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Aquaculture 184 2000. 221231 www.elsevier.

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The culture performance of monosex and mixed-sex new-season and overwintered fry in three strains of Nile tilapia Oreochromis niloticus / in northern Vietnam
Nguyen C. Dan a,b,) , David C. Little b,c
a Research Institute for Aquaculture No.1 (RIA-1), Dinh Bang, Tien Son, Bac Ninh, Viet Nam Aquaculture and Aquatic Resources Management Program, School of Enironment, Resources and Deelopment, Asian Institute of Technology (AIT), P.O. Box 4, Klong Luang, Pathum Thani, 12120, Thailand c Institute of Aquaculture, Uniersity of Stirling, Stirling FK9 4LA, Scotland, UK b

Accepted 4 October 1999

Abstract Two experiments E1, E2. to evaluate the performance of new-season and overwintered fry in three strains Viet, GIFT and Thai. of Nile tilapia Oreochromis niloticus . were conducted at Research Institute for Aquaculture No.1 RIA-1. in northern Vietnam. The performance of both mixed and monosex fish of the three strains was also compared. The experiments were carried out from May to November E1. and August to November 1997 E2. with overwintered and new-season fry, respectively. Each experiment included a comparison of production in fertilized earthen ponds 6 = 300 m2 . and cages 18 = 1.2 m 3 . suspended within a single pond 1200 m2 .. In the pond trials, the three strains were stocked communally as either monosex or mixed-sex groups, whereas treatments strain = sex. were separated into replicate cages. Individual size of fish stocked was 11.8 " 0.8 g E1. and 9.4 " 0.4 g E2.. Floating pellets were fed at 5% b.wt. dayy1, 30% crude protein. initially, and declined to 2% by the third month of culture. Water quality and fish growth were monitored monthly. Overwintered fingerlings, both mixed and monosex, of all three strains performed well. Daily weight gain DWG. ranged between 1.8 and 2.3 g dayy1 in both ponds and cages. Among the three tilapia strains, the GIFT fish attained a significantly P - 0.01. larger individual final weight 387.7 " 21.0 g in ponds; 410.6 " 35.1 g in cages. compared to the Thai 351.6 " 26.1 g in

) Corresponding author. Research Institute for Aquaculture No.1 RIA-1., Dinh Bang, Tien Son, Bac Ninh, Vietnam. Tel.: q 84-4-878-0408; fax: q 84-4-827-3070. E-mail address: ncdan@fpt.vn N.C. Dan..

0044-8486r00r$ - see front matter q 2000 Elsevier Science B.V. All rights reserved. PII: S 0 0 4 4 - 8 4 8 6 9 9 . 0 0 3 2 9 - 4

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ponds; 373.2 " 30.4 g in cages. and Viet strains 359.5 " 33.2 g in ponds; 350.3 " 33.4 g in cages.. Overall, monosex fish of the three strains grew significantly faster than mixed-sex fish P - 0.05.. DWG of new-season fingerlings E2. in ponds ranged from 1.6 to 1.8 g dayy1 , showing slower growth compared to overwintered fingerlings. There was no significant difference in the growth between the three strains, and between mixed and monosex fish in ponds. Growth of all three strains was significantly slower in cages than ponds overall DWG s 0.80.9 g dayy1 ; P - 0.01.. The results suggest that the performance of overwintered seed compare well with new-season fry. These results are discussed in the context of developing appropriate strategies for tilapia seed productions for northern Vietnam. q 2000 Elsevier Science B.V. All rights reserved.
Keywords: Tilapia; Overwintering; Growth; Survival; Sex

1. Introduction One of the problems of tilapia culture in sub-tropical regions is their poor tolerance of seasonally low temperatures, such as occur in northern Vietnam. Nile tilapia in general does not grow at temperatures below 168C and exhibits poor survival if water temperatures fall below 108C for more than few days Chervinski, 1982.. Growth and reproduction are adversely affected at temperatures of less than 208C and 228C respectively Behrends et al., 1990.. The Nile tilapia requires a minimum temperature of 208C to spawn Hauser, 1977.. Seasonally cold temperatures in northern Vietnam thus affect both hatchery and food fish production as, between December and March, water temperatures drop to between 118C and 208C. Under these conditions, mortalities can occur, especially in shallow ponds, and breeding activities are completely curtailed. Conditions are highly suited to raising tilapia between April and December, when temperatures range from 258C to 358C. However, as spawning only begins in April, seed are not normally available until JuneJuly, reducing the duration of the production season. Overwintering of late-spawned tilapia fry may be a possible way to make fingerlings available early in the following grow-out season. Overwintering of tilapia fry has been successfully carried out using heated facilities Behrends et al., 1990. or underground warm water Cruz and Ridha, 1994. and in green houses Jiazhao, 1991., but these resources are generally unavailable in northern Vietnam. Overwintering late-season fingerlings in deep hapas 3.0 m. suspended in deep ponds 3.5 m. has been demonstrated Nguyen, 1996, unpublished data.. However, the long-term effects of overwintering on subsequent growth and reproduction have not been established. Nile tilapia production is still undeveloped in northern Vietnam but recent initiatives may stimulate rapid expansion. Traditional fish culture is based on polyculture of carps in earthen ponds; small-sized tilapias, where present, have had low market acceptance. Recently, several new improved strains have been introduced, as well as monosex technology, to overcome this problem. Interest in the development of cage culture is also growing. The relevance of these innovations to the needs of farmers and consumers in northern Vietnam are considered in the current study. The purpose of this study was to

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evaluate and to compare the performance of mixed-sex and monosex stocks of three strains of Nile tilapia. Their performance as overwintered E1. and new-season fry E2. was investigated in ponds, and cage-in-pond systems.

2. Materials and methods 2.1. Experimental duration and location Two experiments E1, E2. were conducted in both ponds and cages during the growing season of 1997 at Research Institute for Aquaculture No. I RIA-1. located near Hanoi, northern Vietnam. Overwintered Nile tilapia seed E1. were stocked earlier and cultured longer 15 May to 15 November 1997. than new-season seed E2. 5 August to 25 November 1997.. 2.2. Origin of Nile tilapia strains The three strains of Oreochromis niloticus used in the two experiments were Thai T., GIFT G. and Viet V. strains. The Thai strain originated from Egypt Pham et al., 1998. but experimental stocks were derived from fish transferred from the Asian Institute of Technology AIT., Thailand in 1994. The GIFT strain was derived from the International Center for Living Aquatic Resources Management ICLARM., Philippines in May 1994 ICLARM, 1998.. The Viet V. strain has been maintained in northern Vietnam since its introduction from Taiwan, via southern Vietnam, in 1977 ICLARM, 1998.. 2.3. Production of oerwintered and new-season fry Natural spawning in similar hapa-in-pond systems produced the progeny of each strain for both experiments. Eggs and yolk-sac fry were harvested after the same 5-day period and artificially incubated to the first feeding stage Macintosh and Little, 1995.. The number of brood fish used for fry production was 642 214 brood fish strainy1 with the size ranging from 162 to 250 g and the ratio of male:female was 1:1.. The size of breeding hapa was 40 m2 4 hapas. and size of rearing hapa was 5 m2 24 hapas.. Total number of seed harvested for each breeding time was about 60,000 swim-up fry. Batches of fry were then divided into two groups. One group monosex. was fed a fine fry feed treated with 17a-methyltestosterone dissolved in food grade ethanol using standard sex reversal methods Macintosh and Little, 1995.. The second group mixedsex. was treated in the same way, including the addition of ethanol to the food, but no hormone was added. Overwintered fish E1. were produced in early October 1996 and held in deep hapas suspended in a single deep pond from December 1996 to March 1997. The size of E1 fry before overwintering was 1.11 " 0.07 g. During overwintering, fry were fed with a pelleted feed containing 3035% crude protein at a rate of 12% fish biomass dayy1 on days when water temperature exceeded 168C. Survival rate of fry after overwintering in

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hapas-in-ponds was 78.790.7% in the cold season 19961997 Nguyen, 1997, unpublished data.. New-season fry E2. were produced from brood fish spawned at the end of April 1997 and reared until early August to reach a fingerling size of 10 g prior to stocking in ponds and cages. Survival rate, of fry to fingerlings, exceeded 90%. 2.4. Experimental design and set up Twelve earthen ponds used for communal stocking of the three strains were similar in shape and size, with a surface area of 285295 m2 and water depth of 1.5 m. The six ponds stocked with overwintered fry E1. on 15 May 1997, and other six ponds stocked with new-season fry E2. on 5 August 1997. A total of 36 net cages were used for stocking of the strains separately in both experiments. Nylon net, aluminium-frame cages 1.0 = 1.0 = 1.2 m; 1.2 cm mesh. were suspended 40 cm off the bottom of a single large pond area s 1200 m2 ; 1.4 m deep., with a freeboard of 20 cm above the water surface. River water was added to all the ponds monthly, via an irrigation canal, to replace water losses due to seepage and evaporation and maintain 1.0-m water depth in the cages. Fingerlings of the three strains were stocked at 1.5 fishrm2 in earthen ponds, and 30 fishr1 m3 cage. Individual fish of the three strains were identified by fin clipping T, left pelvic; G, right pectoral; V, anal. before communal stocking in ponds. Treatments were assigned based on blocks of two adjacent ponds, stocked with mixed-sex and monosex fingerlings respectively. This has been found to be the optimal design to reduce the effects of heterogeneity in earthen pond experiments Darmi, 1996.. The three strains monosex and mixed-sex. were randomly assigned to 18 3 = 3 = 2. treatment cages, within three blocks located along a central axis of the pond. A randomized block design was used to quantify possible environmental differences between different parts of the large, wind-affected pond. Fish in both pond and cage-in-pond systems were fed twice daily, 6 daysrweek with a floating pelleted feed containing 30% crude protein. Feeding rate was set at 5% of fish biomass dayy1 in the first month, 3% in the second month and 2% thereafter, adjusted biweekly based on sampled mean weight and survival. Recruits produced from reproduction in the communally stocked ponds were removed during monthly sampling by seine net. Fertilizers urea and TSP. were applied weekly to all experimental ponds, after soaking in water and broadcasting, at rates of 4 kg hay1 dayy1 of N and 1 kg hay1 dayy1 of P. At initial stocking, fish were counted and batch weighed. The initial mean weight of fish was also estimated by individually weighing a sample of 30 fish from each strain before stocking in each pond or cage. Monthly and at final harvest, a sample of 30 fish of each strain in each pond was individually weighed. All fish were measured individually in each cage monthly. At final harvest, all fish were batch weighed and counted after draining of ponds. Males and females in each pond or cage were counted and weighed separately. Methods and timing of water quality sampling and are presented in Table 1. 2.5. Data analyses and interpretations Individual fish growth rate g dayy1 . was determined as mean final weight minus initial weight, divided by experimental period days., based on individual measurements.

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Table 1 Methods of analysis of chemical and physical water quality of experimental units ponds, water quality sampled from a minimum of three locations per pond; cages, one sample per cage. Parameters Temperature and dissolved oxygen DO. pH Chlorophyll-a and phytoplankton biomass Alkalinity Method, frequency and timing of sampling DO meter, measured twice daily at 6:00 a.m. and 3:00 p.m. digital pH meter, measured twice weekly at 6:00 a.m. and 3:00 p.m. acetone extraction and measuring absorbance by using spectrophotometer. Ph. biomass s chlorophyll-a=67, measured weekly U volumetric analysis using phenolphthalein and methyl indicators and standard sulphuric acid titrant, measured weekly U Eriochrome black-T indicator, standard CaCO 3 and standard EDTA titrant, measured weekly U phenate method, measured twice monthly U cadmium reduction procedure, measured twice monthly U diazotizing method, measured twice monthly U ascorbic acid procedure, measured twice monthly U persulphate digestion method, measured twice monthly U Source YSI model 58. Pye model 290. APHA 1980.

Boyd and Tucker 1992.

Hardness Concentration of ammonia NH 3 N. Concentration of nitrate NO 3 N. Concentration of nitrate NO 2 N. Dissolved orthorphosphorous Total phosphorous
U

Boyd and Tucker1992. APHA 1980. APHA 1980. APHA 1980. Boyd and Tucker 1992. Boyd and Tucker 1992.

Water samples are taken at 7:00 a.m.

Feed conversion rates were calculated as the weight of feed given divided by biomass of fish harvested. Comparison of growth rate of the treatments was analyzed for both experiments using factorial analysis for which a significance level of P - 0.05 was set. When significant differences between means was detected, a LSD range test was used to compare mean values P - 0.05..

3. Results 3.1. Grow-out of oerwintered fry and new-season fry in ponds Both strain and use of hormonal sex reversal affected individual final size of fish stocked as overwintered seed in ponds P - 0.05.. The GIFT strain grew fastest 2.1 " 0.13 g dayy1 . and attained a final size 11.1% larger than the Thai and Viet strains which reached a similar size. Considering all strains, MT-treatment resulted in a final size of fish 10.7% larger than mixed-sex fish. However, a strain by strain comparison illustrated that benefits of MT-treatment were minor, except for in the Viet strain. Survival in all treatments ranged from 70.1% to 82.5% and was not significantly

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Table 2 Data of stocking and harvest parameters of overwintered I. and new-season II. seed of three strains of Nile tilapia raised communally as either monosex male or mixed-sex groups in earthen ponds mean values"s.e., replicatess 3. Strains At stocking Number Mean weight g. At harvesting Number Mean weight g. 350.0"28.6 ab 375.3"35.7 bc 338.7"37.6 a 353.1"23.5ab 400.0"26.3 c 380.3"28.7 c 199.5"20.4 a 201.3"21.1ab 197.0"22.5a 211.9"16.9 b 194.2"20.4 a 192.0"19.3 a Daily weight gain g dayy1 . Survival %.

Experiment 1 (oerwintered seed 184 days) Thai mixed-sex 435 10.7"0.7 GIFT mixed-sex 435 10.5"0.8 Viet mixed-sex 435 10.5"0.9 Thai monosex 435 10.7"0.7 GIFT monosex 435 10.3"0.8 Viet monosex 435 9.9"1.2 Experiment 2 (new-season seed 112 days) Thai mixed-sex 435 9.0"0.4 GIFT mixed-sex 435 9.2"0.4 Viet mixed-sex 435 9.3"0.4 Thai monosex 435 9.1"0.5 GIFT monosex 435 9.0"0.3 Viet monosex 435 9.1"0.3

305 358 330 320 316 337

1.8"0.15 2.0"0.19 1.8"0.20 1.9"0.12 2.1"0.13 2.0"0.15

70.1a 82.3 a 75.8 a 73.6 a 72.9 a 77.5a 77.3 a 74.0 a 78.6 a 72.4 a 77.9 a 76.6 a

339 322 342 310 339 333

1.7"0.18 1.7"0.17 1.7"0.16 1.8"0.16 1.7"0.16 1.6"0.17

different Table 2.. Factorial analysis indicated that new-season fry stocked in ponds showed no significant differences in final size that were related to strain or hormone
Table 3 Data of stocking and harvest parameters of overwintered I. and new-season II. seed of three strains of Nile tilapia raised communally as either monosex male or mixed-sex groups in cages mean values"s.e., replicatess 3. Strains At stocking Number Experiment 1 (oerwintered seed Thai mixed-sex 90 GIFT mixed-sex 90 Viet mixed-sex 90 Thai monosex 90 GIFT monosex 90 Viet monosex 90 Mean weight g. 184 days) 12.7"0.8 11.3"0.6 12.4"0.5 11.6"0.5 12.0"0.8 12.3"0.8 At harvesting Number Mean weight g. 373.0"34.1a 391.4"33.3 ab 337.1"33.5a 373.3"26.7 ab 429.7"27.5 b 363.5"33.3 a 102.0"10.6 ab 112.5"11.2 b 97.6"9.7 a 103.5"9.4 ab 105.1"10.9 ab 98.9"9.5a DWG g dayy1 . Survival %.

86 87 85 87 86 86

1.9"0.19 2.1"0.18 1.8"0.18 1.9"0.14 2.3"0.15 2.0"0.18

95.6 a 96.7 a 94.4 a 96.7 a 95.6 a 95.6 a 100.0 a 97.8 a 100.0 a 98.9 a 100.0 a 100.0 a

Experiment 2 (new-season seed 112 days) Thai mixed-sex 90 9.5"0.4 GIFT mixed-sex 90 9.5"0.4 Viet mixed-sex 90 9.6"0.4 Thai monosex 90 9.7"0.5 GIFT monosex 90 9.8"0.5 Viet monosex 90 9.7"0.4

90 89 90 89 90 90

0.8"0.09 0.9"0.09 0.8"0.09 0.8"0.07 0.8"0.09 0.8"0.08

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treatment. A paired comparison indicated that the final size of monosex Thai strain was larger than mixed-sex P - 0.05. however. The survival rate of the three strains ranged from 72.4% to 78.6% showing no significant difference between strains. 3.2. Grow-out of oerwintered fry and new-season fry in cages. Overwintered fry raised in cages showed a similar trend in effects of strain and hormone treatment on performance as fish stocked in ponds. GIFT fish achieved a significantly P - 0.05. larger final size compared to Thai and Viet strain. A comparison of mixed-sex fish revealed no differences between strain however. A paired

Table 4 Monthly average of physical and chemical water parameters in experimental ponds and cages mean values"s.e.. at RIA-1 in period from May to November 1997 Parameters Number of Experimental ponds samples per Mean of Range of experimental parameters parameters unit Experimental cages Mean of Range of parameters parameters

Experiment 1 Temperature 8C. 344 DO mg O 2 ly1 . 344 pH 52 Transparency cm. 52 Orthorphosphorus mg ly1 . 12 Total phosphorus mg ly1 . 12 Nitrite mg ly1 . 12 Nitrate mg ly1 . 12 Ammonia mg ly1 . 12 Hardness mg CaCO 3 ly1 . 24 Alkalinity mg CaCO 3 ly1 . 24 Phytoplankton biomass 24 s chlorophyll-a=67. mg my3 . Experiment 2 Temperature 8C. 220 DO mg O 2 ly1 . 220 pH 32 Transparency cm. 32 Orthorphosphorus mg ly1 . 8 Total phosphorus mg ly1 . 8 Nitrite mg ly1 . 8 Nitrate mg ly1 . 8 Ammonia mg ly1 . 8 Hardness mg CaCO 3 ly1 . 16 Alkalinity mg CaCO 3 ly1 . 16 Phytoplankton biomass 16 s chlorophyll-a=67.mg my3 .

24.632.0 29.6"0.8 24.831.3 3.19.6 4.5"0.6 2.46.2 7.79.1 7.59.1 21.3"5.4 15.626.3 21.9"3.3 15.625.0 0.02"0.02 0.010.05 0.02"0.02 0.010.06 0.07"0.06 0.030.22 0.07"0.07 0.030.09 0.07"0.07 0.010.29 0.13"0.13 0.010.2 0.4"0.2 0.040.94 0.6"0.3 0.10.8 1.2"0.5 0.31.67 1.4"0.6 0.61.60 56.8"8.0 33.370.0 66.9"8.6 61.968.5 62.2"7.4 44.682.7 68.8"7.6 62.982.7 2512.2 716.24774.3 2537.1 1838.53223.4

29.4"0.8 4.4"0.7

28.1"1.0 3.8"0.5 21.9"4.9 0.05"0.03 0.09"0.05 0.07"0.06 0.3"0.2 1.1"0.8 57.1"15.7 59.5"11.4 1899.1

24.431.3 29.0"0.9 4.14.3 4.2"0.4 7.57.9 7.9"0.3 17.534.4 22.2"4.3 0.010.09 0.02"0.01 0.020.14 0.09"0.02 0.020.14 0.02"0.02 0.20.5 0.6"0.4 0.51.9 1.2"0.6 24.592.8 62.7"9.7 40.167.3 65.9"10.5 671.35372.7 2149.1

24.431.6 3.14.5 7.57.7 11.526.6 0.010.05 0.030.21 0.020.11 0.160.92 0.61.7 42.782.5 51.868.9 895.82237.8

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comparison of each strain indicated monosex reached a larger final size than mixed-sex fish, but that hormone treatment resulted in a smaller difference in final size than in pond-reared fish. The mean DWG of fish in all treatments ranged from 1.8 " 0.18 to 2.3 " 0.15 g Table 3.. The survival of mixed-sex and monosex stocks in the three strains ranged between 94.4% and 96.7% and was not significantly different. New-season fry stocked in cages grew poorly compared to fish stocked in ponds 0.80.9 g dayy1 .. There was no effect of hormone treatment and the effect of strain was less distinct. The survival of fish in all cages was very high however, ranging from 97.8% to 100%. 3.3. Sexual maturation of fish Fry were found in ponds stocked with mixed-sex, overwintered fish from the fourth month onwards. Mouth brooding females were also detected in cages stocked with mixed-sex fish from the fourth month onwards. In ponds stocked with new-season mixed-sex fingerlings, there were no fry observed during the period of experiment, and no mouth brooding females found at harvest. 3.4. Water quality parameters The physical and chemical parameters of pond water during both experiments Table 4. were maintained within the tolerance range of Nile tilapia, as reported by several authors Jiazhao, 1991; Popma and Lovshin, 1996.. Levels of net primary productivity did decline from August to November Table 4. explaining the relatively poor performance of fish in E2. The new-season fry, reared in cages grew particularly slowly despite similar water quality to ponds stocked at the same time.

4. Discussion The potential role of overwintered tilapia fingerlings in Northern Vietnam was clearly indicated by the first experiment. Fish of all three strains, both mixed or monosex, reached a size of 300400 g within a culture period of 6 months in either cages or ponds. Tilapia of more than 200 g attract premium prices in Hanoi Cao, 1998.. Growth of overwintered individual fish compared well with other studies. Diana et al. 1991, 1994. observed individual growth of 1.2 g dayy1 in hand-sexed male tilapia raised in ponds receiving high rate of fertilizers and 1.72.0 g dayy1 in fertilized ponds with supplemental feeding. Siddiqui et al. 1989. reported growth rates of 1.71.8 g dayy1 in Nile tilapia reared in out-door tanks. Both mixed-sex and monosex overwintered fry grew well in ponds and cages. Among the three strains, GIFT strain reached a larger individual size in both cages and ponds at final harvest than either the Thai or Viet strains. This is perhaps to be expected given that this strain has been selected for growth performance in a wide range of environments. However, this growth differential is much less than reported for this selected strain compared to unselected controls Eknath, 1995.. The comparable performance of the Viet strain with the two recently

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introduced strains suggests that it should have been promoted more widely in the past. The value of monosex tilapia has been linked both to the population control affected by male only stocks and the faster individual growth of males Mair and Little, 1991.. Using monosex seed can control the stunting problem caused by competition for food that otherwise occur between recruits and stocked fish. The comparison of performance between monosex and mixed-sex groups of the three strains E1. showed that on average, monosex fish grew more than 10% faster than mixed-sex fish in both ponds and cages. Faster growth of monosex tilapia has been related to the lack of energy expenditure in egg production and mouth brooding by females and lower energy expenditure on courtship by males Macintosh and Little, 1995.. Stunting of mixed-sex treatment was not observed however. Although the period before the tilapia reach sexual maturity is short, which is about 3 to 6 months Hepher and Pruginin, 1982., the fish in E1 which were at least 7 months old at stocking did not appear to reach sexual maturation until after another 3 months at a size of around 200 g. The absence of a significant decline in growth rate during the onset of reproduction suggests that the biweekly removal of recruits through seining to sample fish was sufficient to control any impact of recruitment on growth of the stocked fish. This indicates that culture of monosex stock was not required to produce fish of 200 g and that even larger fish could be produced relatively easily through intermittent harvest of recruits. Stocking large overwintered seed early in the season indicated potential for double cropping small fish 200 g., or production of a single large sized fish over the whole season. New-season fry stocked as late as August also reached optimal marketable size 200 g. before cool temperatures retarded growth. This stocking and harvest schedule would have important implications for the use of different strains and the value of hormone treatment. New-season fry showed little difference in performance of monosex compared to mixed-sex, or between strains and a comparison of the growth of overwintered seed to a size of 200 g showed a similar situation. The large differences in initial size and age, timing of stocking and culture duration prevent direct comparison of new-season and overwintered seed performance. An assessment of overwintered and new-season fry performance can suggest practical choices for culturist, however. The growth curves of fish in ponds and cages in both experiments were more or less linear indicating that the critical standing crop in either ponds or cages was not exceeded Diana et al., 1994. possibly due to the initially low stocking density and provision of a pelleted feed in addition to fertilization. The linear growth also suggests that overwintered fish were not stunted. New-season fingerlings E2. of the three Nile tilapia strains grew slowly in cages compared to fish stocked in ponds. Several factors may have influenced this result. Heavy rains occurred from August to October 1997 increasing turbidity in ponds, reducing primary productivity, and limiting the availability of natural food. It is possible that fish that had access to the whole pond were less affected by the reduction in natural food, which is considered more critical for juvenile fish Huet, 1994. compared to those confined in cages. The overwintered fish, which were larger at this stage and relatively more dependent on the supplementary feed, continued to grow fast in the same cage-in-pond culture system.

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A further major factor affecting the growth rates of the overwintered and new-season fry was the relative exposure to lower temperatures occurring in the latter part of the growing season. Whereas overwintered fish grew during the period of optimal temperatures early in the season MayAugust, 30.531.98C., new-season fry were stocked shortly before ambient temperature began to decline AugustNovember; 31.924.98C. Table 4.. Survival of fish was higher in the protected environment of cages ) 94%. than ponds 6586%., however. Regular seining of ponds was observed to cause mortality, but overall survival was in the range normal for pond culture of Nile tilapia Diana et al., 1994..

5. Uncited references Coche, 1982 Philippart and Ruwet, 1982

Acknowledgements This study is a part of a research project to improve capacity for tilapia seed production in northern Vietnam under the AIT Aqua-Outreach Programme. The study was conducted at the Research Institute for Aquaculture No.1 RIA-1., Bac Ninh, Vietnam. The authors acknowledge DANIDA for financial support and Dr. Graham C. Mair for his kind assistance. David Little was seconded to the Asian Institute of Technology by the Department for International Development, UK.

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