Photosynthetic performance of giant clams, Tridacna maxima and T. squamosa, Red Sea

Two species of giant clams, Tridacna maxima and T. squamosa, coexist in the Red Sea, but exhibit distinctly different depth distributions: T. maxima mostly occurs in shallow waters (reef flat and edge), while T. squamosa may occur down to the lower fore-reef slope. Giant clams have been described as mixotrophic, capable of both filter-feeding and photosynthesis due to algal symbionts (zooxanthellae), therefore, observed depth preferences were investigated in relation to possible differences in autotrophy vs. heterotrophy. This study was conducted from April to June 2004, at the reef near the Marine Science Station, Aqaba, Gulf of Aqaba, Red Sea, and in May 2007, at a reef near Dahab, Sinai Peninsula, Egypt. In situ measurements using a submersible pulse amplitude modulated fluorometer (Diving PAM), revealed no significant differences in effective PSII quantum yield (ΔF/Fm′) and relative electron transport rates (ETR) between the two species; but rapid light curves (ETR vs. light, photosynthetically active irradiance, PAR) showed significant differences in maximum photosynthetic rates (ETR_max), with 20% higher values in T. maxima. Chamber incubations displayed higher net and gross oxygen production by T. maxima (88.0 and 120.3 μmol O₂ cm⁻² mantle area day⁻¹) than T. squamosa (56.7 and 84.8 μmol O₂ cm⁻² mantle area day⁻¹); even under shading conditions (simulated depth of 20 m) T. maxima still achieved 93% of the surface gross O₂ production, whereas T. squamosa reached only 44%. A correlation was found between ETR and net photosynthesis measured as oxygen production (T. maxima: R ² = 0.53; T. squamosa: R ² = 0.61). Calculated compensation depth (CD) (gross photosynthesis equals respiration) in T. maxima (16 m) matches the maximum depth of occurrence in this study (17 m). By contrast, the CD of T. squamosa (9 m) was much shallower than the maximum vertical range (42 m). Findings suggest T. maxima is a strict functional photoautotroph limited by light, whereas T. squamosa is a mixotroph whose photoautotrophic range is extended by heterotrophy. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Inorganic nitrogen metabolism inUlva rigida illuminated with blue light

Inorganic nitrogen metabolism in blue light was studied for the green algaUlva rigida C. Agardh collected in the south of Spain (Punta Carnero, Algeciras) in the winter of 1987. NH₄ ⁺ has been reported to inhibit NO₃ ^- uptake; however,U. rigida showed a net NO₃ ^- uptake even when the NH₄ ⁺ concentration of the external medium was three or four times greater than the concentration of NO₃ ^-. NO₃ ^- uptake rates were similar in both darkness and in blue light of various photon fluence rates (PFR) ranging from 17 to 160 mol m^-2 s^-1. Since NO₃ ^- uptake is an active mechanism involving the consumption of ATP, respiratory metabolism can provide enough ATP to maintain the energetic requirement of NO₃ ^- transport even in darkness. In contrast, NO₃ ^- reduction inU. rigida was highly dependent on the net photosynthetic rate. After 7 h in blue light, intracellular NO₃ ^- concentrations ([NO₃ ^-] _i ) were higher in specimens exposed to intensities below the light compensation point (LCP) than in those incubated at a PFR above the LCP. When PFR is below the light compensation point, NO₃ ^- reduction is low, probably because all the NADH produced by the cells is oxidized in the respiratory chain in order to produce ATP to maintain a steady NO₃ ^- transport rate. The total nitrogen (TN) and carbon (TC) contents decreased from darkness to 33 mol m^-2 s^-1 in blue light. In this range, catabolic processes prevailed over anabolic ones. In contrast, increases in TN and TC contents were observed above the light compensation point. The C : N ratio increased with light intensity, reaching a stable value of 17 at 78 mol m^-2 s^-1 in blue light. Intracellular NO₃ ^- concentration and NO₃ ^- reduction appear to be directly controlled by light intensity. This external control of [NO₃ ^-]_i and the small capacity ofU. rigida to retain incorporated NO₃ ^-, NO₂ ^- and NH₄ ⁺ ions may explain its nitrophilic character.     

Adaptation of solitary corals and their zooxanthellae to low light and UV radiation

Adaptation of solitary corals, Fungia repanda and F. echinata, and their zooxanthellae to low light and ultraviolet light B (UV-B) was studied with respect to changes in their protein contents, photosynthetic pigment contents and the photosynthesis-irradiance (P-I) curves. The corals were collected from 1 to 50 m depths in the Republic of Belau (Paulau) in 1990 and 1991. The chlorophyll a content in a unit surface area of the coral did not change significantly with the depth of the habitat, whereas cellular chlorophyll a in the algae increased with the depth. Zooxanthellae density and protein content in a unit surface area of Fungia spp. decreased with the depth. Photosynthetic parameters normalized by a unit surface area of the Fungia spp., maximum gross photosynthetic rate (P _gmax area^-1) and dark respiration rate (R area^-1), were negatively correlated with the depth, while initial slope of the P-I curve () did not show significant correlation with the depth. Compensation light intensity (Ic) decreased with the depth. In isolated zooxanthellae, P _max chl a ^-1, and R chl a ^-1 decreased with the depth, while _{chl a} was constant. P _gmax cell^-1 and R cell^-1 did not change significantly but _cell increased with the depth. Ic decreased with the depth as in the intact corals. Reduction of protein content in a unit area of the coral from deeper habitat implies decrease of host animal tissues. Reduction of Ic can be explained by decrease of R area^-1, which may be due to the diminution of animal tissues. The photoadaptational response to low light intensity of intact Fungia spp. was found to be a combination of the photoadaptation of symbiotic algae and the decrease of host animal tissue. In order to study their adaptation to ultraviolet (UV) radiation, P-I curves of Fungia spp. and isolated zooxanthellae were analyzed before and after UV-B irradiation. 1 h UV-B irradiation showed no effect on the photosynthetic rate of the shallow water (1 m) corals, while it inhibited the photosynthesis of the deep water (30 m) corals and zooxanthellae isolated from both shallow and deep water corals. These results indicate that the host, Fungia spp., in shallow water have protective mechanism for intense UV-B in their habitat. These photoadaptational mechanisms seem to allow the Fungia spp. to have wide vertical distribution where light intensity spans more than two orders of magnitude.     

Effects of dissolved ammonium addition and host feeding with Artemia salina on photoacclimation of the hermatypic coral Stylophora pistillata

 Effects of nutrient treatments on photoacclimation of the hermatypic coral Stylophora pistillata (Esper) were studied. Studies on photoacclimation of colonies from different light regimes in the field were evaluated and used to design laboratory experiments. Coral colonies were collected in the Gulf of Eilat (Israel) from January to March 1993. Exterior branches of colonies from different depths (1 to 40 m) displayed different trends in production characteristics at reduced and very low levels of illumination. From 24 ± 3% to 12 ± 2% of incident surface photosynthetic active radiation (PAR_o), zooxanthella population density and chlorophyll a+c per 10⁶ zooxanthellae increased, a trend seen in the range of light levels optimal for coral growth (90 to 30% PAR_o). The P _max of CO₂ per 10⁶ zooxanthellae decreased, while P _max of CO₂ per 10³ polyps increased, indicating an increase in zooxanthella population density at low light levels. Proliferous zooxanthella frequency (PZF, a measure of zooxanthella division) declined significantly at light levels <18 ± 3% PAR_o. At the lowest levels of illumination (<5% PAR_o), zooxanthella population density decreased, as did the PZF; chl a+c per 10⁶ zooxanthellae was unchanged. In 28-d experiments, exterior coral branches from the upper surfaces of colonies from 3 m depth (65 ± 4% PAR_o) were incubated in aquaria under bright (80 to 90% PAR_o), reduced (20 to 30% PAR_o), and extremely low (2 to 4% PAR_o) light intensities. At each light intensity, the corals were maintained in three feeding treatments: sea water (SW); ammonium enriched SW (SW + N); SW with Artemia salina nauplii (SW + A). An increase in P _max of CO₂ per 10³ polyps was found in corals acclimated to reduced light (20 to 30% PAR_o) in nutrient-enriched SW, while in SW, where the increase in zooxanthella population density was smaller, it did not occur. Nutrient enrichments (SW + N at 2 to 4% PAR_o and SW + A at 20 to 30% PAR_o) increased zooxanthella population density, but had no effect on chl a+c per 10⁶ zooxanthellae. Acclimation for 14 d to reduced (10 to 20% PAR_o) and extremely low (1 to 3% PAR_o) light intensities shifted ¹⁴C photoassimilation into glycerol and other compounds (probably glycerides), rather than sugars. Both ammonium addition and feeding with Artemia salina nauplii resulted in an increase in photosynthetic assimilation of ¹⁴C into amino acids. We conclude that acclimation to reduced light consists of two processes: an increase in photosynthetic pigments and in zooxanthella population density. Both processes require nitrogen, the increase in zooxanthella population density needing more; this adaptation is therefore limited in nitrogen-poor sea water. Received: 19 June 1998 / Accepted: 13 June 2000     

Light effects on nutrient-limited,oceanic primary production

Phytoplankton growth dynamics were investigated throughout the photic zone at three stations in the North Central Pacific Gyre south of the Hawaiian Islands. Ambient nutrients, vertical light profiles, phytoplankton biomass, and primary production were measured. Photosynthetically active radiation, measured with a submarine quantaspectrophotometer, illustrates vertical variations in photic spectral quality and is presented as incident quanta flux of visible light. Primary production was determined throughout the photic zone under conditions where the samples were collected, injected with ¹⁴C, and incubated under entirely in situ conditions to eliminate preincubation perturbation and to ensure representative response to both light quality and quantity. Oceanic phytoplankton activity is described as a continuous function of incident light under the prevailing low nutrient conditions, and the important rate constants are calculated based upon field data from oligotrophic regions. The vertical profiles of phytoplankton activity and incident quanta flux are analyzed in terms of a substrate-limited system according to the equqtion A _(z) = A _max (q (_(z) – q _o) [K + (q (_(z)) – q (_o], where q(_z) is the quanta flux at a given depth, and A(_z) is the phytoplankton assimilation number at that depth. This is done on the rationale that systematically declining levels of quanta flux, vertically, represent corresponding declines in the availability of substrate for the photochemical processes of photosynthesis. Comparison of data from isolated oceanic regions with those from a station located 15 miles off Oahu show that although large differences in the phytoplankton parameters are evident throughout the entire photic zones of these regions, the hyperbolic A(_z)-q(_z) relationship describes the data fairly well in both cases. The comparison suggests that this relationship may apply to the general case of an oligotrophic water column. These experiments show trends which may be useful for diagnosing phytoplankton activity in the field where N and P levels are low.Oceanic Institute Contribution No. 77-131.     

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.     

Modelling the growth and harvest yield of the giant kelp Macrocystis pyrifera

The giant kelp Macrocystis pyrifera is one of the largest and fastest growing seaweeds and is dominant over large areas of the west coast of North America. A model of its growth has been developed which describes plant biomass and production over the course of a year as a function of environmental parameters which affect the light flux. Such parameters include water clarity, spacing between plants, bottom depth, latitude, harvesting activity, and photosynthetic response (P _max and I _k ). Model results for a standard set of conditions (latitude 33°N, 3 m plant spacing, water absorbance of 0.115 m^-1 and 12 m depth) yield a peak daily gross production of almost 6 g C m^-2 d^-1, peak daily net production of almost 3 g C m^-2 d^-1, and a peak specific growth rate of about 0.022 d^-1. Annual gross production for this case is 1 567 g C m^-2 yr^-1; annual net production is 537 g C m^-2 yr^-1. These values are comparable to those from field measurements. Size and timing of biomass and production peaks are affected by changes in the parameters describing the light field, with peaks usually occurring later in the year for more adverse circumstances. Inhigher latitudes, the seasonal variation is so extreme that the plant could not last the year at 53° N in 12 m of water, although it is able to survive the year in shallower water. Harvesting has severe effects on biomass and production. Model results suggest that light limitation is a very important constraint on kelp growth that should not be overlooked. This implies that differences in parameters describing two environments must be considered when comparing results obtained at different locales.     

Influence of light on algal symbionts of the deep water coral Leptoseris fragilis

Photoadaptations of zooxanthellae living within the deep water coral Leptoseris fragilis taken from the Gulf of Aqaba (Red Sea) were studied. Specimens-collected in summer 1988 between 110 and 120 m depth —were transplanted to 70 and 160 m. At each depth individuals were exposed in their natural growth position (oral side facing the surface) or in a reverse growth position (oral side facing the bottom). After 1 yr of exposure the corals were collected and the zooxanthellae were isolated. As a function of the availability of light with depth and growth position several algal parameters showed changes which are related to photoadaptations. The relatively low density of zooxanthellae of 0.15x10⁶ cellsxcm^-2 at a natural growth depth of 116 m decreased to 0.0034x10⁶ cellsxcm^-2 (2%) at 160 m in specimens growing with a natural orientation. In corals with a downward-facing oral surface at the same depth (160 m) only degenerated algae could be observed. With respect to depth dependence the volume of the algae decreased from 728 m³ at 116 m to 406 m³ at a depth of 160 m and the content of pigments increased. The augmentation of peridinin per cell was low (two times at 160 m compared to 116 m). Chlorophyll a and in particular chlorophyll c ₂ concentrations per cell were enhanced. Compared to natural amounts at 116 m, chl a was five times and chl c ₂ eight times higher at 160 m. At all depths the chl c ₂ content per cell was higher than for chl a. The formation of chl a/chl c ₂ complexes as light harvestor is discussed. Light harvesting, with chl c ₂ prevailing may be explained as a special type of chromatic adaptation of L. fragilis in a double sense: (1) in the habitat light short wavelengths predominate. This light can be directly absorbed with pigments such as chl a and chl c ₂. (2) Host pigments absorb visible violet light and transform these wavelengths, less suitable for photosynthesis, into longer ones by means of autofluorescence. The emitted longer wavelengths fit the absorption maxima of the algal pigments. Thus the host supports photosynthesis of his symbionts. Corals exposed at 160 m depth with a downward facing oral surface were alive after 1 yr and the host wavelength transforming pigment system was still present, but zooxanthellae were absent or degenerated. The light field at 160 m seems therefore to be critical: the combined photoadaptations of host and symbionts, allowing photosynthesis under barren light conditions, seem to be exhausted. In L. fragilis the photoadaptive strategies of host and symbionts cooperate harmoniously. In addition, the adaptations are interlocked with the particular light situation of the habitat with respect to light quantity and quality. The cooperation of physical and organismic parameters examplifies how evolution and, in particular, coevolution has led to optimal fitness.     

Effects of light intensity on nitrate and nitrite uptake and excretion by Chaetoceros curvisetus

Michaelis-Menten uptake kinetics were observed at all light intensities. With constant illumination, the V_max and K₁ in nitrate uptake over the natural light intensity range of 0 to 2000 E were 0.343 g-at NO₃–N(g)^-1 at protein-N h^-1 and 26 E, respectively. Nitrate uptake was inhibited at higher light intensities. The K_s for nitrate uptake did not vary as a function of light intensity remaining relatively constant at 0.62 g-at NO₃–N 1^-1. With intermittent illumination, the V_mzx for light intensity in nitrate uptake over a light intensity range of 0 to 5000 E was 0.341 g-at NO₃–N(g)^-1-at protein-N h^-1. No inhibition of nitrate uptake was observed at higher than natural light intensities. Chaetoceros curvisetus will probably never experience light inhibition of nitrate uptake under natural conditions.     

Primary production and environmental factors in an oligotrophic biome in the Aegean Sea

The rate of the primary production of the phytoplankton community in the Petalion Gulf, Aegean Sea, was studied from January 1970 to May 1971, at a station situated at approximately Latitude 37°54N; Longitude 24°11E. A variety of physical and chemical parameters such as chlorophyll, primary nutrients (N,P,Si), temperature, salinity, oxygen and light penetration were also studied simultaneously. The rate of the gross primary production varied from 40 to 200 mg C m^-2 day^-1, with a mean value of 90 mg C m^-2 day^-1. The annual gross primary production was calculated to be 33 g C m^-2, which is the minimum known value in the Aegean and Mediterranean Seas. Maximum production was found at the depth of 20 m on the average, mainly due to high light intensities. Petalion Gulf supports a small photosynthetic biomass, as indicated by the low seasonal values of chlorophyll a (0.01 to 0.18 mg m^-2), the highest values being found in the summer. The low production rate noted may have been due to the low nutrient concentrations found: N, 0.04 to 0.32 g-at/1; P, 0.00 to 0.15 g-at/1; Si, 0.45 to 2.25 g-at/1. It is suggested that inorganic phosphorus and nitrogen may alternate in limiting primary production rates in these oligotrophic waters. The temperate waters of the Petalion Gulf are stratified in summer (15.5° to 24.7°C) and well-mixed in winter (12.9° to 15.0°C); they are oxygen-saturated throughout the year, and of high transparency, with 86 m depth for the euphotic zone on the average yearly. The Petalion Gulf is therefore characterized as a typical oligotrophic biome in the Aegean and Eastern Mediterranean Seas.     

Primary production of sands in the lagoon of an atoll and the role of foraminiferan symbionts

Chlorophyll content and in situ oxygen production of sands were measured in the lagoon of Takapoto Atoll, Tuamotu Islands (French Polynesia). Stations were spaced from a depth of 17 m to the water limit on the beach. Photosynthetic pigments and net primary production for the top 3 cm of sediment ranged from 56 to 907 mg chlorophyll a m^-2 and from 115 to 354 mg O₂ m^-2 h^-1, respectively. Responsible organisms were mostly Foraminifera, among which Amphistegina lessoni, containing unicellular symbionts, dominated. The sands are thus significant contributors to the lagoon's primary production, while the role of phytoplankton is comparatively negligible.     

Cell cycle and toxin production in the benthic dinoflagellate Prorocentrum lima

Profiles of diarrhetic shellfish poisoning (DSP) toxins produced throughout the growth cycle and the cell cycle of the toxigenic marine dinoflagellate Prorocentrum lima were studied in triplicate unialgal batch cultures. Cells were pre-conditioned at 18 ± 1 °C, under a photon flux density (PFD) of 90 ± 5 μmol m⁻² s⁻¹ on a 14 h light:10 h dark photoperiod. In exponential growth phase, cultures were synchronized in darkness for 17 d. After dark synchronization, cultures were transferred back to the original photoperiod regime. Cells were harvested for DSP toxin analysis by LC-MS (liquid chromatography with mass spectrometry), and double-stranded (nuclear) DNA was quantified by flow cytometry. The cell populations became asynchronous within approximately 3 d after transition from darkness to the 14 h light:10 h dark photoperiod. This may be due to the prolonged division cycle (5 to 7 d) that is not tightly phased by the photoperiod. Unlike other planktonic Prorocentrum spp., cytokinesis in P. lima occurred early in the dark and ceased by “midnight”. Cellular levels of the four principal DSP toxins, okadaic acid (OA), OA C8-diol-ester (OA-D8), dinophysistoxin-1 (DTX1) and dinophysistoxin-4 (DTX4), ranged from 0.37 to 6.6, 0.02 to 1.5, 0.04 to 2.6, and 1.8 to 7.8 fmol cell⁻¹, respectively. No toxin production was evident during the extended period of dark synchronization nor during the initial period when NH₄ was consumed as the major nitrogen source. Soon after the cells were returned to the 14 h light:10 h dark cycle and they began to take up NO₃, cellular levels of all four toxins gradually increased. This increase in DSP toxins usually occurred in the light, marked by a rise in DTX4 levels that preceded an increase in the cellular concentration of OA and DTX1 (delayed by 3 to 6 h). Thus, DTX4 synthesis is initiated in the G1 phase of the cell cycle and persists into S phase (“morning” of the photoperiod), whereas OA and DTX1 production occurs later during S and G2 phases (“afternoon”). No toxin production was measured during cytokinesis, which happened early in the dark. The evidence indicates that toxin synthesis is restricted to the light period and is coupled to cell cycle events. Received: 3 September 1998 / Accepted: 30 March 1999     

Phytoplankton community structure and primary production in small intertidal estuarine-bay ecosystem (eastern English Channel,France)

From May 2002 to October 2003, a fortnightly sampling programme was conducted in a restricted macrotidal ecosystem in the English Channel, the Baie des Veys (France). Three sets of data were obtained: (1) physico-chemical parameters, (2) phytoplankton community structure illustrated by species composition, biovolume and diversity, and (3) primary production and photosynthetic parameters via P versus E curves. The aim of this study was to investigate the temporal variations of primary production and photosynthetic parameters in this bay and to highlight the potential links with phytoplankton community structure. The highest level of daily depth-integrated primary production Pz (0.02–1.43 g C m⁻² d⁻¹) and the highest maximum photosynthetic rate P ^B _max (0.39–8.48 mg C mg chl a ⁻¹ h⁻¹) and maximum light utilization coefficient α^B [0.002–0.119 mg C mg chl a ⁻¹ h⁻¹ (μmol photons m⁻² s⁻¹)] were measured from July to September. Species succession was determined based on biomass data obtained from cell density and biovolume measurements. The bay was dominated by 11 diatoms throughout the year. However, a Phaeocystis globosa bloom (up to 25 mg chl a m⁻³, 2.5 × 10⁶ cells l⁻¹) was observed each year during the spring diatom bloom, but timing and intensity varied interannually. Annual variation of primary production was due to nutrient limitation, light climate and water temperature. The seasonal pattern of microalgal succession, with regular changes in composition, biovolume and diversity, influenced the physico-chemical and biological characteristics of the environment (especially nutrient stocks in the bay) and thus primary production. Consequently, investigation of phytoplankton community structure is important for developing the understanding of ecosystem functioning, as it plays a major role in the dynamics of primary production.     

Gene flow among population of a teleost (painted greenling,Oxylebius pictus) from Puget Sound to southern California

Net photosynthetic oxygen evolution in Amphiroa anceps (Lamarck) Decaisne is inhibited at high oxygen concentrations. Photosynthesis is highest between pH 6.5 and 7.5. At pH 9 to 10 there is still a significant photosynthetic rate, suggesting that this alga can use HCO _- ³ as a substrate for photosynthesis. At pH 7.0 to 8.5, the photosynthetic rate saturates at a total inorganic carbon concentration (C_i) greater than 3 mM. At pH 8.5 and 8.8, calcification rate continues to increase with increasing concentration of C_i. Between pH 7 and 9, the calcification rate in the light in A. foliacea Lamouroux is proportional to the photosynthetic rate, whereas at higher pH where the photosynthetic rate is very low, the calcification rate is stimulated by the higher concentration of CO _2- ³ ion. At all pH values examined, the calcification rate of living plants in the dark and of dead plants is directly proportional to the CO _2- ³ ion concentration, suggesting little metabolic involvement in calcification processes in the dark, whereas calcification in live A. foliacea in the light is influenced both by the photosynthetic rate and the CO _2- ³ ion concentration in the medium.     

Phytoplankton exudate release as an energy source for the growth of pelagic bacteria

The release of dissolved organic carbon (DOC) from phytolankton during photosynthesis, and the utilization of this carbon by planktonic bacteria, was studied using ¹⁴CO₂ and selective filtration. Natural sea water samples from a coastal area of the Northern Baltic Sea were incubated in the laboratory for detailed studies, and in situ for estimation of annual dynamics. In a laboratory incubation (at +1°C) the concentration of ¹⁴C-labelled dissolved organic carbon increased for about 2 h and then reached a steady state, representing about 0. 1% of the total DOC. Labelled organic carbon in the phytoplankton and bacterial fractions continued to increase almost linearly. The continuous increase in the bacterial fraction is thought to represent almost instantaneous utilization of the DOC released from the phytoplankton during photosynthesis. As an annual average, in 4 h in situ incubations, about 65% of the labelled organic carbon was found in the phytoplankton fraction (>3 m), about 27% in the bacterial fraction (0.2 to 3 m) and the remaining 8% as DOC (<0.2 m). Large variations in these percentages were recorded. The measured annual primary production was 93 g C m^-2 (March to December), and the estimated bacterial production due to phytoplankton exudates 29 g C m^-2. This represents a release of DOC of about 45% of the corrected annual primary production of 110 g C m^-2 (assuming a bacterial growth efficiency of 0.6).     

DCMU-enhanced fluorescence as an index of photosynthetic activity in phytoplankton

Tests have been carried out, both on phytoplankton cultures and in the field, on the relation between the ratio of 3-(3,4-dichlorophenyl)-1,1-dimethyl urea (DCMU)-enhanced to normal fluorescence (F _D /F _N ) and the specific photosynthetic rate (P/B). In all cases observed, significant linear correlations were found between these two values. Differences in this relation were observed according to species and physiological conditions. Nutrient stress that occurs in batch cultures decreases both F _D /F _N and P/B, while lowered light intensity has a different effect on both, increasing F _D /F _N and decreasing P/B. This is interpreted as indicating that F _D /F _N is an index of photosynthetic efficiency at different light levels and an indicator of the specific photosynthetic rate (P/B) when physiological conditions vary at a given light intensity. The practical use of DCMU-enhanced fluorometry to estimate instantaneous P/B values in the field is discussed, stressing the frequent calibrations needed, as in all in situ fluorometry studies.     

Primary production of phytoplankton at a frontal zone located at the northern slope of the Dogger Bank (North Sea)

In July 1988 a survey was made in the Dogger Bank area of the North Sea. As a result of wind stress the area was found to be frequently well mixed. At the northerly slope a transition zone was observed between the stratified central North Sea and the well-mixed Dogger Bank area. Low nutrient concentrations were observed in surface waters; especially for nitrate (<0,1µM). High concentrations of phosphate (>0,5µM), nitrate (>1µM), ammonium (>2µM) and silicate (>2µM) only prevailed below the thermocline. Chlorophylla values were below 1µg l^–1 near the surface. Enhanced values (up to 4µg l^–1) were observed in the deeper layer at the transition zone and just below the thermocline at well-stratified locations. At the transition zone high specific C-fixation rates (up to 100 mg C mg^–1 chla d^–1) at the surface indicated the presence of enhanced productivity. The compensation depth for primary production was found to coincide with a specific C-fixation rate of 5 mg C mg^–1 chla d^–1. At greater depths, phytoplankton was only found where tidally induced vertical mixing allowed a regular exposure to higher light intensities. Storms resulted in a rapid redistribution of chlorophylla and enhancement of the C-fixation rate in the upper layer of the water column.Publication No. 10 of the project Applied Scientific Research Netherlands Institute for Sea Research (BEWON)     

In situ measurement of the urea decomposition rate and its turnover rate in the Pacific Ocean

The in situ decomposition rate of urea was measured using ¹⁴C-labelled urea at 3 areas in the North Pacific Ocean: Sagami Bay on the southern coast of central Japan, the northwestern Pacific central waters and the subarctic Pacific waters. The mean values of the decomposition rates of urea in surface waters of these areas were 44.5, 1.51 and 1.32 mol urea m^-3 d^-1, respectively. These rates decreased with depth. High rates of urea carbon incorporation into particulate matter and the CO₂ liberation from urea carbon into seawater were obtained in light bottles in the euphotic zone, while low rates were found in dark bottles. The turnover rates of urea in the 3 areas were calculated respectively as 12, 113 and 110 d at the surface, and the values increased with depth.     

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.     

Effects of hypoxia on the respiratory and circulatory regulation of Nephrops norvegicus

The burrowing decapod Nephrops norvegicus (L.) was kept under various degrees of hypoxia in order to measure respiration, heart rate, scaphognathite rate, haemolymph oxygen content and pH. An emergence reaction to hypoxia occurred only in dim light (<10^-2 m-c) or darkness, but after 10 d of moderate hypoxia the decapods showed no emergence response at all. The weight specific respiration of quiescent individuals was relatively low and increased only slightly in hypoxia (P_wO₂=40 torr). Heart rate, about 50 beats min^-1, changed little during hypoxia, down to P_wO₂=40 torr, whereas scaphognathite rates rose from about 60 beats min^-1 at normoxia to peak at 120 beats min^-1 at P_wO₂=40 torr. The oxygen extraction efficiency (E) remained at 20 to 30% during the first hour of hypoxia then rose gradually to maximum values of 30 to 40%. A small respiratory alkalosis of the blood became evident only after 4h of hypoxia (P_wO₂=50 torr). Normoxic postbranchial O₂ tensions (P_aO₂) were low (25–30 torr) and showed only a small decline during hypoxia. Over 10 to 13 d in moderate hypoxia an effective biosynthesis of 0.024 mM haemocyanin individual^-1 d^-1 occurred in fed decapods, whereas controls (normoxic) showed no significant change in pigment levels. A linear relationship between oxygen carrying capacity and haemocyanin concentration was found. It is contended that N. norvegicus is better able to cope with periodic exposure to hypoxia when food of sufficient quantity and quality is available.