Le système assimilation-régénération des sels nutritifs dans les eaux superficielles de l'océan Austral

Low primary-productivity levels in high-nutrient environments of the Southern Ocean are not yet well understood. An accurate knowledge of nutrient assimilation by phytoplankton, at the base of the pelagic food-web, therefore appears to be essential. A study of the mesoscale hydrological structure and chemical observations in the upper layers of the Indian sector of the Antarctic Ocean (Cruise MD-25 FIBEX of M.S. Marion Dufresne, January–February 1981) permitted us to develop a method for estimating nutrient consumption by phytoplankton. After strong vertical mixing in winter, the homogeneous Antarctic Surface Water (ASW) is, in summer, divided into two parts by a well-formed pycnocline. In the upper layer, corresponding approximately to the photic zone, we observed nutrient depletion resulting from photosynthetic activity, while the underlaying thermal minimum layer displayed unaltered winter characteristics, including nutrient concentrations typical of winter surface-water conditions. Taking into account the nutrient depletion in summer, we calculated the assimilation ratios for Antarctic phytoplankton as follows: ([NO ₃ ^- lower]-[NO ₃ ^- upper]):([PO ₄ upper])=N:P=11.2±2.4 and N:Si=0.27±0.05. These ratios are lower than the classical Redfield ratios (N:P=16 and N:Si=1), but are in good agreement with the elementary composition of phytoplankton samples collected during the same cruise. Moreover, they agree with previously published data on stoechiometric determinations of particulate matter in surface-water samples. Besides, in the transition layer between the lower layer and Warm Deep Water (WDW), the calculated mineralization ratios were slightly higher than the assimilation ratios: N:P=14.6 and N:Si=0.37. From these ratios, it would appear that, in these surface layers of the Southern Ocean (down to 250 m), assimilation-regeneration mechanisms operate in such a way that phosphate and silicate contents decrease much more than nitrate content during the northward drift of the surface waters. The determination of assimilation ratios taking into account the nutrient depletion of the upper layer allowed us to evaluate the carbon net-production which integrates time-space variations in the photic layer over the whole early summer period. We estimated a production rate (0.4 g C m^-2 d^-1) of the same order of magnitude as the mean value obtained by the ¹⁴C method (0.2 g C m^-2 d^-1). The theoretic silicium consumption by phytoplankton along a meridian transect, calculated using an estimated assimilation ratio of C:Si=1,85, indicates that the marked south-north decrease in silicate concentration in the Antarctic Surface Water would mainly result from biological activity. The silicate concentrations thus calculated were in good agreement with concentrations measured along a previous transect across the Southern Ocean.

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Le système assimilation-régénération des sels nutritifs dans les eaux superficielles de l'océan Austral

     

Oxygen-binding by haemocyanins from an ecological series of amphipod crustaceans

The oxygen-binding properties of haemocyanins (Hc) from three species of gammaridean amphipods, Gammarus locusta (L.) (subtidal), Echinogammarus pirloti (Sexton and Spooner), (intertidal, marine) and E. marinus (Leach) (intertidal, estuarine), one species of hyalid amphipod Hyale nilsonni Rathke (high intertidal, marine) and the talitrid amphipod Orchestia gammarellus (Pallas) (semi-terrestrial) have been studied. All the species were collected from the Firth of Clyde, Scotland, during the spring of 1992. The oxygen-carrying capacity of haemolymph from each species was low, although variable, and was correlated with the low concentration of Hc present. The Hc oxygen-affinity of native gammarid haemolymph was relatively high [partial pressure of oxygen required for half-saturation, P₅₀=4 to 5 torr (0.53 to 0.67 kPa)] at their respective in vivo pH values. At equivalent pH, however, Hc from G. locusta displayed a lower O₂-affinity than either Echinogammarus species. Gammarid Hcs had a large Bohr effect ( log P₅₀/ pH=-1.16 to-1.47). Resuspended Hc isolated from whole H. nilsonni showed similar O₂-binding properties to those of the gammaridean amphipods [P₅₀=6.3 torr (1.44 kPa) at pH=8.0; log P₅₀/pH=-1.20]. Comparable data for haemolymph from O. gammarellus showed that the Hc had a lower affinity for O₂ [P₅₀=14.1 torr (1.87 kPa) at in vivo pH] and exhibited a more moderate Bohr effect ( log P₅₀/ pH=-0.79). To eliminate the possibility that these differences were due to the different haemolymph constituents, each of the Hcs were pelleted and resuspended in physiological saline. The differences noted above persisted, demonstrating that they were due to inherent O₂-binding properties of the Hc molecules themselves. An increase in L-lactate resulted in an increase in Hc oxygen-affinity for both Echinogammarus species but not for O. gammarellus. This study has confirmed that there is a clear difference between Hcs from aquatic and semi-terrestrial amphipod genera. The results lend further support to the hypothesis that the move on to land by amphipod crustaceans is accompanied by a decrease in Hc oxygen-affinity, a decrease in the Bohr effect and a decrease in effector (in this case L-lactate) sensitivity.     

Photosynthetic characteristics of nanoplankton and picoplankton from the surface mixed layer

Nanoplankton and picoplankton primary production has been studied at two oceanic stations in the Porcupine Sea-bight and at one shelf station in the Celtic Sea. At both sites, low wind conditions in June and July 1985 resulted in greatly reduced vertical turbulent mixing and a secondary, temporary thermocline developed in what is usually a well-mixed surface layer; as a result, there was physical separation of the phytoplankton within two zones of the surface mixed layer. The photosynthetic characteristics of three size fractions (>5 m, <5 to >1 m and <1 to >0.2 m) of phytoplankton populations from the two zones have been measured. Phytoplankton was more abundant at the oceanic stations and chlorophyll a values were between 1.3 and 2.2 mg chlorophyll a m^-3, compared with 0.3 to 0.6 mg chlorophyll a m^-3 at the shelf station; at both stations, numbers of cyanobacteria were slightly higher in the lower zone of the surface mixed layer. There was no effect of the temporary thermocline on the vertical profiles of primary production and most phtosynthesis occurred in the surface 10 m. Photosynthetic parameters of the three size fractions of phytoplankton have been determined; there was considerable day-to-day variation in the measured photosynthetic parameters. Assimilation number (P _m ^B ) of all >5 m phytoplankton was lower for the deeper than for the surface populations, but there was little change in initial slope (a ^B ). The small oceanic nanoplankton (<5 to >1 m) showed changes similar to the >5 m phytoplankton, but the same size fraction from the shelf station showed changes that were more like those shown by the picoplankton (<1 m) viz, little change in P _m ^B but an increase in a ^B with depth. Values of a ^B were generally greater for the picoplankton fraction than for the larger phytoplankton, but values of adaptation parameter (I _k )(=P _m ^B /) were not always less. There was little evidence to support the hypothesis that these populations of picoplankton were significantly more adapted to low light conditions than the larger phytoplankton cells. When photosynthetic parameters of the picoplankton were normalised to cell number (P _m ^C /a ^C ) rather than chlorophyll a, P _m ^C was comparable to other published data for picoplankton, but a ^C was much lower. The maximum doubling time of the picoplankton at saturating irradiance is calculated to be ca. 8.5 h for the oceanic population and ca. 6.2 h for the shelf population.     

Dietary effects on oocyte yolk-composition inCapitella sp. I (Annelida: Polychaeta): Fatty acids and sterols

In 1988, immatureCapitella sp. I (initial biomass 80µg dry wt) were raised on four diets: Gerber cereal, TetraMin fish food,Ulva sp., and benthic diatoms. After 2 wk of culturing different populations on these diets, eggs were dissected from gravid females, frozen and analyzed for fatty acid and sterol composition. Eggs produced by worms on different food types were discriminated by fatty acid composition in a principal component analysis (PCA), with 18:1⁹, 18:2, 20:5 and 20:4 fatty acids showing the greatest differences between experimental groups. The sterol profiles of all eggs were dominated by cholesterol (cholest-5-en-3-ol) and cholest-5,24-dien-3-ol (>60%). A PCA of egg sterols discriminated between adult diets with cholesterol, 23,24-dimethyl-cholesta-5-en-3-ol, cholest-5,24-dien-3-ol and a C-29 stenol showing the greatest differences. In field populations ofCapitella sp. I, oocytes produced at different times of the year may have different levels of lipids depending on their dietary availability during vitellogenesis. Variations in oocyte composition may influence larval growth and development and thus have an impact on population dynamics. Alternatively, variations in fatty acid and sterol composition of oocytes may allow us to identify the food resources utilized by this species.     

Primary production as influenced by diel periodicity of phytoplankton photosynthesis

Diel periodicity in parameters of photosynthesis-irradiance (P-I) curves was incorporated into calculations of integral daily phytoplankton production for the Santa Barbara Channel off southern California (USA). Model equations of the relationship between photosynthesis and light were used in combination with observed slope () and asymptote (P _max) values presented in the preceding paper. Primary production was always 19 to 39% less than comparable estimates obtained with the assumption of constant maximum daily and P _max values. Regardless of which P-I formulation was used of 6 tested, observed production (using a temporal series of simulated in situ incubations) ranged from 13% less to 25% more than estimates from constant midday and P _max values. The amplitude and timing of diel oscillations differed somewhat among 3 field stations. Maximum to minimum ratios ranged from approximately 3 to 5 for , and 4 to 6 for P _max. The differences in amplitude and timing of oscillations in P-I curves both contributed to errors in calculating phytoplankton production. Thus, photosynthetic periodicity in the upwelling area of the Santa Barbara Channel influences phytoplankton production. There were oscillations in both and P _max, and the time-dependence of these parameters should be considered to improve the accuracy of predictive models of primary productivity.     

Importance of the fatty acids 20:5ω 3 and 22:6ω 3 in the diet of plaice (Pleuronectes platessa) larvae

The requirement of plaice (Pleuronectes platessa L.) larvae for the fatty acids 20:5 3 and 22:6 3 was studied. Larvae were reared from first feeding to beyond metamorphosis on Artemia sp. nauplii (EG brand), whose nutritional content had been manipulated by enrichment. Some larvae were fed Artemia rich in 20:5 3 and 22:6 3, and others were fed nauplii with no 22:6 3 and low levels of 20:5 3 (1.9% of the total fatty acids). The differences in diet had no significant difference on growth or survival of the plaice larvae. The results indicate that the plaice larvae in this study did not require 22:6 3 in their diet, even when the levels of 20:5 3 were low.     

Ammonium enhancement of dark carbon fixation and nitrogen limitation in zooxanthellae symbiotic with the reef coralsMadracis mirabilis andMontastrea annularis

The nutrient status (limitation vs sufficiency) of dinoflagellates (zooxanthellae) symbiotic with reef corals in Bermuda was assessed in 1989 and 1990 by measuring the enhancement of dark carbon fixation with 20 M ammonium by isolated symbionts. A colony ofMadracis mirabilis was kept in the laboratory and fed daily or starved for one month. Symbionts from fed portions of the colony had ammonium-enhancement ratios (NH _4dark ⁺ ; SW_dark;SW=seawater without added ammonium) similar to those of the original field population (1.2 to 1.3). Ammonium-enhancement ratios increased with starvation of the host (x1.7) as did values forV _D:V _L [(ammonium dark rate-seawater dark rate): light rate in seawater]. Both parameters indicated decreasing nitrogen sufficiency of the algae when the host was not fed, but starvation appeared to affect these algae less than symbionts of sea anemones. Field samples of zooxanthellae fromM. mirabilis (Three Hill Shoals and Bailey's Bay Flats) yielded results similar to those for fed corals, but those taken from Bailey's Bay Flats in May 1990 yielded exceptionally high values for enhancement (>3) andV _D:V _L indicating pronounced nitrogen limitation at the time of sampling. We sampled zooxanthellae from populations ofMontastrea annularis at 8 m (Three Hill Shoals) and 24 m (Soldier's Point) depths. Enhancement andV _D:V _L values for zooxanthellae from the 8 m corals were density-dependent: symbionts from corals with normal symbiont densities displayed the most nitrogen limitation (enhancement values=1.4 to 2.0), while those from bleached corals with lower density exhibited enhancement andV _D:V _L values typical of nitrogen-sufficient algae. Symbionts isolated from the 25 m corals yielded the highest values, and appeared to exhibit the least nitrogen-sufficiency for this species.     

The influence of irradiance on the biochemical composition of three phytoplankton species and their nutritional value for larvae of the Pacific Oyster (Crassostrea gigas)

Three species of phytoplankton grown at high (HL) or low light (LL) were fed as saturating rations to laboratory-reared larval Crassostrea gigas. Larval C. gigas fed diets of HL grown Chaetoceros gracilis and HL grown Isochrysis aff. galbana grew faster than those fed LL grown cells of the same phytoplankton species. Faster growth of C. gigas larvae was consistently associated with increases in the percent composition of short chain saturated fatty acids (FA) 14:0+16:0 in the HL grown cells. There were no consistent and significant differences between HL and LL grown phytoplankton cells in their content of carbon, nitrogen, protein, lipid or carbohydrate. Intraspecific increases in percent composition of essential fatty acids (EFAs), 20:53 and 22:63, in the phytoplankton were not associated with improvements in the growth or survival of the oyster larvae. Oyster larvae fed diets of Phaeodactylum tricornutum with a relatively high proportion of EFAs grew more slowly than those fed C. gracilis. In this experiment the proportion of dietary EFA 20:53 was negatively correlated with oyster growth rates. The faster growing oyster larvae contained relatively more of the FAs 14:0+16:0 which may be useful as measures of larval oyster condition. After a diet of one phytoplankton species for ca. 10 d, oyster larvae acquired distinctive FA profiles resembling that of their phytoplankton prey.     

Activity and metabolism of larval Atlantic cod (Gadus morhua) from Scotian Shelf and Newfoundland source populations

Patterns of activity and metabolism were investigated in larval Atlantic cod (Gadus morhua L.) between December 1991 and July 1992: (1) throughout larval development; (2) between two genetically discrete populations (Scotian Shelf and Newfoundland) and (3) as a function of two different culture temperatures. During the yolk-sac stage (0 to 5 d post-hatch), changes in swimming speed were not related to mass-specific metabolic rates; no portion of the mass-specific oxygen consumption could be explained by changes in activity. In the mixed feeding stage (6 to 14 d posthatch), there was a tendency for oxygen consumption to be related to changes in swimming speed. In the exogenous feeding stage (>14 d post-hatch), oxygen consumption significantly increased with swimming speed. These ontogenetic patterns of activity and metabolism were the same for larvae from the Scotian Shelf and Newfoundland populations. However, over the entire larval life and among ontogenetic stages, the metabolic cost of activity (mass-specific O₂ consumption/swimming speed) of Scotian Shelf larvae was significantly higher than that of Newfoundland larvae. When cod larvae, that had developed at 5°C, were acutely exposed to 10°C, Scotian Shelf larvae had a higher intrinsic cost of activity than Newfoundland larvae, over the entire larval life. During the exogenous feeding stage, the mean metabolic cost of activity for Newfoundland larvae raised at 10°C and tested at 10°C was significantly higher and more variable than that of larvae raised at lower temperatures. However, the metabolic cost of activity of larvae raised and tested at 10°C was not significantly different between source populations. Together these findings suggest that differences in swimming energetics reflect changing energy requirements for activity among ontogenetic stages, and reflect adaptation to regional environments among genetically discrete populations.     

Chlorophyll a fluorescence,an alternative method for estimating primary production

The in vivo chlorophyll a fluorescence index (F_+DCMU-F_-DCMU/F_+DCMU) of natural waters was compared to the ¹⁴C-determined primary production, and the fluorescence intensity in the presence of 3-(3,4-dichlorophenyl)-1,1-dimethylurea (F_+DCMU) was studied as a function of extracted and spectrophotometrically determined chlorophyll concentrations. Samples were taken every second week from May through October, 1979, at the station Systrarna situated in a coastal area of the Bottnian Sea. In addition, samples from the Archipelago Sea of the Baltic were collected on board the Finnish research vessel R/S Aranda during the September cruise 1979. The correlations between the fluorescence index and the ¹⁴C-determined primary production and between F_+DCMU and total chlorophyll concentration were good when samples taken over short time intervals were compared. The shortcomings of both the fluorescence and the ¹⁴C methods are discussed. It is concluded that the fluorescence method is useful if it is desirable to follow with high time resolution any changes in the potential for photosynthesis (or primary production) in a water mass over relatively short time periods; e.g. during an algal bloom. The fluorescence method can furthermore be technically developed for automatic monitoring with a high time resolution. Efforts are being made in our laboratory to develop the method further to give information about the in situ rates of photosynthesis rather than the potential for photosynthesis in a photoplankton population.     

Diel oscillations of the photosynthesis-irradiance (P-I) relationship in natural assemblages of phytoplankton

Diel oscillations in the photosynthesis-irradiance (P-I) relationship are described for marine phytoplankton assemblages at 6 stations in an upwelling area off the southern California coast (USA) between May and August 1980. The initial slope () and asymptote (P _max) of P-I curves changed significantly over the day; both parameters were in phase and had similar changes in amplitude. The diel oscillations in photosynthesis appeared unrelated to changes in chlorophyll a concentrations. Amplitudes of daily variations in photosynthesis ranged from approximately 3 to 9, as measured by the maximum to minimum ratio for photosynthetic capacity (P _max). Diatom-rich samples collected during an upwelling event and those dominated by dinoflagellates both had midday to early afternoon maxima in and P _max. Samples from other locations had peak photosynthetic activity later in the afternoon. The relationship between and P _max was consistent in all phytoplankton samples analyzed, with a surprisingly high correlation considering the spatial and temporal scales encompassed in this study. These results indicate that the photosynthesis-irradiance (P-I) relationship is time-dependent and, moreover, that changes in and P _max are closely coupled for a variety of natural phytoplankton assemblages.     

Sterols in coral-reef animals

The sterol constituents of both coral and non-coral-reef animals (mollusks, coelenterates, echinoderms) have been investigated, and determined by gas-liquid chromatography and spectral analyses. All coral-reef animals examined contained larger amounts of C₂₈-sterols, such as 24-methylcholesterol, than non-coral-reef animals. The seastar Acanthaster planci contains the same sterol constituents as some coral-reef coelenterates and mollusks, differing only in the position of the double bond in a steroid ring. This observation suggests the transport of dietary ⁵-sterols to ⁷-sterols through the food-chain in A. planci.     

Chitinolytic enzymes in the integument and midgut-gland of the shrimpPalaemon serratus during the moulting cycle

Chitinase andN-acetyl--D-glucosaminidase activity were quantified inPalaemon serratus (Pennant) integument and midgut gland during the moulting cycle. Studies were performed on specimens collected near Concarneau, France, in July 1989. The changes in specific activity are different in the two organs: in the midgut gland enzymatic activity is high throughout the whole moulting cycle with a weak peak at the early premoult Stage D1, whereas in the integument the activity of both enzymes is very low throughout post- and intermoult stages and rises only after premoult Stage D1. The highest specific activity is reached in D1 for chitinase and somewhat earlier (D1) forN-acetyl--D-glucosaminidase. The increase in specific chitinolytic activity coincides with an increase in ecdysteroids.     

Seasonal photosynthetic characteristics of Ascoseira mirabilis (Ascoseirales,Phaeophyceae) from King George Island,Antarctica

Monthly variation in photosynthesis, dark respiration, chlorophyll a content and carbon: nitrogen (C:N) ratios in different lamina sections of adult plants of Ascoseira mirabilis Skottsberg from King George Island, Antarctica, was investigated between September 1993 and February 1994. Light saturated net photosynthesis (P _max) showed maximum values in September (12 to 25 mol O₂ g^-1 fr wt h^-1), and decreased towards the summer to values ranging between 2.0 and 5.0 mol O₂ g^-1. In the distal section, however, a second optimum occurred in December (25 mol O₂ g^-1 fr wt h^-1). Dark respiration rates were also highest in October and November and decreased strongly in December to February (6.0 and 1.0 mol O₂ g^-1 fr wt h^-1, respectively). Gross photosynthesis exhibited high values between September and December. Concomitant with the seasonal decrease of photosynthetic efficiency () from mean values of 1.2 mol O₂ g^-1 fr wt h^-1 (mol photons cm^-2 s^-1)^-1 in September to 0.3 mol O₂ g^-1 fr wt h^-1 (mol photons cm^-2 s^-1)^-1 in January, the initial light saturating point (I _k) gradually increased from 19 to 60 mol photons m^-2 s^-1. Likewise C:N ratios were low in spring (12 to 13) and increased in summer (20). In general, the photosynthetic parameters P _max, gross photosynthesis, and Chl a concentrations were significantly higher in the distal section of the thallus. In contrast, C:N ratios were lower in the distal section of the lamina. The results show that photosynthesis obviously strongly supports growth of the alga in late winter to spring, as it does in some morphologically related brown algae from temperate and polar regions. The question whether growth is additionally powered     

A Modelling Study of Transient,Buoyancy-Driven Exchange Flow over a Descending Barrier

Results are presented from a series of model studies of the transient exchange flow resulting from the steady descent of an impermeable barrier separating initially-quiescent fresh and saline water bodies having density ₀ and ₀+()₀, respectively. A set of parametric laboratory experiments has been carried out (i) to determine the characteristic features of the time-dependent exchange flow over the barrier crest and (ii) to quantify the temporal increase in the thickness and spatial extent of the brackish water reservoir formed behind the barrier by the outflowing, partly-mixed saline water. The results of the laboratory experiments have been compared with the predictions of a theoretical model adapted from the steady, so-called maximal exchange flow case and good qualitative agreement between theory and experiment has been demonstrated. The comparisons indicate that head losses of between 7% and 3% are applicable to the flow over the ridge crest in the early and late stages, respectively, of the barrier descent phase, with these losses being attributed to mixing processes associated with the counterflowing layers of fresh and saline water in the vicinity of the ridge crest. The experimental data show (and the theoretical model predictions confirm) that (i) the dimensionless time of detection t _det(g/H _b)^1/2 of the brackish water pool fed by the dense outflow increases (at a given distance from the barrier) with increasing values of the descent rate parameter g'H _b/(dh _b/dt)² and (ii) the normalised thickness (x,t)/H _b of the pool at a given reference station increases monotonically with increasing values of the modified time (t–t _det)/(H _b/g)^1/2, with the rate of thickening decreasing with increasing values of the descent rate parameter g'H _b(dh _b/dt)². Here, g=(g/₀)()₀ is the modified gravitational acceleration, H _b is the mean depth of the water and dh _b/dt denotes the rate of descent of the barrier height h _b with elapsed time t after the two water bodies are first brought into contact.     

Photosynthetic characteristics of marine phytoplankton from contrasting physical environments

Two suites of phytoplankton samples have been collected in consecutive years at various times over a day from selected depths within vertically mixed and stratified water columns in the western Irish Sea, in order to provide a range of possible light histories within the populations collected. Values for the maximum rate of ¹⁴C retention (P _max) and the initial slope of the ¹⁴C retention: light intensity curve () were obtained. Supra-thermocline samples from the stratified water exhibited higher P _max values than corresponding subthermocline samples. Higher values of were also generally associated with samples from the supra-thermocline zone of the stratified region. Differences in the depth distribution of P _max and in the mixed water were small, except in the presence of a shallow thermocline. In one suite of samples from the stratified water, a diurnal increase in the P _max values of the supra-thermocline samples was observed. P _max values obtained from the samples from the mixed water were interpreted in relation to the distribution obtained from the samples from the stratified zone. Data from both the contrasting sites visited for one sample suite demonstrated a two-phase relationship between the chlorophyll a concentration and both P _max and . The rates of ¹⁴C retention of the first suite of samples were estimated by two techniques. The average differences in the retention were greater in samples from the sub-as opposed to suprathermocline zone. No trends were apparent in the smaples from the mixed waters.     

Influence of irradiance on the nutritional value of two phytoplankton species fed to larval Japanese scallops (Patinopecten yessoensis)

Japanese scallop (Patinopectin yessoensis Jay) larvae grew faster and were larger after 18 d when fed a diet of high-light(HL)-grown Chaetoceros simplex or HL Pavlova lutheri relative to diets of the same phytoplankton species grown at low light (LL). When provided as saturating rations to larval scallop, these diets could be ranked: HL C. simplex>LL C. simplex>HL P. lutheri>LL P. lutheri. In both phytoplankton species, HL-grown cells contained more of the short-chain saturated fatty acid (FA), 16:0 than LL-grown cells. Scallop growth rates were a significant function of the amounts (mg g^-1 dry wt) and the proportions (as percentage of total FAs) of the FAs 14:0 and 14:0+16:0 (total saturated FAs) in their diet. The proximate biochemical composition of HL- versus LL-grown phytoplankton showed no significant differences in protein, total lipid, carbon, carbohydrate or nitrogen per cell which were consistently associated with the greater nutritional value of HL cells. In spite of this high variability in proximate composition, the larval growth rate was a significant function of the average carbon content, nitrogen content and cell volume of the phytoplankton cells. Increased amounts of the essential polyunsaturated FAs 20:5 3 and 22:6 3 in the phytoplankton were negatively correlated with larval scallop growth rates. Thus HL-grown phytoplankton cells were nutritionally superior to LL-grown cells. This nutritional superiority seems to be determined by the fatty acid composition of the cells which, in turn, is controlled by variation in irradiance. The general tendency of predator FA profiles to resemble that of their prey was not observed in larvae fed P. lutheri. The much greater amounts of 18:4 3, 20:5 3, and 22:6 3 FA in P. lutheri relative to C. simplex were not evident in the scallop larvae fed these cells.     

Incorporation and utilization of bacterial lipids in theSolemya velum symbiosis

We undertook a detailed analysis of the lipid composition ofSolemya velum (Say), a bivalve containing endosymbiotic chemoautotrophic bacteria, in order to determine the presence of lipid biomarkers of endosymbiont activity. The symbiont-free clamMya arenaria (L.) and the sulfur-oxidizing bacteriumThiomicrospira crunogena (Jannasch et al.) were analyzed for comparative purposes. The ¹³C ratios of the fatty acids and sterols were also measured to elucidate potential carbon sources for the lipids of each bivalve species. Both fatty acid and sterol composition differed markedly between the two bivalves. The lipids ofS. velum were characterized by large amounts of 18: 17 (cis-vaccenic acid), 16:0, and 16 : 17 fatty acids, and low concentrations of the highly unsaturated plant-derived fatty acids characteristic of most marine bivalves. Cholest-5-en-3-ol (cholesterol) accounted for greater than 95% of the sterols inS. velum. In contrast,M. arenaria had fatty acid and sterol compositions similar to typical marine bivalves and was characterized by large amounts of the highly unsaturated fatty acids 20 : 53 and 22 : 63 and a variety of plant-derived sterols. The fatty acids ofT. crunogena were similar to those ofS. velum and were dominated by 18:17, 16:0 and 16:17 fatty acids. Thecis-vaccenic acid found inS. velum is almost certainly symbiontderived and serves as a potential biomarker for symbiontlipid incorporation by the host. The high concentrations ofcis-vaccenic acid (up to 35% of the total fatty acid content) in both symbiont-containing and symbiont-free tissues ofS. velum demonstrate the importance of the endosymbionts in the lipid metabolism of this bivalve. The presence ofcis-vaccenic acid in all the major lipid classes ofS. velum demonstrates both incorporation and utilization of this compound. The ¹³C ratios of the fatty acids and sterols ofS. velum were significantly lighter (–38.4 to –45.3) than those ofM. arenaria (–23.8 to – 24.2) and were similar to the values found for the fatty acids ofT. crunogena (–45); this suggests that the lipids ofS. velum are either derived directly from the endosymbionts or are synthesized using endosymbiontderived carbon.Woods Hole Oceanographic Institution Contribution No. 7356Please address all correspondence and reprint requests to Dr Conway at her present address: Department of Biological Sciences, University of Pittsburgh, Pennsylvania 15260, USA     

Interactions between nutritional status and long-term responses to ultraviolet-B radiation stress in a marine diatom

Influences of nutritional status on the photoinhibitory effects of ultraviolet-B radiation (UVBR: 290 to 320 nm) on the specific growth rates (_obs) and biomass of Phaeodactylum tricornutum were determined using nutrient-replete batch cultures and nutrient-limited continuous cultures. P. tricornutum cultures were exposed to UVBR doses representative of current mid-latitude and ozone-depletion intensities. Specific growth rates and biomass were inhibited from 2 to 16% by UVBR during nutrient-replete growth. However, no effect of UVBR on _obs or biomass was detectable when nutrient limitation exceeded the potential for limitation by UVBR. Thus, a competitive interaction appears to occur between macronutrient stress and UVBR stress, such that _obs and biomass will be determined by the most limiting factor. Our results suggest that measurable decreases in phytoplankton _obs and biomass from UVBR are most likely in nutrient-rich areas of the ocean, while these parameters may not be appropriate for measuring UVBR stress in regions of nutrient limitation.     

Availability of colloidal ferric oxides to coastal marine phytoplankton

Cell growth of a coastal marine diatom, Phaeodactylum tricornutum (stock cultures), and two red tide marine flagellates, Heterosigma akashiwo and Gymnodinium mikimotoi (stock cultures), in the presence of soluble chelated Fe(III)-EDTA (1:2) and of four different phases of ferric oxide colloids were experimentally measured in culture experiments at 20°C under 3000 lux fluorescent light. Soluble Fe(III)-EDTA induced the maximal growth rates and cell yields. The short-term uptake rate of iron by H. akashiwo in Fe(III)-EDTA medium was about eight times faster than that in solid amorphous hydrous ferric oxide (Fe₂O₃·xH₂O) medium. In culture experiments supplied with four different ferric oxide forms, the orders of cell yields are amorphous hydrous ferric oxide>-FeOOH (lepidocrocite)>Fe₅O₇(OH)·4H₂O (hydrated ferric oxyhydroxide polymer >-FeOOH (goethite). The specific growth rates () at logarithmic growth phase in Fe(III)-EDTA, amorphous hydrous ferric oxide and -FeOOH media were significantly greater than those in Fe₅O₇ (OH)·4H₂O and -FeOOH media. The thermodynamically stable forms such as Fe₅O₇(OH)·4H₂O and -FeOOH supported a little or no phytoplankton growth. The iron solublities and/or proton-promoted iron dissolution rates of these colloidal ferric oxides in seawater at 20°C were determined by simple filtration techniques involving -activity measurements of ⁵⁹Fe. The orders of solubilities and estimated dissolution rate constants of these ferric oxides in seawater were consistent with that of cell yields in the culture experiments. These results suggest that the availability of colloidal iron to provide a source of iron for phytoplankton is related to the thermodynamic stability and kinetic lability of the colloidal ferric oxide phases, which probably control the uptake rate of iron by phytoplankton.