Influence of temperature on the growth rate of Skeletonema costatum in response to variations in daily light intensity

Daily light intensities (I _o) can vary 10-fold during the winter-spring and late-summer diatom blooms in New England, USA, coastal waters. Laboratory cultures and natural populations incubated in dialysis sacs were examined to determine the time course of growth rate in Skeletonema costatum (Greville) Cleve in response to variations in daily light intensity during two bloom periods in Narragansett Bay, Rhode Island, USA. Log-phase cultures of S. costatum require 2 d to attain maximum growth rates at 2°C following transfer to saturating intensities. At 20°C, only 1 d is required. As temperature increases, Detonula confervacea (Cleve) Gran, Thalassiosira nordenskiöldii Cleve and Ditylum brightwellii (West) Grunow also exhibit rapid increases in mean daily division rates (K) following transfer to saturating light intensities. Thalassiosira pseudonana Hustedt, however, did not alter the time required to achieve maximum K as temperature varied. Natural populations of S. costatum did not show a well-defined relationship between K and light. Throughout a winterspring bloom, K was limited by low temperatures and exhibited no clear response to variations in I _o. A change in K in response to variation in I _o may occur on a daily basis during the summer, when temperatures are near 20°C; this has yet to be verified for in situ populations.     

Interactions between nitrate and ammonium uptake: variation with growth rate,nitrogen source and species

The interaction between nitrate and ammonium uptake was examined as a function of preconditioning growth rate and nitrogen source by adding nitrate, ammonium, or both to nitrogen-sufficient,-deficient, and-starvedSkeletonema costatum (Grev.) Cleve and nitrogen-deficientChaetoceros debilis Cleve. By simultaneously measuring the internal accumulation of intermediates of nitrogen assimilation and the rates of nitrogen assimilation, the metabolic control of nitrogen uptake could be assessed. After the simultaneous addition of nitrate and ammonium to culture, both nitrate and ammonium uptake rates were decreased in comparison with the rates observed when each was added alone, although nitrate uptake was usually decreased more than ammonium uptake. Since both nitrate and ammonium uptake rates vary with time, preconditioning growth conditions, nitrogen sources present, and species, it was necessary to use several different indices to quantify inhibition. In general, ammonium inhibition of nitrate uptake inS. costatum was greatest in cultures preconditioned to ammonium and those at low growth rates, whereas ammonium uptake was inhibited most in cultures preconditioned to nitrate. In nitrogen-deficientC. debilis, nitrate uptake was more inhibited by ammonium, but uptake returned to normal rates more quickly than inS. costatum, whereas inhibition of ammonium uptake was similar. These results explain why the interaction between nitrate and ammonium uptake in the field can be so variable. Inhibition of uptake is not controlled by internal ammonium or total amino acids, nor is it related to the inability to reduce nitrate. Instead, inhibition must be determined in part by the external concentration of nitrogen compounds and in part by some intermediate(s) of nitrogen assimilation present inside the cell.Bigelow Laboratory Contribution No. 82022     

Concentration Requirement of glycine as nitrogen source for supporting effective growth of certain marine microplanktonic algae

Eighteen species of marine phytoplankton from 9 algal classes were tested under axenic-culture conditions for their capacity to improve phototrophic-growth performance on glycine with increasing concentration of this amino-acid serving as sole nitrogen source. Whereas all these species showed poor-to-no growth on 0.5 mM glycine, 14 species manifested considerable gain in growth-rate and yield at higher concentration levels. The highest level tested (25 mM) was generally not inhibitory and produced the best yields in these cases. Some species (notably Skeletonema costatum) showed marked reduction in the adaptation-lag period with increased glycine concentration, but Chlamydomonas palla required a very long adaptation period, which was little affected by concentration. Dunaliella tertiolecta, Emiliania huxleyi and Chroomonas salina showed sharp (albeit elevated) concentration thresholds for efficient glycine utilization. Rhodomonas lens and two cyanophycean strains, Agmenellum quadruplicatum and Anacystis marina, appeared incapable of successful growth on glycine at all concentrations tested, while the rhodophyte Porphyridium marinum displayed marginal growth. The results were insufficient to draw taxonomically general conclusions on algal-class potential for glycine utilization, but they showed unequivocally that more than 83% of phytoplankton species from 9 algal classes can utilize glycine for growth if the appropriate substrate concentration is provided.     

Nitrate uptake in marine phytoplankton: Energy sources and the interaction with carbon fixation

Field studies of whole natural phytoplankton communities from Knight Inlet, B. C., Canada and laboratory cultures of the diatom Skeletonema costatum indicate inorganic carbon fixation may be temporarily suppressed following 10 to 15% enrichment with NO ₃ ^- or NH ₄ ⁺ . (This effect is suggested to be due to competition between inorganic carbon and nitrogen for adenosine triphosphate (ATP), and is reduced when chlorophyll a is increased intracellularly after 6 to 8 h.) Results imply that the source of ATP for nitrate uptake is primarily from Photosystem I (cyclic photophosphorylation) in the presence of light. It would appear that a transient nutrient-adaptive response occurs upon addition of extracellular nitrogen.     

Polychlorinated biphenyl (PCB) effects on marine phytoplankton photosynthesis and cell division

Inhibition of photosynthesis and cell division by polychlorinated biphenyls (PCBs) was studied using 7 marine phytoplankton species representing 4 algal classes. PCB concentrations as low as 1.0 g l^-1 reduced cell division of Thalassiosira pseudonana 3H and Isochrysis galbana. Both photosynthesis and cell division of T. pseudonana 3H, Chaetoceros socialis, Skeletonema costatum, T. pseudonana 13-1, Monochrysis, lutheri and I. galbana were inhibited at a PCB concentration of 10.0 g l^-1. The effects on photosynthesis were immediate and probably resulted in reduced rates of cell division. Interspecific differences in susceptibility were observed. These differences have significance with respect to primary production and the species composition of phytoplankton communities. The initial slopes of photosynthesis-irradiance (P-I) curves for the diatoms S. costatum and T. pseudonana 3H were reduced in the presence of PCBs. These results suggest that PCBs affect the photosynthetic light reactions.     

Diel variation in chlorophyll a,carbohydrate and protein content of the marine diatom Skeletonema costatum

Skeletonema costatum (Greville) Cleve isolated from Narragansett Bay, USA, was incubated at 3 light intensities (ca. 0.008, 0.040 and 0.075 ly min^-1) under a 12 h light: 12 h dark (12L:12D) photoperiod at 2°, 10° and 20°C. Cellular chlorophyll a increased at intensities less than ca. 0.040 ly min^-1; increases occured within one photoperiod at temperatures above 10°C. Cellular carbohydrate increased with light intensity at all temperatures; increases during the photophase were due to net production of the dilute acid-soluble fraction. Cellular protein increased during the photoperiod at 10° and 20°C; there was little difference in cellular protein among all cultures after one photoperiod. The rate at which cellular chlorophyll a increased in response to a decrease in light suggests that diel variation in cellular chlorophyll a is temperature-dependent in S. costatum. Protein: carbohydrate ratios ranged from ca. 0.5 to 2.0 over a diel cycle; ratios increased at lower intensities and higher temperatures. The diel range in protein:carbohydrate ratios equals that in cultures developing nitrogen deficiency; thus, use of this ratio as an index to phytoplankton physiological state must account for diel light effects.     

Spring bloom sedimentation in a subarctic ecosystem

Results from a 5-yr study (1985 to 1989) in Auke Bay, Alaska show that termination of the spring bloom consistently occurred at limiting nitrate concentrations. Following nutrient exhaustion, phytoplankton sinking rates increased and displayed greater temporal variability. Threshold nitrate concentrations, approximating K_s values of the species present, were found to signal initiation of increased sedimentation. For Thalassiosira aestivalis, the threshold was ～2 μmol l^-1, while for Skeletonema costatum the threshold was ～1 μmol l^-1, suggesting genus-specific differences in sinking-rate sensitivity to nitrate exhaustion. Overall, sinking rates of the three principal genera ranked (high to low) Thalassiosira spp.> S. costatum>Chaetoceros spp., while the nitrate sensitivities of the sinking rates of the genera ranked (high to low) Thalassiosira spp.> Chaetoceros spp.> S. costatum. Thalassiosira spp. showed the most consistent sinking rate increases following nutrient impoverishment. During a bloom dominated by T. aestivalis, a decrease of cell sinking rate with depth coincided with a decrease in short-term nutrient stress as measured by intracellular nitrate pools. In addition, no correlation was found between chain length or aggregate formation and sinking rate for this species. Though we measured only small-scale cell-cell adhesion, not larger-scale marine snow formation, this supports the notion that the sinking rates of Thalassiosira spp. were controlled primarily by cell physiology. For S. costatum, however, shorter chains sank faster. The sinking behavior of the species studied here figures prominently in their pelagic ecology and in the carbon flux of coastal ecosystems, both of which are driven by short-term variability.     

Effect of nitrogen source on short-term light and dark CO2 uptake by a marine diatom

Based on a series of short-term incubations involving the marine diatom Chaetoceros simplex (Bbsm), precultured in NH ₄ ⁺ -, NO ₃ ^- -and urea-limited continuous cultures at several dilution rates, we found that both the short-term specific rate of ¹⁴CO₂ uptake and the amount of CO₂ fixed after 8- and 16-min incubations were unaffected by enrichment with NH ₄ ⁺ , urea, or NO ₃ ^- when NH ₄ ⁺ or urea were the preconditioning forms of N, but were slightly suppressed when the cells were first grown on NO ₃ ^- . Similar enrichments in the dark, however, led to significant CO₂ uptake under all conditions of NH ₄ ⁺ enrichment and to similarly enhanced CO₂ uptake, but only at high growth rates, when urea was the source of enrichment nitrogen. Our light results are contrary to some contemporary findings, but there does seem to be agreement that photosynthetic rates of rapidly growing phytoplankton will not be affected by exposure to pulses of nitrogen. Enhanced dark uptake, in contrast, appears to be characteristic of phytoplankton under all degrees of N limitation, and, as such, may be useful as an “all or nothing” index of the nitrogen status of natural waters. There is some indication that the index may be useful in determining both the form of and the degree of N limitation as well.     

Coagulation efficiency and aggregate formation in marine phytoplankton

Flocculation of phytoplankters into large, rapidly sinking aggregates has been implicated as a mechanism of vertical transport of phytoplankton to the sea floor which could have global significance. The formation rate of phytoplankton aggregates depends on the rate at which single cells collide, which is mainly physically controlled, and on the probability of adhesion upon collision (=coagulation efficiency, stickiness), which depends on physico-chemical and biological properties of the cells. We describe here an experimental method to quantify the stickiness of phytoplankton cells and demonstrate that three species of diatoms grown in the laboratory (Phaeodactylum tricornutum, Thalassiosira pseudonana, Skeletonema costatum) are indeed significantly sticky and form aggregates upon collision. The dependency of stickiness on nutrient limitation and growth was studied in the two latter species by investigating variation in stickiness as batch cultures aged. In nutrient repleteT. pseudonana cells stickiness is very low (< 5 × 10^?3), but increases by more than two orders of magnitude as cell growth ceases and the cells become nutrient limited. Stickiness ofS. costatum cells is much less variable, and even nutrient replete cells are significantly sticky. Stickiness is highest (> 10^?1) forS. costatum cells in the transition between the exponential and the stationary growth phase. The implications for phytoplankton aggregate formation and subsequent sedimentation in the sea of these two different types of stickiness patterns are discussed.     

Measurements of electron transport activities in marine phytoplankton

A simple and sensitive spectrophotometric method of measuring electron transfer in the transport system (ETS) in marine phytoplankton has been developed and characterized. The assay is based on the reduction of the tetrazolium salt 2-(p-iodophenyl)-3-(p-nitrophenyl)-5-phenyl tetrazolium chloride (INT) by homogenates in the presence of the nonionic detergent Triton X-100. The simplicity and sensitivity of this assay have considerable advantage over existing methods of measuring ETS activity in marine samples where numerous assays must be rapidly carried out on samples which often demonstrate low activities. Several established substrates and inhibitors of electron transport in the ETS were investigated, and each is discussed in relation to the site of INT reduction in the diatoms Skeletonema costatum and Chaetoceros debilis and in the unicellular green alga Dunaliella tertiolecta. The results from these studies suggest that the maximum rate of electron transport, V _max, is measured in each case.     

Grazing rates of the Atlantic menhaden Brevoortia tyrannus as a function of particle size and concentration

The feeding behavior of adult Atlantic menhaden (Brevoortia tyrannus) upon 5 species of phytoplankton and 2 species of zooplankton has been studied. Four recognizable feeding stages which were a function of the concentration and size of the food particles were observed. During rapid feeding the fish swam at a constant speed for a prolonged period over a wide range of particle concentrations. Particle and food carbon-concentrations at the threshold for initiation and termination of feeding were inversely related to particle size. Carteria chuii (13.2 μ) was not grazed at a significant rate, while two-cell chains of Skeletonema costatum (16. 5 μ) were filtered from the water, indicating a minimum-size threshold for filtration of between 13 and 16 μ. The most rapid filtering rates were observed for the copepod Acartia tonsa ( \(\bar x\) volume swept clear = 24.8 l/fish/min). The maximum food-particle size acceptable to a menhaden appears to be between Acartia tonsa (1200 μ) and adult Artemia salina (10 mm). These results suggest that the large schools of menhaden found in Atlantic coastal waters could have a significant effect on the plankton, selectively grazing zooplankton, larger phytoplankton, and the longer chains of chain-forming diatoms.     

Feeding responses of the tentaculate deposit-feederEupolymnia nebulosa (Annelida: Polychaeta): Influence of sexual maturity

Feeding responses of the tentaculate depositfeeding polychaeteEupolymnia nebulosa (Montagu) were studied by measuring rates of uptake of three different¹⁴C-labelled diatoms (unialgal cultures ofNavicula incerta Grunow,Nitzschia acicularis Wm Smith, andNitzschia sp.). Worms used during this study were collected in the harbor of Port-Vendre (Western Mediterranean) during August 1986 (immature worms) and December 1987 (mature worms). Uptake rates were affected both by the length of the experiments and by the nature of the food offered. The highest rate of uptake (2.98 10^–4 mg ashfree dry wt of algae mg^–1 dry wt of worms h^–1) was obtained during short-term experiments (4 h) with the smallest diatom (Nitzschia sp.). The lowest rate of uptake (0.21 10^–4 mg ash-free dry wt of algae mg^–1 dry wt of worms h^–1) was also obtained withNitzschia sp., but for a long-term (48 h) experiment. There was no significant difference between rates of uptake of immature and mature worms.     

Consumption of diatoms and diatom filtrates by the tentaculate deposit-feederEupolymnia nebulosa (Annelida: Polychaeta)

Rates of organic uptakes of three live diatoms (Nitzschia acicularis, Nitzschia sp. andNavicula incerta), and of three corresponding filtrates by the deposit-feeding polychaeteEupolymnia nebulosa (Montagu) were measured under similar experimental conditions. Worms used during this study were collected by SCUBA diving at Port-Vendres in shallow water (7 m deep) during the summer of 1986. Uptake rates of live diatoms were affected both by length of the experiments and by the nature of the food offered. The highest rate of uptake (11.8 10^–4 mg algal ash-free dry wt mg^–1 worm dry wt h^–1) was recorded during a short-term experiment (4 h) with the smallest diatom (Nitzschia sp.). The lowest rate (1.1 10^–4 mg algal ash-free dry wt mg^–1 worm dry wt h^–1) was recorded during a long-term experiment (48 h) with the largest diatom (Nitzschia acicularis). Filtrates ofN. acicularis were more readily utilized than those ofNitzschia sp. andNavicula incerta. Because of differences in uptake rates of algal filtrates as a function of species, it is not possible to evaluate the bias due to interaction with dissolved substances in experimental studies assessing ingestion of live benthic diatoms byE. nebulosa.     

Sinking rate response to depletion of nitrate,phosphate and silicate in four marine diatoms

The effect of nutrient (N, P, and Si) depletion on sinking rates was studied for two small (Skeletonema costatum (Grev.) Cleve and Chaetoceros gracile Schütt) and two large [Ditylum brightwellii (West) Grun and Coscinodiscus wailesii (Gran et Angst)] centric diatoms obtained from stock cultures. Each diatom was examined under conditions of (1) nutrient repletion (=log growth phase), (2) nutrient depletion (48 h without a given substrate), and (3) recovery (24h after addition of limiting substrate to nutrient-deplete populations). All nutrient-replete cultures displayed low sinking rates despite large differences in cell size. In nutrient-deplete populations, sinking rate was related to the kind of nutrient depleted and varied among species. Silicate depletion elicited by far the greatest increase in sinking rates in all 4 species, indicating that biochemical aspects of silicon metabolism are more important to buoyancy regulation than density-related variations in the amount of silicon per cell. Since N- and P-depletion caused lower sinking rates in 3 of the species, this observation calls for re-evaluation of the axiom that nutrient depletion necessarily causes increased sinking rates. The exception was Coscinodiscus wailesii, which sank faster under all types of nutrient limitation. In most cases, sinking rates typical of log-phase cultures were not regained within 24 h after the addition of limiting nutrient to nutrient-depleted populations. Ultimately, the length of the recovery period may be useful in identifying the metabolic processes responsible for buoyancy regulation in actively growing cells.     

Simultaneous nitrogen and silicate deficiency of a phytoplankton community in a coastal jet-front

The nutritional status of a phytoplankton community was investigated in a coastal jet-front located in the Gulf of St. Lawrence, Canada, in 1987. During the sampling period, the frontal community was mainly composed of the diatomsChaetoceros debilis, Skeletonema costatum, Thalassiosira gravida andC. pelagicus. As previously reported for the St. Lawrence, some frontal stations were depleted both in nitrate and silicate. At stations impoverished in nitrate, internal nitrate pool concentrations were low or undetectable, suggesting that cells had not, recently, been exposed to a nitrate flux which exceeded the nitrate assimilation rate. At these impoverished stations, however, ambient and intracellular concentrations of ammonium and urea were high, suggesting that the community was not nitrogen-deficient. The comparison between the ambient silicate concentrations and the silicate requirement (K _s ) of the dominant diatoms suggests thatC. debilis andS. costatum were Si-deficient. This is further supported by the low silicate uptake rates and intracellular concentrations measured at the silicate impoverished stations. The silicate deficiency also resulted in a decrease in the seston and phytoplankton N:C ratios.Please address all correspondence and requests for reprints to Dr Levasseur at his new address: Maurice Lamontagne Institute, 850 Route de la Mer, Mont-Joli, Québec G5H 3Z4, Canada     

Ecological growth strategies in the seaweeds Gracilaria foliifera (Rhodophyceae) and Ulva sp. (Chlorophyceae): Soluble nitrogen and reserve carbohydrates

The seaweeds Gracilaria foliifera (Rhodophyceae) and Ulva sp. (Chlorophyceae) were grown in an outdoor continuous-flow system at both ambient incident light (I₀) and 0.13 I₀. During the winter, both species accumulated substantial soluble nitrogen reserves (up to 1020 g-at N·g dry wt^-1 in G. foliifera and 630 g-at N·g dry wt^-1 in Ulva sp.). The rate at which these N reserves were depleted was proportional to the growth rate. Seaweeds grown at 0.13 I₀ had lower growth rates and higher levels of soluble tissue N than plants grown at I₀. During the spring-summer growing season, peaks in tissue N followed nutrient peaks in the ambient seawater. Ulva sp. had higher nutrient uptake and growth rates than G. foliifera and showed greater fluctuations in soluble tissue N. This may characterize opportunistic seaweed species with high biomass turnover rates. At I₀, the levels of starch (up to 340 mg·g dry wt^-1 in G. foliifera and 170 mg·g dry wt^-1 in Ulva sp.) were highest during the spring and summer. During this period, fluctuations in starch content were inversely related to growth rate and soluble tissue N. Seaweeds grown at 0.13 I₀ did not accumulate starch. Neither species was found to overwinter with starch reserves.     

Production of glycine betaine and dimethylsulfoniopropionate in marine phytoplankton. I. Batch cultures

The quantitative significance of the nitrogenous compound glycine betaine (GBT) and its sulfur analog dimethylsulfoniopropionate (DMSP) to intracellular pools in marine phytoplankton is not well known. In a series of experiments conducted in August 1993, we measured these compounds, as well as total organic sulfur, carbon, and nitrogen, over the growth cycle in six isolates of marine phytoplankton, Amphidinium carterae Hulburt, Chrysochromulina sp. Lackey, Emiliania huxleyi Hay et Mohler, Prorocentrum minimum (Pavillard) Schiller, Skeletonema costatum (Greville) Cleve, and Tetraselmis sp. At the same time, we measured cellular concentrations of protein, amino acids, chlorophyll, and inorganic nutrients. All six species produced DMSP, while three produced GBT at lesser levels. In the Chrysochromulina sp. isolate, levels of GBT were greater than DMSP during the exponential phase of growth, but declined sharply as the culture approached stationary phase. This change appeared to coincide with the onset of nitrogen limitation. Other nitrogenous osmolytes were produced in five of the six species but in much smaller quantities. DMSP contributed significantly to cellular sulfur throughout the growth cycle although, in some algae, the proportion of dissolved DMSP increased substantially during stationary growth. When present, GBT formed a sizeable fraction of the cellular nitrogen only during exponential growth. A significant percentage (ca. 50%) of the organic nitrogen could not be accounted for even when cellular pools of protein, amino acids, inorganic nitrogen, and nitrogenous osmolytes were combined. Based on these experiments, there does not appear to be a reciprocal relationship between DMSP and GBT production, although GBT production does appear to be correlated with nitrogen availability. Received: 5 January 1998 / Accepted: 29 June 1999     

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     

Uptake and release of phosphorus by phytoplankton in the Chesapeake Bay estuary,USA

The phytoplankton uptake and release rates for inorganic phosphate, dissolved organic phosphate and polyphosphate were estimated during 5 cruises on the Chesapeake Bay over a 9-month period. Phosphorus in all pools turned over in several minutes to 100 h, and each soluble pool appeared to contain fractions which were metabolically useful to the phytoplankton. Maximal uptake rates (V _m ) for orthophosphate ranged from 0.02 to 2.95 μg-at P (1.h)^-1 with half saturation constants (K _s ) between 0.09 and 1.72 μg-at P l^-1. At low soluble reactive phosphorus concentrations, the uptake rate of trace ³²P orthophosphate was initially rapid, but declined after 15 to 60 min incubation. The data suggest that the initial uptake phase was dominated by exchange of ³²PO₄ ^≡ for ³¹PO₄ ^≡ in the membrane transport systems whereas the subsequent phase represented the net incorporation of orthophosphate into phytoplankton cells.     

Photoadaptation of photosynthesis in Gonyaulax polyedra

Gonyaulax polyedra Stein exhibited a combination of photoadaptive strategies of photosynthesis when only a single environmental variable, the light intensity during growth, was altered. Which of several biochemical/physiological adjustments to the light environment were employed depended on the level of growth irradiance. The photoadaptive strategies employed over any small range of light levels appeared to be those best suited for optimizing photosynthetic performance and not photosynthetic capacity. (Photosynthetic performance, P _i, is defined as the rate of photosynthesis occurring at the level of growth irradiance.) Among all photosynthetic parameters examined, only photosynthetic performance showed a consistent correspondence to growth rates of G. polyedra. Above 3500 to 4000 W cm^-2, where photosynthetic performance was equal to photosynthetic capacity, cells were not considered light-limited in either photosynthesis or growth. At these higher light levels, photosynthetic perfomance, cell volume, growth rates and respiration rates remained maximal; photosynthetic pigment content varied only slightly, while the photosynthetic capacity of the cells declined. At intermediate light levels (3000 to 1500 W cm^-2), photosynthesis, not growth, was light-limited, and photoadaptive strategies were induced which enhance absorption capabilities and energy transfer efficiencies of chlorophyll a to the reaction centers of G. polyedra. Photosynthetic capacity remained constant at about 280 mol O₂ cm^-3 h^-1, while photosynthetic performance ranged from 100 to 130 mol O₂ cm^-3 h^-1. Major increases in photosynthetic pigments, especially peridinin-chlorophyll a-proteins and an unidentified chlorophyll c component, accompanied photoadaptation to low irradiances. Maximal growth rates of 0.3 divisions day^-1 were maintained, as were respiration rates of about-80 mol O₂ cm^-3 h^-1 and cell volumes of about 5.4×10^-8 cm^-3 cell^-1. Below about 1250 W cm^-2, photosynthesis in G. polyedra was so light-limited that photosynthetic performance was unable to support maximal growth rates. Under these conditions, G. polyedra displayed photostress responses rather than photoadaptive strategies. Photostress was manifested as reduced cell volumes, slower growth, and drastic reductions in pigmentation, photosynthetic capacity, and rates of dark respiration.