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Short communication: The effect of different pH and photoperiod regimens on the survival rate and developmental period of the larvae of Portunus pelagicus (, Brachyura, )

Item Type article

Authors Ravi, R.; Manisseri, M.K.

Download date 28/09/2021 09:20:50

Link to Item http://hdl.handle.net/1834/37376 Iranian Journal of Fisheries Sciences Short Communication 12(2) 490-499 2013 ______The effect of different pH and photoperiod regimens on the survival rate and developmental period of the larvae of Portunus pelagicus (Decapoda, Brachyura, Portunidae)

Ravi R.*; Manisseri M. K. Received: December 2011 Accepted: August 2012 -Marine Biodiversity Division, Central Marine Fisheries Research Institute, Post Box No. 1603, Kochi-682 018, India *Corresponding author’s e mail: [email protected]

Keywords: Portunus pelagicus, Photoperiod, Larval rearing, Larval development, Larval survival.

Crustacean larval development occurs (Waddy and Aiken, 1991), cannibalism within a narrow range of environmental (Hecht and Pienaar, 1993) and swimming parameters (Sastry, 1983). Among the activity and hence metabolism (Gardner, abiotic characteristics that influence the 1996). developmental period and survival rate of The effects of photoperiod on the larvae, pH and photoperiod are of great larval survival have been studied in several importance. High and low levels of pH are such as the common known to adversely affect the physiology crangon (Dalley, 1980), red king of the due to the increased Paralithodes camtschaticus amounts of ammonia and nitrite (Oanh et (Nakanishi, 1987), red frog crab Ranina al., 2000). High values and sudden ranina (Minagawa, 1994), spiny rock fluctuations in pH are found to cause edwardsii (Bermudes and Downloaded from jifro.ir at 15:49 +0330 on Thursday February 15th 2018 disease outbreaks (Cheng and Chen, 1998) Ritar, 2008) and Japanese and stress (Oanh et al., 2000), respectively japonicus (Matsuda et al., 2012). in larvae. A pH around 8.5 was Changes in survival rate and found to be the optimum for larval rearing developmental period of decapod larvae in of the mud crab paramamosain different lighting regimes are often simply (Nghia et al., 2007). Another important attributed to effects on feeding. In the giant parameter that decides the survival and crab Pseudocarcinus gigas, it was developmental period of crustacean larvae observed that continuous light or dark is the period of exposure to light or regimes should be avoided for better darkness. Other aspects of behaviour and overall larval survival (Gardner, 2001). In physiology that are influenced by lighting many decapod species, larvae are less includes entrainment of physiological severely affected by ‘0’ hour light and are processes to circadian cycles (Dalley, able to capture prey at a reduced rate as in 1980), endocrine control of metamorphosis R. ranina (Minagawa, 1994) or they may (Eagles et al., 1986), initiation of ecdysis actually feed at a rate equal to or greater 491 Ravi and Manisseri, Photoperiod and pH variations in Portunus pelagicus larval rearing... ______than in light as in the commercial basis. Knowledge on the Homarus americanus (Eagles et al., 1986) influence of abiotic characteristics like pH and P. camtschaticus (Nakanishi, 1987). A and photoperiod on the rate of survival and 24 hour light period caused a relatively developmental period of the larvae of P. rapid development of larvae in P. gigas pelagicus is of extreme significance in (Gardner and Maguire, 1998) and P. standardising the hatchery technology and camtschaticus (Nakanishi, 1987). developing a conservation management Similarly, longer light hours promoted strategy for the species. The studies on the smooth and regular metamorphosis, effect of these parameters on the larvae of against delayed or suppressed P. pelagicus from Indian waters are metamorphosis from phyllosoma stage to meagre. The present study attempts to puerulus stage caused by longer dark hours trace out the effects of pH and photoperiod in P. japonicus (Matsuda et al., 2012). on the survival rate and developmental Though there are a number of studies on period of the larvae of P. pelagicus from the effect of light on the activity, the south-east coast of India. The study swimming or phototaxis of crab larvae (for also attempts to find out the optimal range example, Sulkin (1975); Gardner and of these parameters for each larval stage. Maguire(1998), those on the effect of pH Ten female P. pelagicus of size and photoperiod on crab larval range 140 - 160 mm carapace width with development period and survival are light yellow berry were collected from the relatively less (Minagawa, 1994; Gardner Gulf of Mannar off Mandapam (9°09' N, and Maguire, 1998). The marine blue 78°44' E). The animals were brought live swimmer crab Portunus pelagicus is one to the wet laboratory, given a prophylactic of the candidate species for culture dip in 200 ppm formalin for 30 minutes because of its fast growth, attractive (Parado-Estepa et al., 2002) and stocked in

Downloaded from jifro.ir at 15:49 +0330 on Thursday February 15th 2018 appearance and taste. The species is 1000 l FRP (Fibreglass Reinforced Plastic) available throughout the coast of India tanks with black interior for especially in the south-east and south-west acclimatization (2 days). Three of the most region and breeds round the year (Pillai active animals were weighed (to the and Nair, 1973). Since commercial nearest 1 g) and stocked separately in 500 l hatcheries are not available for the species FRP tanks for the study. The animals were (Soundarapandian and Tamizhazhagan, fed to satiation with raw clam (Meritrix 2009), the farmers are forced to depend on meritrix) and cuttlefish (Sepia pharaonis) the wild for juveniles for culture practices. meat (1:1). Excess feed, faeces, discarded Difficulty in obtaining juveniles from wild, eggs and other debris were siphoned out their non-uniform size and concerns of from the tank daily, accompanied by 10% stock depletion due to overexploitation water exchange. The temperature, salinity, have encouraged research activities to pH and photoperiod were maintained at 28 frame out a cost effective and foolproof ± 1 °C, 35 ‰, 8.1 ± 0.1 and 12hL:12hD, technology for producing crab seeds on a respectively. Dissolved oxygen level was Iranian Journal of Fisheries Sciences 12(2) 2013 492 ______maintained above 5 mg/l throughout the Shinkarenko, 1979) were calculated by experiment. A sandy substratum (100 mm daily random sampling ( 10 samples were thickness) was provided in all the tanks for taken each time, twice daily, till the end of refuge and to prevent egg loss (Davis et al., the experiment) (mortality due to any 2004). The pH was adjusted by adding 1 reason other than pH or photoperiod normal solution (N) sodium carbonate or variation is assumed to be the same in all 1N hydrochloric acid when required. the replicates of the values tested under Tanks were covered with black sheets (200 each parameter). At first crab instar (C1), µm thickness) after the photophase. they were harvested using 500 μm mesh Photoperiod adjustments were carried out nets and counted to estimate the final using fluorescent lamps and the light survival rate. The periods of development intensity maintained was 750 lux. The of the larvae (from Z1 to C1) for each berry colour was examined daily and at replicate under the temperature and dark grey colour stage they were salinity treatments were also recorded. transferred to cylindro-conical FRP tanks Zoeae 1 and 2 were fed twice daily of 500 l capacity with white interior for with Skeletonema costatum and hatching. Brachionus plicatilis (cell densities Once the eggs were hatched, the maintained at 50,000 no ml-1 and 25 no zoea 1 (identified as per Shinkarenko, ml-1, respectively). The larvae were also 1979) were collected using 150 μm mesh fed with formulated feed (Frippak 2 sieve and the most active ones (identified CD - INVE, Belgium) at a rate of 0.5 g by their vigorous swimming activity in 1000 l-1 (4 times daily). Zoeae 3 and 4 opposite direction in a whirl), from the were also fed with S. costatum, B. plicatilis same brood were stocked after rinsing in and formulated prawn feed (at cell fresh seawater in 200 l rectangular FRP densities 50,000 cells ml-1, 30 no ml-1 and tanks at a density of 25 no l-1 (i.e. 5000 at the rate 0.5 g 1000 l-1, respectively). The Downloaded from jifro.ir at 15:49 +0330 on Thursday February 15th 2018 larvae per tank, as this stocking density frequency of feeding was maintained the gave better results in earlier trials). Dead same as in the case of Z1 and Z2. larvae, feed waste and other debris were Megalopae were fed daily once with siphoned out prior to the daily water Moina macrura and newly hatched exchange (70%). Abiotic parameters were Artemia nauplii (cell densities maintained monitored regularly and continuous were 3 no ml-1 and 5 no ml-1, respectively). aeration was provided to maintain the They were also fed twice daily with dissolved oxygen level above 5mg/l in all freshly prepared egg custard at the rate of the tanks throughout the experiment. The 0.5 g 1000 l-1. Rotifers and M. macrura experiments were carried out in triplicate were collected from the wild and mass (n = 3) with controls (3 tanks were stocked cultured in 1000 l FRP tanks on initially). The survival rates at each of the microalgae. Microalgal (Chlorella sp. and five larval stages such as zoea 1 (Z1), zoea S. costatum) inoculums were obtained 2 (Z2), zoea 3 (Z3), zoea 4 (Z4) and from the live feed laboratory of the Central megalopa (M) (identified as per Marine Fisheries Research Institute, 493 Ravi and Manisseri, Photoperiod and pH variations in Portunus pelagicus larval rearing... ______Mandapam and scaled up in 1000 l FRP The survival rates (stage wise and overall) tanks. and the overall larval developmental Two parameters viz. pH and periods of each replicate under different photoperiod were investigated separately. pH and photoperiod treatments were One set of experiment investigated the statistically analyzed with one-way effect of three different pH values and ANOVA. The data with significant another set investigated the effect of three difference (P<0.01) were then subjected to photoperiod regimens. The pH values Post Hoc Test of multiple comparison selected for the study were 7.5 ± 0.1 using ‘Fisher's Least Significant (T1/treatment 1), 8.1 ± 0.1 (T2/treatment Difference (LSD)’ (Payne et al., 1993). 2/control) and 8.5 ± 0.1 (T3/treatment 3). Among results, the overall survival rates For increasing or decreasing pH, 1N were expressed as a percentage of the total sodium hydroxide or 1N hydrochloric acid larvae stocked and the larval was added, when required. The developmental period in number of days. temperature, salinity and photoperiod Complete larval development took regimen maintained were 28.0 ± 0.1 ºC, place in all the replicates under the 35 ‰ and 12hL:12hD, respectively. parameters studied. The survival rate fell Temperatures were maintained by keeping almost gradually with time and stage of the the tanks in chilled room and then larvae in all the pH and photoperiod increasing the water temperature to the treatments studied. The differences in the desired levels using aquarium heaters with average overall survival rates and thermostats (AZOO, USA, 100W). developmental period among different pH The photoperiod regimens chosen treatments investigated were not for the experiment were 6hL:18hD (T1), significant (P>0.01). The overall survival 12hL:12hD (T2/control) and 18hL:6hD rate at pH 7.5 ± 0.1, 8.0 ± 0.1 and 8.5 ± 0.1

Downloaded from jifro.ir at 15:49 +0330 on Thursday February 15th 2018 (T3). Tanks were covered with black were 6.28  0.12%, 6.76  00.91% and sheets (200 µm thickness) after the 5.59  0.53%, respectively. When photophase and the light intensity was compared with the larval survival rates at maintained at 750 lux (Gardner and the control pH 8.0 ± 0.1, the survival rates Maguire, 1998; Hoang et al., 2002). The at other pH values such as 7.5 ± 0.1 and temperature, salinity and pH maintained 8.5 ± 0.1, showed significant variance were 28.0 ± 0.1ºC, 35 ‰ and 8.1 ± 0.1, (P<0.01) at some of the larval stages respectively. Observations on feeding, (Table 1, Fig. 1). The average larval growth, cannibalism, activity and survival developmental period was 16 days. were recorded every morning and evening.

Iranian Journal of Fisheries Sciences 12(2) 2013 494 ______Table 1: Preferred pH and photoperiod values at each larval stage of Portunus pelagicus and the corresponding survival rates

Larval Preferred Corresponding Preferred Corresponding stages pH survival rates (%) photoperiods survival rates (%)

(±SD) (hL/hD) (±SD) Z1 7.5 ± 0.1 73.06 ± 02.57 12:12 73.87 ± 01.67

Z2 8.0 ± 0.1 84.81  04.22 06:18 84.51 ± 02.95

Z3 8.5 ± 0.1 65.60  04.68 18:06 85.10 ± 18.37

Z4 8.0 ± 0.1 42.47  09.29 18:06 55.03 ± 11.52

M 7.5 ± 0.1 43.70  07.61 12:12 39.04 ± 05.17

Downloaded from jifro.ir at 15:49 +0330 on Thursday February 15th 2018 Figure 1: The survival rates of P. pelagicus larvae at each stage of development under different pH (Z1- zoea 1, Z2- zoea 2, Z3-zoea 3, Z4-zoea 4, M- megalopa) (*P<0.1)

The differences in the average overall photoperiod regimen 12hL:12hD, the survival rates and developmental period survival rates at other photoperiod among the photoperiod treatments regimens such as 18hL:06hD and investigated were not significant (P>0.01). 06hL:18hD, showed significant variance The overall survival rate at photoperiods (P<0.01) at some of the larval stages 18hL:06hD, 12hL:12hD and 06hL:18hD (Table 1, Fig. 2). The average larval were 5.55 ± 0.65%, 7.74 ± 0.37% and 6.63 developmental period was 16 days. ± 0.97%, respectively. When compared with the larval survival rates at the control

495 Ravi and Manisseri, Photoperiod and pH variations in Portunus pelagicus larval rearing... ______

Figure 2: The survival rates of P. pelagicus larvae at each stage of development under different photoperiods (Z1-zoea 1, Z2- zoea 2, Z3-zoea 3, Z4-zoea 4, M-megalopa) (*P<0.1)

Rearing techniques, disease and nutrition are Low values of pH are considered to be the three main areas of research that have harmful for crustacean larvae. A pH below 5 supported commercial production of marine has been found to cause decreased activity, fish and crustacean larvae (Sorgeloos and gill damage, moulting difficulty and death in Leger, 1992). One of the serious Macrobrachium rosenbergii larvae (New consequences of ambient pH variation is the and Singholka, 1985). Such harmful effects change in the ammonia level, which is the were not observed in any of the replicates principal excretory product in under the pH treatments followed here. (Kinne, 1976). In the present study, the pH Sudden fluctuations in pH and rearing in values investigated the influence of larval low pH levels have lead to mass mortality survival rate in an almost similar manner. due to stress in the larvae of blue crab However, since the highest average survival (Sandoz and Rogers, Downloaded from jifro.ir at 15:49 +0330 on Thursday February 15th 2018 was obtained at 8.0 ± 0.1, that pH was 1944) and M. rosenbergii (Oanh et al., 2000). considered as the most suitable for larval However, even the lowest pH tested (7.5 ± rearing in the present study. A similar range 0.1) did not cause any significant adverse of pH was found to be the optimum for effect in the present study. larval rearing in the crab Ucides Intensity variations, spectral cordatus, (Ventura et al., 2008) and white composition, polarization and photophase leg shrimp Litopenaeus vannamei (Van-Tri duration are some of the characteristics of and Hoa, 2004). Furthermore, commercial light that influence the physiological seed production of P. pelagicus was attained activities and behaviour of animals (Dalley, with an overall survival rate of 4.3% at a pH 1980). Among these, the daily cycle of light range of 7.5 - 8.0 (Soundarapandian et al., and darkness, and the seasonal changes in 2007). The survival rates under a particular the proportions of light-dark period are of pH varied significantly with stage. This more significance in the life of crustaceans indicates that the influence of a particular (Dalley, 1980). In the present study, the pH varies considerably from stage to stage. different photoperiod regimens investigated Iranian Journal of Fisheries Sciences 12(2) 2013 496 ______had a statistically insignificant influence on somewhat parallel in all the replicates the survival rate and developmental period studied. Decapod larvae use methods other of P. pelagicus larvae. Concurrent than sight to spot prey (Kumlu, 1999), insignificant influence was shown also in the probably this is the reason why the larval first three states of phyllosoma larvae in J. growth was not adversely affected by more edwardsii when reared under photoperiod dark hours in the 6hL:18hD regimen. In P. regimes with 0, 6, 12, 18 or 24 h light argentinus, long light hours increases (Bermudes and Ritar, 2008). Likewise, the moulting rate subsequently resulting in survival rates of juvenile F. merguiensis did lower biomass incorporation due to the high not show significant difference (P>0.01) expenditure of energy for moulting (Diaz et when reared under photoperiod regimens al., 2003). However, marked or abrupt 12hL:12hD and 10hL:14hD (Hoang et al., differences in growth were not observed in 2003). the present study. However, since 12hL:12hD showed Cannibalism is another characteristic the highest survival rate and lowest that is known to be influenced directly by developmental period, that regimen is the light. A significant increase in cannibalism, most favourable photoperiod for P. proportional to the length of light hours was pelagicus larval rearing here. Similar reported in the zoeae P. gigas (Gardner and photoperiod regimens were reported to be Maguire, 1998). The authors also opined optimum for larval rearing in the tiger that in dim light, P. gigas larvae might have shrimp (Khalik et al., difficulty in sighting small feed like Artemia 1992), mud crab (Quinitio et sp. and therefore would switch over to al., 2001) and palaemonid shrimp younger zoeae, which was displayed by an Palaemonetes argentinus (Diaz et al., 2003). increase in cannibalism. In M. rosenbergii, The significant stage-specific preference for the larvae aggregated in the brighter parts of photoperiod observed here shows that the the tank were more vulnerable to Downloaded from jifro.ir at 15:49 +0330 on Thursday February 15th 2018 most ideal photoperiod values, where the cannibalism (Kutty et al., 2000). In the survival rate is expected to be the highest present study, the larvae were provided with and the moulting rate optimum, do change enough feed in terms of both variety and with larval stage. quantity. In addition, the larvae might have Zoeal stages of P. pelagicus in the used some method other than sight for present study showed phototaxis as spotting prey during dim light conditions. So generally shown by zoeae of many other cannibalism for the purpose of feeding is (e.g. Sulkin et al., 1979; Gardner, doubtful. However, an enhanced rate in 1996), but an apparent movement towards cannibalism was indeed noticed in the light at megalopae stage was absent as brighter regions of the tank where the larvae against the reports in the portunid crab aggregated. Genetic factor appears to be the Macropipus sp. and common shore crab reason for this tendency. Carcinus maenas (Rice, 1966). In the present study, feeding was found to be normal and consequently the growth was 497 Ravi and Manisseri, Photoperiod and pH variations in Portunus pelagicus larval rearing... ______Acknowledgements Iheringia, Série Zooologia, 93(3), 249- We express our sincere thanks to Dr. G. 254. Syda Rao, Director, Central Marine Eagles, M. D., Aiken, D. E. and Waddy, S. Fisheries Research Institute (CMFRI), L., 1986. Influence of light and food on Cochin, for his guidance and encouragement. larval American Homarus Acknowledgement is also due to Dr. G. americanus. 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