Variations in cardiac performance and heat shock protein expression to thermal stress in two differently zoned limpets on a tropical rocky shore

Understanding variation in physiological adaptations to thermal stress is vital when investigating intertidal species?? distribution patterns. The thermal sensitivities of two limpets, Cellana grata and C. toreuma, differed in accordance with their vertical distributions. Cardiac performance was maintained at higher temperatures (~47°C) for the high-zone C. grata than the mid-zone C. toreuma (~42°C). At 40°C, C. grata maintained regular heart function for ~4?h, while heart function of C. toreuma decreased rapidly. Heat shock protein expression revealed that C. toreuma had two constitutive isoforms, Hsp77 and Hsp72, and C. grata one inducible form, Hsp75, which was upregulated at 40°C, suggesting C. grata has a more effective heat shock response than C. toreuma. The temperature-adaptive differences in cardiac thermal tolerance and Hsp expression match observed differences in thermally induced mortalities with the onset of summer and may help predict differential effects of climate change on the two congeners.     

Assessment of factors affecting heart rate of the limpet Patella vulgata on the natural shore

 Heart rate variations of a population of the limpet Patella vulgata were monitored in the natural environment (Lough Hyne, southern Ireland) by non-invasive, optoelectronic recording. The heart rates of 145 limpets of different sizes, living on vertical and horizontal substrata, were measured both in air and water at different environmental temperatures, while the animals were inactive on their home scars. The heart rates of emersed, inactive limpets were positively related to air temperature and negatively related to limpet size. These relationships were similar for limpets on vertical and horizontal substrates. In contrast, no significant relationship between heart rate and temperature was found in submerged limpets, probably due to the narrow thermal range of the water during the study period. During submersion, a significant negative relationship between heart rate and size was evident for limpets on vertical surfaces but not for limpets on horizontal surfaces. In general, submerged limpets had a higher heart rate, 1.16 times that of limpets exposed to air. Moreover, the heart rates of nine animals were recorded while they were moving and while inactive on their home scars. Active limpets had a faster heart rate, 1.6 times that of limpets resting on their home scars. The dependence of heart rate on environmental temperature, size, respiratory medium and activity, as observed in limpets on the shore, agrees well with laboratory data and with previous findings of the correlation of oxygen consumption with the same factors. Such in situ measurements may, therefore, prove useful in attempts to determine natural levels of energy expenditure in models on the behaviour of foraging molluscs. Received: 5 September 1999 / Accepted: 17 April 2000     

Analyses of limpet defense and predator offense in the field

Between November 1982 and August 1984 diving observations of two intertidal limpets' defensive behaviours in response to two predatory species were made near the Seto Marine Biological Laboratory of Kyoto University, in Wakayama Prefecture, Japan, for over 800 h. The focus of the observations was a simple isolated rock, 0.9 m² in area with an inclination of 75°. The pulmonate homing limpet Siphonaria sirius responded by fleeing from its home in response to the whelk Thais clavigera and by calmping tightly to its home scar in response to the starfish Coscinasterias acutispina. Both behaviours were successful for avoiding predation. After a whelk moved far away, limpets returned to their homes. The patellid limpet Cellana toreuma showed mantle folding behaviour or fled for an average distance of 8 cm vertically upwards in response to the whelk and for an average distance of 21 cm vertically upwards in response to the starfish. The amount of time predators spent foraging increased in the low intertidal zone relative to the mid and high zones, and most predator attacks were observed during the limpets' resting periods (submersion in late afternoon or nighttime). The upward directional flight displayed by C. toreuma was related to the upward orientation of the limpets' heads when they are in a resting position. Half of the limpets observed fleeing downward away from a starfish, a movement elicited by a downward attack, received a second attack. These limpets were preyed upon at the second encounter 67% of the time. The upward fleeing behaviour of C. toreuma resulted in the shifting of its resting site higher up the study site where fewer attacks occurred and where the mortality rate was at its lowest. It is suggested that the homing behaviour of S. sirius and the upward fleeing behaviour of C. toreuma are mechanisms by means of which interspecific competition between limpets is reduced and coexistence on a small rock surface is maintained.     

Genetic differentiation in Sargassum polyceratium (Fucales: Phaeophyceae) around the island of Curaçao (Netherlands Antilles)

The seaweed Sargassum polyceratium Montagne inhabits a broad spectrum of subtidal and intertidal habitats. Genetic diversity and spatial genetic structure were examined within and among 12 stands using random amplified polymorphic DNA (RAPD) phenotypes. Data were analyzed using analysis of molecular variance (AMOVA) and Shannon's information measure. In both analyses, 60-75% of the variation occurred within stands and 25-40% between stands. These values are consistent with out-crossing, high-dispersal species. Significant differentiation was found among bays ca. 25 km apart (Shannon's G'_st averaged 0.37 and pairwise AMOVA K_st values averaged 0.272) and among stands 150-200 m apart within bays (AMOVA K_st values averaged 0.149). Effects of shore (windward vs. leeward), depth, and bay on population structure were tested. These analyses revealed that the factor depth is confounded with shore, and that bays show significant differentiation from each other but are not completely isolated from one another. Mantel tests for differentiation-by-distance were significant along both sides of the island but stronger along the windward side. A neighbor-joining analysis of genetic distances among stands showed that the effects of currents around both tips of the island were especially important for shallow populations. For S. polyceratium, depth and bay promote population differentiation along shores, yet dispersal around the tips of the island simultaneously connects these populations to varying degrees. This study highlights the importance of investigating the relative contribution of habitat factorsin relation to island-scale population structure.     

Strategies for carbon acquisition in the red marine macroalga Coccotylus truncatus from the Baltic Sea

Marine macroalgae need carbon-concentrating mechanisms because they have only limited access to CO₂ in their natural environment. Previous studies have shown that one important strategy common to many algae is the activity of periplasmic carbonic anhydrases that catalyse the dehydration of HCO₃^- into CO₂. The latter can then cross the plasma membrane by passive diffusion. We hypothesised that an active (energy-consuming) mechanism might also be involved in the membrane transport of CO₂, as is the case in a number of microalgae. Coccotylus truncatus was chosen as a model organism for this study because it belongs to a group of algae that usually lack direct HCO₃^- uptake: sublittoral red algae. The method used to study carbon uptake was pH drift of the seawater medium surrounding the algae in a closed vessel, with and without the addition of specific inhibitors or proton buffers. Measured parameters included pH, total inorganic carbon and alkalinity of the seawater medium. Our results suggest that, in C. truncatus, periplasmic carbonic anhydrase as well as H⁺ extrusion, probably driven by a vanadate-sensitive P-type H⁺-ATPase (proton pump), are involved in CO₂ uptake. No direct uptake of HCO₃^- was discovered. This paper also presents data on the buffer capacity of several proton buffers and the carbon-uptake inhibitors acetazolamide, 4,4'-diisothiocyano-stilbene-2,2'-disulfonate (DIDS) and orthovanadate in Baltic Sea water with a salinity of 6.5 psu.     

Oxygen and carbon stable isotopic profiles of the fan mussel, Pinna nobilis, and reconstruction of sea surface temperatures in the Mediterranean

Stable oxygen and carbon isotope profiles ('¹⁸O_skeletal and '¹³C_skeletal), taken along the direction of growth from the umbo to the shell margin in shells of the pinnid Pinna nobilis, were used to reconstruct sea surface temperatures (SST) in the south-east Mediterranean and ontogenetic records of metabolic CO₂ incorporation. Comparison of the seasonal cycle of SST, predicted from the '¹⁸O_skeletal record of a small (young) rapidly growing pinnid and temperature measured with a continuous in situ recorder showed that P. nobilis calcifies under isotopic equilibrium with surrounding seawater, thus indicating that P. nobilis shells can be used as a reliable predictor of SST. A 10-year SST record for the south-east Mediterranean was reconstructed from the shell profiles of four pinnid shells of different sizes and ages collected in 1995 and 1996. Reliable resolution of the seasonal SST could only be achieved during the first 4 years of shell growth. As the pinnids grew older, the temperature record was poorly resolved because the shell growth had diminished with age, resulting in time-averaging of the record. The amplitude of the generated seasonal temperature cycle compared favourably (DŽ°C) with a long-term temperature record from northern Mediterranean waters. Clear seasonal cycles in '¹³C_skeletal were observed with an amplitude of ~1.0‰, similar to the calculated seasonal changes in '¹³C of seawater (0.6‰) overlying seagrass meadows. An ontogenetic trend towards less positive '¹³C_skeletal values was too large to be attributed to any decrease in '¹³C in seawater resulting from the invasion of anthropogenic CO₂. It is suggested that the temporal changes of '¹³C_skeletal are due to incorporation of respiratory CO₂ into the extrapallial fluid and reflect changes in the metabolic activity of the pinnid rather than changes in the isotopic composition of dissolved inorganic carbon within the surrounding seawater.     

Possible involvement of the glycocalyx in the ichthyotoxicity of Chattonella marina (Raphidophyceae): immunological approach using antiserum against cell surface structures of the flagellate

Although the ichthyotoxic mechanism of Chattonella marina is still unknown, several lines of evidence suggest that the reactive oxygen species (ROS), such as superoxide anion (O₂^-), hydrogen peroxide (H₂O₂)_, and hydroxyl radical (·OH), produced by C. marina are involved in the mortality of fish exposed to this flagellate. Recently, we found that the cell-free supernatant prepared from C. marina, which is considered to contain the glycocalyx, showed NADPH-dependent O₂^- generation. In this study, we prepared antiserum against the crude glycocalyx of C. marina. Using indirect immunofluorescence, it was confirmed that the antiserum specifically reacted with C. marina cells. In addition to C. marina, the antiserum also reacted with other raphidophycean flagellates such as Heterosigma akashiwo, Olisthodiscus luteus, and Fibrocapsa japonica, whereas no reactivity was observed against six other flagellate species tested. These results suggest that raphidophycean flagellates have common epitopes recognized by the antiserum. Interestingly, immunohistochemical analysis of paraformaldehyde-fixed gill lamellae from yellowtail exposed to C. marina revealed that the antiserum stained the surface of gill lamellae, while no such staining pattern was observed in control gill lamellae. These results suggest that the glycocalyx may be discharged when C. marina cells are inhaled into the fishes' mouths and then come into contact with the gill surface. Based on the present results, together with our previous findings, we propose that continuous accumulation of the discharged glycocalyx on the gill surface occurs during C. marina exposure, which may be responsible for the ROS-mediated severe gill tissue damage leading to fish death.     

Characterization of urease activity in three marine phytoplankton species,<Emphasis Type="Italic"> Aureococcus anophagefferens</Emphasis>,<Emphasis Type="Italic"> Prorocentrum minimum</Emphasis>, and<Emphasis Type="Italic"> Thalassiosira weissflogii</Emphasis>

The availability of different forms of nitrogen in coastal and estuarine waters may be important in determining the abundance and productivity of different phytoplankton species. Although urea has been shown to contribute as much as 50% of the nitrogen for phytoplankton nutrition, relatively little is known of the activity and expression of urease in phytoplankton. Using an in vitro enzyme assay, urease activities were examined in laboratory cultures of three species: Aureococcus anophagefferens Hargraves et Sieburth, Prorocentrum minimum (Pavillard) Schiller, and Thalassiosira weissflogii (Grunow) Fryxell et Hasle. Cultures of P. minimum and T. weissflogii were grown on three nitrogen sources (NO₃^m, NH₄⁺, and urea), while A. anophagefferens was grown only on NO₃^m and urea. Urease was found to be constitutive in all cultures, but activity varied with growth rate and assay temperature for the different cultures. For A. anophagefferens, urease activity varied positively with growth rate regardless of the N source, while for P. minimum, urease activity varied positively with growth rate only for cultures grown on urea and NH₄⁺. In contrast, for T. weissflogii, activity did not vary with growth rate for any of the N sources. For all species, urease activity increased with assay temperature, but with different apparent temperature optima. For A. anophagefferens, in vitro activity increased from near 0-30°C, and remained stable to 50°C, while for P. minimum, increased in vitro activity was noted from near 0-20°C, but constant activity was observed between 20°C and 50°C. For T. weissfloggii, while activity also increased from 0°C to 20°C, subsequent decreases were noted when temperature was elevated above 20°C. Urease activity had a half-saturation constant of 120-165 wg atom N l^у in all three species. On both an hourly and daily basis, urease activity in A. anophagefferens exceeded nitrogen demand for growth. In P. minimum, urease activity on an hourly basis matched the nitrogen demand, but was less than the demand on a daily basis. For T. weissflogii, urease activity was always less than the nitrogen demand. These patterns in urease activity in three different species demonstrate that while apparently constitutive, the regulation of activity was substantially different in the diatom. These differences in the physiological regulation of urease activity, as well as other enzymes, may play a role in their ecological success in different environments.     

Seasonal variation in ecophysiological patterns in macroalgae from an Arctic fjord. I. Sensitivity of photosynthesis to ultraviolet radiation

Marine macroalgae inhabiting Arctic coastal ecosystems are exposed to pronounced seasonal variations in the radiation regime, including harmful UVB radiation. This study presents the first data on the seasonal changes in the sensitivity of macroalgal photosynthesis towards UV exposure by comparing under-ice, clear-water and turbid-water conditions characteristic for late winter, spring and summer. Various brown (Laminaria saccharina, L. digitata, L. solidungula, Saccorhiza dermatodea, Desmarestia aculeata), red (Palmaria palmata, Devaleraea ramentacea) and one green macroalgal species (Monostroma aff. arcticum) were collected at the same water depth throughout the seasons in the Kongsfjord (Spitsbergen, Svalbard, Norway). Maximum quantum yield (F_v/F_m) and maximum photosynthetic electron transport rates (ETR_max) were determined immediately after collection, after 2 h exposure to artificial UV radiation and after 18 h recovery in dim white light. Photosynthesis of the studied species showed different responses depending on their morpho-functional and physiological characteristics, their life strategies, phenology and depth distribution. Within the genus Laminaria, maximum quantum yield of adult specimens of the deep-water species L. solidungula was most strongly UV sensitive. Adult L. saccharina exhibited a lower UV sensitivity than a 6-month-old specimen. Inhibition of photosynthesis after UV exposure remained at the same level throughout the study period, both in adult L. saccharina and S. dermatodea. However, adult specimens of L. saccharina collected in May showed partial recovery only, whereas photosynthesis of specimens from both species collected later recovered fully. D. aculeata exhibited a remarkable decrease of UV sensitivity during the study period. Photosynthesis of specimens collected under the ice was strongly inhibited by UV, but the degree of inhibition decreased during spring and summer. Concomitantly ETR_max values were low after UV exposure in specimens collected in June, but increased later in the season. P. palmata exhibited a relatively flexible response. Photosynthesis in specimens collected under the ice in June or in turbid water in July/August was relatively strongly inhibited; specimens collected during sunny periods and in clear water in spring showed a much lower degree of photoinhibition after UV exposure. The seasonal pattern of low/high ETR_max values in spring/summer is probably a characteristic of the life strategy of this species. The UV sensitivity of D. ramentacea exhibited a similar seasonal pattern. In M. aff. arcticum, UV sensitivity increased and ETR_max values decreased during the study period, reflecting the life strategy of this annual late winter/spring species. The physiological basis for the seasonal changes in UV sensitivity of photosynthesis is presented in a companion paper (this issue).     

Small-scale spatial and seasonal differences in the distribution of beach arthropods on the northwestern Tunisian coast. Are species evenly distributed along the shore?

Spatial distributions of sand beach arthropods were studied at Zouara (Nefza), a beach dune system along the northwestern coast of Tunisia. Two transects with pitfall traps, perpendicular to the shoreline and placed from sea towards land, captured spontaneously crawling arthropods. The pitfalls were used to obtain data on horizontal zonations for 2 days during different seasons (April, October). In April, in order to assess local, long-shore distributions of surface-active beach invertebrates, ten transects with pitfall traps were placed every 100 m, for 2 days. Core samples were also taken in correspondence with each of the ten transects to obtain the distribution of burrowing individuals. Sand samples were taken for successive laboratory analyses (salinity, grain size and organic contents). Local climatic conditions were also recorded. One isopod (Tylos europaeus, subdivided into three different size classes), two amphipods (Talitrus saltator, Talorchestia brito and indistinguishable juveniles) and four coleopteran species (Eurynebria complanata, Scarites laevigatus, Phytosus nigriventris, Phaleria acuminata) were considered. Mean zonations perpendicular to the shoreline showed that crustaceans were found closer to the sea-line limits than coleopterans and that between seasons some species changed their mean zonation more than others. Local, long-shore distributions of active and burrowing individuals showed that most species were unevenly distributed along the shore. These discontinuous distributions differed according to the species and age class and were mainly related to specific microclimatic conditions and to nutrition.     

Genetic variability and level of differentiation in North Sea and Baltic Sea populations of the green alga<Emphasis Type="Italic"> Cladophora rupestris</Emphasis>

Cladophora rupestris is a perennial filamentous macroalga belonging to the Chlorophyta. It is widely distributed on both sides of the northern Atlantic Ocean and penetrates into the brackish Baltic Sea down to ca. 4 psu salinity. In this paper we present evidence for genetic differentiation of a Baltic form of this marine alga. We assessed genetic structure within and among 11 populations ranging along a salinity gradient from the Norwegian coast to the northern Baltic Sea proper. Samples of 328 individuals were studied using starch-gel protein electrophoresis to evaluate genetic variability and interpopulation differentiation based on allozymes. Of 11 loci examined, only one was polymorphic. For this locus, encoding superoxide dismutase (SOD-3), a total of seven alleles were distinguished. We found two genetically differentiated groups of populations of C. rupestris, one Baltic Sea group and one North Sea group, with a distinct border in the southern Kattegat near the entrance to the Baltic Sea. The genetic differentiation for SOD-3, expressed as pairwise F_ST values between the populations, was generally higher within the Baltic Sea group (0.10-0.43) than within the North Sea group (0.05-0.10); in the latter group also fewer pairs of populations differed significantly. Pairs of populations from different groups had the highest F_ST values (0.20-0.60). Hierarchical analysis of variance showed that 29.6% of the total variation in the SOD-3 locus was explained by variation between the two groups, while only 4.2% was explained by variation among the populations within the groups. The remaining variation (66.2%) was found within the populations.     

How hot for how long? The potential role of heat intensity and duration in moderating the beneficial effects of an ecosystem engineer on rocky shores

Studies of the importance of ecosystem engineers have focused on their benefit to biodiversity through ameliorating environmental stress, without understanding the exact benefits gained by associated organisms. On monsoonal tropical shores, species experience strong seasonality in environmental conditions from almost temperate winters to tropical summers when mass mortalities can occur during daytime emersion. The limpet, Cellana grata, associates with an ecosystem engineer, the barnacle Tetraclita japonica, in summer, but not in winter. To understand the benefits of this association, physiological responses (heart rates and osmotic responses) of the limpet either amongst Tetraclita or on open rock surfaces were investigated under three environmental conditions in a laboratory mesocosm: awash (non-stressed), low (30 °C) and high thermal stress (40 °C) of varying durations (3 or 6 h). In general, at 30 °C, limpets showed similar physiological responses under all conditions for both exposure durations. After 6 h at 40 °C, however, all limpets on open rock surfaces died, whereas those associated with barnacles survived. The surviving limpets experienced similar levels of stress as those exposed to 40 °C on open rock surfaces for half the time (3 h), showing that both the level and duration of stress were important. Limpets, therefore, gain benefits from engineering species when conditions are extreme or stress is prolonged. Under low temperatures or for short durations (e.g. winter), associating with barnacles does not provide physiological benefits. Understanding how, and to what extent, associates benefit from ecosystem engineers highlights how close the margin between survival and mortality can be when conditions are extreme.     

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.     

Studies on pigments,proteins and photosynthetic rates in some mangroves and mangrove associates from Bhitarkanika,Orissa

Pigment contents, proteins and net photosynthesis were investigated in fully developed leaf of 1-year-old seedlings of six mangroves (Bruguiera gymnorrhiza, Rhizophora apiculata) and mangrove associates (Caesalpinia bonduc, Cerbera manghas, Derris heterophylla, Thespesia populnea), collected from Bhitarkanika, located on the east coast of India. Large variations in the photosynthetic rates (P_N) among the six species were observed, ranging from 10.16 µmol CO₂ m^-2 s^-1 in C. bonduc to 15.28 µmol CO₂ m^-2 s^-1 in R. apiculata. The total leaf protein content ranged from 12.09 mg g^-1 dry wt in T. populnea to 51.89 mg g^-1 dry wt in B. gymnorrhiza. The chlorophyll a/b ratio was typically about 3.0 in all the studied species, except C. bonduc (2.8). Photosynthetic rates and chl a/b ratio in the leaves were found to be correlated. Analysis of chlorophyll and xanthophyll spectra suggested: (1) variations in different forms and amounts of carotenes as well as xanthophylls and (2) the presence of high amounts of near-UV-absorbing substances in leaves, particularly in the two mangroves (B. gymnorrhiza, R. apiculata) and a mangrove associate (T. populnea), which appears to be an adaptive feature. Estimation of the chl a/b ratios in isolated thylakoids yielded a low value of 1.8 for R. apiculata and >2.6 for other species. The total protein/chlorophyll ratios in thylakoids varied considerably from 3.14 (D. heterophylla) to 10.88 (T. populnea) among the mangrove associates and from 16.09 to 18.88 between the members of the Rhizophoraceae. The chlorophyll/carotenoid ratios in thylakoids of the six species were more or less similar. The absorption spectra for washed thylakoids of C. manghas and D. heterophylla exhibited absorption characteristics typical for C₃-plant thylakoids. However, thylakoids isolated from R. apiculata, B. gymnorrhiza, C. bonduc and T. populnea exhibited an unusual increase in absorption in the blue region (380-410 nm) of the absorption spectrum. The presence of high-absorbing (in the short-wavelength, near-UV region) pigments appears to be closely associated with the thylakoids in R. apiculata and T. populnea. Our results, therefore, suggest a wide range of variation, not only in protein and pigment contents of photosynthetic tissues, but also in the spectral characteristics and composition of the pigments in mangrove species. An understanding of the nature of these pigments in mangroves and their associates, under their natural conditions and especially in relation to eco-physiological adaptations, is necessary, not only in relation to conservation, but also to allow propagation under different salinity conditions.     

Cardiac output during exposure to <Emphasis Type="Italic">Chattonella</Emphasis><Emphasis Type="Italic">marina</Emphasis> and environmental hypoxia in yellowtail (<Emphasis Type="Italic">Seriola</Emphasis><Emphasis Type="Italic">quinqueradiata</Emphasis>)

The experiments were undertaken to measure, for the first time, cardiac output in yellowtail (Seriola quinqueradiata) during exposure to a harmful red tide flagellate (Chattonella marina). The responses were compared with those during exposure to environmental hypoxia to evaluate the significance of the drop of arterial oxygen partial pressure (PaO₂) in the fish-kill mechanisms by C. marina. PaO₂ immediately decreased, whereas heart rate (HR) was maintained until shortly before death during exposure to C. marina. Suffocation developing during the exposure resulted from a decrease in blood oxygen content, but not from lowered blood flow to the tissue. Although exposure to both C. marina and hypoxia immediately decreased PaO₂, arterial oxygen content (CaO₂) and pH (pHa) were significantly lower, whereas HR and cardiac output (&Qdot;) remained significantly higher, for the C. marina-exposed fish than for hypoxia-exposed fish. Although the drop in PaO₂ appears to play a pivotal role in the mechanisms of fish death by C. marina, other physiological response(s) should also be considered.     

Seasonal fluctuations in the photosynthetic capacity of photosystem II in symbiotic dinoflagellates in the Caribbean reef-building coral <Emphasis Type="Italic">Montastraea</Emphasis>

The photosynthetic capacity of photosystem II (PS II) in symbiotic dinoflagellates (Symbiodinium sp.), as measured by analysis of chlorophyll fluorescence, was investigated in the primary Caribbean reef-building corals, Montastraea annularis and Montastraea faveolata, for 5 years and Montastraea franksi over 2 years in the Bahamas. Significant seasonal fluctuations in the quantum yield of charge separation (F_v/F_m) of PS II were found in all species at all depths, with the highest photosynthetic capacity consistently recorded between mid-winter and early spring and the lowest photosynthetic capacity occurring in the mid to late summer. Corals residing in shallow depths of 1-2 m showed the greatest fluctuations in F_v/F_m, whereas deeper corals (3-4 and 14 m depths) had consistently higher values of F_v/F_m. Densities of symbiotic dinoflagellates and photosynthetic pigments followed a similar pattern. Fluctuations of photosynthetic capacity showed a strong correlation with seasonal patterns of water temperature and light. Such seasonal shifts in photosynthetic capacity are most likely due to several biochemical processes in the algae that lead to alterations of both photoprotection and photodamage. While symbiont density changed significantly on a seasonal basis, visual signs of coral bleaching were noted only in the fall of 1995 and the spring and summer of 1998. Comparisons of photosynthetic capacity and the decrease in the number of symbionts and their subsequent recovery indicated that symbiont populations in this study had the ability to recover quickly following bleaching events, as long as continued physical perturbation (e.g. thermal stress) did not shorten the recovery phase. Large-scale bleaching events are best viewed as the end points of seasonal physiological variation in which photosynthetic capacity and density of symbiotic dinoflagellates are reduced to a lower level than during "non-bleaching" years.     

A novel Vibrio sp. pathogen of the coral Pocillopora damicornis

A coral pathogen was isolated from the diseased tissue of Pocillopora damicornis in Zanzibar. The pathogenic bacterium, referred to as Vibrio coralyticus YB, was classified as a member of the genus Vibrio. Based on its 16S rDNA sequence, V. coralyticus is probably a new species. In controlled aquaria experiments at 26-29°C, inoculation of pure cultures of V. coralyticus YB either into the seawater or by direct contact onto the coral caused tissue lysis of P. damicornis fragments. At 29°C, lysis began as small white spots after 3-5 days, rapidly spreading so that by 2 weeks the entire tissue was destroyed, leaving only the intact bare skeleton. When an infected diseased coral was placed in direct contact with a healthy one, the healthy coral lysed in 2-4 days, further indicating that the disease was contagious. Inoculation with as few as 30 bacteria ml^-1 was sufficient to infect and lyse corals. Seawater temperature was a critical variable for the infectious process: infection and lysis occurred rapidly at 27-29°C, slowly at 26°C and was not observed at 25°C. The data suggest that the presence of V. coralyticus YB, even in low numbers, in seawater surrounding a coral reef will lead to tissue destruction of P. damicornis, when seawater temperatures rise.     

Movement patterns of the limpet Cellana grata (Gould) observed over a continuous period through a changing tidal regime

Time-lapse videography of limpets mounted with light-emitting diodes was used to monitor the movements of a population of the non-homing Cellana grata on a vertical gully wall in Hong Kong. Animals were monitored for >7 days to examine spatial and temporal variation in their behaviour as the tides transited from a semi-diurnal to an almost diurnal pattern. Movement was synchronised with the tides, irrespective of the day–night cycles. Limpets rested low on the shore and were stimulated to move by the rising tide. Individuals moved up shore with the flooding tide, maintaining themselves in the awash zone, and then down shore on the ebbing tide, until they reached a resting height when the tide then fell beneath them, exhibiting ‘zonal shuttling’. A tight coupling of limpet position to tide height persisted through the changing tidal pattern, and almost all animals displayed the same organisation of activity over all tides. Initiation of activity and maximum height reached were probably controlled by the tides, but the cessation of activity may have been controlled by an internal clock. The pattern observed is consistent with the threat of attack from aquatic predators coupled with the need to minimise physical stresses while exposed. It is also consistent with the avoidance of grazing lower on the shore where interspecific competition may be more intense. Limpets showed two peaks of activity per tide, corresponding to when the rate of change of tide height was the greatest, except when tides became much reduced during the transit to a diurnal pattern. Movement was triphasic: an initial rapid phase, followed by a slower phase in the high shore and then a rapid phase before the limpets stopped moving. This structure is common in limpets and in this case is likely to be a consequence of animals maintaining themselves within the awash zone. Tide height appears to determine foraging activity, but with modifications in the behaviour in response to factors operating at more local temporal and spatial scales.     

Influence of environmental factors on burrow irrigation and oxygen consumption in the mudflat invertebrate Urechis caupo

We examined burrow irrigation activity by the mudflat worm Urechis caupo in response to suspended food, ambient hypoxia (down to 3.3 kPa PO₂), hydrogen sulfide exposure (up to 100 µmol l^-1), and short-term temperature change (range 10-22°C). In normoxic, nutrient-free water at 14°C, O₂ consumption ( [(M)\dot]O₂ ) \left( {\dot M{\rm O}_2 } \right) was 45 nmol min^-1 g^-1, water flow rate ( [(V)\dot]_W ) \left( {\dot V_{\rm W} } \right) was 27 ml min^-1 (0.66 ml min^-1 g^-1), frequency of peristaltic waves (F_P) was 2.6 contractions min^-1, stroke volume (SV) was 11 ml, and O₂ extraction coefficient (EO₂) was 0.27. Adding suspended food to the burrow water occasionally elicited stereotypical feeding behavior but had no effect on any measured variables during nonfeeding periods. Hypoxia greatly decreased [(M)\dot]O₂ \dot M{\rm O}_2 (75% reduction at 3.3 kPa PO₂) but did not affect [(V)\dot]_W \dot V_{\rm W} , F_P, SV, or EO₂. Sulfide at 50 µmol l^-1 or less had no effect on burrow irrigation activity, whereas 100 µmol l^-1 sulfide decreased [(V)\dot]_W \dot V_{\rm W} by 58% and F_P by 50% but had no effect on SV. Temperature strongly affected [(V)\dot]_W \dot V_{\rm W} (Q₁₀ of 1.9 from 10°C to 22°C). We propose that U. caupo's ability to live in the hypoxic, sulfidic mud of productive mudflat environments, combined with its very efficient mucous net, allows it to process much less water for feeding than other suspension-feeding invertebrates. This, in turn, necessitates an efficient O₂ extraction mechanism, which is provided by the water lung activity of U. caupo's unique hindgut.     

Long-term trends in abundance of cladocerans in the Central Baltic Sea

Long-term dynamics (1960-1997) of the cladoceran species Bosmina coregoni maritima, Evadne nordmanni and Podon spp. are described for the Gdansk Deep and the Gotland Basin (Central Baltic Sea). By using correlation analyses on seasonal time-series, the influence of temperature and salinity on the abundance of cladoceran species was investigated. A clear affinity to higher temperature was found for B. coregoni maritima in summer as well as for E. nordmanni and Podon spp. in spring. In addition to temperature, association tests with salinity revealed besides species-specific preferences, regional and temporal differences. Contrary to B. coregoni maritima, both other species were positively associated to salinity in summer and autumn in the Gdansk Deep. In the Gotland Basin only E. nordmanni was positively correlated to salinity in autumn. Differences in the response to hydrographic variables are possibly stage specific, i.e. between resting eggs and adults, or due to a different adaptation to the abiotic environment.