Seasonal variation of membrane fluidity of the naturally acclimatized Baltic Sea amphipods Gammarus spp. and Monoporeia affinis

The membrane fluidity of the Baltic Sea crustaceans Gammarus spp. and Monoporeia affinis was studied in different seasons. Gammarus spp. were collected at a location with stable salinity and with temperature fluctuations from about 0 to 20 °C, and M. affinis at a deep location with stable salinity and a constant low temperature of about 1.5 to 4.5 °C. The membrane fluidity was measured from preparations enriched with plasma and mitochondrial membranes employing a fluorescence polarization technique using 1,6-diphenyl-1,3,5-hexatriene (DPH) as a probe. The measurements were carried out without any preceding acclimation of the animals. In Gammarus spp., the membrane fluidity increased in the order: summer animals < autumn animals < spring animals. The differences between seasons were significant, with pronounced homeoviscous adaptation, and correlated well with the water temperature. It is suggested that in Gammarus spp., temperature is the determining factor for membrane fluidity. In M. affinis, the membranes of summer animals were the most fluid, and there was a significant difference only between summer and spring animals. In M. affinis the difference was not connected to the water temperature and no homeoviscous adaptation was noticed. The deep-water M. affinis experience a long period of fresh food deficiency, which probably affects the membrane fluidity. Although there are seasonal differences in fluidities between Gammarus spp. and M. affinis, it is interesting to note that all data obtained from M. affinis settled between the polarization lines of spring and summer Gammarus spp., regardless of the prevailing temperatures. Received: 30 June 1999 / Accepted: 7 June 2000     

Foraging strategies of the marine iguana,Amblyrhynchus cristatus

Summary Two foraging strategies were found in marine iguanas (Amblyrhynchus cristatus); (1) subtidal feeding: the animals swam out to sea and dived for algae in the subtidal zone; (2) intertidal feeding: the animals foraged around low tide in the intertidal zone on more or less exposed algae. Most marine iguanas were very consistent in their foraging strategy and so could be classified as subtidal feeders (SFs) or intertidal feeders (IFs). Feeding strategy was weight-related (Fig. 1), not sexspecific. Animals 1,200 g were IFs, animals >1,800 g SFs. Some iguanas in between followed a mixed foraging strategy. SFs foraged independently of the tides, IFs always around low tide (Figs. 2, 3). Feeding time patterns of IFs and SFs are described (Table 1). Sea motion seemed to have little effect on the foraging pattern of SFs, but strongly influenced that of IFs (Fig. 2). The smaller a marine iguana, the faster it cooled when immersed in water (Fig. 4). The difference between water temperature and core temperature of animals returning from foraging was significantly greater in IFs than SFs (Fig. 5). SFs swimming in very cold water regulated their body temperature to prevent excessive cooling. Possible costs and benefits of the two foraging strategies are discussed. Only part of a marine iguana population lives really amphibiously and only ca. 5% of a 24 h day is spent close to or in the water. All social activities, including mating, take place on land. These life history characteristics preclude those adaptations to an amphibious way of life that would at the same time reduce the iguanas' ability to be maximally active at their typical terrestrial body temperature of 35° C.     

Factors affecting the feeding rate of the winkle Littorina littorea

The winkle Littorina littorea (L.) feeds mainly when moistened by the tide and is quiescent when exposed to the drying effects of the wind and sun. Upper shore winkles thus experience a reduced feeding time compared with individuals living lower in the intertidal zone, but are able to completely compensate by an increased rate of radular activity when immersed. The rate of radular activity is influenced by the body size of winkles, by experimental temperature and by acclimation temperature, but is mainly modified by the duration of exposure to air which directly controls the rate of radular activity on immersion.     

Distribution and temperature adaptation in the teleost fish genus Gibbonsia

Along the California coast there are three intertidal species of the genus Gibbonsia (G. elegans, G. metzi, G. montereyensis). These species have distinct but overlapping distributions. Collections were made seasonally from three areas. Each collection was divided into two groups — a critical thermal maximum and minimum were determined for one group and the other group was tested after a 3 week acclimation period. The three species showed genetically different adaptation abilities and these were correlated with species differences in latitudinal distributions. The species that experiences the widest seasonal temperature range proved capable of anticipatory adjustment to temperature while the two experiencing small seasonal changes showed only reactive adjustments. the species with the widest distribution showed the greatest ability to adapt to temperature extremes. After 1 week acclimation all three species demonstrated different mechanisms for heat and cold adaptation.     

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.     

Distribution of Juvenile,subadult and adult Scylla serrata (Crustacea: Portunidae) on tidal flats in Australia

In Deception Bay, northern Australia, during 1979–1981, a study was made of the distribution of Scylla serrata (Forskal) in an area having a broad intertidal zone. Juveniles (20 to 99 mm carapace width) were resident in the mangrove zone, remaining there during low tide. The majority of subadult crabs (100 to 149 mm) migrated into the intertidal zone to feed at high tide and retreated to subtidal waters at low tide. Adults (150 mm and larger) were caught mainly subtidally and only small numbers were captured in the intertidal at high tide. Few crabs were captured in the coolest months (May to August). Adults were captured on the flats mainly in the warmest months (January to April), but subadults could be captured over the entire summer (September to March). Juveniles were found in the upper intertidal throughout the year.     

Role of temperature on lipid/fatty acid composition in Pacific cod (Gadus macrocephalus) eggs and unfed larvae

During early development, oviparous fish species must use finite lipid and fatty acid (FA) reserves for both catabolism and structural components. In cold environments, developing fish have the additional constraint of maintaining membrane fluidity for metabolic efficiency (homeoviscous adaptation), resulting in further demand on lower melting point FAs like n-3 polyunsaturated fatty acids (PUFAs). To examine whether marine fish embryos physiologically adapt to changing temperature environments, we incubated Pacific cod (Gadus macrocephalus) eggs at 5 temperatures (0, 2, 4, 6, and 8 °C) in the laboratory and sampled them repeatedly during development to measure changes in lipid/FA composition. Pacific cod embryos increased n-3 PUFA content during the egg stage in all temperature treatments, with the possible exception of 0 °C, where poor survival and hatch success limited our ability for continued sampling. At the beginning of the hatch cycle, free-swimming embryos shifted from lipogenesis to lipid catabolism. The rates of lipogenesis and catabolism were temperature dependent, and the distinct increase in unsaturated fatty acids at temperatures <8 °C was consistent with homeoviscous adaptation theory. However, with the possible exception of embryos at 0 °C, the relative amounts of essential fatty acids (e.g., EPA, DHA, AA) were conserved in a similar manner across incubation temperatures. Collectively, these data suggest Pacific cod are capable of homeoviscous adaptation but cannot tolerate temperatures approaching 0 °C despite their possible ability to biosynthesize PUFAs from other energetic sources.     

Diatoms of the microphytobenthic community in a tropical intertidal sand flat influenced by monsoons: spatial and temporal variations

Seasonal variations in the microphytobenthic diatom community were investigated in an intertidal sand flat of a tropical marine environment influenced by monsoons. Cores of sediments were collected along the beach gradient: low tide, mid tide and high tide zone up to a depth of 15 cm.. Diatom abundance was lowest during the monsoons and highest during the post-monsoons and the early pre-monsoon season throughout the intertidal transect. Diatom diversity was highest at the mid tide, followed by the high and low tide zones. Diatoms were viable up to a depth of 15 cm throughout the intertidal transect. The diatom community included the pennates, the permanent residents of this area, centric genera, which lead an attached mode of life and also some planktonic genera, brought in from ambient waters. Among the pennates, Navicula and Amphora were the dominant genera whereas in the case of centrics, Thalassiosira dominated the community throughout the intertidal transect down to a 15 cm depth. . Grain size fractions, which served as predictors of some diatom genera changed with tidal zones. The effect of winds on the resuspension of the pennate diatoms was evident only at the low tide zone down to a depth of 5 cm . Chlorophyll a concentration proved to be a good predictor of both pennate and centric diatom abundance at the low tide zone down to a depth of 10 cm and at the mid tide zone down to a depth of 5 cm.. However, even though chlorophyll a concentrations failed to reveal any positive correlation with the diatom abundance at both the deeper sediment layers and the high tide zone, the fact that viable cells were present at these areas reveal that the diatoms adopt survival strategies, contributing significantly to the carbon budgets of such unstable habitats.     

Combining heat-transfer and energy budget models to predict thermal stress in Mediterranean intertidal mussels

Recent studies have emphasised that organisms can experience physiological stress well within their geographic range limits. Developing methods for mechanistically predicting the presence, absence and physiological performance of organisms is therefore important because of the ongoing effects of climate change. In this study, we merged a biophysical–ecological (BE) model that estimates the aquatic (high tide) and aerial (low tide) body temperatures of Mytilus galloprovincialis with a Dynamic Energy Budget (DEB) model to predict growth, reproduction and mortality of this Mediterranean mussel in both intertidal and subtidal environments. Using weather and chlorophyll-a data from three Mediterranean sites along the Italian coasts, we show that predictions of sublethal and lethal (acute) stress can potentially explain the observed distribution (both presence and absence) of mussels in the intertidal and subtidal zones, and the maximum size of animals in the subtidal zones. Importantly, our results suggest that different mechanisms limit the intertidal distribution of mussels, and that these mechanisms do not follow a simple latitudinal gradient. At the northernmost site (Palermo), M. galloprovincialis appears to be excluded from the intertidal zone due to persistent exposure to lethal aerial temperatures, whereas at the southernmost sites (Porto Empedocle and Lampedusa) sublethal stress is the most important driver of mussel intertidal distribution. Our predictions provide a set of hypotheses for future work on the role of climate change in limiting intertidal distribution of mussels in the Mediterranean.     

Lysosomal,tissue and cellular alterations in the gills,palps and intestine of the mussels Mytilus galloprovincialis,in relation to pollution

Lysosomal, tissue and cellular alterations of the gill, the palp and the intestine epithelium of the mussel Mytilus galloprovincialis (L.) collected from six stations along a closed estuarine system (Thermaikos Gulf, northern Greece) were monitored in May, August and December. Semi-quantitative evaluation of histopathological alterations was performed under the light microscope, while certain of the latter alterations were confirmed electron microscopically. In addition, morphometrical evaluation of the volume density of the lysosomes under the electron microscope was conducted. The observed alterations, as the detachment of gill epithelial cells from the basement membranes and the dilated extracellular spaces formed either between the lateral plasma membranes or between the infoldings of the basal plasma membrane in the palps and the intestine are possibly related to the degree of pollution at the examined stations. The particularly expanded extracellular spaces formed by basal plasma membrane infoldings in the intestine in spring may be attributable to red tide algal infection. On the other hand, differences in morphometric parameters of lysosomal structures indicated no direct relationships with pollution levels.Communicated by O. Kinne, Oldendorf/Luhe     

Upper temperature tolerances of some European molluscs

The upper temperature tolerances of three European species of Cardium [C. (Cerastoderma) glaucum Bruguière, C. tuberculatum L., C. (Cerastoderma) edule L.] are compared by means of median lethal temperature (LT₅₀) and median burial temperature (BT₅₀) determinations for periods of exposure up to 96 h. The influence of previous acclimation temperatures on these determinations is also considered. C. glaucum from shallow water (0.5 to 1 m), lagoonal, habitats in the Mediterranean Sea shows a greater thermal tolerance than C. tuberculatum from deeper water (4 to 5 m) in the Mediterranean. The thermal tolerance of C. edule from the intertidal zone, but at higher latitudes, is intermediate. Young C. glaucum show a greater resistance to short exposures to high temperatures than adults, but for longer exposures their tolerance is the same. All 3 species exhibit a significant effect of previous acclimation temperature on the burrowing response, and C. glaucum and C. tuberculatum a significant but lesser effect on LT₅₀. LT₅₀ determinations with C. edule showed secondary mortality effects which may have obscured any effect of acclimation temperature.     

Depletion of heavy isotopes of oxygen and hydrogen in tissue water of intertidal plants: implications for water economy

Stable oxygen and hydrogen isotope ratios of leaf and thallus water of the intertidal seagrasses Phyllospadix scouleri and P. torreyi and the marine algae Egregia menziesii, Gelidium coulteri, and Corallina vancouverensis from three locations in California, USA, were determined in 1987. Compared with subtidal seawater, most plant-water samples were depleted in the heavy isotopes ¹⁸O and deuterium. Depletion of heavy isotopes was greatest in plants growing at the highest intertidal elevations. This was an unexpected result, because enrichment of heavy isotopes occurs during evapotranspiration in terrestrial plants. Two possible mechanisms for this isotopic depletion are proposed: direct uptake of heavy isotope-depleted water vapor and preferential diffusion of ¹⁶O and ¹H into littoral plants from water remaining in the intertidal zone at low tide.     

Improved survival under heat stress in intertidal embryos (<Emphasis Type="Italic">Fucus</Emphasis> spp.) simultaneously exposed to hypersalinity and the effect of parental thermal history

Intertidal organisms exposed to thermal stress normally experience other stresses simultaneously, but how these combined stresses modify tolerance to heat, especially for embryos, is poorly understood. Tolerance of fucoid algal embryos to heat, with and without acclimation to a sublethal temperature and with simultaneous exposure to hypersaline media, was examined. Embryos of Fucus vesiculosus L. (mid-intertidal zone) were less tolerant than embryos of Fucus spiralis L. (upper intertidal zone); without acclimation and with a growth temperature of 14°C, about half of the embryos survived 3 h exposure to 33°C in F. vesiculosus and of 35°C in F. spiralis. Conditions experienced by parental thalli (4°C versus 14°C storage) significantly affected the heat tolerance of embryos grown for 24 h post-fertilization at 14°C in F. vesiculosus, a result that is important for biologists using fucoid algae as model systems. Acclimation to a sublethal temperature (29°C) or exposure to the LT₅₀ (33°C, F. vesiculosus; 35°C, F. spiralis) in 100 psu seawater (2850 mmol kg^–1 osmolality) resulted in 30–50% higher levels of embryonic survival. Higher levels of HSP60s were found in embryos exposed to 29–33°C than to 14°C; lower levels of HSP60s were present in embryos exposed to the LT₅₀ under hypersaline conditions than in normal seawater. Contemporaneous studies in 1995–1996 of substratum temperature and desiccation levels were made at Schoodic Point, Maine (USA) underneath F. spiralis and F. vesiculosus canopies and in Semibalanus balanoides patches. This study extends the bioindicator utility of heat-shock proteins in studies of intertidal organisms and demonstrates the importance of integrated stress responses in survival of a single stress factor (e.g. temperature).Electronic Supplementary Material Supplementary material is available in the online version of this article at     

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.     

Substrate size mediates thermal stress in the rocky intertidal

Variation in physical factors, such as slope, orientation, and wind exposure, shapes thermal conditions. Variation in substrate size is common in many habitats, but its thermal consequences for organisms are not well characterized. Larger substrates should remain more thermally stable and act as thermal refuges for associated organisms during short, thermally stressful periods such as midday temperature peaks or tidal exposure. In observations and a transplant and thermal integration experiment, we found that larger rock substrates stayed cooler and facilitated greater survival of the barnacle Semibalanus balanoides in the high intertidal relative to small substrates during the hot summer months in southern New England, USA. However, in thermally benign northern New England, rock substrate size had no effect on barnacle distributions, indicating that the thermal effects of substrate size are mediated by regional climate.     

Ultrastructural changes in the gill epithelium of the crab Chasmagnathus granulatus (Decapoda: Grapsidae) in diluted and concentrated seawater

Chasmagnathus granulatus Dana, 1851 is an intertidal estuarine crab that experiences acute salinity changes ranging from <1‰ to full-strength seawater and even hypersaline waters in tide pools concentrated by evaporation. Ultrastructural changes induced by salinity in the posterior gills were examined in crabs collected from the Rio de la Plata estuary Argentina during March 1999. The posterior gills of C. granulatus are involved both in ion uptake and ion secretion depending on the acclimation medium. These organs are mostly lined with a thick tissue, which presents the characteristics of a typical salt-transporting epithelium. Electron microscopy analysis of gill tissue from crabs acclimated to dilute, full, and concentrated seawater (12‰, 34‰, and 45‰ salinity) showed significant development of basolateral membrane interdigitations, with numerous mitochondria and conspicuous apical membrane infoldings. Morphometrical analysis indicated that the subcuticular space delimited by the infolding of the apical membrane was significantly increased in the gills of high-salinity acclimated crabs. Septate junctions, which are thought to define the paracellular permeability, were significantly shorter in high-salinity acclimated crabs, suggesting a possible role of the paracellular pathway in salt secretion.     

Genetic evidence for the cryptic species pair, <Emphasis Type="Italic">Lottia digitalis </Emphasis>and <Emphasis Type="Italic">Lottia austrodigitalis</Emphasis> and microhabitat partitioning in sympatry

It has been proposed that the common West Coast limpet, Lottia digitalis, is actually the northern counterpart of a cryptic species duo including, Lottia austrodigitalis. Allele frequency differences between southern and northern populations at two polymorphic enzyme loci provided the basis for this claim. Due to lack of further evidence, L. austrodigitalis is still largely unrecognized in the literature. Seven additional enzyme loci were examined from populations in proposed zones of allopatry and sympatry to determine the existence of L. austrodigitalis as a sibling species to L. digitalis. Significant allele frequency differences were found at five enzyme loci between populations in Laguna Beach, southern California, and Bodega Bay, northern California; strongly supporting the existence of separate species. Both species exhibit two microhabitat morphotypes, a gooseneck barnacle morph in the mid-intertidal zone and a rock morph in the high-intertidal zone. In sympatry, L. austrodigitalis was more abundant higher in the intertidal on rocks, whereas L. digitalis was more abundant lower in the intertidal on barnacles. This finding supports earlier claims of microhabitat partitioning in this sibling species pair. In addition to this finding, the transition zone between the species was found to have shifted substantially northward in only two decades, from Monterey Peninsula, CA to near Pigeon Point, CA, where L. digitalis previously dominated.     

Critical thermal maxima of two species of estuarine fish

Critical thermal maxima (CTM) were determined for winter flounder (Pseudopleuronectes americanus, Walbaum) and scup (Stenotomus chrysops, L.), acclimated to ambient water temperature or constant increments above ambient. The acclimation regimes simulated a natural environment by tracking the temperature fluctuations in Point Judith Pond, a small Rhode Island estuary. An elevation in CTM from 30.2° to 35.6°C was observed for scup as the mean acclimation temperature increased from 14.8° to 22.2°C. CTM for flounder increased from 26.4° to 32.4°C over an acclimation range of 4.2° to 23.0°C. The use of CTM studies for predicting the effects of heated effluents is discussed.     

Field observations on the rhythmic behaviour of Nerita textilis (Gastropoda: Prosobranchia)

Field observations on Nerita textilis Dillwyn inhabiting rocky cliffs at Sar Uanle (Southern Somalia) revealed that during the resting phase some of the snails are scattered on the upper cliff and some are clustered at lower levels. During feeding they all migrate downward, but to different extents. Maximum activity of both groups occurs during low water (LW) between midday and midnight. During spring tides (ST), the scattered snails reside higher up and make smaller feeding excursions than during neap tides (NT). During ST, some clustered snails join the belt of scattered individuals, while the trend reverses during NT. A diurnal and a tidal component appear in their periodicity.Publication of the Centro di Studio per la Faunistica ed Ecologia Tropicali del C.N.R. Researches on the coast of Somalia. The shore and the dune of Sar Uanle, No. 14.     

Seasonal abundance,distribution and growth of postlarval and juvenile grass shrimp (Palaemonetes pugio) in a Georgia,USA, salt marsh

Postlarval and juvenile grass shrimp (Palaemonetes pugio Holthuis) ≦15 mm total length (TL) were abundant at low tide in shallow aquatic microhabitats (i.e. puddles and films of residual tidal water) in the intertidal zone of a salt marsh on Sapelo Island, Georgia, USA from 1982 to 1984. The highest concentrations of young P. pugio occurred at 190 to 200 cm above mean low water, ∼10 to 20 cm below mean high water. The intertidal distributions of young grass shrimp expanded and contracted with changes in tidal amplitude. Postlarval grass shrimp (6 to 8 mm TL) continuously recruited into the intertidal marsh population from May until October, but densities varied in a regular pattern with peaks in abundance occurring at ∼2-week intervals, corresponding to spring tide periods in the lunartidal cycle. Although present nearly year-round in the intertidal marsh, juveniles (9 to 15 mm TL) were most abundant from August to October. Apparent growth rates of individuals up to 15 mm TL averaged 0.268±0.026 (mean±95% C.I.) mm d^-1 from May to October and 0.070±0.032 mm d^-1 in November and December. Unlike larger aquatic organisms, which can forage in the emergent marsh only when it is flooded by the tide, juvenile grass shrimp have constant access to intertidal resources. Although potentially important predators in this system, the role of young P. pugio in the trophic organization of salt marsh benthic invertebrate assemblages has yet to be examined. Contribution No. 576 of the University of Georgia Marine Institute