx.doi.org/10.1016/j.aquatox.2015.06.010
 
Peixoto S., Soares R., Wasielesky W., Cavalli R.O., and Jensen L., 2004 Morphometric relationship of weight and length of cultured Farfantepenaeus paulensis during nursery, grow out, and broodstock production phases, Aquaculture, 241(1-4): 291-299
http://dx.doi.org/10.1016/j.aquaculture.2004.08.008
 
Robinson M.L, Gomez R.L, Rauwb W.M, Peacocka M.M., 2008, Fulton’s body condition factor K correlates with survival time in a thermal challenge experiment in juvenile Lahontan cutthroat trout (Oncorhynchusclarki henshawi), Journal of Thermal Biology, 33:363-368
http://dx.doi.org/10.1016/j.jtherbio.2008.05.004
 
Rochet M.J., 2000, May life history traits be used as indices of population viability? Journal of Sea Research 44(1):145-157
http://dx.doi.org/10.1016/S1385-1101(00)00041-1
 
Ruan X.H., Luo K., Luan S., Kong J.,Xu S.Y., Chen R.J, Chen G.L., 2013, Evaluation of growth performance in Litopenaeus vannamei populations introduced from other nations, Journal of Fisheries of China, 37(1): 34-42
http://dx.doi.org/10.3724/SP.J.1231.2013.38268
 
Sun M.M., Huang J.H., Jiang S.G., Yang Q.B., Zhou F.L., WEN W.G., and JIANG S.G., 2011, Comparison on characteristics of growth and resistance to ammonia among 13 families of Penaeus monodon, Journal of Shanghai Ocean University. 20(4):510-516
 
Sun M.M., Huang J.H., Jiang S.G., Yang Q.B., Zhou F.L., Zhu C.Y., Yang L.S., and Su T.F., 2015, Estimates of heritability and genetic correlations for growth-related traits in the tiger prawn Penaeus monodon, Aquaculture Research. 46: 1363-1368
http://dx.doi.org/10.1111/are.12290
 
Tang J., Liu W.G., Lin J.S., and He M.X., 2011., Evaluation on mid-term growth of 9 families of pearl oyster Pinctada facata, South China Fisheries Science, 7(5):30-36
 
Zheng H.P., Zhang G.F., and Liu X., 2004, Different responses to selection in two stocks of the bay scallop, Argopecten irradians Lamarck (1819), Journal of Experimental Marine Biology and Ecology, 313: 213-223
http://dx.doi.org/10.1016/j.jembe.2004.04.015

 

International Journal of Aquaculture
• Volume 5
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