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     

Tidal variations in the photosynthesis of estuarine phytoplankton isolated in a tank

In 1981 two large (1 200 1) seawater samples from the St. Lawrence Estuary were kept under constant temperature and light conditions for periods of 50 and 68 h, respectively. In both tank experiments, semidiurnal variations in NH₄ were observed that could be related to cyclical NH₄ uptake by the phytoplankton. Semidiurnal cycles in photosynthetic efficiency (^B) and intracellular chlorophyll a in the tank, phased on tides at sea, were also evidenced in both experiments. These results support the hypothesis that variations in phytoplankton photosynthetic activity, which are possibly endogenous, can be phased on semidiurnal variations in vertical tidal mixing (variations of the mean light conditions in the mixed layer). In addition, observed variations in intracellular chlorophyll a suggest the possibility of endogenous cycles of phytoplankton light and shade adaptation.Contribution to the program of GIROQ (Groupe interuniversitaire de recherches océanographiques du Québec)     

The theoretical basis for estimating phytoplankton production and specific growth rate from chlorophyll, light and temperature data

Phytoplankton maximum specific growth rate, μ_max, and maximum photosynthetic quantum yield, Φ_max, can be related mathematically via the photosynthetic light curve, P(I). A model is presented in which maximum quantum yield defines the initial slope of the light curve and is assumed to be a known constant, while maximum specific growth rate defines the light-saturated region of the curve and is assumed to be a known function of temperature. The effect of introducing μ_max(T) into P(I, Φ_max) is to replace the unknown, temperature-dependent light saturation parameter with a term involving the ratio μ_max/Φ_max. The advantage of writing P(I) in terms of both μ_max and Φ_max is that those parameters are particularly well documented in the literature. Consequently, estimates of nutrient-unlimited phytoplankton growth and production rates can be based solely on the constants μ_max, Φ_max and k_c (light absorption per unit of chlorophyll) and the free variables light, temperature and chlorophyll concentration. Rate estimates appear to be accurate to within a factor of two for an extremely wide range of conditions.One particularly significant result of introducing μ_max into P(I, Φ_max) is that the carbon : chlorophyll ration, θ, appears explicitly. It is possible to derive an expression for optimum θ based on the assumption that adaptive changes in carbon/chlorophyll occur so as to maximize the specific growth rate for given conditions of light and temperature. Laboratory and field data are compiled from the literature to test the formulae presented here.     

Mélange vertical et capacité photosynthétique du phytoplancton estuarien (estuaire du Saint-Laurent)

Time series of chlorophyll a, photosynthetic capacity and many physical parameters were sampled hourly for 167 h in August, 1975, at an anchor station located in the Middle Estuary of the St. Lawrence River, Canada. Sampling was carried out during the transition from neap tides to spring tides. The long-and short-term variations in chlorophyll a are coupled with the advection of water masses which depends on tidal currents. Vertical mixing also influences the chlorophyll a concentration of the cells, since it modifies the physiological state of the phytoplankton. Furthermore, circadian periodicities were observed in the photosynthetic capacity, suggesting that the phytoplankton of this area have a homogeneous light history due to strong vertical mixing. Under these conditions, the photosynthetic capacity is adapted to the mean light intensity in the mixed layer. The semimonthly (M _f) variations of the mean light intensity in the mixed layer depend on the M _f variations in the vertical mixing, whereas in the short-term, the variations in mean light intensity in the mixed layer are circadian.

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Mélange vertical et capacité photosynthétique du phytoplancton estuarien (estuaire du Saint-Laurent)

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Contribution au programme du Groupe interuniversitaire de recherches océanographiques du Québec (GIROQ)     

Picophytoplankton responses to changing nutrient and light regimes during a bloom

The spring bloom in seasonally stratified seas is often characterized by a rapid increase in photosynthetic biomass. To clarify how the combined effects of nutrient and light availability influence phytoplankton composition in the oligotrophic Gulf of Aqaba, Red Sea, phytoplankton growth and acclimation responses to various nutrient and light regimes were recorded in three independent bioassays and during a naturally-occurring bloom. We show that picoeukaryotes and Synechococcus maintained a “bloomer” growth strategy, which allowed them to grow quickly when nutrient and light limitation were reversed. During the bloom picoeukaryotes and Synechococcus appeared to have higher P requirements relative to N, and were responsible for the majority of photosynthetic biomass accumulation. Following stratification events, populations limited by light showed rapid photoacclimation (based on analysis of cellular fluorescence levels and photosystem II photosynthetic efficiency) and community composition shifts without substantial changes in photosynthetic biomass. The traditional interpretation of “bloom” dynamics (i.e., as an increase in photosynthetic biomass) may therefore be confined to the upper euphotic zone where light is not limiting, while other acclimation processes are more ecologically relevant at depth. Characterizing acclimation processes and growth strategies is important if we are to clarify mechanisms that underlie productivity in oligotrophic regions, which account for approximately half of the global primary production in the ocean. This information is also important for predicting how phytoplankton may respond to global warming-induced oligotrophic ocean expansion.     

Diel changes in phytoplankton photosynthetic efficiency in Brackish waters

From 18 to 23 September 1974, investigations on the diel changes in phytoplankton were carried out in the Baltic Sea. Every 4 h, water samples were collected from 2 and 15 m, and PO₄, chlorophyll a, temperature, salinity, pH, phytoplankton composition and phytoplankton light photosynthesis relationship were determined. Continuous measurements of surface irradiance and some estimations of zooplankton were also made. P ^B (photosynthesis per unit chlorophyll a at low light levels of 2·10^-2 cal cm^-2 min^-1) revealed only random variation during the sampling period, i.e., 1.0 to 1.6 mg C (mg chlorophyll a)^-1 h^-1. P _m ^B (Light-saturated photosynthesis per unit of chlorophyll a) displayed pronounced diel fluctuations with the highest value of about 6 mg C (mg chlorophyll a)^-1 h^-1 around noon, and the lowest value of about 2.5 mg C (mg chlorophyll a)^-1 h^-1 during the night, during which latter period the value of P _m ^B was more or less constant. Reasons for the diel fluctuations are discussed, and an equation which describes these fluctuations is proposed. Using this equation, the daily phytoplankton production estimated in incubators by a previously described method can be corrected for the time of day at which samples are collected.     

Benthic foraminiferal distribution on the southeastern Brazilian shelf and upper slope

Foraminiferal data were obtained from 66 samples of box cores on the southeastern Brazilian upper margin (between 23.8°–25.9°S and 42.8°–46.13°W) to evaluate the benthic foraminiferal fauna distribution and its relation to some selected abiotic parameters. We focused on areas with different primary production regimes on the southern Brazilian margin, which is generally considered as an oligotrophic region. The total density (D), richness (R), mean diversity ([`(H^￠ )] \overline{{H^{\prime } }} ), average living depth (ALD _X ) and percentages of specimens of different microhabitats (epifauna, shallow infauna, intermediate infauna and deep infauna) were analyzed. The dominant species identified were Uvigerina spp., Globocassidulina subglobosa, Bulimina marginata, Adercotryma wrighti, Islandiella norcrossi, Rhizammina spp. and Brizalina sp.. We also established a set of mathematical functions for analyzing the vertical foraminiferal distribution patterns, providing a quantitative tool that allows correlating the microfaunal density distributions with abiotic factors. In general, the cores that fit with pure exponential decaying functions were related to the oligotrophic conditions prevalent on the Brazilian margin and to the flow of the Brazilian Current (BC). Different foraminiferal responses were identified in cores located in higher productivity zones, such as the northern and the southern region of the study area, where high percentages of infauna were encountered in these cores, and the functions used to fit these profiles differ appreciably from a pure exponential function, as a response of the significant living fauna in deeper layers of the sediment. One of the main factors supporting the different foraminiferal assemblage responses may be related to the differences in primary productivity of the water column and, consequently, in the estimated carbon flux to the sea floor. Nevertheless, also bottom water velocities, substrate type and water depth need to be considered.     

Dynamique du phytoplancton et matière organique particulaire dans la zone euphotique de l'Atlantique Equatorial

At two fixed stations in the Equatorial Atlantic Ocean (0°–4° W), the physical, chemical and biological properties of the euphotic layer were determined for 14 d (Station A: 5–18 February, 1979) and 13 d (Station B: 20 October–7 November, 1979), respectively. The stability of the water column allowed comparison of 3 different “systems”: (i) a well-illuminated and nitrate-depleted mixed layer; (ii) a chlorophyll maximum layer (chl a _max) in the thermocline which is poorly illuminated (6.3% of surface irradiance); (iii) a well-illuminated but nitrate-rich (>0.9 μg-at l^-1) mixed layer. In each layer the particulate organic carbon (COP), nitrogen (NOP) and phosphorus (POP) contents were measured and compared with the phytoplankton biomass. In the chlorophyll maximum layer, the phytoplankton biomass contributed significantly to the total particulate organic matter (between 55 and 75%). In the nitrate-depleted mixed layer, the results varied according to whether the regression technique [COP=f(chl a)] was used, or the chl a synthesis during the incubation of the samples. With the former technique, the phytoplankton carbon (C _p) content appeared minimal, because the y intercept, computed using all the data of the water column, was probably overestimated for this layer. POP would be more associated with living protoplasm than with carbon and nitrogen in the three layers. In the chlorophyll a maximum layer it constitutes a valuable detritus-free biomass measurement, since 80% of the POP consist of phytoplankton phosphorus. The assimilation numbers (NA=μg C μg chl a ^-1 h^-1) were high in all three layers, but the highest values were recorded in the nitrate-depleted mixed layer (NA=15 μg C μg chl a ^-1 h^-1). In the chlorophyll maximum layer, light would be a limiting factor during incubation: between 10²⁵ and 8.10²⁴ quanta m^-2 d^-1 NA and light are positively correlated independant of nitrate concentration. The growth rates of phytoplankton (μ) were estimated and compared to the maximum expected growth rate. Our main conclusion was that despite very low biomass and nutrient content, the mixed layer was in a highly dynamic state, as evidenced by high rates of phytoplankton growth and short nutrient turnover times (1 d or less for PO-P₄ in the mixed layer versus 3 d in the thermocline). The presence of nitrate in the water column allows the development of a higher phytoplankton biomass but does not increase growth rate.     

Relationship between photosynthesis and non-photochemical quenching of chlorophyll fluorescence in two red algae with different carotenoid compositions

Algae are continuously exposed to short-term fluctuations in irradiance. We investigate how two red algae species regulate photosynthetic efficiency to cope with such changes and identify some strategies that differ from higher plants. Two red algae, Gracilaria domingensis and Kappaphycus alvarezii, with antheraxanthin and lutein as major xanthophylls, respectively, reacted to the onset of low light (below E _k) with a substantial decrease of NPQ. This is different from higher plants, but similar to previous observations in, e.g. cyanobacteria where it indicates an increase in the effective absorbance cross-section of Photosystem II (PSII) by state transition. Kinetic studies in continuous light revealed a high susceptibility of PSII to light stress ((1-q _P)/NPQ) in K. alvarezii immediately after the sudden onset of high light, followed by a decrease. This was caused by a slower onset of NPQ in K. alvarezii, followed by acclimation. In G. domingensis, susceptibility of PSII to light stress was stable with time, but absolute values of (1-q _P)/NPQ were higher than in K. alvarezii. These observations suggest that K. alvarezii may be better adapted to high light levels, but is less well prepared for large sudden changes in irradiation. In K. alvarezii, photosynthesis continued to increase with increasing irradiation when NPQ was saturated. As (1-q _P) and NPQ were still balanced in this situation, most likely, processes other than photosynthetic oxygen release are responsible for the increasing net O₂ production observed.     

Aspects of water column primary productivity in the Chukchi Sea during summer

Measurements of primary productivity, chlorophyll a, incident solar radiation, phosphate-P, silicate-Si, nitrate-N, nitrite-N, ammonium-N, temperature and salinity were made in the Marginal Ice Zone of the Chukchi Sea in summer 1974. Low to moderate levels of primary productivity (0.07 to 0.97 g C m^-2 half-day^-1) were observed; primary productivity exceeded 3 g C m^-2 half-day^-1 at two stations. Surface primary productivity was nitrogen-limited at most stations. Mean chlorophyll a concentration in the photic zone varied from 0.4 to 17.8 mg m^-3. Higher concentrations and significant subsurface accumulation of chlorophyll a, reaching 40 mg m^-3, were observed in July at stations near the ice-edge than those in open water. No chlorophyll maximum was noted in September, when values ranged from 0.4 to 2.2 mg m^-3. It is postulated that the contribution of sea-ice algae to the total chlorophyll content can be substantial, but that the stay of these cells in the water column may not be long. Non-linear regression estimates from solar radiation and chlorophyll-specific primary productivity data showed a maximal photosynthetic rate of 18 mg C mg chlorophyll a ^-1 half-day^-1, an optimal light intensity of 54 langleys half-day^-1, and markedly reduced primary productivity at moderately higher light intensities. These features indicate that phytoplankton was shade-adapted.     

Effects of wave-induced lightflecks in the intertidal zone on photosynthesis in the macroalgae Postelsia palmaeformis and Hedophyllum sessile (Phaeophyceae)

Intertidal algae are exposed to a highly variable photic regime because of crashing waves. We measured photosynthetic rate of whole fronds of the seaweeds Postelsia palmaeformis (Ruprecht) (at Bodega Marine Laboratory, 1991) and Hedophyllum sessile (Setchell) (Phaeophyceae) (at Friday Harbor Laboratories, 1990) in flashing and steady-state photosynthetically active radiation (PAR) of equal irradiance, using a recirculating metabolism chamber designed to minimize the thickness of diffusional boundary layers on the surfaces of algal tissues. The dimensionless ratio of photosynthetic rate under flecking PAR (P_fleck) and the sum of photosynthetic rate under steady state PAR (P_s) of high and low irradiance was computed for lightfleck periods from 0.2 to 100 s. P_fleck:P_s is a measure of the light-flash utilization-efficiency in flashing light, and was greater than unity at periods between lighflecks of 2 to 30 s, with a peak at 10 s. We used a novel optical fiber irradiance meter to measure PAR incident on fronds of P. palmaeformis as they were washed about by waves breaking in the intertidal zone, and compared the light records to that obtained by a stationary sensor under the canopy. PAR flashing period was closely correlated with the period of breaking waves in stands of P. palmaeformis. We estimated the seasonal spectrum of period between light flashes in stands of this species by examining the spectral density of ocean waves at Bodega Marine Laboratory, Bodega Bay, California. The wave spectrum peaks at a period (10 s) where light-flashes may theoretically enhance the light-flash utilization-efficiency of seaweeds. We calculate that the enhanced light-flash utilization-efficiency wrought by wave-induced light-flashes may contribute to significant gains in primary productivity of these macroalgae under some conditions.     

Diel vertical migrations and nocturnal feeding of a dense coastal krill scattering layer (Thysanoessa raschi and Meganyctiphanes norvegica) in stratified surface waters

To study the nocturnal feeding of euphausiids during vertical migrations and its impact on the phytoplankton, a phytoplankton-rich water mass (drogue marked) drifting over a dense krill scattering layer (acoustic 104kHz) in the lower St. Lawrence estuary was monitored over 46 h in July 1982. Phytoplankton >20 m was abundant and mostly concentrated at the bottom of the photic layer above the pycnocline. Less than 42% of the particulate carbon was due to phytoplankton. The krill scattering layer was about 2 to 3 km in width, elongated along the 100-m bathymetric contour, and absent when the bottom was shallower than 50 m. Its upper day depth was 50 m. At deeper depths, its vertical distribution frequently changed from unimodal to polymodal shapes and viceversa, often with large concentrations of zooplankton just above the bottom. Typical vertical migrations were observed on both days. At night the scattering layer had a lower scattering strength. Most of it was below the thermocline but net catches showed that large concentrations of euphausiids (up to 57 individuals m^-3) crossed it. Stomach pigment content of Thysanoessa raschi was generally low, but mean stomach fullness was always high. They were more opportunistic than herbivorous. From stomach fullness and the presence of a food bolus in mouth parts, feeding in surface waters appeared to be intensive, but gut content indicated that food was not processed there. It is therefore suggested that individuals underwent vertical interchanges across the thermocline while feeding during the night. Meganyctiphanes norvegica had significant herbivorous activity during the night. The grazing pressure impact of the scattering layer on phytoplankton was negligible.Contribution to the program of GIROQ (Groupe interuniversitaire de recherches océanographiques du Québec)     

A general solution of Benjamin-type gravity current in a channel of non-rectangular cross-section

We consider the steady-state propagation of a high-Reynolds-number gravity current in a horizontal channel along the horizontal coordinate x. The bottom and top of the channel are at z =?0, H, and the cross-section is given by the quite general form ?f ₁(z) ≤?y ≤?f ₂(z) for 0 ≤?z ≤?H, where f _1,2 are piecewise continuous functions and f ₁ +?f ₂ >?0 for ${z \in(0,H)}$ . The interface of the current is horizontal, the (maximum) thickness is h, its density is ρ _c . The reduced gravity g′ =?|ρ _c /ρ _a ? 1|g (where ${- g\hat{z}}$ is the gravity acceleration and ρ _a the density of the ambient) drives the current with speed U into the stationary ambient fluid. We show that the dimensionless Fr =?U/(g′ h)^1/2, the rate of energy dissipation (scaled with the rate of pressure work), and the dimensionless head-loss Δ/h, can be expressed by compact formulas which involve three integrals over the cross-section areas of the current and ambient. By some standard manipulations these integrals are simplified into quite simple line-integrals of the shape-function of the channel, f(z) =?f ₁(z) +?f ₂(z), and of z f(z). This theory applies to Boussinesq and non-Boussinesq currents of “heavy” (bottom) and “light” (top) type. The classical results of Benjamin (J Fluid Mech 31:209–248, 1968) for a rectangular channel are fully recovered. We also recover the Fr results of Marino and Thomas (J Fluid Eng 131(5):051201, 2009) for channels of shape y =?±b z ^α (where b, α are positive constants); in addition, we consider the energy dissipation of these flows. The results provide insights into the effect of the cross-section shape on the behavior of the steady-state current, in quite general cases, for both heavy-into-light and light-into-heavy fluid systems, Boussinesq and non-Boussinesq. In particular, we show that a very deep current displays ${Fr = \sqrt{2}}$ , and is dissipative; the value of Fr and rate of dissipation (absolute value) decrease when the thickness of the current increases. However, in general, energy considerations restrict the thickness of the current by a clear-cut condition of the form h/H ≤?a _max ?< 1.     

Age-specific responses to elevated salinity in the coastal marsh plant black needlerush (Juncus roemerianus Scheele) as determined through polyphasic chlorophyll a fluorescence transients (OJIP)

This study employed polyphasic chlorophyll a fluorescence transients (OJIP), a non-invasive marker of environmental stress in plants, to evaluate salt tolerance in three different Juncus roemerianus age classifications (6-, 24-, and 60-months). Following exposure to elevated salts (30 psu), the younger plants sustained growth, which was comparable to freshwater controls. While older (60-month) plants receiving only freshwater also grew over the 8-week study, the older salt-treated plants did not increase in size. Similarly, there were significant declines in variable chlorophyll a fluorescence parameters (F _v/F _m and F _v/F _o), electron transport flux per reaction center (ET_o/RC), and photosystem II performance index (PI_ABS) for 60-month J. roemerianus following salt treatment. These responses were not evident in the two younger salt-treated age classifications. Our results suggest that older J. roemerianus are less tolerant to rapid and sudden increases in salinity relative to younger plants and that this age-specific response may help explain observed discrepancies in salt tolerance in J. roemerianus.     

Optimization of exergy and implications of body sizes of phytoplankton and zooplankton in an aquatic ecosystem model

Size appears to be an important parameter in ecological processes. All physiological processes vary with body size ranging from small microorganisms to higher mammals. In this model, five state variables — phosphorus, detritus, phytoplankton, zooplankton and fish are considered. We study the implications of body sizes of phytoplankton and zooplankton for total system dynamics by optimizing exergy as a goal function for system performance indicator. The rates of different sub-processes of phytoplankton and zooplankton are calculated, by means of allometric relationships of their body sizes. We run the model with different combinations of body sizes of phytoplankton and zooplankton and observe the overall biomass of phytoplankton, zooplankton and fish. The highest exergy values in different combinations of phytoplankton and zooplankton size indicate the maximum biomass of fish with relative proportions of phytoplankton and zooplankton. We also test the effect of phosphorus input conditions corresponding to oligotrophic, mesotrophic, eutrophic system on its dynamics. The average exergy to be maximized over phytoplankton and zooplankton size was computed when the system reached a steady state. Since this state is often a limit cycle, and the exergy copies this behaviour, we averaged the exergy computed for 365 days (duration of 1 year) in the stable period of the run. In mesotrophic condition, maximum fish biomass with relative proportional ratio of phytoplankton, zooplankton is recorded for phytoplankton size class 3.12 (log V μm³ volume) and zooplankton size 4 (log V μm³ volume). In oligotrophic condition the highest average exergy is obtained in between phytoplankton size 1.48 (log V μm³ volume) and zooplankton size 4 (log V μm³ volume), whereas in eutrophic condition the result shows the highest exergy in the combination of phytoplankton size 5.25 (log V μm³ volume) and zooplankton size 4 (log V μm³ volume).     

Diel periodicity of photosynthesis in marine phytoplankton

Short-term changes in photosynthesis were documented for 17 of 24 marine phytoplankton species, representing a range of taxonomic groups. Periodicity in phytoplankton photosynthesis on light-dark cycles (diel periodicity) was widespread but not universal for the species studied. The centric diatoms Lauderia borealis, Ditylum brightwellii, Stephanopyxis turris, Coscinodiscus rex, Chaetoceros gracile, and Biddulphia mobiliensis had strong diel periodicity in photosynthetic capacity (P _max). Amplitudes of the daily variations ranged from 2.9 to >50, with maxima in the morning or near midday, and with minima during the dark period, and these variations were not dependent on changes in cell pigmentation. There was some evidence for sustained photosynthetic periodicity in constant conditions in several diatoms, and an endogenous rhythm may have been present. The photosynthesis-irradiance (P-I) relationship was time-dependent for representative marine diatoms, with both the initial slope () and the asymptote (P _max) of P-I curves exhibiting significant synchronous diel oscillations. Moreover, detailed studies of the amplitude and timing of photosynthetic periodicity for the diatoms L. borealis and D. brightwellii demonstrated large temporal variations in photosynthesis with morning maxima. These P-I oscillations are discussed with reference to models of primary production which use the relationship between photosynthesis and light as a component of predictive equations for phytoplankton growth in the sea.     

Effects of light intensity and nutrient availability on diel patterns of cell metabolism and growth in populations of Synechococcus spp.

Between July 21 and August 8, 1984, phytoplankton were collected from the surface (2 m) and/or chlorophyll maximum of a neritic front, warm-core eddy 84-E and Wilkinson's Basin in the Northwest Atlantic Ocean and incubated up to 38 h in 200-liter vats. Effects of light intensity and nutrient availability on diel patterns of cell metabolism were analyzed in a 0.6- to 1-m fraction, where Synechococcus spp. represented 80 to 100% of the total photoautotrophs. Populations held under in situ conditions exhibited daytime peaks in photosynthetic potential (P_max) that were an order of magnitude higher than nighttime P_max values. Daytime phasing of P_max peaks had no relationship to asynchronous fluctuations in cellular activities of ribulose 1,5 bisphosphate carboxylase (RUBPCase) or phosphoenol pyruvate carboxylase (PEPCase), or to variations in chlorophyll content. Daytime P_max peaks were about 12 h out of phase with nighttime maxima in the frequency of dividing cells (FDC). The phase relationship between P_max and FDC could be altered by manipulating environmental conditions. High light exposure of depp populations did not affect timing of the P_max peak, but its magnitude increased and coincided with increased RUBPCase activity and chlorophyll photobleaching. In the eddy population, a major shift in the timing of peak P_max was induced when increased light intensity was accompanied by nutrient enrichment. This change coincided with major increases in cellular chlorophyll and carboxylating enzyme activity. Lowering irradiance and/or increasing nutrient availability elicited different diel pattern in cellular metabolism in surface populations from the eddy and from Wilkinson's Basin that appeared linked to differences in the nutrient status of the cells. Rates of cell division estimated from the percentage of dividing cells in preserved samples were 0.83 divisions d^-1 in surface warm-core eddy populations, supporting the view that carbon and nitrogen turnover rates in oligotrophic waters can be sufficient to promote near optimal growth of Synechococcus spp.     

Short-term variability of photosynthesis in natural marine phytoplankton populations

Harmonic regression analysis has been used to determine the short-term variability in the photosynthetic rate (mgC/mg chlorophyll a/h) of phytoplankton in three inlets of Japan. In natural water without large zooplankton present, the photosynthetic rate [log P=log (100xmgC/mg chlorophyll a/h)] can be expressed as (B+A cos T). Factor B represents the average photosynthetic rate, of which the maximum is usually designated as P _max, and Factor A corresponds to the slope of the regression line. The phase of the periodicity, represented by T, is adjusted to give the highest correlation: usually T is expressed as [360/24 x (local time + 4)] in degrees. The correlation between Factors A and B is very high (r=0.95, P<0.001), indicating that Factor A may depend upon Factor B (potential activity of chlorophyll a). Both Factors A and B decrease with decreasing irradiance, but the slope of each regression between Factor A and irradiance varies with season. Continuous darkness reduces the phase of the periodicity to one cycle a day when phytoplankton has multiple cycles of photosynthetic rate per day. Adequate nutrient supply from zooplankton regeneration may cause an increase in Factor B; however, excess density of zooplankton decreases Factor A.     

Influence des conditions hivernales sur les productions phyto- et zooplanctoniques en Méditerranée Nord-Occidentale. II. Biomasse et production phytoplanctonique

Hydrological observations, and measurements of nutrient chemistry, plankton biomass, and production were carried out during the Médiprod I cruise of the R.V. “Jean-Charcot”. The March cruise was characterized by almost winter conditions, exhibiting strong vertical mixing of water masses in the offshore region and nutrient transport up to the photic zone. According to the working hypothesis, the strong vertically mixed area (e.g. Station 15) and the surrounding oligotrophic area are separated by an intermediate zone, where both nutrients and stability have produced phytoplankton-bloom conditions. During the April cruise, highest biomass and production rates were encountered everywhere in the offshore region, and especially in the previously mixed area of central divergence. Production was as high as 2 gC·m^-2 day^-1, and the standing crop of chlorophyll was 3 mg·m^-3; such values are rather important for the so called “poor” Mediterranean Sea. Salinity-phosphate and chlorophyll-phosphate diagrams are presented. Biomass and production rates are in agreement with the potential fertility based on the nutrient content of the waters. The disappearance of 1 μatg P·PO₄ by photosynthetic uptake corresponds to 7.7 mg chlorophyll a, which represents the autotrophic biomass remaining after grazing by the simultaneous zooplankton bloom. Biomass and production features are analyzed in regard to interaction of both nutrient availability and the stability of water masses. Stability conditions can be created either by intrusion of local mixing in a stratified oligotrophic area (“winter bloom”), or by thermal stratification of the upper layer (“spring bloom”). In the latter case, the highest biomasses are present in the zone where the nutrients were previously introduced by mixing. The oligotrophic situation remained constant during the two crunises in the surrounding coastal area, which is characterized by low-salinity water and, therefore, absence of vertical nutrient transport into the photic zone. Chlorophyll pigment concentration and photosynthetic rates in the “Cote d'Azur” region are similar to those in the “Provence” region; this situation may result more from upwelling of nutrient-rich intermediate water than from the mixing process which predominates in the latter region.     

Mesocosm studies of DCMU-enhanced fluorescence as a measure of phytoplankton photosynthesis

When measurements of in-vivo fluorescence are used to estimate photosynthesis in the field, the marked temporal and spatial variations in phytoplankton populations, and their nutrient and light histories, have produced varied results. Natural phytoplankton populations in large, flow-through mesocosms with different controlled nutrient and sewage sludge additions were sampled weekly from June to September 1984. Good correlations were observed between the increase in fluorescence upon the addition of DCMU (F) and both in-situ production and the parameters (P_m and ) of the photosynthesis-irradiance curve for these phytoplankton populations. Good correlations were also obtained between DCMU-enhanced fluorescence (F_DCMU) and chlorophyll a concentrations. The relationship between F and in-situ ¹⁴C production was consistent among mesocosms even in the face of major shifts from diatom-dominated to dinoflagellate-dominated populations. On the other hand, the F_DCMU:Chl a relationship was significantly different between mesocosms and related to species composition. It was concluded that F offers the possibility of rapidly and accurately indexing both in-situ production and the photosynthetic capacity of mixed phytoplankton populations.This study was supported by EPA cooperative agreement 810265-03 and NOAA grant NA-83-ABD-00008