Stromatolitoid microbial nodules from Bermuda —a special micro habitat for meiofauna

In Bermuda, stromatolitoid microbial nodules are found in dense groups in seagrass beds and on subtidal sandy or rocky bottoms. They develop between January and the beginning of August and may reach 15 cm in diameter and 6 cm in height. Nodules grow on top of the sediment surface and consist of convex interwoven mats of the cyanobacterium Phormidium corium and fine sediment particles trapped in these mats. Nodules were studied in the field and laboratory (in 1989–1992) with respect to their structure, microbial community, sediment chemistry and associated meiofauna. Vertical profiles taken with microelectrodes showed steep gradients of oxygen and sulfide. While free oxygen was only detected in the upper 2 mm, the sulfide concentrations increased with depth and reached maximal 1250 moll^-1 inside the nodules. On the basis of microscopic observations and sediment chemistry, vertical sections through a nodule reveal four distinct layers: (A) an oxic green surface layer of growing cyanobacteria mats, (B) an anoxic yellow-white laminated layer of sediment particles, mucus and unpigmented cyanobacteria sheaths, (C) an anoxic sulfidic (sulfide <50 moll^-1) interface where colour changes from yellow to gray and (D) an anoxic sulfidic (sulfide can increase to 1250 moll^-1) gray core composed of sediment and decomposing cyanobacteria. The following meiofauna taxa were found in the nodules: Ciliata, Turbellaria, Gnathostomulida, Gastrotricha, Nematoda, Kinorhyncha, Polychaeta, Ostracoda and Harpacticoida. The distribution of the meiofauna was significantly different (p<0.05) within the four layers. The highest density of individuals was found in the sulfidic interface and core. Nematodes represented the dominant group in general. Thiobiotic organisms, such as the Gnathostomulida, Solenofilomorphidae, and Stilbonematinae were primarily found in the anoxic sulfidic layers of the nodules.     

Oxygen/sulfide regime and nematode fauna associated with Arenicola marina burrows: new insights in the thiobios case

Profiles of oxygen and sulfide around the burrows of the lugworm, Arenicola marina, from a North Sea tidal flat were examined with microelectrodes, and the steep gradients were related to the microdistribution of nematodes. Around the tail shaft free oxygen penetrated only 2 mm into the burrow wall, coinciding with a bright zone sharply limited by the ambient black sediment. Contrastingly, in normal bottoms of the tidal flat (controls) only the surface of the bright zone was supplied with free oxygen. Here, the dark colouration coincided with the presence of free hydrogen sulfide. Around the tail shaft the nearest free hydrogen sulfide was detected 6 mm from the burrow wall leaving several millimetres of black sediment without measurable free sulfide. We discuss how these divergencies may relate to the stability of the oxygen/sulfide gradients and the course of time involved in their formation. A total of 54 nematode species were identified. Based on non-metric Multidimensional Scaling Ordination, four nematode assemblages corresponded to four microhabitats of the A. marina burrow: the funnel, the feeding pocket, the tail shaft and the feacal cast. The tail shaft assemblage (oxic plus partly anoxic zones) was similar to that of the anoxic zone of the control sediment. It was dominated by the most abundant nematode in the present study, Metalinhomoeus biformis (mean abundance in tail shaft 202 indx10 cm^-3). Adults of common nematode species from sulfidic microhabitats had a significantly higher length/diameter ratio than those inhabiting the oxic zone of the control sediment (p<0.001). The chemical recordings and metric analysis indicate that these slender nematodes around the A. marina tail shaft and in the reduced horizons of the reference sites represent thiobiotic assemblages, as compared to the shorter and stouter oxybiotic species characterising the assemblages from the surface zone and (partly) the funnel.     

Symbiont photosynthesis increases both respiration and photosynthesis in the symbiotic sea anemone Anemonia viridis

Dark respiration of the symbiotic sea anemone Anemonia viridis (Forskäl) was observed to increase by 34% when anemones were exposed to hyperoxic sea water (150% oxygen saturation) overnight, and by 39% after exposure to 6 h in the light at a saturating irradiance of 300 E m^-2 s^-1 at normoxia (100% oxygen saturation). No increase due to light stimulation was observed in aposymbiotic control anemones. In darkness, the oxygen concentration of the coelenteric fluid was hypoxic. However, within 10 min of anemones being illuminated, coelenteric fluid was hyperoxic, and it remained elevated throughout a 12 h light period. When measured over a 24 h period (12 h light: 12 h dark), the dark respiration rate increased gradually over the first 6 h of the light period until it was 35% above the dark night-time resting rate. It remained elevated throughout the remaining light period and for 2 h into the following dark period, after which it fell back to the resting rate. Gross photosynthesis (P ^gross) increased significantly when anemones were exposed to either hyperoxia (150% oxygen saturation) or 300 E m^-2 s^-1 at normoxia. This increase was not observed when symbiotic anemones were illuminated at a low-light intensity of 100 E m^-2 s^-1. The results of this study suggest that respiration in the dark is limited by oxygen diffusion and that normal respiration is restored in the daytime by utilisation of the oxygen released by photosynthesis. Furthermore, it appears that the increased respiration following exposure to high-light intensities provides a CO₂-rich intracellular environment which further enhances the photosynthetic rate of the zooxanthellae.     

The methane mussel: roles of symbiont and host in the metabolic utilization of methane

Methane mussels (Bathymodiolus sp., undescribed; personal communication by R. Turner to CRF) were collected in September 1989 and April 1990 from offshore Louisiana in the Gulf of Mexico. These mussels contain endosymbiotic methane-oxidizing bacteria and are capable of utilizing environmental methane as a source of energy and carbon. Oxygen consumption, methane consumption, and carbon dioxide production were measured in mussels with intact symbionts, functionally aposymbiotic mussels, and separated symbiont preparations under controlled oxygen and methane conditions, in order to study the roles of the symbionts and the hosts in methane utilization. The association was found to be very efficient in fixing methane carbon (only 30% of CH₄ consumed is released as CO₂), and to be capable of maximal rates of net carbon uptake of nearly 5 mol g^-1 h^-1. Rates of oxygen and methane consumption were dependent upon oxygen and methane concentrations. Maximal consumption rates were measured at 250 to 300 M O₂ and 200 to 300 M CH₄, under which conditions, oxygen consumption by the gill tissues (containing symbionts) had increased more than 50-fold over rates measured in the absence of methane. A model is proposed for the functioning of the intact association in situ, which shows the symbiosis to be capable of achieving growth rates (net carbon assimilation) in the range of 0.003 to 0.50% per day depending upon oxygen and methane concentrations. Under the conditions measured in the seep environment (200 M O₂, 60 M CH₄), a mussel consuming methane at rates found to be typical (4 to 5 mol g^-1 h^-1) should have a net carbon assimilation rate of about 0.1% per day. We suggest that the effectiveness of this symbiosis arises through integration of the morphological and physiological characteristics inherent to each of the symbiotic partners, rather than from extensive specialization exhibited by other deep-sea chemotrophic associations.     

Metabolic and blood characteristics of the hydrothermal vent tube-worm Riftia pachyptila

Specimens of the hydrothermal vent pogonophoran Riftia pachyptila Jones were collected by submersible at a depth of 2 600 m at the 21°N hydrothermal vent site on the East Pacific Rise (20°50N, 109°06W) in April and May of 1982. The worms were maintained in pressurized aquaria for up to 45 d for metabolic studies. Consumption of O₂ was regulated down to low P_{O 2} (oxygen partial pressure) values; O₂ consumption rates were 0.63 and 1.12 mol g^-1 wet wt h^-1 at 2.5° and 8°C, respectively; such rates were comparable to those previously measured for other pogonophorans. Intact specimens of R. pachyptila (including bacterial symbionts) did not consume significant amounts of CH₄ from the environment. The respiratory quotients, in the absence of added sulfide, indicated that metabolism was mainly heterotrophic. High rates of uptake of dissolved amino acids were recorded for one specimen. The total [CO₂] in the vascular blood and the Hb-containing coelomic fluid were high. Under anaerobic conditions, there were equilibrium distributions of pH, total [CO₂] and sulfide concentrations between the vascular blood and the coelomic fluid, apparently because these metabolites were readily exchanged between the two compartments. The vascular blood bound neither CH₄ nor H₂. However, sulfide was reversibly bound by both the vascular blood and coelomic fluid; because this binding depended strongly on pH (with a maximum at about 7.5), HS^- was probably the molecular species bound. Under anaerobic, but not aerobic conditions, the trophosome bound substantial amount of sulfide; thus, the high concentrations of sulfide in the trophosome may have resulted mainly from sulfide bound to sulfide oxidases under anaerobic conditions. The coelomic fluid had a relatively low buffering capacity (2.2 mmol CO₂pH^-1).     

Experimental ecology of the temperate scleractinian coral Astrangia danae I. Partition of respiration,photosynthesis and calcification between host and symbionts

Calcification, photosynthesis and respiration of the scleractinian coral Astrangia danae were calculated from the changes in total alkalinity, pH, calculated total CO₂, and oxygen concentration produced by colonies incubated in glass jars. A correction for changes in ammonia, nitrate and nitrite was taken into account and the method evaluated. The fluxes of oxygen and CO₂ were highly correlated (r=0.99). The statistical error of alkalinity determinations was less than 10% of the changes observed in the slowest calcifying samples. Metabolism of polyparium alone was estimated by difference after removal of tissue and reincubation of bare corallum. Zooxanthellae concentration in the polyps was obtained from cell counts made on homogenates of polyp tissue. The calculated photosynthetic rate of the zooxanthellae in vivo was 25 mol O₂ (10⁸ cell)^-1 h^-1 at a light intensity of 120 Ein m^-2 s^-1. In corals having 0.5x10⁹ zooxanthellae/dm² of colony area up to 8% of the total photosynthesis was attributed to the corallum microcosm. Polyp respiration, photosynthesis, and CaCO₃ uptake rates were all much higher than rates previously reported from A. danae, apparently because in these experiments the organisms were better fed. This increased photosynthesis in turn enhanced calcification still further. The symbiosis therefore appears to provide a growth advantage even to fed corals, under the conditions of these experiments.     

Oxygen dependent sulfide detoxification in the lugwormArenicola marina

The lugwormArenicola marina L. oxidizes entering sulfide to thiosulfate. After 8 h of normoxic incubations with sulfide concentrations of 0.2 to 1.0 mmoll^-1 thiosulfate in the coelomic fluid amounted up to about 4 mmoll^-1 whereas sulfite concentrations were 100-fold lower and no accumulation of sulfate in the coelomic fluid was found. The sulfide oxidation was highly oxygen dependent. An increase of oxygen partial pressure ( ) in the medium was followed by enhanced thiosulfate production and by a decrease of sulfide concentration in the coelomic fluid. Under normoxia, the sulfide oxidation rate was sufficient to compensate the influx of sulfide into the coelomic fluid when the sulfide concentration in the medium was below 0.33 mmoll^-1. When external sulfide was raised beyond this level, sulfide up to 5 moll^-1 in the coelomic fluid appeared. Succinate in the body wall tissue was low as long as no sulfide appeared in the coelomic fluid, indicating the maintenance of an aerobic metabolism. The oxidation of sulfide to thiosulfate was localized in the mitochondria of the body wall tissue. The oxygen consumption of mitochondria was stimulated by the addition of sulfide. The mitochondrial sulfide oxidation rate depended on the amount of mitochondrial protein and followed a Michaelis-Menten kinetic. An apparentK _m of 0.68±0.29 moll^-1 and aV _max of 41.9±22.3 nmol min^-1 mg^-1 protein was calculated. Sulfide was stoichiometrically oxidized to thiosulfate with 1 mol sulfide consuming 1 mol oxygen. Sulfide oxidation was not inhibited by sulfide concentrations as high as 100 moll^-1. At low concentrations of cyanide or azide, when respiration without sulfide was already inhibited, sulfide oxidation could still be stimulated, tentatively indicating the existence of an alternative terminal oxidase. Specimens examined in the present study were collected near St. Pol de Leon, France, from 1989 to 1992.     

Physiological energetics of the intertidal sea anemone Anthopleura elegantissima

Energy budgets were calculated for individuals of the sea anemone Anthopleura elegantissima (Brandt), collected in 1981 and 1982 from Bodega Harbor, California, USA. Rates of ammonium excretion were measured in high-and low-intertidal, symbiotic and aposymbiotic sea anemones within 24 h of collection. Among symbiotic and aposymbiotic individuals, no differences in excretion rate were found on the basis of intertidal height. However, rates of ammonium excretion in aposymbiotic anemones (2.14 mol NH ₊ ⁴ g^-1 h^-1) were significantly higher than in symbiotic ones (0.288 mol NH ₊ ⁴ g^-1 h^-1). Rates of excretion were used with estimated rates of oxygen uptake to calculate nitrogen quotients (NQ). NQ and RQ values from the literature were used to calculate an oxyenthalpic equivalent [501 kJ (mol O₂)^-1 for R+U], and mass proportions of protein (54%), carbohydrate (44%) and lipid (2%) catabolized during routine metabolism in this species 24 h after feeding. Integrated energy budgets of these experimental anemones were calculated from data on ingestion, absorption and growth, and estimates of translocated energy from the symbiotic algae. Contribution of zooxanthellae to animal respiration based on translocation=90% and RQ=0.97 are 41 and 79% in high-and low-intertidal anemones, respectively. Calculated scope for growth is greater than directly measured growth in both high-and low-intertidal individuals. The deficit, estimated as 30% of assimilated energy in high-intertidal anemones, is attributed to unmeasured costs (specific dynamic effect) or production (mucus). Low-intertidal anemones lost mass during the experiment, implying that the magnitude of the deficit was greater in these anemones than in upper intertidal individuals. Anemones from both shore levels lost zooxanthellae during the experiment, which contributed to energy loss since the contribution of the zooxanthellae is greater in low-intertidal anemones. Scope for growth is preserved in high-intertidal anemones (29% of assimilated energy) because metabolic demands are lower due to aerial exposure, and prey capture rate is higher compared to lowshore anemones. Although possibly underestimated, lower scope for growth in low-shore anemones may result from continuous feeding and digestion processes that are less efficient than those of periodically feeding high-intertidal anemones.     

Uptake of glucose by bacteria in the sediment

A method is described for the incubation of undisturbed sediment cores under in situ conditions with the addition of low concentrations of ¹⁴C-glucose. Data are presented for respiration, gross uptake and actual uptake rate of glucose by bacteria in sandy, wave-washed beaches of the Baltic Sea. On the average, the bacteria respired 8% of the total glucose taken up. The gross uptake measured was between 2.3×10^-3 and 6.8×10^-3 g ¹⁴C-glucose g sediment^-1 (dry weight) h^-1 (average 4.7×10^-3 g g^-1 h^-1). Minima in the gross uptake rate corresponded with maxima in the concentration of natural free dissolved glucose. For the actual uptake rate, however, very similar uptake rates were calculated for the sediments examined (between 1.4×10^-1 and 1.9×10^-1 g glucose g^-1 h^-1, average 1.7×10^-1 g g^-1 h^-1).Publication No. 183 of the Joint Research Program at Kiel University (Sonderforschungsbereich 95 der Deutschen Forschungsgemeinschaft).     

Respiration and excretion by the ctenophore Mnepiopsis leidyi

Respiration (dissolved oxygen and carbon dioxide) and excretion (dissolved organic carbon, inorganic and organic nitrogen and phosphorus) rates were measured for a variety of sizes of Mnemiopsis leidyi over a temperature range of 10.3° to 24.5°C. Both respiration and excretion rates were a direct linear function of animal weight and very temperature sensitive (Q₁₀4). Oxygen uptake ranged from 155 to 489 g at O/(g dry weight) day^-1 and carbon dioxide release from 43 to 166 M. Organic carbon made up about 38% of the total carbon released. Inorganic nitrogen excretion, exclusively in the form of ammonium, comprised 54% of the total nitrogen release and ranged from 10 to 36 M NH₄/(g dry weight) day^-1. Average release of dissolved primary amines (expressed as glycine equivalents) equaled 43% of the organic nitrogen fraction. Inorganic phosphorus release ranged from 2.0 to 4.9 M/(g dry weight) day^-1 and made up about 72% of the total phosphorus loss. The turnover of elements in the body was calculated as 5 to 19% per day for carbon and nitrogen, depending on the temperature, and an even higher 20 to 48% per day for phosphorus. These values are comparable to rates observed for small, active zooplankton.     

The respiratory metabolism of Tilapia mossambica (Teleostei)

Tilapia mossambica (Teleostei) weighing 5 to 80 g were acclimated at 30°C to salinities of 0.4 (tap water), 12.5 (50% sea water) and 30.5 (100% sea water). Their respiration was measured at routine activity and the partial pressure of ambient oxygen gradually reduced from 250 to 50 mm Hg. Respiration is salinity-dependent; the proportionate ability to use oxygen in any one salinity is — above the critical pO₂ —the same in all experimental groups. This ability is a function of temperature and increases from 15° to 30°C, becoming temperature independent from 30° to 40°C as long as the pO₂ remains above 150 mm Hg. At 50 mm Hg pO₂, the limiting effect of oxygen causes a decrease in metabolic rate. This limiting effect is minimal in 80 g fish kept in an isotonic medium (12.5 S), allowing greater scope for activity and a higher rate of oxygen uptake.     

Contrasting effects of sulfide and thiosulfate on symbiotic CO2-assimilation of Phallodrilus leukodermatus (Annelida)

Experiments were conducted in order to specify the reductants responsible for the carbon dioxide fixation of the symbiotic sulfur bacteria in the gutless marine oligochaete Phallodrilus leukodermatus (Annelida) from shallow calcareous sediments in Bermuda. Carbon dioxide-uptake rates were suppressed by S⁼ and stimulated by S₂O₃ ⁼. Individuals which hosted bacteria containing reserve energy substances maintained a high short-term CO₂-uptake activity, while bacteria in worm homogenates and in worms treated with an antibiotic (Baypen) did not show any significant metabolic activity. Absolute uptake rates in P. leukodermatus were usually considerably higher than those reported for other animals harbouring prokaryotic sulfuroxidizing symbionts. Utilization of thiosulfate rather than sulfide is compatible with the preferred occurrence of the worms around the redox discontinuity layer and has been confirmed in other thiobiotic animals. Sulfur stored in the symbiotic bacteria appears to be oxidized to sulfate and be excreted when the worms are held under energy-limited conditions. The data emphasize the complexity of the possible metabolic pathways involved in the oxidation of reduced-sulfur compounds by bacterial symbionts in marine invertebrates.     

Effect of temperature and food on the heart beat,ventilation rate and oxygen uptake of Mytilus edulis

The heart beat, ventilation rate and oxygen uptake of Mytilus edulis L. were measured simultaneously, in response to changes in temperature and food level. There was no thermal acclimation of heart-beat frequency or amplitude to temperatures from 5° to 25°C. Oxygen consumption and ventilation rate acclimated to 10°, 15° and 20°C, but not to 25°C. Starvation reduced the rate of oxygen uptake and heart-beat frequency to a standard level and, in response to food, the ventilation rate and oxygen consumption immediately increased to an active level. Feeding was maintained after the initiation of active metabolism, and during the following 10 days the heart-beat frequency gradually increased to the level characteristic of fed individuals. There was no direct correlation between the rate of oxygen consumption and heart rate, and an apparent absence of a close nervous coupling between ventilation rate and heart rate in M. edulis.     

The role of chemosensory behavior of Symbiodinium microadriaticum,intermediate hosts,and host behavior in the infection of coelenterates and molluscs with zooxanthellae

Symbiotic dinoflagellates, Symbiodinium microadriaticum (=zooxanthellae), may gain access to aposymbiotic hosts (i.e., those lacking zooxanthellae) by chemosensory attraction of the motile algae by the potential host or via an intermediate host. Laboratory experiments showed that motile zooxanthellae were attracted to intact aposymbiotic host animals, but not to starved symbiotic hosts. Fed symbiotic hosts and brine shrimp (Artemia sp.) nauplii also attracted motile zooxanthellae. The attraction of these zooxanthellae was directly correlated with nitrogen levels in the seawater surrounding the hosts; thus ammonia and possibly nitrate could be atractants. Brine shrimp nauplii, acting as intermediate hosts actively ingested both motile and non-motile zooxanthellae. the ingested zooxanthellae tended to remain morphologically unaltered during and after passage through the gut of the brine shrimp. Capture and ingestion of brine shrimp containing zooxanthellae by aposymbiotic scyphistomae of the jellyfish Cassiopeia xamachana led to infection of the scyphistomae with zooxanthellae. Zooxanthellae isolated from 17 different species of coelenterates and molluscs could be transferred via brine shrimp to the endodermal cells of the scyphistomae. However only 10 of these isolates persisted to establish a permanent association with C. xamachana. Scyphistomae in suspensions of motile zooxanthellae responded by a classical coelenterate feeding response, which may facilitate ingestion of the potential symbionts and establishment of a symbiosis.     

Fluctuations and interactions of bacterial activity in sandy beach sediments and overlying waters

Fluctuations and interactions of the following microbiological variables and sediment properties were investigated on samples from sandy beaches of the Baltic Sea: bacterial number and biomass, net uptake and respiration rate of ¹⁴C-glucose (U), concentration of natural free dissolved glucose and fructose, actual uptake rate and turnover time of glucose, sand-grain size and shape, water and organic matter content of the sediment. Spearman rank correlation analysis demonstrated significant relationships between cell number, biomass and actual uptake rate of glucose. The concentration of natural glucose varied with sand-grain shape, and the uptake rates of glucose were inversely correlated with the water content of the sediment. In the overlying water, cell number and/or biomass were significantly correlated with both concentration and uptake rates of glucose. Partial correlation analysis, however, indicates that, in the water overlying the sediment at least one of the standing crop variables (bacterial number or biomass) is independently variable with uptake activity. The sediment bacterial standing crop does not necessarily reflect metabolic activity. Various significant interactions were noted between mainly sediment properties (water content, organic matter content, grain size) and variables in the water above (cell number, biomass, concentration and uptake rates of glucose).Publication No. 206 of the Joint Research Program at Kiel University (Sonderfor-schungsbereich 95 der Deutschen Forschungsgemeinschaft).     

Carbon content,dark respiration and mortality of the mixotrophic planktonic ciliate Strombidium capitatum

Carbon content and rate of dark respiration was determined on individual Strombidium capitatum (Leegard) Kahl cells uniformly labelled with ¹⁴C in culture. Isolated individuals were incubated in sterile medium in the dark for periods of up to 24h, and cumulative respired ¹⁴CO₂ was retained in an alkaline trap. Cell carbon varied by more than an order of magnitude and followed a bi-modal distribution. Small cells of 2 to 7 ng C cell^-1 respired at specific rates of 3 to 5% cell C h^-1, whereas large cells of 7 to 25 ng C cell^-1 respired at 1 to 2% cell C h^-1. Mortality was greater for small cells than for large ones, and was greatest during the first few hours. Small cells accounted for 40% of all cells at initial time, T ₀, whereas none of these survived a 16 h incubation. It is proposed that the rates of carbon-specific dark respiration observed for small cells compromise their ability to survive more than a few hours in the dark without food. The combination of influence of size (carbon content) together with differential proportions of small cells resulting from mortality contributed to considerable variance in carbon-specific respiration rates. When smaller and larger modal groups were considered separately, this variance was significantly reduced for both groups. Using this refined data, there was no significant starvation-induced reduction in carbon-specific rates. The mean rate remained at between 1.1 and 1.4% cell C h^-1 for large cells over the 24 h period, and between 3.8 and 4.1% cell C h^-1 for small cells over the 8 h of their survival. This observation for a planktonic mixotrophic ciliate contrasts with published observations for heterotrophic protists which have reported reduction of carbon-specific respiration rate with starvation.     

Photosynthesis-irradiance responses and photosynthetic periodicity in the sea anemone Aiptasia pulchella and its zooxanthellae

Sea anemones (Aiptasia pulchella) containing zooxanthellae (Symbiodinium microadriaticum) were maintained in a long-term laboratory culture on a 12 h light (100 E m^-2 s^-1):12 h dark cycle. Photosynthetic oxygen production was measured for the symbiotic association and for freshlyisolated zooxanthellae. Light utilization efficiencies () were similar for both sets of zooxanthellae, suggesting negligible shading of zooxanthellae by animal tissue in this association. Whereas freshly-isolated zooxanthellae were photoinhibited at high irradiances (800 to 1 800 E m^-2 s^-1), zooxanthellae in the host continued to function at photosynthetic capacity. Time of day may influence photosynthetic measurements in symbiotic organisms, as it was found that photosynthesis in A. pulchella followed a diel periodicity at both light-saturating (1 200 E m^-2 s^-1) and subsaturating (150 E m^-2 s^-1) irradiances. There was a peak period of photosynthesis between 12.00 and 14.00 hrs. Light stimulated dark respiration rates of A. pulchella. Dark respiration of sea anemones increased somewhat towards the end of the light cycle and was always greater after exposure to high irradiances.     

The nature of the symbiosis between Indo-Pacific anemone fishes and sea anemones

Under the general heading of symbiosis, defined originally to mean a living together of two dissimilar species, exist the sub-categories of mutualism (where both partners benefit), commensalism (where one partner benefits and the other is neutral) and parasitism (where one partner benefits and the other is harmed). The sea anemone-fish (mainly of the genus Amphiprion) symbiosis has generally been considered to benefit only the fish, and thus has been called commensal in nature. Recent field and laboratory observations, however, suggest that this symbiosis more closely approaches mutualism in which both partners benefit to some degree. The fishes benefit by receiving protection from predators among the nematocyst-laden tentacles of the sea anemone host, perhaps by receiving some form of tactile stimulation, by being less susceptible to various diseases and by feeding on anemone tissue, prey, waste material and perhaps crustacean symbionts. The sea anemones benefit by receiving protection from various predators, removal of necrotic tissue, perhaps some form of tactile stimulation, removal of inorganic and organic material from on and around the anemone, possible removal of anemone parasites, and by being provided food by some species of Amphiprion.     

Chemical composition,metabolic rates and feeding behavior of the midwater ctenophore Bathocyroe fosteri

Individuals of the midwater ctenophore Bathocyroe fosteri (0.01 to 1.6 g dry weight, DW) were collected from Bahamian waters by the submersible Johnson-Sea-Link during May and September/October 1983 and October/November 1984 from 530 to 700 m depth. Metabolic rates were measured and showed oxygen consumption to be in the range of 0.01 to 0.18 mg O₂ g^-1 DW h^-1 at temperatures ranging from 9° to 12°C. Ammonium excretion (0.01 to 0.14 g-at N g^-1 DW h^-1) was typically low. Energy expenditures estimated from respiration data (ca. 7% body C d^-1) indicated that one to three midwater crustacean prey (ca. 150 g C d^-1) could provide the daily maintenance ration required by a 40 mm ctenophore. These metabolic characteristics complemented in situ observations of poor locomotor ability and passive feeding behavior.     

Shallow-water benthic and pelagic metabolism:

Pelagic primary production and benthic and pelagic aerobic metabolism were measured monthly at one site in the estuarine plume region of the nearshore continental shelf in the Georgia Bight. Benthic and water-column oxygen uptake were routinely measured and supplemented with seasonal measures of total carbon dioxide flux. Average respiratory quotients were 1.18:1 and 1.02:1 for the benthos and water column, respectively. Benthic oxygen uptake ranged from 1.23 to 3.41 g O₂ m^-2 d^-1 and totalled 756 g O₂ m^-2 over an annual period. Water column respiration accounted for 60% of total system metabolism. Turnover rates of organic carbon in sediment and the water column were 0.09 to 0.18 yr^-1 and 6.2 yr^-1, respectively. Resuspension appeared to control the relative amounts of organic carbon, as well as the sites and rates of organic matter degradation in the benthos and water column. Most of the seasonal variation in benthic and pelagic respiration could be explained primarily by temperature and secondarily by primary productivity. On an annual basis, the shelf ecosystem appeared to be heterotrophic; primary production was 73% of community metabolism, which was 749 g C m^-2 yr^-1. The timing of heterotrophic periods through the year appeared to be closely related to both river discharge and the periodicity of growth and death of marsh macrophytes in the adjacent estuary. The results of this study support the estuarine outwelling hypothesis of Odum (1968).This is Contribution No. 530 from the University of Georgia Marine Institute. This work was supported by the Georgia Sea Grant College Program maintained by the National Oceanic and Atmospheric Administration, US Department of Commerce