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.     

A comparative study of metabolic energy expenditure in the limpets Patella cochlear,P. oculus and P. granularis

Oxygen consumption of 3 species of Patella was measured in air and water at various temperatures. Measurements at constant temperature over a full tidal cycle showed no tidal or light-dark rhythms. Measurements under conditions simulating natural tidal, temperature and day-night cycles allowed calculation of daily respiratory energy budgets. P. cochlear occurs low on the shore, but experiences a food shortage due to intense intraspecific competition. Its rate of respiration is moderate, but metabolic expenditure is kept low because exposure to air is brief and body temperatures seldom rise above 23°C. P. cochlear has a respiratory rate-temperature (R-T) curve which peaks at 20°C and forms a plateau between 20° and 32.5°C. The midshore P. oculus has abundant food and adopts an exploitative strategy. Growth rate is very high, and this high turnover of energy is linked with a high metabolic rate, high Q10 (temperature coefficient) values, high body temperatures during the day-time low tide, and a respiratory R-T curve peaking at 32.5°C. Small P. oculus occur mainly in intertidal pools and respire faster in water, while larger individuals occur on bare rocks and respire faster in air over the upper temperature range. In contrast, the upper-shore P. granularis has little food, and conservation of energy is essential, particularly as its growth rate is moderate and its reprocurve output high. Respiratory losses are reduced by suppression of the R-T curve and low Q10 values, resulting in relative independence of temperature. Small P. granularis occur low on the shore and respire slower in water. Larger individuals occur at high levels due to migration, and respire slower in air. This further reduces respiratory energy losses. The patterns of respiration in these 3 species are thus related to food availability, resulting in exploitative or conservationist strategies.     

Seasonal oxygen consumption of the saltmarsh lsopod Sphaeroma rugicauda

Oxygen consumption of the saltmarsh isopod Sphaeroma rugicauda (Leach) was measured on 10 occasions during 1973. It was found that the metabolism depended upon both body size and exposure temperature. For most of the year, slope values relating oxygen consumption with size were between 0.5 and 0.9, but during August they were greater than 1.0. Within the environmental temperature range the R-T curve was strictly temperature-dependent, but during the summer there was a translation and rotation of the upper part of the curve to the right. A plateau of temperature-independent metabolism occurred between 30° and 35°C, which are temperatures approaching the critical maximum for the species. Thermal acclimation for 7 days at 20°C resulted in an extended plateau of temperature-independent metabolism between 15° and 35°C.     

Sex ratio and hermaphroditism in Oregon intertidal populations of the echinoid Strongylocentrotus purpuratus

Examination of large numbers of the echinoid Strongylocentrotus purpuratus (Stimpson) in the years 1969, 1970 and 1971 indicated that the sex ratio at Yaquina Head, Oregon (USA) was significantly different from 1:1, with males predominating. During 1968, 1970 and 1971 at Boiler Bay and Whale Cove, Oregon, the sex ratio did not differ significantly from 1:1 but, in 1969 at Boiler Bay, the proportion of males was 0.61. Two functionally hermaphroditic individuals were found among the 1,354 Yaquina Head echinoids examined, but none were found at Boiler Bay in an equal number of individuals. The hypothesis is proposed that S. purpuratus is a labile gonochorist, with a multiple, autosomal, sex-determining mechanism whose expression may be influenced by the environment.     

Comparative studies on the metabolism of shallow-water and deep-sea marine fishes. II. Red-muscle metabolism in shallow-water fishes

Maximal rates of oxygen consumption in vitro have been measured under standardized conditions at three test temperatures (5°, 15°, 25°C) on minced preparations of red muscle from 10 species of shallow-water marine teleost fishes. These fishes came from three different geographic areas, two with cool average water temperatures (near 15°C: coastal southern California, Galápagos Islands) and one with warm average water temperatures (near 25°C: Hawaiian Islands). The group is made up of post-juvenile or adult epipelagic fishes, which are moderately or very active in terms of their locomotor activities. A large part of the range of phylogenetic diversity among the teleosts is represented, as is the body weight range from a few grams to several kilograms. The purpose of the work is to provide part of a set of tissue-metabolism data on shallow-water fishes for future comparison with similar results from deep-sea species. Of 8 complete curves for oxygen uptake rate versus temperature (R-T curves), 6 are normal in shape (Q₁₀1.5), 1 is normal but with a low Q₁₀, and 1 is partly flat, partly normal. The differences between the species in terms of both absolute positions and slopes of the R-T curves are not related in any consistent way to any of the three testable variables: phylogenetic position, long-term adaptation temperature, and body size. The red muscles of a variety of adult epipelagic fishes, at ecologically realistic temperatures, are shown to be exceptions to the general rule that tissues of ectothermous lower vertebrates have lower metabolic rates than comparable tissues of non-torpid endothermous higher vertebrates. This circumstance probably is a major factor in the great capacities for sustained high-speed swimming shown by most epipelagic fishes. Other physiological and ecological implications of the results are discussed.     

Some aspects of the thermal tolerance of Trichomya hirsuta (Mollusca: Bivalvia) of Eastern Australia

The upper temperature tolerance range for Trichomya hirsuta L. is described for acclimation temperatures of 10° to 32°C at 30% S. Changes in thermal resistance with time are described along with the incipient lethal temperatures. The eurythermicity for the species is 240.7 C°-squared. The ultimate upper incipient lethal temperature is 35.1°C. Definition of the incipient sublethal temperature is given, and is equal to 32°C. Diet effects are shown not to be significant in short-term lethal experiments, but indicate that unfed mussels reduce their resistance after prolonged exposure to high temperatures. Size was also shown to have no effect on resistance and tolerance. A comparison of the tolerance triangle of T. hirsuta with those of the temperate species Mytilus edulis and Mya arenaria revealed that the 24-h LT₅₀ minus 2.2 C° approximates the incipient lethal temperature. The eurythermicities of the temperate species are Mytilus edulis, 363 C°-squared; and Mya arenaria, 415 C°-squared.     

Metabolic adaptation of Cancer irroratus developmental stages to cyclic temperatures

Metabolic-temperature responses of the developmental stages of the sublittoral crab Cancer irroratus cultured at 10° to 20°C daily cyclic and 15°C constant temperatures were determined. Generally, the metabolic rate increased with temperature in the lower range with Q₁₀'s (temperature coefficients) above 2, compensated in the midrange with Q₁₀'s between 1 and 2, and declined at the higher temperatures with Q₁₀ values less than 1. For the larvae cultured at a constant temperature of 15°C, the compensatory response range narrowed with development from first zoeae to the later zoeal stages. In contrast, the compensatory response of the first zoeae, megalops, and crab stages within the range 10° to 25°C was interrupted by a zone of thermal sensitivity between 15° and 20°C for those individuals cultured in the 10° to 20°C cyclic regime. The compensatory response range is narrower for the third stage zoeae and broader for the second, fourth, and fifth stage zoeae. Metabolic rate-temperature (M-T) patterns of C. irroratus developmental stages cultured under the cyclic regime varied from those held at constant temperature by increased respiration and metabolic rate compensation between 20° and 25°C, and by an extension of the metabolically active range towards higher temperatures.     

Temperature tolerance of the polychaete worms Diopatra cuprea and Clymenella torquata

The resistance to high temperatures of two species of intertidal tube-dwelling polychaete worms has been tested to show seasonal and geographic variations. The summer 50% survival temperature at Beaufort, North Carolina, was 42.5 °C for Diopatra cuprea (Bosc) and 40.5 °C for Clymenella torquata (Leidy). Winter 50% survival temperatures for both species at Beaufort were approximately 4.0 C° lower. Both species showed a geographic difference in 50% survival temperature of more than 4.0 C° between North Carolina and Massachusetts in summer. D. cuprea from the Mississippi coast showed a lower survival temperature, probably due to combined temperature and salinity effects. Laboratory acclimation of C. torquata from Beaufort at low temperatures, during summer months, produced experimental results similar to those from winter animals. The seasonal differences in temperature tolerance are phenotypic expressions of a physiological response which can be related to environmental temperature patterns.     

Temperature adaptation in sea anemones: Physiological and biochemical variability in geographically separate populations of Metridium senile

Latitudinally separate populations of the sea anemone Metridium senile (L.) are very similar genetically by electrophoretic criteria, yet respond differently to temperature. Anemones from southern and northern California (USA) have different oxygen consumption patterns in response to acclimatory and acute changes in temperature. Northern anemones show a pronounced increase in Q₁₀ at temperatures just above the normal environmental range, but southern anemones do not. The two populations also differed in the extent of metabolic compensation to temperature following several weeks of acclimation. This acclimation regime resulted in changes in the activities of several enzymes of intermediary metabolism, yet the extent and direction of these changes did not display a consistent trend with regard to acclimation temperature or population. The biochemical concomitants of acclimatory and acute temperature effects were studied further by measuring the concentrations of adenylates (ATP, ADP, and AMP) in anemones from the two populations exposed to different temperature regimes. During cold acclimation for several weeks, total adenylate concentrations (A_T) increased in both the southern and northern populations, possibly due to metabolic rate compensation, since A_T is positively correlated with tissue metabolic rate in many species. Moreover, the extremely low weight-specific oxygen consumption rates of M. senile are probably related to its very low A_T values. Acute temperature decreases had no effect on adenylate concentrations and adenylate energy charge (AEC); in contrast, acute temperature increases led to large changes in adenylate concentrations. The effects of starvation on adenylate concentrations are pronounced, and the effect is temperature-dependent. In starved individuals held at 20°C, AEC values fell to 50% of normal values after 8 d, while those held at 10°C maintained normal AEC values.     

Comparative thermal metabolic patterns in Euterpina acutifrons dimorphic males

Thermal metabolic acclimation patterns have been determined for cold-and warm-acclimated dimorphic males of the copepod Euterpina acutifrons. The copepods were acclimated either to 15° or 25°C. Metabolic measurements were made at the two acclimation temperatures. At 25 °C, the small males had a higher rate than the large ones. At 15 °C, the large males had the higher rate. The metabolic pattern of thermal acclimation is also distinctive in the two forms. Small males showed metabolic adaptation at the lower acclimation temperature (15 °C), but not at the higher acclimation temperature (25 °C). In the large males the acclimation patterns were reversed, i. e., these males acclimated at the higher temperature, not at the lower. The acclimation patterns based on Precht's (1958) scheme of acclimation are entirely different in the two forms.Supported by PHS grant 5-SO5-FR-07070-02.     

Some effects of temperature on the growth and metabolic rate of juvenile blue crabs,Callinectes sapidus,in the laboratory

Juvenile blue crabs, Callinectes sapidus Rathbun, were grown in the laboratory at different temperatures, and metabolic-rate determinations were made. Growth is shown to be dependent upon temperature. Crabs kept at high temperatures (34° and 27°C) grow faster than those kept at lower temperatures (13°, 15°, and 20°C). Increase in size per molt is less at higher temperatures than at lower ones. Mortality is directly proportional to temperature between 13° and 34°C and is very high during ecdysis at elevated temperatures. Metabolic rate increases with temperature, but various degrees of acclimation are seen after 4 weeks exposure. No acclimation of general activity to temperature was found. The findings are applied theoretically to crabs living in the region of heated discharge canals of electrical generators: the motile blue crab could extend its growing season without decreasing size at maturity by active selection of thermal surroundings.In part based on a thesis submitted in partial fulfillment of the requirements for the Degree of Master of Science at the University of Florida, USA.     

Respiratory metabolism of crabs from marine and estuarine habitats: an interspecific comparison

The gill-surface area and R-T (metabolic rate-temperature) response during aquatic and aerial respiration of three different species of crabs, the subtidal Scylla serrata, the intertidal Sesarma quadratum, and the supratidal Ocypoda platytarsis, were compared. Scylla serrata has the highest, Sesarma quadratum the second highest, and O.platytarsis the lowest gill-surface area. Oxygen-consumption values in water under different ambient partial pressures of oxygen indicate that all three species display equally efficient aquatic respiration despite variation in gill area and number. There is no coherent R-T response either inter- or intra-specifically. However, the ability to extract oxygen from the surrounding medium (respiratory efficiency) conditions the R-T response in the different species. The R-T trends in aquatic respiration of Scylla serrata and aerial respiration of O.platytarsis reveal a normal response to temperature. The proportion of aquaticoxygen uptake to total respiration in all sizes in both Scylla serrata and Sesarma quadratum remains more or less constant (between 88 and 92%), whereas in the supratidal O.platytarsis, this proportion decreases with increasing weight (from 88 to 64 %), indicating a progressive loss of gill respiration to total respiration. All three species are metabolically equally efficient; what one lacks in respiratory surface it compensates by respiratory efficiency.     

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.     

Effects of temperature on photosynthesis and respiration in hermatypic corals

Photosynthesis and respiration rates of the reef corals Pocillopora damicornis (Linn.), Montipora verrucosa (Lamarck), Porites compressa Dana and Fungia scutaria Lamarck were measured under controlled temperatures. Results indicate that coral metabolism is closely adapted to ambient temperature conditions. Tropical corals measured at Enewetak, Marshall Islands, showed greater primary production compared to maintenance requirements at elevated temperatures than did subtropical varieties of the same species in Hawaii. Photosynthesis: respiration (P:R) ratios were significantly and negatively related with temperature between 18° and 31°C for all Hawaiian corals, whereas at Enewetak this ratio generally showed a curvilinear relationship for this temperature range. Extrapolations of P:R regressions on temperatures to a value of 2.0 (estimated as a minimum required for long-term functional autotrophy) coincide for Hawaiian specimens with published upper lethal temperatures. Extrapolation of P:R regressions for Enewetak specimens at temperatures above 25°C suggests lethal temperatures for these corals to be 2 to 5 C° higher than for Hawaiian corals, in good agreement with recent experimental findings. Interspecific differences in P:R temperature regressions for Hawaiian corals correlating with upper lethal temperature tolerances are described.Contribution No. 505 of the Hawaii Institute of Marine Biology.     

Prey defense,predator preference,and nonrandom diet: The interactions between Pycnopodia helianthoides and two species of sea urchins

Interactions between the predatory sea star Pycnopodia helianthoides (Brandt, 1835) and two of its natural prey, the sea urchins Strongylocentrotus purpuratus (Stimpson, 1857) and S. franciscanus (Agassiz, 1863), are examined with regard to predator preference, predator diet, and prey defenses. The sea star is able to detect both species of sea urchin upstream in a Y-trough, but does not consistently choose one over the other (i.e., no preference). However, when the sea star is presented with equal numbers of similar-sized specimens of the two species of sea urchin, its diet is markedly nonrandom, since S. purpuratus is eaten almost 98% of the time. The defensive responses of the two species of sea urchin differ in form and effectiveness. S. franciscanus employs its long spines as defensive weapons, pinching the rays of an attacking sea star. This defensive response is more effective than the pedicellarial response used by S. purpuratus. The nonrandom diet of the predator seems to result primarily from prey defensive responses that differ in effectiveness, rather than from an intrinsic, behavioral preference of the predator at an earlier stage in the predator/prey interaction.     

Experimentelle Untersuchungen zur Ökologie der marinen Kieselalge Thalassiosira rotula. I. Temperatur und Licht

Nearly 500 crabs, Scylla serrata (Forskal) (family Portunidae), ranging in wet weight from 0.2 to 14.0 g, were acclimated to 27° and 35°C and their respiratory metabolism under water and on exposure to air at test temperatures ranging from 16° to 38°C was studied. In aquatic respiration, the response to temperature of crabs acclimated to a temperature of 16°C is statistically significant, and directly related to their weight. Smaller crabs did not survive at the warm acclimation level of 35°C. The metabolic rates of cold-adapted S. serrata are higher than those of warm-adapted ones. The effect of acclimation to aerial respiration on crabs acclimated to cold temperature varied slightly between large and small crabs. The aerial respiration rate was less than a tenth of the aquatic rate for all sizes. The response of S. serrata to warm acclimation in air has been found to be almost opposite to its response in water.This paper formed part of a thesis approved for the award of the degree of Doctor of Philosophy by the Madras University, India.     

Heat tolerance,growth and regeneration in three North Sea bryozoans exposed to different constant temperatures

Three species of bryozoans—Membranipora membranacea (L.), Electra pilosa (L.) and Conopeum reticulum (L.) — are capable of acclimating to elevated temperatures, above the normal range experienced in nature, when exposed to a gradual increase in ambient temperature. Conspicuous differences in LD 50 values, as a consequence of acclimation, occur between representatives of the same species acclimated and grown at constant temperatures in the laboratory. The tolerance range of these species is influenced by their thermal history in the laboratory. While increased ambient temperatures accelerate growth rate, final colony size attained after prolonged exposure declines at higher temperatures. The size of zooecia attained is inversely proportional to the test temperature. Colonies of E. pilosa maintained at 22°C develop erect branches. Hence, it is probable that E. pilosa forma erecta is only a growth form of normally encrusting colonies of E. pilosa. Temperature affects rate of regeneration.     

Removal of glycine from solution by the sea urchin Strongylocentrotus purpuratus

Radioautography and liquid-scintillation counting of juvenile sea urchins — Strongylocentrotus purpuratus (Stimpson) — exposed to ³H- and ¹⁴C-labeled glycine showed that the ³H is rapidly translocated throughout the animal and lost, while the ¹⁴C is slowly translocated into the interior of the urchin and incorporated into insoluble material.     

Thermal tolerance of Strongylocentrotus purpuratus early life history stages: mortality, stress-induced gene expression and biogeographic patterns

In this study, we examined the differential thermal tolerance of Strongylocentrotus purpuratus early life history stages by comparing high temperature–induced mortality and the relative levels of the stress-induced gene, hsp70, between S. purpuratus embryos and larvae from adults collected throughout the species range. There was no significant difference between gastrulae and 4-arm plutei mortality from all sites examined. Furthermore, there was little variability in temperature tolerance across the biogeographic range as southern gastrulae and 4-arm plutei exhibited similar tolerances to northern individuals. Relative levels of hsp70 mRNA expression did not differ overall between the two developmental stages at each site. Across sites, all gastrulae and 4-arm plutei exhibited maximum hsp70 expression at approximately 25°C; however, the range of hsp70 expression was narrower in southern individuals, suggesting they are living closer to their upper thermal limit than northern individuals.     

Energetics of early development for the sea urchinsStrongylocentrotus purpuratus andLytechinus pictus and the crustaceanArtemia sp.

Embryos and larvae of two species of sea urchin,Strongylocentrotus purpuratus andLytechinus pictus, and larvae of the brine shrimpArtemia sp. (San Francisco brand) were cultured to investigate the contribution of dissolved organic material in seawater to the energetics of early development. When embroys ofS. purpuratus were reared in artificial seawater, a net loss in dry organic mass was observed. In contrast, when sibling embryos were reared to Day 2 under identical conditions in natural seawater, there was either a net increase in dry organic mass or no change. A net decrease in mass was observed in only one of five cultures reared in filtered natural seawater. Energy budgets for each species were determined by giving energy equivalents to the changes in carbohydrate, lipid and protein, and to the rate of oxygen consumption for each day of development. In the case ofS. purpuratus, the use of endogenous reserves accounted for either 0 or 38% of the metabolic demand for two independent cultures reared from Days 0 to 2. For larvae ofL. pictus, reared to 8 d, only 66% of the metabolic demand could be accounted for by the use of endogenous reserves. Sea urchins are capable of transporting dissolved organic material from seawater. Calculations revealed that the energy deficit during the early development of sea urchins (S. purpuratus) could be accounted for by the uptake of dissolved organic matter from seawater. However, for a species that cannot use this resource (Artemia sp.), the metabolic needs during development are supplied through the use of endogenous reserves.