Settlement and Growth of the Green Mussel Perna Viridis (L.) in Coastal Waters: Influence of Water Velocity
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Aquatic Ecology 32: 313–322, 1998. 313 © 1998 Kluwer Academic Publishers. Printed in the Netherlands. Settlement and growth of the green mussel Perna viridis (L.) in coastal waters: influence of water velocity S. Rajagopal1, V. P. Venugopalan2,K.V.K.Nair2, G. Van der Velde1 andH.A.Jenner3 1Department of Ecology, Laboratory of Aquatic Ecology, University of Nijmegen, Toernooiveld, 6525 ED Nijmegen, The Netherlands (E-mail [email protected]); 2Marine Biology Programme, Water and Steam Chem- istry Laboratory, BARC Facilities, Kalpakkam 603 102, India; 3KEMA Power Generation, P.O. Box 9035, 6800 ET Arnhem, The Netherlands Accepted 20 November 1998 Key words: Perna viridis, flow velocity, growth rate, larval occurrence, spat settlement, spawning periods Abstract Green mussels Perna viridis were observed to be a major foulant in the seawater intake tunnel of a coastal power station. Field experiments were carried out to ascertain what factors were responsible for the successful colonisation by mussels. Two adjacent stations (25 m apart) were selected, one representing the coastal waters and the other representing the intake screens (with higher water velocity). Gonadal activity, larval abundance, spat settlement and growth rate of the mussels were monitored at monthly intervals for a total period of two years. The results showed that the breeding activity of the mussels at the study area is influenced largely by temporal distribution of seawater temperature. However, ensuing larval availability in the coastal waters is more dependent on food availability. On the other hand, spat settlement and growth rate are predominantly influenced by water flow, probably as a result of increased propagule and food flux rate at higher water velocities. Higher water velocity at the intake screens also contributed to mussel dominance by preventing settlement of many potential competitors. Introduction ing seawater (Rajagopal et al., 1997). On an earlier appraisal (Rajagopal et al., 1991), it was found that Green mussels, Perna viridis are widely distributed out of 570 tons of fouling debris lodged inside the in the Indo-Pacific region; their distribution extends concrete intake tunnel of a power station, P. viridis from Japan to New Guinea and from the Persian Gulf alone constituted 411 tons. Since this was the first time to South Pacific Islands (Siddall, 1980). They are such massive colonisation of marine mussels has been a characteristic species of midlittoral and sublittoral observed in the cooling circuits of an Indian power zones where they often constitute dense populations station, we were interested to know what aspects of on rocky substrata. In spite of their wide distribu- the ecology of the mussels make them such successful tion and their importance in the ecology of rocky colonisers. This was important as earlier workers who shore ecosystems, detailed works on their biology are studied fouling phenomena on the east coast had not only a few (Lee, 1985). These mussels are also im- indicated the dominance of P. viridis among the nat- portant from the point of view of animal protein for ural sessile communities (Paul, 1942; Daniel, 1954; human consumption and some aspects of their biol- Renganathan et al., 1982; Rao, 1990). Moreover, in- ogy relevant to fishery and culture have, therefore, formation regarding the breeding activity of P. viridis been studied by other workers (Qasim et al., 1977; from different localities of the Indian peninsula was Sivalingam, 1977; Parulekar et al., 1982; Rivonkar inconsistent. Larval availability and growth rate are et al., 1993; Rajagopal et al., 1998). However, these important parameters influencing successful colonisa- mussels also deserve serious attention on account of tion by sessile species. Earlier workers had indicated their potential to foul industrial cooling systems us- the importance of water flow on the population ecol- 314 Figure 1. (a) Map showing the Kalpakkam. (b) Schematic represen- tation of the Madras Atomic Power Station seawater intake tunnel showing 2 sampling stations (not drawn to scale). ogy of mussels (Nixon et al., 1971; Perkins, 1974; Venugopalan et al., 1991; Wildish & Kristmanson, Figure 2. Seasonal variations in the hydrographic parameters ((a) 1997). It was possible that greater water flow ex- temperature, (b) salinity, (c) dissolved oxygen and (d) chloro- perienced within the cooling water circuit could be phyll-a) in Kalpakkam coastal waters from April 1988 to March responsible for the successful colonisation of mus- 1990. Data are presented as mean SD. sels. Bearing this in mind, the population ecology of P. viridis was monitored at two locations which repre- Materials and methods sented (a) their natural habitat (coastal waters) and (b) cooling intake point of the power station experienc- Site description ing high water velocity. The present study investigates whether flow regimes are significant in influencing the Kalpakkam is situated (12◦320 N and 80◦110 E) about population ecology viz., growth rate, breeding activity 65 km south of Madras (Figure 1a). Madras Atomic and spat settlement of P. viridis in coastal waters of Power Station (MAPS), Kalpakkam is a seawater Kalpakkam, east coast of India. cooled station, and uses a 468 m long sub-seabed tun- nel to draw cooling water (35 m3 s−1) for its twin (2 × 235 MWe) reactors (for details refer Rajagopal, 315 1997). The seawater flows by gravity from the intake Larval abundance (Figure 1b) via the tunnel to the forebay pump house, from where it is pumped (12 pumps) to the condensers. Mussel larvae were concentrated from 500 l of sea- The coolant seawater flow in the tunnel when all the water using a 22 µm mesh net (De Wolf, 1973), 12 pumps are running, works out to be about 3 m s−1 every month from May 1988 to May 1990. The larvae (Madras Atomic Power Station Design Manual, 1975). were subsequently fixed in 5% buffered formalin and The intake point is guarded by steel weld mesh screens counted in a Sedgwick rafter counter. to prevent the entry of large objects into the cooling circuit. Spat settlement × × Sampling stations Concrete blocks (20 20 20 cm) were used to sam- ple spat fall in coastal waters (Sta 1), as described by Two stations were selected for the study (Figure 1b). Nair et al. (1988) and Rajagopal et al. (1997). Three Station 1 represents the coastal waters which is 8 m test blocks were suspended at 1 m, 4 m and 7 m using deep and experiences coastal currents of velocity in nylon ropes and retrieved after 30 d to estimate spat the order of 0.2–0.3 m s−1. Station 2 is the seawater fall. At Sta 2, spat samples were collected from the intake point which is characterised by high water ve- steel intake screens at 2 m, 4 m and 6 m. Earlier tri- locity (as high as 3 m s−1, depending on the flow). als had shown that mussel settlement on steel surfaces The distance between Sta 1 and Sta 2 is about 25 m. were comparable to those on concrete. The samples The physicochemical characteristics of the water are, (in triplicate) were collected at each depth, and the −2 therefore, identical at both stations, except for water data were averaged and presented as numbers dm 2 2 velocity. (dm D 100 cm ). Hydrographical features Growth rate Hydrographical features of the study site were studied Growth rate measurements were initiated by suspend- by collecting surface water samples at fortnightly in- ing test blocks at 1 m (Sta 1) at the beginning of spat tervals during the period April 1988 to March 1990. settlement (April 1988). Every month about 30 mus- Parameters like temperature, salinity, dissolved oxy- sels were randomly collected from the concrete blocks gen (DO) and chlorophyll-a were monitored (Strick- (1 m depth at Sta 1) and from the intake screens (2 m land & Parsons, 1972) to understand their variation depth at Sta 2). Collection of mussels from test blocks and possible influence on the breeding pattern and involved sampling of a different subset of the mus- growth rate of the mussels. sel population every month. In order to monitor the growth increment of the same population over a period Gonad observations of time, mussels (11 0.6 mm shell length, n D ca. 100) were confined in cages (75 × 75 × 75 cm, 0.5 cm Mussels were collected every month (April 1988 to mesh size) and left suspended at 1 m depth at Sta 1 March 1990) from the two stations and were used for 375 d (Page & Hubbard, 1987). Their shell growth for gonadal studies. About 60–65 mussels (approx. increment was monitored at monthly intervals during 30–40 mm shell length) were collected from each the period October 1988 to September 1989. station. In the laboratory, the gonadal tissues were removed from the mantle lobes and fixed in Bouin’s Statistical analysis fluid for 24 h and later transferred to 40% alcohol. Sections (10–15 µm) were made from wax-embedded A 2-factor analysis of variance (ANOVA) was used to tissues and stained with haematoxylin and eosin. The examine variability in gonad index and spat settlement method of Seed (1969) was adopted to categorise of P. viridis taking account of season (sampling time) the gonads into four groups viz., spent/resting, de- and station as two independent variables (Sokal & veloping/redeveloping, ripe and spawning. Gonad in- Rohlf, 1981). Spat settlement of P. viridis at different dex (GI) was also determined based on the method depths was tested by 1-factor ANOVA. For post-hoc described by King et al. (1989). comparison of monthly means, we used student t-tests for comparison of two means and Student-Neuman- Keuls (SNK) tests for comparison of multiple means 316 (Zar, 1984). Prior to the analysis the data were tested with spawning and GI data, with two peaks in a year, for normality and homogeneity of variance.