Effect of freezing on photosynthesis of intertidal macroalgae: relative tolerance of Chondrus crispus and Mastocarpus stellatus (Rhodophyta)

The effect of freezing on photosynthetic metabolism was studied in the red algae, Chondrus crispus and Mastocarpus stellatus. Plants of both species were collected from the intertidal at Chamberlain or Kresge Point, Maine, USA (43°56N, 69°54W) between February and March 1987. Photosynthetic rates were measured immediately after freezing at-20°C and following recovery periods in seawater. Photosynthesis in C. crispus declined rapidly following freezing, falling to 70% of control values within 1 h and 30% after 3 h exposure. Minimum photosynthetic rates (7 to 9% of controls) occurred following freezing exposures of 12 h or more. Full photosynthetic recovery in C. crispus after 3 h at-20°C required 48 h. Photosynthesis in C. crispus did not fully recover in plants frozen for 6 h or more. In contrast, photosynthesis in M. stellatus was relatively unaffected by freezing exposures of <12 h. Twelve hours or more at-20°C reduced photosynthesis to 55% of controls. Photosynthesis in M. stellatus fully recovered from 24 h at-20°C within 24 h. In both species the reduction of photosynthesis by freezing was associated with damage to the plasma membrane and reduced efficiency of energy transfer from phycobilisomes to chlorophyll a, but did not appear to involve ribulose-1,5-bisphosphate carboxylase oxygenase activity. The freezing tolerance of C. crispus and M. stellatus positively correlates with their respective intertidal distributions, suggesting that freezing may be involved in controlling the distributions of these species on the shore.     

Tolerance to freezing and supercooling of interstitial turbellaria and polychaeta from a sandy tidal beach of the island of Sylt (North Sea)

Tolerance experiments on freezing and supercooling (without ice formation) were designed to determine correspondence between tolerance to low temperatures and spatio-temporal distribution of one turbellarian and five polychaete species from sandy beaches of the North Sea island of Sylt. Freezing tolerances are always less than supercooling tolerances. Dinophilus gyrociliatus is significantly more sensitive to freezing (LD₅₀ value after 30 min of freezing:-3°C) than the others, whereas Stygocapitella subterranea is significantly less sensitive (50% mortality at-15.7°C after 30 min). The supercooling tolerances differ considerably among the species. The sequence of tolerances (LD₅₀ values) is as follows: Microphthalmus sczelkowii (-2.9°C after 4 h); D. gyrociliatus (<1 h at-8°C); M. listensis (5.6 h at-8°C); Protodriloides symbioticus (8.2 h at-8°C); Notocaryoplanella glandulosa (66 h at-8°C); S. subterranea (72 h at-8°C). Species of sand flats (d. gyrociliatus, M. listensis, P. symbioticus) have lower tolerances than those of the beach slope (N. glandulosa, S. subterranea). Among the latter, tolerances increase with distance of the distributional area from low tide level. S. subterranea, a species occurring at the uppermost position in the intertidal, proves to be best adapted to both freezing and supercooling. Species preferring deeper regions of the beach (M. sczelkowii) show lower supercooling tolerances than surface dwelling forms. Northern species usually have higher tolerances to cold than southern ones, reaching their distribution limits near the island of Sylt.     

Effect of environmental factors on byssal thread formation

The effects of various factors on byssal thread formation have been examined in the laboratory using Modiolus demissus and Mytilus edulis. With M. demissus thread formation (threads/mussel/h0 and proportion of mussels forming threads decreased with increasing size; prior exposure to air enhanced subsequent thread formation; mechanical agitation reduced thread formation. In addition, low salinity acclimated mussels adapted more rapidly to 32 ppt than high salinity acclimated mussels did to 16 ppt; threads were not formed in the absence of calcium and/or magnesium; and there was no reduction in thread formation at temperatures as high as 27° to 28°C. M. edulis died and thread formation approached zero at temperatures exceeding 26°C.     

Effects to perfluorooctane sulfonate (PFOS) on the mollusk Unio ravoisieri under laboratory exposure

To understand acute toxicity and oxidative stress of perfluorinated compounds in the freshwater ecosystems, we exposed freshwater mussels (Unio ravoisieri) to perfluorooctane sulfonate (PFOS), over a range of concentrations from 10 to 100?mg/L, in a laboratory experiment. Lethal concentration (LC50) was of about 65.9?mg/L after 96?h of exposure. The oxidative stress was assessed in gill and digestive gland of the freshwater mussels after 7 days of exposure to different nominal PFOS concentrations (C1=?2?mg/L, C2?=?6?mg/L and C3=?10?mg/L). C1 and C2 increased significantly (p?<?.05) the superoxide dismutase activity in both tissues compared, while the highest C3 decreased the enzyme activity. This implements an unfavourable response that highlights the excess of reactive oxygen species produced after contamination. The Catalase activity was also increased by about 40.05% and 66.63%, respectively, in gill and digestive gland after exposure to C3. The Malondialdehyde (MDA) level was increased in both gill and digestive gland in a concentration-dependent pattern. In contrast, the contamination of U. ravoisieri by PFOS did not affect the acetylcholinesterase activity in both organs (p?>?.005). These results provided information on potential biomarkers that could be effectively applied for the monitoring of freshwater ecosystem using indicator species such as U. ravoisieri.     

Oil-induced disruption of foraging behaviour of the asteroid keystone predator, Coscinasterias muricata (Echinodermata)

A prey-localization test allowed the observation of the foraging behaviour of Coscinasterias muricata in the presence of Mytilus edulis and the disruption of this behaviour following oil exposure. Asteroids were exposed to dilutions of water-accommodated fractions (WAF) of Bass Strait stabilised crude oil (control, 2%, 10%). Effects of exposure (4 d) and depuration (20 d) were quantified using circular statistical analyses. Observations suggested that disruption of behaviour might be concentration-dependent. Control asteroids successfully located mussels during tests. A small proportion (3 of 16) of asteroids exposed to 2% WAF also located the mussels, while asteroids exposed to 10% WAF did not. Following depuration, exposed asteroids recovered their chemoreception capacities. It is concluded that (1) Coscinasterias muricata is able to locate prey mussels through chemoreception, (2) exposure to oil disrupts its foraging behaviour, and (3) the effect is reversible. Received: 4 March 1998 / Accepted: 25 November 1998     

Relationships between effects of environmental temperature and seston on growth and mortality of Mytilus edulis in a temperate northern estuary

The growth and mortality of experimentally-rafted Mytilus edulis L. of known age at 7 locations in a northern estuary (Damariscotta River, Maine, USA) were related to environmental temperatures and to the presence or absence of various potential food sources. All particles were regarded as potential food substrates. Growth decreased appreciably at sites where water temperatures exceeded 20°C, but mussels survived a wide range of elevated temperature exposures, ranging from 0 to 149 degree-days in excess of 20°C. The maximum temperature was 25°C. Mortalities of mussels at all sites but did affect the extent of mortality, which increased abruptly in late summer, when water temperatures were declining. Differences in degree-days of exposure to elevated temperatures did not influence the timing of mortality, which occurred synchronously at all times, but did affect the extent of mortality, which ranged from 35 to 90%. The period of high mortality was preceded by a rapid decline in phytoplankton standing crop. Total particle concentrations decreased during this period, but the shift toward larger particles suggests that there was little, if any, decrease in total volume of material in the seston, at least through August. It is suggested that living phytoplankton provided the critical energy source for these mussels, and that the mortalities were caused by rapidly reduced ration at a time of metabolic stress. The role of temperature and the possible role of the gametogenic cycle are discussed. In addition, the feeding efficiency of M. edulis may have decreased during this period due to an increase in mean particle diameter. Chlorophyll was divided into a nannoplankton fraction (<-20 m) and a larger fraction (>20 m). A smaller size criterion for this distinction is proposed for future studies. The importance of nannoplankton to the primary production of this estuary and the role of nannoplankton and nannoplankton-sized particles in the diet of mussels in nature are discussed.     

Niche overlap in mud snails (hydrobiidae): freezing tolerance

Tolerance to freezing was measured in four species of hydrobiid snails exposed to temperatures from 0° to-8°C and salinities from 0 to 30 S four up to 7 d. Experiments showed the following increasing order of tolerance to freezing: Potamopyrgus jenkinsi (Smith), Hydrobia neglecta Muus, H. ulvae (Pennant) and H. ventrosa (Montagu). Survival decreased with increasing salinity in P. jenkinsi, while the opposite was found in Hydrobia species. H. ulvae survived only slightly better than H. neglecta, while H. ventrosa was very tolerant to freezing even at low salinities in contrast to the other species of Hydrobia. H. neglecta suffered higher winter mortality than H. ventrosa during the severe winter of 1985 in an estuary.     

Biological effects of oil on early development of the Baltic herring Clupea harengus membras

The effects of petroleum hydrocarbons from two crude oils and one fuel oil (No. 1) were studied on the ontogenic development of the Baltic herring Clupea harengus membras L. Herring eggs exposed to water-soluble fractions of the oils at the time of fertilization showed no decrease in numbers of fertilized eggs compared to eggs exposed 6 or 72 h after fertilization. During embryongenesis, treatment with 3.1 to 8.9 ppm or 3.3 to 11.9 ppm total oil hydrocarbons from light fuel oil and the two crude oils respectively, gave rise to alterations in embryonic activity, decreased heart rate, and premature or delayed hatching. Although many larvae hatched from eggs exposed to contaminated water (3.1 to 11.9 ppm total oil hydrocarbons), the majority of the (70 to 100%) were malformed or dead 1 day after hatching. Exposure of eggs to 5.4–5.8 ppm total oil hydrocarbons resulted in significantly (P<0.001) decreased lengths of the larvae. Increased temperature (from 9° to 14°C) aggravated the effects of the oils. The results are discussed in relation to the potential effects of oil spills and chronic oil pollution on fish eggs and larvae in the Baltic Sea.     

Photosynthetic response to temperature and desiccation of the intertidal alga Mastocarpus papillatus

When exposed to air during low tide, intertidal macroalgae experience a terrestrial environment and often encounter extreme levels of heating and desiccation. Two aspects of photosynthesis may be influenced by this increase in temperature and decrease in water content during exposure to air: (1) the rate of aerial photosynthesis itself, and (2) the rate at which aquatic photosynthesis recovers upon immersion in water at high tide. This laboratory study examines the effect of air temperature and desiccation on photosynthesis of the intertidal macroalga Mastocarpus papillatus Kützing. Plants were collected at Hopkins Marine Station, California, USA (36°37N; 121°54W) between July and December 1990. When apical tips were exposed to 15 to 25°C air for 2 h, photosynthesis was rapidly recovered upon reimmersion in seawater. Recovery was delayed, but complete, when tissue was exposed to 30°C air, but did not occur after exposure to 35°C air. Desiccation did not influence either the rate or the ultimate level of recovery upon reimmersion. Photosynthesis in air generally decreased with increasing desication, with no net photosynthesis occurring below 25% relative water content. Net photosynthesis of hydrated thalli increased with air temperature from 15 to 30°C, then decreased at 35°C. Dark respiration of hydrated thalli increased over the entire temperature range. This study indicates that thallus heating and desiccation during periods of exposure to air can potentially influence the total carbon budget of M. papillatus.     

Effects of various temperature cycles on the larval development of the gastropod molluscCrepidula fornicata

Veligers ofCrepidula fornicata (L.) were reared for 12 days at constant temperatures of 15°, 20°, 25°, 30° and 35°C, and at 5 C° daily cycles of equal periodicity (COEP) over the temperature ranges 15° to 20°C, 20° to 25°C, 25° to 30°C and 30° to 35°C. COEP consisted of equal periods (6 h) of maximum temperature, minimum temperature, and uniformly increasing and decreasing temperature each 24 h period. Survival was high and not influenced by cyclic or constant temperature from 15° to 30°C. At 35°C and COEP 30° to 35°C, all larvae died before Day 6. Shell growth rate increased markedly over the range 15° to 25°C, and growth rates at cyclic temperatures in this range were intermediate between growth rates at the corresponding constant temperatures. Larvae reared at COEP 15° to 20°C and COEP 30° to 35°C had discontinuities in their shells due to inhibition of shell secretion during the adverse part of each temperature cycle. Groups ofc. fornicata veligers were exposed for 2 days to daily temperature cycles of equal and unequal periodicity in the critical 30° to 35°C range. [Cycles of unequal periodicity (COUP) consisted of unequal periods (varying between 3 and 15 h) of maximum and minimum temperature and uniformly increasing and decreasing temperature each 24 h period.] These veligers showed shell growth although their body tissue declined, as indicated by decreasing carbon content per larva. Least shell growth and most body tissue loss occurred in those cycles with the longest exposure to higher temperature. Larvae exposed for arious days to the mildest 30° to 35°C COUP (15 h at 30°C, 3 h increasing temperature, 3 h at 35°C and 3 h decreasing temperature) recovered and resumed normal growth when transferred to constant 30°C, but their growth was retarded in proportion to the number of days in the temperature cycle. Rates of shell growth of veligers in temperature cycles show an immediate effect of environmental temperature, while changes in carbon content per larva better reflect the effects of temperature on general metabolism and survival.     

Desiccation,higher temperatures and upper intertidal limits of three species of sea mussels (Mollusca: Bivalvia) in New Zealand

Distribution, abundance, and resistance adaptations to higher temperature and desiccation of three species of intertidal mussels (Mytilus edulis aoteanus, Perna canaliculus and Aulacomya maoriana) were studied in New Zealand. M. edulis aoteanus generally was more abundant upshore, with P. canaliculus dominating downshore. M. edulis aoteanus was more common than P. canaliculus on the outside of mixed-species clumps. Abundance of A. maoriana was variable, with individuals favouring damp habitats such as inside mussel clumps. In moving air at 75% relative humidity and at 20°C or 30°C, median lethal levels of water loss were similar for P. canaliculus and M. edulis aoteanus but lower for A. maoriana. Rates of desiccation varied inversely with size and were higher for P. canaliculus, due mainly to valve gaping with resultant loss of water from the mantle cavity. M. edulis aoteanus was more tolerant of higher water temperatures than were the other species. Success in colonizing upshore or more aerially exposed habitats seems to be related to ability of small mussels to tolerate desiccation, especially during hot, windy weather.     

Adjustment of the components of energy balance in the gastropod Crepidula fornicata in response to thermal acclimation

Synchronous measurements were made of the routine rate of oxygen consumption and the clearance rate of Phaeodactylum tricornutum at different exposure temperatures by specimens of the suspension-feeding gastropod Crepidula fornicata which had been acclimated to temperatures between 10° and 25°C. The results show that the cost of activity (l O₂ consumed h^-1/ml seawater cleared h^-1) increases dramatically in individuals exposed to short-term increases of temperature up to 30°C, especially in limpets acclimated to 10°C. The process of thermal acclimation, however, results in two compensatory adjustments in energy expenditure and uptake which profoundly affect the energetics of water transport. Firstly, the routine oxygen consumption shows lateral translation of the rate-temperature curve which results in the maintenance of a relatively uniform energy expenditure despite an increase in acclimation temperature from 10° to 25°C. Secondly, because of the form of the rate-temperature curve for filtration by C. fornicata, lateral translation in response to warm acclimation results in an increase in the maximal clearance rate. Lateral translation of the rate-temperature curves for feeding rates and for oxygen consumption in response to thermal acclimation may thus be linked to maintain a balance between energy gain and expenditure. In this way, the greatly increased cost of activity which would occur with increase of temperature in the absence of acclimation is evaded. The minimal maintenance energy requirement, and hence the greatest scope for growth and reproduction, is then adjusted to coincide with temperatures prevailing in the environment.     

Freezing rate and duration determine the physiological response of intertidal fucoids to freezing

Differential thermal analysis (DTA) was used to measure the freezing temperature of nine species of red brown intertidal macroalgae from the coast of Maine, USA in 1991. Using slow and rapid cooling rates approximating those found in the field for Ascophyllum nodosum (L.) Le Jol. we found that, for a given rate, the freezing points of all species were similar: -7.06 to -8.02°C for slow cooling (ca. 0.25°C min^-1) and -3.42 to -4.56°C for rapid cooling (ca. 5.0°C min^-1). In the low shore species, Fucus evanescens C. Ag., photosynthesis was inhibited to a greater extent when plants were frozen or thawed rapidly than after slow freezing or thawing. However, in the upper shore species, F. spiralis (L.), photosynthesis recovered rapidly and completely regardless of freezing rate. Rapidly frozen F. evanescens also experienced greater loss of plasmalemmal integrity, evidenced by a greater loss of cellular contents on re-immersion, than those frozen slowly. Light-limited photosynthesis following freezing was more severely inhibited than light-saturated photosynthesis. Respiration was generally enhanced immediately after freezing, but then declined to rates below those of unfrozen controls within 2 h following re-immersion, with control rates of respiration being achieved after a 24 h recovery period. Our data suggest that the physiological consequences of winter emersion at sub-zero temperatures may vary widely between individual plants of freezing-susceptible species, due to the wide variations in freezing rate associated with microhabitat effects.     

Factors affecting methyl and inorganic mercury dynamics in mussels and shrimp

Radiotracer experiments were designed to study the effect of certain factors on the accumulation and loss of methyl and inorganic mercury in mussels (Mytilus galloprovincialis) and benthic shrimp (Lysmata seticaudata). Methyl mercury was accumulated from both food and water to a greater degree than inorganic mercury by both species. There was a tendency for small mussels to concentrate more mercury than larger individuals, but the reason for this difference remains unclear. A trend towards greater mercury accumulation at higher temperatures was noted for mussels, but the temperature effect was relatively small over a 10 C^o range between 8° and 19°C. Methyl mercury residues were eliminated by both species more slowly than those of the inorganic form. Loss from mussels was somewhat more rapid at higher temperatures; however, as in the case of accumulation, the effect of temperature was relatively small. Loss rates for small mussels were not significantly different from those for large individuals. Enhanced methyl mercury elimination was noted for the group of mussels maintained in their natural environment. The more rapid metal turnover in these individuals compared with mussels held in the laboratory was attributed to differences in the availability of natural particulate food matter and, hence, subsequent growth of the animals in the two experimental systems. The observed differences underscore the need for caution in predicting in situ flux of metals such as mercury in certain species based solely on data derived from laboratory experiments.     

Freezing tolerance in the intertidal red algaeChondrus crispus andMastocarpus stellatus: Relative importance of acclimation and adaptation

The effect of repeated daily freezing on photosynthesis, growth and phenotypic acclimation to freezing was studied in the red algaeChondrus crispus Stackhouse andMastocarpus stellatus (Stackhouse in With.) Guiry. Algae used for experiments were collected from Chamberlain, Maine, between March and August 1987, and field observations and experiments were carried out at Chamberlain and Kresge Point, Maine between March 1987 and March 1989. After ca 30 d of daily freezing for 3 h at –5°C photosynthesis ofC. crispus was reduced to 55% of control values. Growth rates ofC. crispus were also reduced in fronds frozen daily compared to unfrozen controls, and eventually fronds became bleached and fragmented resulting in biomass losses. Fronds ofC. crispus, frozen daily, had higher photosynthetic rates following freezing events than unfrozen controls indicating that this species can acclimate to freezing conditions. Acclimation to freezing involves the light-harvesting reactions of photosynthesis. In contrast, photosynthesis and growth inM. stellatus were unaffected by repeated daily freezing for 3 h at –5°C for 36 d. No differences in photosynthesis following freezing were observed between frozen and control fronds suggesting thatM. stellatus does not phenotypically acclimate to freezing. The greater freezing tolerance ofM. stellatus relative toC. crispus results, in part, from genetic adaptations associated with plasma membranes and the light-harvesting reactions of photosynthesis.     

High-temperature tolerance of photosynthesis in the red alga Chondrus crispus

Chondrus crispus (Stackhouse) is a perennial red seaweed, common in intertidal and shallow sublittoral communities throughout the North Atlantic Ocean. In the intertidal zone, C. crispus may experience rapid temperature changes of 10 to 20C° during a single immerison-emerision cycle, and may be exposed to temperatures that exceed the thermal limits for long-term survival. C. crispus collected year-round at Long Cove Point, Chamberlain, Maine, USA, during 1989 and 1990, underwent phenotypic acclimation to growth temperature in the laboratory. This phenotypic acclimation enhanced its ability to withstand brief exposure to extreme temperature. Plants grown at summer seawater temperature (20°C) were able to maintain constant rates of lightsaturated photosynthesis at 30°C for 9 h. In contrast, light-saturated photosynthetic rates of plants grown at winter seawater temperature (5°C) declined rapidly following exposure to 30°C, reached 20 to 25% of initial values within 10 min, and then remained constant at this level for 9 h. The degree of inhibition of photosynthesis at 30°C was also dependent upon light intensity. Inhibition was greatest in plants exposed to 30°C in darkness or high light (600 mol photons m^-2s^-1) than in plants maintained under moderate light levels (70 to 100 mol photons m^-2s^-1). Photosynthesis of 20°C-acclimated plants was inhibited by exposure to 30°C in darkness or high light, but the degree of inhibition was less than that exhibited by 5°C-grown plants. Not only was light-saturated photosynthesis of 20°C plants less severely inhibited by exposure to 30°C than that of 5°C plants, but the former also recovered faster when they were returned to growth conditions. The mechanistic basis of this acclimation to growth temperature is not clear. Our results indicate that there were no differences between 5 and 20°C-grown plants in the thermal stability of respiration, electron transport associated with Photosystems I or II, Rubisco or energy transfer between the phycobilisomes and Photosystem II. Overall, our results suggest that phenotypic acclimation to seawater temperature allows plants to tolerate higher temperatures, and may play an important role in the success of C. crispus in the intertidal environment.     

Influence of high temperature and residual chlorine on marine phytoplankton

Marine phytoplankton forms are frequently exposed to sudden biological changes such as rapid rise in water temperature and chlorine content of their environment, resulting from the use of sea water for cooling purposes by electric generators. The direct influence of these effluents, i.e. inhibitory effects of high temperature and residual chlorine on growth and photosynthesis of Chlamydomonas sp. and Skeletonema costatum, were investigated experimentally. Chlamydomonas sp. and S. costatum exposed to high temperatures were affected in their growth from 43° and 35°C, respectively, by immersion of the respective cultures in a warm bath for 10 min. Treatment at high temperatures of 40 °C and 30° 35°C for 10 min, influenced their photosynthetic activities, which were completely inhibited immediately after 10 min exposure at 42° and 37 °C, respectively. S. costatum was killed by chlorine at a concentration of 1.5 2.3 ppm when exposed for exactly 5 or 10 min, while Chlamydomonas sp. was not irreversibly damaged even at 20 ppm chlorine or more with the same exposure period. These results lead to the conclusion that the high temperature of, and residual chlorine in, effuents from a power plant discharging into the open sea, should not cause great damage to marine phytoplankton in that area.     

The significance of temperature,salinity and zinc as lethal factors for the mussel Mytilus edulis in a polluted estuary

Mussels, Mytilus edulis L., were subjected to high temperatures, low salinities and dissolved zinc in order to investigate possible environmental hazards of a discharge of heated effluent near Newport on the Yarra River estuary, Victoria, Australia. Exposure to zinc at 0.8 mg l^-1 for 14 d in otherwise favourable conditions significantly increased mortality resulting from subsequent exposure to temperatures between 29° to 31°C for 24 h without added zinc. Mussels collected from water of temporarily lowered salinity (8–16 S) showed significantly lower thermal resistance than controls collected from marine salinities (35 S). Mussels taken from a marine environment and exposed to 10 S died at a rate which increased with temperature. Mussels acclimated for 14 d to combinations of 10°, 16° and 22°C and 22 and 35 S, and subsequently exposed to increased zinc concentrations accumulated zinc to levels which were independent of temperature and salinity. The zinc was lethal more quickly at 22°C and 35 S than at the lower temperatures and salinities. The modes of toxic action of the salinity, zinc and temperature factors are discussed and it is argued that zinc which has been found accumulated in mussels near Newport could be reducing their resistance to raised temperatures and perhaps other stresses, probably as a result of effects on lysosomal functioning. The evidence suggests that the heated effluent will accelerate any toxic effects of zinc or low salinities which occur near Newport and so poses a hazard in winter as well as in summer.     

Rearing Pandalus borealis larvae in the laboratory

Larvae of the northern shrimp Pandalus borealis (Krøyer) are pelagic. In the Estuary and Gulf of St. Lawrence, Canada, the early stages are found in the upper 25-m of the water column in spring and early summer and are expected to experience a range of water temperatures from as low as 0°C to as high at 6°C. Little is known of the impact of water temperature on metabolic requirements of northern shrimp larvae. In this study, routine respiration (VO₂), maximum respiration (electron transport system activity, ETSA) and metabolic scope for growth (MS, ETSA–VO₂) of northern shrimp larvae were measured as a function of temperature (3, 5 and 8°C), developmental stage (I–V at 3°C, I–VII at 5°C and 8°C) and growth rate in dry mass. After logarithmic transformation, all three metabolic variables were linearly related to dry mass. The increase in VO₂ with body mass was faster at 5°C than at 3 or 8°C, whereas with ETSA this increase was slower. As a result, MS increased more slowly with dry mass at 5°C than at 3 and 8°C. However, MS did not limit growth in this study, since it explained only 39% of the variability in growth. All three metabolic variables as well as growth varied together as a function of temperature and ontogeny. Q₁₀ of all three metabolic variables ranged from 1.6 and 2.2 for stages I–V larvae, except for VO₂ at stage I (3.9) and stage III (2.9).     

External synchronizers of tidal activity rhythms in the prawns Penaeus indicus and P. monodon

Phasing of persistent circatidal rhythmicity to an artificial tidal cycle was assessed in the prawns Penaeus indicus Milne Edwards and P. monodon (Fabricius) collected from the Vellar estuary, South India, in the period between June and December 1984. Simulated 6 h cycles of 20 and 30 S, and 6 h cycles of 20° and 30°C induced a persistent tidal rhythmicity after 20 cycles. The imposed 6 h cycles of 25 and 30 S, and 25° and 30°C induced tidal rhythms after 30 cycles. In both cases, re-established tidal activity rhythms were evident for at least 48 h — higher activity occurring during the higher salinity and lower temperature phases of the simulated tidal cycles. Artificial tidal cycles of still water and running water synchronized the tidal rhythm after 20 cycles. Combined 30 S, 20°C, for 6 h and 20 S and 30°C for 6 h established a persistent tidal rhythm after 10 cycles, whereas wave action had no influence on tidal synchronization. The influence of possible interactions of tidal rhythms and in situ tidal variables on circatidal activity is discussed.