Depth-dependent changes in chlorophyll fluorescence number at a Sargasso Sea station

Chlorophyll a-specific in vivo fluorescence exhibited depth-dependent changes in a Sargasso Sea phytoplankton community, decreasing from a maximum value at the surface to a minimum at 90m, and then increasing again below 90 m. This distribution pattern was not explained by irradiance conditions, diurnal variability, senescence in the deep population, or changes in light-absorption efficiency of chlorophyll a. However, a significant positive correlation was found between mean phytoplankton cell size and fluorescence number in the upper euphotic zone, where nutrient concentrations were low. We hypothesize that the direct cause for this observed correlation was nutrient limitation. In this picoplankton-dominated community, packaging effect was minimal. Under nutrient-limiting conditions, as mean cell size increases photosynthetic efficiency decreases and therefore fluorescence number increases. In the lower euphotic zone where nutrients were not limiting, changes in fluorescence number exhibited weak size-dependence and appeared to be related to species compositional changes.     

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.     

Photosynthesis and chlorophyll a fluorescence rhythms of marine phytoplankton

Circadian rhythms in photosynthesis were defined in field populations of phytoplankton. Measurements of carbon-dioxide fixation rates demonstrated that a diurnal periodicity of photosynthesis in samples incubated under natural light-dark (LD) cycles also were observed to continue in similar samples which had been photoadapted to constant dim light (LL) for 48 h. These changes in photosynthetic rates preceded sunset and sunrise, had daily amplitudes that ranged from 1.5 to 2.0, appeared to be independent of light-intensity, and displayed maxima about midday, while rates of dark fixation of carbon dioxide and the photosynthetic pigment content per cell were constant over the circadian cycle. Similar rhythmicity also was detected in room-temperature (22°C) chlorophyll a fluorescence yield, in both the obsence and presence of the photosynthesis inhibitor DCMU [3-(3,4-dichlorophenyl)-1, 1-dimethylurea]. However, the magnitude and timing of the fluorescence rhythm maxima seem to depend on wavelengths monitored and, in part, on the measuring technique used. Also, the circadian changes in the fluorescence intensity were abolished at low temperature (-60°C), and the shape of the emission spectra of chlorophyll fluorescence of cells in LD and LL did not change over time. The significance of the fluorescence rhythms with regard to chlorophyll a determinations and photosynthetic rates is discussed. It was concluded that there was sufficient similarity between circadian rhythms of photosynthesis in natural phytoplankton populations and in laboratory cultures of dinoflagellates to suggest that the mechanism of regulation may be the same for both of them.     

In situ characterization of phytoplankton from vertical profiles of fluorescence emission spectra

Vertical profiling of the upper ocean with a laser/fiber optic fluorometer enabled the determination of fluorescence emission spectra of photosynthetic pigments over small vertical scales. Simultaneous acquisition of phycoerythrin (PE) and chlorophyll (chl) emission spectra allowed in situ differentiation between PE-containing cells (cryptomonads and cyanobacteria) and other chl-containing autotrophs. Further, fluorescence spectral peak shifts associated with different species of PE-containing cells resulted in even finer scale in situ taxonomic differentiation. We found that the phycoerythrin fluorescence emission maxima shifted from 578 nm near the surface, to 585 m at the base of the shallow thermocline (30% light level), and to 590 nm below the thermocline at the base of the euphotic zone (1% light level). These shifts in peak emission coincided with a taxonomic change in the PE-containing cells (as determined from analysis of discrete bottle samples) from a greater proportion of Synechococcus spp. in the upper water column to a greater proportion of cryptomonads at the base of the euphotic zone. These results indicate that the composition of the phytoplankton assemblage may be assessed in situ without sample collection.     

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.     

A spatial model of phytoplankton patchiness

The one-dimensional theory of critical-length scales of phytoplankton patchiness is developed to include phytoplankton growth and herbivore grazing as functions of time and space. The critical-length scale L _c for the pathch is then determined by the initial spatial distribution and concentration of the limiting nutrient and herbivores in addition to the daily averaged values of the growth and loss processes. The response of an initial phytoplankton patch to the stresses of turbulent diffusion, nutrient depletion, light periodicity, and nocturnal or continuous herbivore grazing is investigated numerically for several oceanic conditions. Nocturnal grazing, while less stressful on primary production than continous grazing, results in lower phytoplankton standing stocks. Increase in biomass of vertically migrating zooplankton results in a net loss of nutrient which might otherwise be egested, recycled, and utilized in the euphotic zone under continuous grazing conditions. The Ivlev constant is shown via sensitivity analysis to be a significant parameter ultimately influencing phytoplankton production. It is demonstrated numerically that diffusion of phytoplankton cells from areas of high concentration to low concentration prevents the local extinction of the standing stock, thereby rendering a positive herbivore grazing-threshold unnecessary for ecosystem stability.     

Effects of light intensity on photosynthesis and dark respiration in six species of marine phytoplankton

Using an oxygen polarographic electrode, the shapes of photosynthetic curves and the effects of light on dark respiration in 6 species of marine phytoplankton wer examined. The species used were Skeletonema costatum, Ditylum brightwellii, Cyclotella nana (Thalassiosira pseudonana) (all Bacillariophyceae), Dunaliella tertiolecta (Chlorophyceae), Isochrysis galbana (Haptophyceae), and Gonyaulax tamarensis (Dinophyceae). A hysteresis was observed in all species examined with respect to increasing and decreasing light. Compensation light intensities varied by over 4 orders of magnitude, suggesting that the 1% light depth is an ambiguous measure of the euphotic zone. The data suggest that dark respiration accounts for ca. 25% of gross photosynthesis, but is species-dependent. In addition, respiration versus cell size does not describe an inverse exponential function over the size scales examined.This research was performed under the auspices of the United States Energy Research and Development Administration under Contract No. EY-76-C-02-0016.     

Light-shade adaptation by the oceanic dinoflagellates Pyrocystis noctiluca and P. fusiformis

Species-specific rates of photosynthetic carbon uptake (P), chlorophyll a content and P versus irradiance (P-I), have been measured for cells of Pyrocystis noctiluca and P. fusiformis isolated from natural populations collected in the euphotic zone within and below the surface mixed layer in the Sargasso Sea. These same measurements and the assay for ribulose bis-phosphate carboxylase (RuBP-Case), have been made for cultures of P. noctiluca in a 12 h L: 12 h D photoperiod at 9 different constant or at changing light intensities. In nature chl a cell^-1 was constant throughout the euphotic zone. The photosynthetic capacity (P_max), of cells captured below the surface mixed layer was lower by a factor of 10 compared with cells collected from the surface mixed layer. The P_max for P. noctiluca collected and incubated within the surface mixed layer was the same as for cell cultures grown under high light, non nutrient-limiting conditions, suggesting that photosynthesis in the natural system was not nutrient limited. In laboratory cultures under constant low light intensities, chl a cell^-1 increased by a factor of 5 while both P_max and RuBPCase activity decreased by a factor of ca 4 compared with high light intensities. In changing light intensities both P_max and RuBPCase activities were decreased by factors of 4 during low light intervals while chl a cell^-1 approached a constant intermediate value. The change in chl a cell^-1 in response to prolonged exposure to constant low light intensities was first order with a rate constant of 0.33 d^-1. For all irradiance conditions in culture, the P-I dependence could be described by the simple Michaelis-Menten formula. The ratio of P_max to K_I, (the light intensity where P=P_max/2) was a constant with a Coefficient of Variation of 12%: The constancy of this ratio, the parallel changes in RuBPCase activity with P_max and the constant chl a cell^-1 in the Sargasso Sea imply that for P. noctiluca and presumably P. fusiformis in nature, a dark enzymatic step rather than changes in photosynthetic pigment concentrations may regulate the photosynthetic capacity in the changing photic environment.Contribution no. 1141 from McCollum-Pratt Institute and Department of Biology, The Johns Hopkins University. Supported by DOE contract no. EY 76S20 3278, NSF no. OCE 76-02571 and ONR no. N300014-81-C-0062     

Une approche vers l'estimation de la production potentielle du phytoplancton par analyse des cinétiques d'induction de fluorescence

In vivo chlorophyll fluorescence is particularly interesting ot ecologists because of various concepts (biomass, productivity, physiological state) associated with it. Using a modified spectrophotofluorometer, we have studied the kinetics of fluorescence in unialgal cultures and in a natural population of marine phytoplankton. Our apparatus did not achieve satisfactory results with cell suspensions having a chlorophyll concentration less than 10 g l^-1. We have also tested a method for estimating kinetics of diluted cultures and marine phytoplankton using cells collected on glass-fibre filters. For unialgal cultures in the exponential growth phase, the method proved satisfactory, and results obtained from both cell suspensions and filters were in good agreement. However, for aged cultures (principally diatoms) and natural marine phytoplankton the method proved unsuitable. The kinetics of fluorescence induction vary according to taxonomic position of the cells, light intensity of the measuring excitation beam and productiveness of the culture medium. The importance of the kinetics of fluorescence induction for characterization of phytoplankton activity is discussed.     

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     

Time-course of photoadaptation in the photosynthesis-irradiance relationship of a dinoflagellate exhibiting photosynthetic periodicity

Cultures of the marine dinoflagellate Glenodinium sp. were light-shifted and rates of photoadaptation determined by monitoring changes in cell volume, growth rate, pigmentation, parameters of the photosynthesisirradiance (P-I) curves and respiration. To approximate physiological conditions of field populations, cells were cultured on an alternating light-dark cycle of 12hL:12hD, which introduced a daily periodicity of photosynthesis. One result of the present study was to demonstrate how specific parameters of the P-I relationship influenced by periodicity of the light: dark cycle are distinguished from photosynthetic parameters influenced by changes in light level. Under steady-state conditions, rates of both light-saturated (P_max) and light-limited photosynthesis changed in unison over the day; these changes were not related to pigmentation, and displayed their maxima midday. This close relationship between P_max and the slope (a) of the cellular P-I curves in steadystate conditions was quickly adjusted when growth illumination was altered. Rates of light-limited photosynthesis were increased under low light conditions and the periodicity of cellular photosynthesis was maintained. The short-term responses of the P-I relationship to changing light level was different, depending on (1) whether the light shift was from high to low light or vice versa, and (2) whether the high light levels were sufficient to promote maximal photosynthesis rates. Major increases in the photosynthetic carotenoid peridinin, associated with a single type of light-harvesting chromo protein in the chloroplast, was observed immediately upon shifting high light cultures to low light conditions. Following pigment synthesis, significant increases in rates of light-limited photosynthesis were observed in about one-tenth the generation time, while cellular photosynthetic potential was unaffected. it is suggested that general results were consistent with suggested that general results were consistent with earlier reports that the major photoadaptive strategy of Glenodinium sp. is to alter photosynthetic unit (PSU) size. Photoadaptive response times to high light were light-dependent, but appeared to be shower than photoadaptive responses to low light. If light intensities were bright enough to maximize growth rates, photosynthetic response times were on the order of a generation period and pigmentation fell quickly as cells divided at a faster rate. If light-intensities were not sufficient to maximize growth rates, then pigment content did not decline, while rates of light-limited photosynthesis declined quickly. In all cases, photoadaptation was followed best by monitoring fast changes in half saturation constants for photosynthesis, rather than fluctuating changes in pigmentation. Results compared well with time-course phenomena reported for other groups of phytoplankton. Overall, results suggest phytoplankton can bring about photo-induced changes in photosynthesis very quickly and thus accommodate widely fluctuating light regimes over short periods of time.     

Effects of symbiont elimination and reinfection on the life processes of the planktonic foraminifer Globigerinoides sacculifer

Laboratory experiments were carried out to determine the influence of symbiotic dinoflagellates (zooxanthellae) on the shell growth, longevity, and reproductive potential of Globigerinoides sacculifer (Brady). Its symbionts were eliminated by 72-h treatment with a photosynthetic inhibitor (DCMU). Symbiont elimination resulted in earlier gametogenesis (shortened survival time) and smaller shell sizes of G. sacculifer when compared to untreated foraminifera grown in sea water. Individuals kept in continuous darkness in untreated sea water also exhibited early gametogenesis, short survival times and small shell sizes. Aposymbiotic foraminifera formed on the average one or two chambers fewer per individual and their rate of shell size increase is slower than symbiont-bearing foraminifera. Symbionts were lysed within perialgal vacuoles of G. sacculifer when subjected to DCMU treatment or kept in continuous darkness. One DCMU-treated group was reinfected with symbionts from crushed G. sacculifer donors. Soon after reinfection, these foraminifera resumed a shell growth rate and exhibited developmental stages that were nearly equivalent to those of untreated individuals, as deduced from their shell size, frequency of sac-like chambers, rate of gametogenesis, and survival time. Our experiments indicate that the symbionts aid in calcification and that elimination of symbionts triggers gametogenesis, thus shortening the life span of the foraminiferal host. The results imply that shell growth in symbiont-bearing planktonic foraminifera occurs mainly in the euphotic zone and that they do not survive for long periods below it.     

Response of marine phytoplankton to nitrogen deficiency: Decreased nitrate uptake vs enhanced ammonium uptake

The uptake of nitrate and ammonium was measured separately in uni-algal, nitrogen-deficient cultures of four species of marine phytoplankton. Nitrogen-deficient phytoplankton took up ammonium at initial rates which greatly exceeded those measured for nitrogen-sufficient phytoplankton. However, nitrate uptake by nitrogendeficient cultures was generally much slower than either nitrate or ammonium uptake by nitrogen-sufficient cultures or ammonium uptake by nitrogen-deficient cultures. Considerable species differences were observed in the degree to which nitrogen deficiency increased ammonium uptake or decreased nitrate uptake. Loss of ability to take up nitrate, but enhanced ability to take up ammonium, as a result of nitrogen deficiency may be an adaptation to the different mechanisms by which nitrate and ammonium are supplied to the euphotic zone. In areas with an intermittent supply of nitrogen, changes in the ability of some species to take up nitrogen as a result of nitrogen starvation will influence species composition and complicate interpretations of measurements of nitrogen uptake.Contribution no. 1249 from the Department of Oceanography, University of Washington, and contribution no. 82006 from the Bigelow Laboratory for Ocean Sciences     

Short-term variability and control of phytoplankton photosynthetic activity in a macrotidal ecosystem (the Strait of Dover, eastern English Channel)

Short-term changes in phytoplankton photosynthetic activity were studied during different periods of the years 2009 and 2010 in the coastal waters of a macrotidal ecosystem (the Strait of Dover, eastern English Channel). During each sampling period, samples were taken every 1.45 h., from sunrise to sunset, during at least 5 days distributed along a complete spring–neap tide cycle. The photosynthetic parameters were obtained by measuring rapid light curves using pulse amplitude modulated fluorometry and were related to environmental conditions and phytoplankton taxonomic composition. The maximum quantum yield (F _v/F _m) showed clear light-dependent changes and could vary from physiological maxima (0.68–0.60) to values close to 0.30 during the course of 1 day, suggesting the operation of photoprotective mechanisms. The maximum electron transport rate (ETR_m) and maximal light utilization efficiency (α) were generally positively correlated and showed large diel variability. These parameters fluctuated significantly from hour to hour within each day and the intraday pattern of variation changed significantly among days of each sampling period. Stepwise multiple linear regressions analyses indicated that light fluctuations explained a part of this variability but a great part of variability stayed unexplained. F _v/F _m, ETR_m and α were not only dependent on the light conditions of the sampling day but also on those of the previous days. A time lag of 3 days in the effect of light on ETR_m and α variation was highlighted. At these time scales, changes in phytoplankton community structure seemed to have a low importance in the variability in photosynthetic parameters. The photoacclimation index E _k showed a lower variability and was generally different from the incident irradiance, indicating a limited acclimation capacity with a poor optimization of light harvesting during the day. However, in well-mixed systems such as the Strait of Dover, the short-term photoacclimation is disrupted by the high level of variability in environmental conditions. Also, the variability observed in the present study can be associated with a particular kind of photosynthetic response: the “E _k-independent” variability. The physiological basis of this photosynthetic response is largely unresolved and further researches on this subject are still required to better explain the dynamics of phytoplankton activity in the Strait of Dover.     

The effects of ammonium and phosphate enrichments on clorophyll a,pigment ratio and species composition of phytoplankton of Vineyard Sound

Seawater containing natural phytoplankton populations from Vineyard Sound, USA was enriched in the laboratory with three levels each of ammonium and phosphate and with a combination of ammonium and phosphate which provided three different N:P ratios. The addition of ammonium produced more cells and chlorophyll a than the control or the phosphate enrichments. However, enrichment with ammonium and phosphate, regardless of the N:P ratio, yielded the most cells and chlorophyll a. Thus, nitrogen seems to be the primary limiting nutrient, with phosphate showing secondary limiting effects. The ratios of photosynthetic pigments decreased with the increased chlorophyll a production in the enriched cultures. There were no significant changes in the species composition within the cultures, so that the observed changes in pigment ratio and chlorophyll a content were due to physiological responses.     

Development of the ability to take up L-lysine by the diatom Phaeodactylum tricornutum

Freshly harvested cells of Phaeodactylum tricornutum Bohlin grown with nitrate, ammonium or lysine as a sole nitrogen source had a low ability to take up lysine or arginine, but this ability increased when cells were deprived, over 48 h, of either nitrogen or carbon. The effects of nitrogen and carbon deprivation were additive, and the uptake ability was greatest in cells incubated in darkness in nitrogen-free medium. Uptake ability increased in cells illuminated in the presence of 10^-5 M 3-(3,4-dichlorophenyl)-1,1-dimethylurea (DCMU) an inhibitor of photosynthetic electron transport. An inverse relationship between rate of development of the uptake system and rate of photosynthesis was also established. Development of the uptake system was prevented by cycloheximide or by anaerobiosis. Following transfer to a normal nitrate medium, illuminated cells lost the lysineuptake system by dilution as the cells grew. There was a linear and positive correlation between the initial rate of uptake of lysine and the maximum concentration which was maintained in the cells when equilibrium was reached, indicating that transinhibition of lysine uptake may occur and that the extent of this inhibition is related to the size of the internal amino acid pool. The relevance of the findings to the growth of phytoplankton in natural waters is discussed.     

Chlorophyll a fluorescence in marine centric diatoms: Responses of chloroplasts to light and nutrient stress

Observations at sea of large variations in the cellular fluorescence of phytoplankton prompted a study of the fluorescence responses in marine diatoms to light and nutrient stress. When older cultures of Lauderia borealis were exposed to intense light, the in vivo fluorescence of chlorophyll a declined within the first 2 min of exposure. This initial response to light stress appeared to be correlated with a contraction of the chloroplasts. Continued exposure led to a second decline in fluorescence, which required 30 to 60 min for completion. A movement of chloroplasts to the valvar ends of the cell caused this secondary response. Both the contraction and intracellular movement of chloroplasts appeared to be related to both photoinhibition of photosynthesis and diel fluctuations in cellular fluorescence. An investigation of continuous cultures of Cyclotella nana showed that in vivo chlorophyll a fluoresced more strongly in nitrogen-starved cells than in enriched ones. Photoinhibition of cellular fluorescence also increased with the cell's state of nitrogen deficiency.     

Light-dependency of nitrate uptake by phytoplankton over the spring bloom in Auke Bay,Alaska

The effect of light intensity on nitrate uptake by natural populations of phytoplankton was examined by ¹⁵N traceruptake experiments during the spring (March–May 1987) in Auke Bay, Alaska. The data were fit to a rectangular hyperbolic model which included a term for dark uptake. Three types of curves described nitrate uptake as a function of light intensity. The first (Type I) had a low half-saturation light intensity (K _I), low chlorophyll-specific uptakes rates, no dark uptake and occasional photoinhibition. These were observed during a period of biomass decrease, accompanied by low daily light and strong wind, prior to the major bloom. The second type (Type II) had relatively high K _I, high chlorophyll-specific uptake rates, and no dark uptake. Type II curves were observed during most of the period prior to nitrate depletion in the surface waters. Types I and II both appeared prior to nitrate depletion in the water and reflected variations in the light history of the phytoplankton population. The third type (Type III) occurred in nitrate-deplete conditions, when nitrate uptake was less dependent on light intensity (i.e., high rates of dark uptake and lower K _I). Decreased light-dependency during this period was coupled with physiological nitrogen deficiency in the population. Comparing these parameters to those of photosynthetic carbon fixation, K _Ivalues of nitrate uptake were generally higher than those of photosynthesis prior to nitrate depletion, and lower during nutrient-deplete conditions.     

Vitamine B12 et phytoplancton dans l'Océan Antarctique. Distribution et approche expérimentale

In late summer (Antiprod I cruise, March 1977), antarctic waters of the southern part of the Indian Ocean contain low amounts of dissolved Vitamin B₁₂: 1 ng l^-1 on average. However, higher concentrations were recorded in two areas: in the subantarctic convergence zone and between 50° and 57°S, with values greater than 3,5 ng l^-1. Despite nutrient-rich waters, primary production was low and the phytoplankton sparse. Abundance of diatoms was related, to a certain extent, to the concentration of Vitamin B₁₂. An experimental approach (Antiprod II cruise, February and March 1980) was applied to (1) artificially enriched natural sea water, and (2) cultures of two diatom species isolated from antarctic waters, Nitzschia turgiduloides and Chaetoceros sp. The experiments demonstrated that antarctic diatoms in general do not require any of the three investigated vitamins for growth. However, Vitamin B₁₂ has a slightly stimulating effect on growth. This may explain the observed relation between phytoplankton density and vitamin concentration in the Antarctic Ocean. This experimental approach elucidates the minor role of vitamins in the primary production of antarctic waters and provides indirect evidence of the major importance of turbulence as a limiting factor.     

Photoadaption in marine phytoplankton: Response of the photosynthetic unit

Some species of phytoplankton adapt to low light intensities by increasing the size of the photosynthetic unit (PSU), which is the ratio of light-harvesting pigments to P700 (reaction-center chlorophyll of Photosystem I). PSU size was determined for 7 species of marine phytoplankton grown at 2 light intensities: high (300 E m^-2 s^-1) and low (4 E m^-2 s^-1); PSU size was also determined for 3 species grown at only high light intensity. PSU size varied among species grown at high light from 380 for Dunaliella euchlora to 915 for Chaetoceros danicus. For most species grown at low light intensity, PSU size increased, while the percentage increase varied among species from 13 to 130%. No change in PSU size was observed for D. euchlora. Photosynthetic efficiency per chlorophyll a (determined from the initial slope of a curve relating photosynthetic rate to light intensity) varied inversely with PSU size. In contrast, photosynthetic efficiency per P700 was enhanced at larger PSU sizes. Therefore, phytoplankton species with intrinsically large PSU sizes probably respond more readily to the rapid fluctuations in light intensity that such organisms experience in the mixed layer.Contribution No. 1180 from the Department of Oceanography, University of Washington, Seattle, Washington, USA