Respiration,photosynthesis and carbon metabolism in planktonic ciliates

Release of¹⁴C-labelled carbon dioxide from uniformly labelled cells was used to measure respiration by individual ciliates in 2-h incubations in 1989 and 1990. In a strictly heterotrophic ciliate,Strobilidium spiralis (Leegaard, 1915), release of labelled carbon dioxide was equivalent to ca. 2.8% of cell C h^–1 at 20°C, and there was no difference between rates in the dark and light. In the chloroplast-retaining ciliatesLaboea strobila Lohmann, 1908,Strombidium conicum (Lohmann, 1908) Wulff, 1919 andStrombidium capitatum (Leegaard, 1915) Kahl, 1932, release of labelled carbon dioxide was less in the light than in the dark in experiments done at 15°C. InL. strobila release of radiolabel as carbon dioxide was equivalent to ca. 2.4% of cell C h^–1 in the dark but ca. 1% at 50µE m^–2 s^–1, an irradiance limiting to photosynthesis. InS. conicum release of radiolabel as carbon dioxide was equivalent to ca. 4.4% of cell C h^–1 in the dark, but at an irradiance saturating to photosynthesis (250 to 300µE m^–2 s^–1) there was no detectable release of labelled carbon dioxide. InS. capitatum release of radiolabel as carbon dioxide was equivalent to ca. 4.3% of cell C h^–1 in the dark but at an irradiance saturating to photosynthesis was ca. 2.4% of cell C h^–1. These data, combined with data from photosynthetic uptake experiments, indicate that¹⁴C uptake underestimates the total benefit of photosynthesis by 50% or more in chloroplastretaining ciliates.Contribution no. 7510 from the Woods Hole Oceanographic Institution     

Growth patterns of Codium fragile ssp. tomentosoides in response to temperature,irradiance, salinity,and nitrogen source

Seasonal patterns of growth, reproduction, and productivity of Codium fragile spp. tomentosoides (van Goor) Silva were monitored at 3 locations in Rhode Island. Maximal growth occurred during the summer and was more significantly correlated with temperature than any other factor measured in this study. Multiple correlation models suggested an interaction between temperature, irradiance, and available nitrogen. Maximal reproduction occurred in late summer and early fall. The maximal productivity, based on harvested quadrats, was 2. 10 g dry weight m^-2 day^-1. A large amount (up to 87.3%) of the annual production entered the detrital food chain during the winter by fragmentation of the thallus. Culture studies examined the effects of temperature (6° to 30°C), irradiance (7 to 140 E m^-2 sec^-1), daylength (8 h light: 16 h dark to 24 h light: O h dark) and salinity (6 to 48) on growth. Differentiated thalli grew over a broad range of experimental conditions, with maximal growth at 24°C, 24 to 30 S, a minimal irradiance of 28 E m^-2 sec^-1, and 16 h daylength. The effect of increasing daylength was due to increased total daily irradiance rather than to a true photoperiodic effect. Undifferentiated sporelings survived and grew in a narrower range of environmental conditions than thalli. c. fragile spp. tomentosoides grew equally well with nitrate, nitrite, ammonium, and urea as a nitrogen source. The addition of NaHCO₃ stimulated growth at levels of 2.4 to 4.8 mM, suggesting an inorganic carbon limitation in static cultures. This study supports the hypothesis that the in situ seasonal growth pattern of c. fragile spp. tomentosoides is primarily due to the interaction of temperature and irradiance.     

Seasonal and depth related variation in the photosynthesis–irradiance response of<Emphasis Type="Italic"> Ecklonia radiata</Emphasis> (Phaeophyta,Laminariales) at West Island,South Australia

The photosynthesis–irradiance response of Ecklonia radiata (C. Agardh) J. Agardh, a common kelp in the temperate southern hemisphere, was investigated in situ throughout the year and across a depth profile at West Island, South Australia. Temperature and irradiance environment altered throughout the year, varying at 3 m between 14–20°C and 279–705 mol photons m^–2 s^–1. Photosynthetic capacity (P_m) varied throughout the year between 177–278 mol O₂ g^–1 dry wt h^–1 at 3 m and 133–348 mol O₂ g^–1 dry wt h^–1 at 10 m. The irradiance required for sub-saturation of photosynthesis (E_k) varied between 97–152 and 81–142 mol photons m^–2 s^–1 for 3 m and 10 m respectively, and the respiration rate varied between 15–36 and 13–20 mol O₂ g^–1 dry wt h^–1 for 3 m and 10 m. A clear seasonal change in photokinetic parameters was detected and provided strong evidence for a seasonal acclimation response. During winter an increase in the efficiency of light utilisation at low irradiance () was accompanied by a decrease in both E_k and that required for photosynthetic compensation. P_m also increased during the winter and autumn months and respiratory requirements decreased. These changes enable E. radiata to display an optimal photosynthetic performance throughout the year despite significant changes in the surrounding environment.Communicated by P.W. Sammarco, Chauvin     

Growth and herbivory by heterotrophic dinoflagellates in the Southern Ocean,studied by microcosm experiments

Growth and herbivory of heterotrophic dinoflagellates (Gymnodinium sp.) from the Weddell Sea and the Weddell/Scotia Confluence were studied in 1988 in 100-liter microcosms. The microcosms were screened through 200-µm or 20-µm mesh nets and incubated for 12 d at 1 °C under artificial light. Mean cell volume of dinoflagellates was 1 000 to 1 500µm³, and that of their phytoplankton prey 360 to 430µm³. Dinoflagellate growth rate followed a Holling type II functional response, with a maximum growth rate of 0.3 d^–1 and half-saturation food concentrations of 1.0µg chlorophylla l^–1, 50µg C l^–1, or 1 500 cells ml^–1. Carbon budgets based on¹⁴CO₂ assimilation and biomasses of phytoplankton and heterotrophic dinoflagellates suggested a balance between phytoplankton grazing loss and dinoflagellate consumption, assuming a dinoflagellate carbon conversion efficiency of 40%. Applying this to the functional response yielded estimates of maximum ingestion rate (0.8µg Cµg^–1 C d^–1, or 6 pg C dinoflagellate^–1 h^–1) and maximum clearance (0.8 to 1.2 × 10⁵ body volumes h^–1, or 80 to 120 nl ind.^–1 h^–1). The microcosm experiments suggested that heterotrophic dinoflagellates may contribute significantly to maintenance of low phytoplankton biomass in the Southern Ocean.     

Respiration rates of<Emphasis Type="Italic"> Beroe ovata</Emphasis> in the Black Sea

Metabolic rates of the ctenophore Beroe ovata within the length range from 0.4 mm (newly hatched larvae) to 60 mm were investigated. At 20° the respiration rates (Q, µg O₂ ind.^–1 h^–1) of individuals with wet weights (W, mg) less than or greater than 100 mg changed according to the equations Q=0.093W^0.62 and Q=0.016W^0.99, respectively. The weight-specific respiration rate of the juvenile ctenophores with wet body weights of 0.021–100 mg diminished approximately 20-fold (from 0.35 to 0.017 µg O₂ mg^–1 h^–1, respectively), but did not change within the range from 100 to 30,000 mg. The difference in the slope of the regression lines seems to be attributable to the ontogenetic changes in B. ovata metabolism. For the tested temperature range of 10–28°, the mean Q₁₀ coefficient was equal to 2.17±0.5. The basal metabolism of B. ovata narcotised by chloral hydrate was 4.5±0.9 times lower than total metabolism. Such a metabolic range is considered to be characteristic of aquatic invertebrates with high levels of locomotory activity.Communicated by O. Kinne, Oldendorf/Luhe     

Energy content at metamorphosis and growth rate of the early juvenile barnacle<Emphasis Type="Italic"> Balanus amphitrite</Emphasis>

The energetic cost of metamorphosis in cyprids of the barnacle Balanus amphitrite Darwin was estimated by quantification of lipid, carbohydrate and protein contents. About 38–58% (4–5 mJ individual^–1) of cypris energy reserves were used during metamorphosis. Lipids accounted for 55–65%, proteins for 34–44% and carbohydrates for <2% of the energy used. Juveniles obtained from larvae fed 10⁶ cells ml^–1 of Chaetoceros gracilis were bigger (carapace length: 560–616 µm) and contained more energy (5.56±0.10 mJ juvenile^–1) than their counterparts (carapace length: 420–462 µm; energy content: 2.49±0.20 mJ juvenile^–1) obtained from larvae fed 10⁴ cells ml^–1. At water temperatures of 30°C and 24°C and food concentrations of 10⁴ and 10² cells ml^–1 (3:1 mixture of C. gracilis and Isochrysis galbana) as well as under field conditions (26.9±3.1°C and 2.2±0.8 µg chlorophyll a l^–1), juveniles obtained from larvae fed the high food concentration grew faster than juveniles obtained from larvae fed low food concentration until 5 days post-metamorphosis. Laboratory experiments revealed a combined effect of early juvenile energy content, temperature and food concentration on growth until 5 days post-metamorphosis. After 10 days post-metamorphosis, the influence of the early juvenile energy content on growth became negligible. Overall, our results indicate that the energy content at metamorphosis is of critical importance for initial growth of juvenile barnacles and emphasize the dependency of the physiological performance of early juvenile barnacles on the larval exposure to food.Communicated by O. Kinne, Oldendorf/LuheAn erratum to this article can be found at     

Energetic cost of development through metamorphosis for the seastar<Emphasis Type="Italic"> Mediaster aequalis</Emphasis> (Stimpson)

Rates of oxygen consumption were measured for embryos, larvae and juveniles of the seastar Mediaster aequalis for 76 days post-fertilization. The rate increased from 0.65 nmol O₂ ind^–1 h^–1 at 6 h after fertilization to 2.8 nmol O₂ ind^–1 h^–1 at day 35. Larvae became competent to metamorphose around day 35 post-fertilization and began to decrease their metabolic rate after this time. Metamorphosed juveniles consumed 0.74 nmol O₂ ind^–1 h^–1. Eggs contained 138.6 µg lipid ind^–1 and 12.1 µg protein ind^–1. Lipid levels decreased in concentration throughout development while protein levels increased slightly. The lipid levels decreased by 88.5 µg from eggs to day 76 larvae, accounting for 3.5 J of energy. Total oxygen consumption to this point was 3.74 µmol O₂ ind^–1, accounting for 1.84 J. The energetic demand up to day 76 was met completely through the use of lipid reserves. Metamorphosed juveniles expended 0.5 J more than larvae at the same age. Tubes of the polychaete Phyllochaetopterus prolifica were able to induce metamorphosis in M. aequalis larvae and a non-polar extract of these tubes also triggered metamorphosis. Larvae that are delayed to metamorphose can sustain their metabolic rate with lipid reserves for a limited, yet undetermined, period.Communicated by P.W. Sammarco, Chauvin     

Photosynthetic characteristics of Prochloron sp./ascidian symbioses

The prokaryotic green alga Prochloron sp. (Prochlorophyta) is found in symbiotic association with colonial didemnid ascidians that inhabit warm tropical waters in a broad range of light environments. We sought to determine the light-adaptation features of this alga in relation to the natural light environments in which the symbioses are found, and to characterize the temperature sensitivity of photosynthesis and respiration of Prochloron sp. in order to assess its physiological role in the productivity and distribution of the symbiosis. Colonies of the host ascidian Lissoclinum patella were collected from exposed and shaded habitats in a shallow lagoon in Palau, West Caroline Islands, during February and March, 1983. Some colonies from the two light habitats were maintained under conditions of high light (2 200 E m^–2 s^–1) and low light (400 E m^–2 s^–1) in running seawater tanks. The environments were characterized in terms of daily light quantum fluxes, daily periods of light-saturated photosynthesis (H_sat), and photon flux density levels. Prochloron sp. cells were isolated from the hosts and examined for their photosynthesis vs irradiance relationships, respiration, pigment content and photosynthetic unit features. In addition, daily P:R ratios, photosynthetic quotients, carbon balances and photosynthetic carbon release were also characterized. It was found that Prochloron sp. cells from low-light colonies possessed lower chlorophyll a/b ratios, larger photosynthetic units sizes based on both reaction I and reaction II, similar numbers of reaction center I and reaction center II per cell, lower respiration levels, and lower P_max values than cells from high-light colonies. Cells isolated from low-light colonies showed photoinhibition of P_max at photon flux densities above 800 E m^–2 s^–1. However, because the host tissue attenuates about 60 to 80% of the incident irradiance, it is unlikely that these cells are normally photoinhibited in hospite. Collectively, the light-adaptation features of Prochloron sp. were more similar to those of eukaryotic algae and vascular plant chloroplasts than to those of cyanobacteria, and the responses were more sensitive to the daily flux of photosynthetic quantum than to photon flux density per se. Calculation of daily minimum carbon balances indicated that, though high-light cells had daily P:R ratios of 1.0 compared to 4.6 for low-light cells, the cells from the two different light environments showed nearly identical daily carbon gains. Cells isolated from high-light colonies released between 15 and 20% of their photosynthetically-fixed carbon, levels sufficient to be important in the nutrition of the host. Q₁₀ responses of photosynthesis and respiration in Prochloron sp. cells exposed briefly (15–45 min) to temperatures between 15° and 45°C revealed a discontinuity in the photosynthetic response at the ambient growth temperatures. The photosynthetic rates were found to be more than twice as sensitive to temperatures below ambient (Q₁₀=3.47) than to temperatures above ambient (Q₁₀=1.47). The Q₁₀ for respiration was constant (Q₁₀=1.66) over the temperature range examined. It appears that the photosynthetic temperature sensitivity of Prochloron sp. may restrict its distribution to warmer tropical waters. The ecological implications of these findings are discussed in relation to published data on other symbiotic systems and free-living algae.     

Response of the microbial food web to manipulation of nutrients and grazers in the oligotrophic Gulf of Aqaba and northern Red Sea

The control mechanisms within the pelagic microbial food web of the oligotrophic Gulf of Aqaba and the northern Red Sea were investigated in the spring of 1999. Nutrient conditions and potential grazer impact were manipulated in a series of dilution experiments. Ambient nutrient concentrations and autotrophic biomass were very low (0.23–1.21 µmol NO₃ l^–1, 0.06–0.98 µmol NH₄ l^–1, 1.08–1.17 µmol Si l^–1, 0.08–0.12 µmol P l^–1, 0.15–0.36 µg chlorophyll a l^–1). The planktonic community was characterized by low abundances [3.0–5.5×10⁵ heterotrophic bacteria ml^–1, 0.58–7.2×10³ ultraphytoplankton <8 µm ml^–1 (small eukaryotic photoautotrophs and Prochlorococcus sp., excluding Synechococcus sp.), 0.45–4.4×10⁴ Synechococcus sp. ml^–1, 0.32–1.2×10³ heterotrophic nanoflagellates ml^–1, 1.3–3.8×10³ phytoplankton >8 µm l^–1, 0.93–5.4×10² microzooplankton l^–1] and dominated by small forms (0.2–8 µm). Dinoflagellates and oligotrichous ciliates were the most common groups in initial samples among the phytoplankton >8 µm and microzooplankton, respectively. Results show that bottom-up and top-down control mechanisms operated simultaneously. Small organisms were vulnerable to grazing, with maximum grazing rates of 1.1 day^–1 on heterotrophic bacteria and 1.3 day^–1 on ultraphytoplankton. In contrast, algae >8 µm showed stronger signs of nutrient limitation, especially when the final assemblages were dominated by diatoms. Synechococcus sp. were not grazed and only showed moderate to no response to nutrient additions. The high spatial and temporal variation of our results indicates that the composition of the planktonic community determines the prevailing control mechanisms. It further implies that, at this transitional time of the year (onset of summer stratification), the populations fluctuate about an equilibrium between growth and grazing.Communicated by O. Kinne, Oldendorf/Luhe     

Interactive accumulation of mercury and selenium in the sea starAsterias rubens

Interactions between mercury and selenium accumulation and subcellular binding inAsterias rubens (L.), collected in 1987 from Lille Bælt at Middelfart, Funen, Denmark, were investigated in laboratory experiments. Sea stars exposed to 10µg Hg l^–1 for 30 d accumulated mercury in body wall, tube feet and stomach linearly with time at 1.2, 1.2 and 0.5µg Hg g^–1 dry wt d^–1, respectively. Mercury was accumulated in pyloric caeca and coelomic fluid initially at 1.4µg Hg g^–1 dry wt d^–1 and 9.4 ng Hg ml^–1 d^–1, respectively; after 10 d uptake rates decreased. Sea stars exposed to 75µg Se-SeO ₃ ^{- -} l^–1 accumulated selenium linearly with time over 30 d in the stomach, pyloric caeca, tube feet and body wall at 2.0, 1.2, 1.2 and 0.6µg Se g^–1 dry wt d^–1. Sea stars exposed to 75µg Se-SeO ₄ ^{- -} l^–1 maintained selenium levels in the coelomic fluid at 75µg Se l^–1 over 30 d. Exposure to selenate did not alter the selenium concentrations in the tissues. Sea stars exposed concurrently to 75µg Se-SeO ₃ ^{- -} and 10µg Hg l^–1 accumulated more mercury and selenium in tube feet and body wall than did sea stars exposed to the two elements alone. In pyloric caeca and stomach concurrent exposure reduced accumulation of both elements. Mercury was bound predominantly in the insoluble fraction of the tissues, and soluble mercury was bound in proteins of high (> 70 kilodaltons) or very low (< 6000 daltons) molecular weight. Ca. half of the selenium recovered was bound in the insoluble fraction, and soluble selenium was bound in proteins of high (> 70 kilodaltons) or very low (< 6000 daltons) molecular weight. Interaction between the two elements was exerted predominantly in the insoluble fraction of the tissues.     

Diurnal gut pigment rhythm and metabolic rate of<Emphasis Type="Italic"> Calanus euxinus</Emphasis> in the Black Sea

The vertical distribution, diel gut pigment content and oxygen consumption of Calanus euxinus were studied in April and September 1995 in the Black Sea. Gut pigment content of C. euxinus females was associated with diel vertical migration of the individuals, and it varied with depth and time. Highest gut pigment content was observed during the nighttime, when females were in the chlorophyll a (chl a) rich surface waters, but significant feeding also occurred in the deep layer. Gut pigment content throughout the water column varied from 0.8 to 22.0 ng pigment female^–1 in April and from 0.2 to 21 ng pigment female^–1 in September 1995. From the diel vertical migration pattern, it was estimated that female C. euxinus spend 7.5 h day^–1 in April and 10.5 h day^–1 in September in the chl a rich surface waters. Daily consumption by female C. euxinus in chl a rich surface waters was estimated by taking into account the feeding duration and gut pigment concentrations. Daily carbon rations of female C. euxinus, derived from herbivorous feeding in the euphotic zone, ranged from 6% to 11% of their body carbon weight in April and from 15% to 35% in September. Oxygen consumption rates of female and copepodite stage V (CV) C. euxinus were measured at different temperatures and at different oxygen concentrations. Oxygen consumption rates at oxygen-saturated concentration ranged from an average of 0.67 g O₂ mg^–1 dry weight (DW) h^–1 at 5°C to 2.1 g O₂ mg^–1 DW h^–1 at 23°C for females, and ranged from 0.48 g O₂ mg^–1 DW h^–1 at 5°C to 1.5 g O₂ mg^–1 DW h^–1 at 23°C for CVs. The rate of oxygen consumption at 16°C varied from 0.62 g O₂ mg^–1 DW h^–1 at 0.65 mg O₂ l^–1 to 1.57 g O₂ mg^–1 DW h^–1 at 4.35 mg O₂ l^–1 for CVs, and from 0.74 g O₂ mg^–1 DW h^–1 at 0.57 mg O₂ l^–1 to 2.24 g O₂ mg^–1 DW h^–1 at 4.37 mg O₂ l^–1 for females. From the oxygen consumption rates, daily requirements for the routine metabolism of females were estimated, and our results indicate that the herbivorous daily ration was sufficient to meet the routine metabolic requirements of female C. euxinus in April and September in the Black Sea.Communicated by O. Kinne, Oldendorf/Luhe     

Nitrate-reductase activity and in vivo nitrate-reduction rate in Ulva rigida illuminated by blue light

In the marine green alga Ulva rigida C. Agardh, nitrate reductase (NR) is synergetically induced by blue light and nitrate. The present study examines the effect of blue light and a large NO ₃ ^– pulse (0.3 mM) on relevant variables of NO ₃ ^– -assimilation such as NO ₃ ^– -uptake, intracellular NO ₃ ^– -storage, NR activity, in vivo NO ₃ ^– -reduction rate and NO ₂ ^– and NH ₄ ⁺ -accumulation. Nitrate uptake started immediately upon addition of NO ₃ ^– , suggesting the presence of a constitutive carrier, however in the first 1.5 to 2 h, periods of net NO ₃ ^– efflux were frequent. After this time, NO ₃ ^– -uptake and intracellular NO ₃ ^– -accumulation proceeded linearly with time, suggesting the existence of a different NO ₃ ^– -uptake mechanism, which seems to be inducible. Our results indicate that in vivo NO ₃ ^– -reduction is not exclusively dependent on the potential NR activity. In U. rigida, during the first 2 h after a NO ₃ ^– pulse (300 M) there were clear indications that the induction state of the NO ₃ ^– -carrier limits the reduction rate of NO ₃ ^– . Once the induction of the NO ₃ ^– -transporter had been completed (1.5 to 2 h), the NO ₃ ^– -assimilation pathway reached a steady state, NO ₃ ^– -uptake rate, NO ₃ ^– -reduction rate and NO ₂ ^– and NH ₄ ⁺ -accumulation being linear with time. Since the reduction of NO ₃ ^– leads mainly to the accumulation of NH ₄ ⁺ , we conclude that, after the NO ₃ ^– -reduction itself, NH ₄ ⁺ -fixation into carbon skeletons is the limiting step in the assimilation of NO ₃ ^– by U. rigida under blue light.     

Daily energy requirements and trophic positioning of the sand shrimp<Emphasis Type="Italic"> Crangon septemspinosa</Emphasis>

Sand shrimp, Crangon septemspinosa Say, are important to the trophic dynamics of coastal systems in the northwestern Atlantic. To evaluate predatory impacts of sand shrimp, daily energy requirements (J ind.^–1 day^–1) were calculated for this species from laboratory estimates of energy losses due to routine (R_R), active (R_A), and feeding (R_SDA) oxygen consumption rates (J ind.^–1 h^–1), coupled with measurements of diel motile activity. Shrimp used in this study were collected biweekly from the Niantic River, Connecticut (41°33N; 72°19W) during late spring and summer of 2000 and 2001. The rates of shrimp energy loss due to R_R and R_A increased exponentially with increasing temperature, with the magnitude of increase greater between 6°C and 10°C (Q₁₀=3.01) than between 10°C and 14°C (Q₁₀=2.85). Rates of R_R doubled with a twofold increase in shrimp mass, and R_SDA was 0.130 J h^–1+R_R, irrespective of shrimp body size. Shrimp motile activity was significantly greater during dark periods relative to light periods, indicating nocturnal behavior. Nocturnal activity also increased significantly at higher temperatures, and at 20°C shifted from a unimodal to a bimodal pattern. Laboratory estimates of daily metabolic expenditures (1.7–307.4 J ind.^–1 day^–1 for 0.05 and 1.5 g wet weight shrimp, respectively, between 0°C and 20°C) were combined with results from previous investigations to construct a bioenergetic model and make inferences regarding the trophic positioning of C. septemspinosa. Bioenergetic model estimates indicated that juvenile and adult shrimp could meet daily energy demands via opportunistic omnivory, selectively preying upon items of high energy content (e.g. invertebrate and fish tissue) and compensating for limited prey availability by ingesting readily accessible lower energy food (e.g. detritus and plant material).Electronic Supplementary Material Supplementary material is available in the online version of this article at Communicated by J.P. Grassle, New Brunswick     

Ammonium excretion by the ophiuridOphiothrix fragilis as a function of season and tide

Ammonium excretion of a dense population (~1 500 individuals m^–2) of the ophiuridOphiothrix fragilis (Abildgaard) was measured in the Dover Straits (French coast) between May 1989 and March 1990: the excretion rate varied from 4.8 µg N g^–1 dry wt h^–1 in November to 12.8 µg N g^–1 dry wt h^–1 in June. Mean individual ammonium excretion,E, wasE=0.019t +1.26 (whereE=µg N individual^–1 andt=time in min;r=0.80;N=81). Variations in the ammonium excretion rate during a tidal cycle appeared to arise from variations in the duration of the suspension-feeding activity ofO. fragilis, which was governed by the strength of the tidal current. During short-term starvation, excretion was low (E=0.009t+1.47;r=0.91;N=17), increasing with increasing length of starvation [E=4.62lnt–2.5;r=0.95;N=17], as observed for other echinoderms; this could be due to catabolism of tissue. The daily ammonia flux from thisO. fragilis population to the water column was estimated at 41 mg N m^–2 d^–1.     

The influence of macrofauna on estuarine benthic community metabolism: a microcosm study

Effects of benthic macrofauna (Corophium volutator, Hydrobia sp., Nereis virens) on benthic community metabolism were studied over a 65-d period in microcosms kept in either light/dark cycle (L/D-system) or in continuous darkness (D-system). Sediment and animals were collected in January 1986 in the shallow mesohaline estuary, Norsminde Fjord, Denmark. The primary production in the L/D-system after 10 d acted as a stabilizing agent on the O₂ and CO₂ flux rates, whereas the D-system showed decreasing O₂ and CO₂ flux throughout the period. Mean O₂ uptake over the experimental period ranged from 0.38 to 1.24 mmol m^–2 h^–1 and CO₂ release varied from 0.80 to 1.63 mmol m^–2 h^–1 in both systems. The presence of macrofauna stimulated community respiration rates measured in darknes, 1.4 to 3.0 and 0.9 to 2.0 times for O₂ and CO₂, respectively. In contrast, macrofauna lowered primary production. Gross primary production varied from 1.06 to 2.26 mmol O₂ m^–2 h^–1 and from 1.26 to 2.62 mmol CO₂ m^–2 h^–1. The community respiratory quotient (CRQ, CO₂/O₂) was generally higher in the begining of the experiment (0–20 d, mean 1.89) than in the period from Days 20 to 65 (mean 1.38). The L/D-system exhibited lower CRQ (ca. 1) than the D-system. The community photosynthetic quotient varied for both net and gross primary production from 0.64 to 1.03, mean 0.81. The heterotrophic D-system revealed a sharp decrease in the sediment content of chlorophyll a as compared to the initial content. In the autotrophic L/D-system, a significant increase in chlorophyll a concentration was observed in cores lacking animals and cores with C. volutator (The latter species died during the experiment). Due to grazing and other macrofauna activities other cores of the L/D-system exhibited no significant change in chlorophyll a concentration. Community primary production was linearly correlated to the chlorophyll a content in the 0 to 0.5 cm layer. Fluxes of DIN (NH₄ ⁺+NO₂ ^–+NO₃ ^–) did not reveal significant temporal changes during the experiment. Highest rates were found for the cores containing animals, mainly because of an increased NH₄ ⁺ flux. The release of DIN decreased significantly due to uptake by benthic microalgae in the L/D-system. No effects of the added macrofauna were found on particulate organic carbon (POC), particulate organic nitrogen (PON), total carbon dioxide (TCO₂) and NH₄ ⁺ in the sediment. The ratio between POC and PON was nearly constant (9.69) in all sediment dephts. The relationship between TCO₂ and NH₄ ⁺ was more complex, with ratios below 2 cm depth similar to those for POC/PON, but with low ratios (3.46) at the sediment surface.     

Photosynthetic performance of Laminaria solidungula measured in situ in the Alaskan High Arctic

Photosynthetic performance in the kelp Laminaria solidungula J. Agardh was examined from photosynthesis irradiance (P-I) parameters calculated from in situ ¹⁴C uptake experiments, using whole plants in the Stefansson Sound Boulder Patch, Alaskan Beaufort Sea, in August 1986. Rates of carbon fixation were determined from meristematic, basal blade, and second blade tissue in young and adult sporophytes. Differences in saturating irradiance (I _k, measured as photosynthetically active radiation, PAR), photosynthetic capacity (P _max), and relative quantum efficiency () were observed both between young and adult plants and between different tissue types. I _k was lowest in meristematic tissue (20 to 30 E m^–2 s^–1) for both young and adult plants, but consistently 8 to 10 E m^–2 s^–1 higher in young plants compared to adults in all three tissues. Average I _k for non-meristematic tissue in adult plants was 38 E m^–2 s^–1. Under saturating irradiances, young and adult plants exhibited similar rates of carbon fixation on an area basis, but under light limitation, fixation rates were highest in adult plants for all tissues. P _max was generally highest in the basal blade and lowest in meristematic tissue. Photosynthetic efficiency () ranged between 0.016 and 0.027 mol C cm^–2 h^–1/E m^–2 s^–1, and was highest in meristematic tissue. The relatively lower I _k and higher exhibited by L. solidungula in comparison to other kelp species are distinct adaptations to the near absence of light during the eight-month ice-covered period and in summer when water turbidity is high. Continuous measurement of in situ quantum irradiance made in summer showed that maximum PAR can be less than 12 E m^–2 s^–1 for several days when high wind velocities increase water turbulence and decrease water transparency.The Univeristy of Texas Marine Science Institute Contribution No. 695     

Annual abundance and biomass of aloricate ciliates in tropical neritic waters off Kingston,Jamaica

The daily abundance of aloricate ciliates at Lime Cay, Jamaica, a shallow neritic site, ranged from 29 to 118 × 10⁶ m^–2 (0.97 to 3.93 × 10⁶ m^–3) between November 1985 and November 1986. Biomass was converted to kilojoules (1 kcal=4.1855 kJ) assuming 42% carbon, 20.15 kJ (g dry wt)^–1, and 20% cell shrinkage. Biomass ranged from 0.40 to 3.00 kJ m^–2 (13.3 to 100 J m^–3; 0.28 to 2.08µg C l^–1) with an annual mean of 1.11 kJ m^–2 (36.8 J m^–3; 0.764µg C l^–1). Nanociliates (<20µm equivalent spherical diameter, ESD) dominated abundance, but microciliates (> 20µm ESD) dominated biomass.Strombidium, Strobilidium, Tontonia andLaboea species were conspicuous taxa. Annual production estimates of the aloricate assemblage, based on literature growth rates, ranged from 404 kJ m^–2 yr^–1 (37 J m^–3 d^–1) to 1614 kJ m^–2 yr^–1 (147 J m^–3 d^–1). A compromise estimate of 689 kJ m^–2 yr^–1 (i.e., 63 J m^–3 d^–1) is comparable to other estimates from tropical and subtropical regions. A model of annual energy flow through 11 planktonic compartments suggests the total ciliate assemblage (aloricates and tintinnines) to be as productive as metazoan herbivores and metazoan carnivores.     

Effect of elevated temperature on aerobic respiration of coral recruits

Metabolic rates provide a valuable means to assess the condition of early life stages of scleractinians, but their small biomass creates a signal-to-noise problem in a confined respirometer. To avoid this problem, measurements of the oxygen diffusion boundary layer (DBL) and Ficks first law were used to calculate the respiration rate of coenosarc tissue on recruits (i.e., colonies 5–14 mm diameter) of Porites lutea (Edwards and Haime, 1860) exposed to two temperatures at a flow speed of 0.6 cm s^–1. All experiments were completed in Moorea, French Polynesia, between November and December 2003. At 26.8°C, the DBL was 565±55 µm thick, the oxygen saturation adjacent to the tissue was 80±3%, and the mean respiration of the coenosarc was 1.2±0.1 µl O₂ cm^–2 h^–1 (all values mean ± SE, n=10). Exposure to 29.7°C for 24–48 h did not affect the DBL thickness but significantly reduced the oxygen saturation adjacent to the tissue (to 74%) and increased the mean respiration rate by 35%. As the small corals differed slightly in size, in a uniform flow speed they experienced dissimilar flow environments as characterized by the Reynolds number (Re), thereby creating the opportunity to test the flow dependency of respiration. At 26.8°C, respiration and Re were unrelated, but at 29.7°C, the relationship was positive and statistically significant. Thus, respiration of small corals may not be mass transfer limited at low temperature, but relatively small increases in temperature may result in an increased metabolic rate leading to mass transfer limitation and flow-dependent rates of respiration.Communicated by J.P. Grassle, New Brunswick     

Photosynthetic carbon acquisition in the red algaGracilaria conferta

Photosynthetic properties of the common red algaGracilaria conferta, collected from the eastern Mediterranean Sea were investigated in 1989, in order to begin evaluating its adaptative strategies with regard to the inorganic carbon composition of seawater, and to test whether the alleged C₄ photosynthesis of anotherGracilaria species is common within the genus. Net photosynthetic rates ofG. conferta were, under ambient conditions of inorganic carbon (ca. 10µM, CO₂ and 2.2 mM HCO ₃ ^- ), not sensitive to O₂ over the range 10 to 300µM, and the CO₂ compensation point was low (ca. 0.005µM). Ribulose-1,5-bisphosphate carboxylase/oxygenase was the major carboxylating enzyme, with a crude extract activity of 175µmol CO₂ g^–1 fresh wt h^–1 while phosphoenolpyruvate carboxylase and phosphoenolpyruvate carboxykinase were present at 70 and 20%, respectively, of that activity. No activities of the decarboxylases NAD-and NADP-malic enzyme could be detected. The¹⁴C pulse-chase incorporation pattern showed thatG. conferta fixes inorganic carbon via the photosynthetic carbon reduction cycle only, with no evidence for photosynthetic C₄ acid metabolism. Photosynthesis at the natural seawater pH of 8.2 was, at 25°C and saturating light, saturated at the ambient inorganic carbon concentration of 2.5 mM. It is proposed that, under ambient inorganic carbon conditions, a CO₂ concentrating system other than C₄ metabolism provides an internal CO₂ concentration sufficient to suppress the O₂ effect on ribulose-1,5-bisphosphate carboxylase/oxygenase and, thus, on photorespiration, in a medium where the external free CO₂ concentration is lower than theK _m(CO₂) of the carboxylating enzyme. Since inorganic carbon, under natural saturating light conditions, seems not to be a limiting factor for photosynthesis ofG. conferta, it likely follows that other nutrients limit the growth of this alga in nature.     

Egg production ofCalanus finmarchicus at low temperature

Reproduction ofCalanus finmarchicus Gunnerus collected in June 1988 in Polar water and in April 1989 in Atlantic water was studied. Single females were kept at 0°C in the laboratory for 22 d (Polar) and 77 d (Atlantic) with superabundant food concentration (> 400µg Cl^–1) of the diatomThalassiosira antarctica. There was no significant difference between the two populations, although more spent females were found in Polar water, probably due to the different dates of collection. The hypothesis of low temperature determining the geographic range ofC. finmarchicus via reproductive failure is not supported. Mean daily egg production rate of all females from Atlantic water over a 60 d period was 24.4, corresponding to 5.5% body C female^–1 d^–1, when an egg carbon content of 0.23µg is assumed. Coefficient of variation was 25%. Maximum values were 53.2 eggs female^–1 d^–1, corresponding to 12.1% body C d^–1. The highest number of eggs spawned by a single female was 3101, corresponding to a seven-fold turnover of body C during the investigation period; >20% of females produced > 2000 eggs. Body carbon content did not change significantly during the experiment; the C:N ratio increased slightly, indicating lipid accumulation. Delay of response to starvation periods of 2, 4 and 7 d duration was always 2 d: egg production ceased 2 d after the onset of starvation and continued 2 d after onset of feeding.