Marine diatoms grown in chemostats under silicate or ammonium limitation. II. Transient response of Skeletonema costatum to a single addition of the limiting nutrient

Skeletonema costatum was grown at different steady-state growth rates in ammonium or silicate-limited chemostats. The culture was perturbed from its steady-state condition by a single addition of the limiting nutrients ammonium or silicate. The transient response was followed by measuring nutrient disappearance of the liliting perturbation experiment indicate that three distinct modes of uptake of the limiting nutrient can be distinguished; surge uptake (V _s ), internally controlled uptake (V _i ), and externally controlled uptake (V _e ). An interpretation of these three modes of uptake is given and their relation to control of uptake of the limiting nutrient is discussed. The uptake rates of the non-limiting nutrients were shown to be depressed during the surge of the uptake of the limiting nutrient. Kinetic uptake parameters, K _s and V _max, were obtained from data acquired during the externally controlled uptake segment, V _e . The same V _max value of 0. 12 h^-1, was obtained under either silicate or ammonium limitation. Estimates of K _s were 0.4 g-at NH₄-N l^-1 and 0.7 g-at Si l^-1. Short-term ¹⁵N uptake-rate measurements conducted on nitrogen-limited cultures appear to be a combination of V _s or V _i , or at lower substrate concentrations V _s and V _e . It is difficult to separate these different uptake modes in batch or tracer experiments, and ensuing problems in interpretation are discussed.Contribution No. 882 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA. This work represents portion of three dissertations submitted to the Department of Oceanography, University of Washington, Seattle, in partial fulfillment of the requirements for the Ph.D. degree.     

Physioecology of zooplankton. II. Effects of phytoplankton concentration,temperature, and body size on the development and molting rates of Calanus pacificus and Pseudocalanus sp.

Developmental time and stage duration for Calanus pacificus Brodsky and Pseudocalanus sp. and the rate of loss of body carbon by molting for C. pacificus were estimated for copepodite stages cultured under various combinations of phytoplankton concentration and temperature. Mean development time and stage duration for C. pacificus decreased hyperbolically with increasing food concentration, and the minimum time required for reaching a given stage decreased logarithmically with a logarithmic increase in temperature. Low temperature retarded the development of early stages proportionally more than that of late stages, and stage duration increased logarithmically with increasing body weight. Therefore, copepodite development was not isochronal. The rate of loss of body carbon by molting was small, ranging from 0.2 to 2% day^-1. This rate increased hyperbolically with food concentration and was linearly related to the growth rate. The critical food concentration for the rates of development and molting increased with temperature and stage of development, but these rates were less dependent on food concentration than the growth rate. The development rate of Pseudocalanus sp. was higher than that of C. pacificus, and was less influenced by changes in food concentration and temperature. It is postulated that the inverse relationship between temperature and body size results from a differential effect of temperature and body size on the rates of growth and development. That is, with increasing body size the growth rate tends to become temperature-independent, but the development rate remains proportional to temperature. Thus, copepodites growing at low temperature can experience a greater weight increment between molting periods than individuals growing at high temperature, because the growth rate is similar at all temperatures but stage duration is longer at low temperature.Contribution No. 1128 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Life history of the sandy-beach amphipod Dogielinotus loquax (Crustacea: Dogielinotidae) from the outer coast of Washington,USA

Dogielinotus loquax Barnard, a common intertidal macroinvertebrate on exposed ocean beaches near Grays Harbor, Washington, USA, is distributed from mid to mean high tidal levels, with a mean density of 1 830 m^-2 in 1975–1976. Location of maximum density within the intertidal zone is related to surf intensity and varies seasonally. The species is iteroparous and has an average sex ratio of 1:1. Temperature constraints on growth and egg development rates apparently influence the timing of reproduction. Two dominant recruitments occur per year, one in early spring (the summer generation) and another in late summer (the overwintering generation). The sampling design and density estimates permitted estimation of the mortality rate for the summer generation. Shorebird predation is suspected to affect summer generation abundance. Dogielinotus loquax will soon be elevated to the type of a new genus, Proboscinotus (Bousfield and Tzvetkova, in press)Contribution no. 1271 from the Department of Oceanography, University of Washington; Seattle, Washington 98195, USA     

Blooms of surf-zone diatoms along the coast of the Olympic Peninsula,Washington. XI. Regeneration of ammonium in the surf environment by the Pacific razor clam Siliqua patula

A 20 month field study was conducted on ammonium excretion rates of the Pacific razor clam Siliqua patula Dixon along the beaches of Washington State, USA. Excretion rates of all those nutrients likely to be regenerated in sufficient quantity to affect surf diatom growth were measured; ammonium appeared to be the most important metabolite. Excretion of ammonium by razor clams far exceeded that by other beach fauna. Ammonium excretion rates of razor clams were positively correlated with shell length, but no correlation between ammonium excretion rate and water temperature was evident. This may be an artifact or may represent some degree of seasonal acclimation of the species to temperature. Weight-specific ammonium excretion rates were negatively related to clam size, indicating a possible large (and unknown) contribution of regenerated ammonium by smaller clams in their first year of growth; smaller clams were rarely captured during this study.Contribution No. 1048 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA.     

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     

Persistent blooms of surf diatoms along the Pacific coast,USA

Productivity was studied in two diatom species, Chaetoceros armatum T. West and Asterionella socialis Lewin and Norris, which form persistent dense blooms in the surf zone along the Pacific coast of Washington and Oregon, USA. Past observations have shown that surf-diatom standing stock usually declines in summer along with concentrations of nitrate and ammonium. Using the ¹⁴C method, photosynthetic rates in natural surf samples were measured monthly for one year (October 1981 through September 1982) at a study site on the Washington coast. Also measured were temperature, salinity, dissolved nutrients, particulate carbon and nitrogen (used as estimates of phytoplankton C and N), and chlorophyll a. Assimilation numbers (P _max) were higher in summer (5 to 8 g C g^-1 chl a h^-1) than in winter (3 to 4gC). Specific carbon incorporation rates (µ_max) showed no obvious seasonality, mostly falling within the range of 0.09 to 0.13 g C g^-1 C(POC) h^-1. The discrepancy between the seasonal trends for chlorophyll-specific and carbon-specific rates reflects a change in the carbon-to-chlorophyll ratio. Because of seasonal differences in daylength and light intensity, daily specific growth rates () are thought to be higher in summer than in winter. Neither ammonium enrichment assays nor particulate carbon-to-nitrogen ratios provided convincing evidence for nitrogen limitation during summer, and the observed changes in diatom abundance cannot be explained on this basis. Both the high diatom concentrations and their seasonal variations probably are due mainly to factors other than growth rates; two factors considered important are diatom flotation and seasonal changes in wind-driven water transport. C. armatum usually dominates the phytoplankton biomass in the surf zone, and evidence suggests that this species is strongly dominant in terms of primary production.Contribution No. 1391 of the School of Oceanography, University of Washington, Seattle, Washington, USA     

Physioecology of zooplankton. III. Effects of phytoplankton concentration,temperature, and body size on the metabolic rate ofCalanus pacificus

Weight-specific rates of oxygen consumption of actively feeding copepodite stages ofCalanus pacificus Brodsky were measured under various combination of phytoplankton concentration and temperature. The rate decreased logarithmically with a logarithmic increase in dry body weight of copepods, and the relationship between these variables was described using a log-transformed allometric equation. The body-size dependence of the metabolic rate was independent of changes in food concentration and temperature, but the metabolic level increased linearly with a logarithmic increase in temperature and was not significantly affected by changes in food concentration. Respiration rates measured in this study forC. pacificus were about twice as high as rates reported for unfed closely related species of the same genus. An analysis of the metabolic cost of feeding processes suggests that metabolic models derived from feeding models may be of little ecological value at present.Contribution No. 1129 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Switching between herbivory and carnivory by the planktonic marine copepod Calanus pacificus

Adult females of the omnivorous copepod Calanus pacificus, collected from the plankton off La Jolla, California, USA (June, 1978), fed disproportionately on the prey in greatest relative abundance when given mixtures of diatoms (Thalassiosira fluviatilis) and copepod (C. pacificus) nauplii as food. This switch from herbivorous to carnivorous behavior may be significant in nature during the decline of phytoplankton blooms. More generally, the widespread omnivorous habit among pelagic animals suggests a responsive and flexible trophic organization which contributes to the resiliency of planktonic communities in a dynamic physical environment.Contribution No. 1223 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Ingestion-independent rates of ammonium excretion by the copepod Calanus pacificus

The relationship between food ingested and NH ₊ ⁴ excretion rate was investigated for female Calanus pacificus collected in August, 1982, from the San Juan Archipelago, Washington State, USA. The copepods were preconditioned to 6 densities of the diatom Thalassiosira weissflogii (0 to 10⁴ cells ml^–1) for 30 h before the experiment. The experiment was conducted with nutrients added in excess to maintain equal rates of NH ₊ ⁴ uptake by the diatoms at all densities. Although ingestion rates of C. pacificus varied from 0 to over 20% of body N d^–1 at the different food levels, excretion was a constant 6.6 nM NH ₊ ⁴ copepod^–1 h^–1 or about 10% of body N d^–1. This ingestion-excretion relationship, which is consistent with previous respiration and fecundity studies, suggests that the ecological dominance of C. pacificus only under conditions of high food abundance may be due to a dramatic increase in its growth efficiency as ingestion increases above the level supporting a constant metabolic rate. The maintenance of a constant level of metabolism during relatively short periods of low food abundance may be advantageous if it allows the copepod to exploit more effectively short-term variability in its food resulting from environmental heterogeneity or vertical migration.Contribution No. 1360 from the School of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Utilization of glutamate and glucose for heterotrophic growth by the marine pennate diatom Nitzschia laevis

Nitzschia laevis Hustedt grew in the dark in the presence of either glutamate or glucose as substrate. Complex mixtures of yeast extract or tryptone plus lactate also supported good heterotrophic growth, while tryptone alone only supported very slow growth in the dark. The observed growth rates of N. laevis in the dark at different concentrations of glutamate or glucose could be accounted for by the measured uptake rates of these compounds. The affinity of the uptake systems for glutamate and glucose (K _s =0.03 mM for each) was quite high, and similar for dark- and light-grown cells. The lack of a lag-phase when cells were transferred from photoautotrophic to heterotrophic growth conditions can be explained by the presence of uptake systems for glutamate and glucose in ligh--grown cells, as well as in dark-grown cells. However, the uptake capacity was generally higher in the latter than the former. N. laevis also took up alanine and lactate according to Michaelis-Menten kinetics, with a K _s for alanine of 0.02 mM and for lactate of 0.4 mM. Malate and glycerol were not taken up to a significant extent by the cells. Cells grown in continous light had a doubling time of 18 h. The shortest doubling time observed in the dark on glutamate was 48 h and on glucose 24 h. Glutamate was used for heterotrophic growth with an efficiency of 43% and glucose with an efficiency of 48%.Contribution No., 945 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA.     

Physioecology of zooplankton. I. Effects of phytoplankton concentration,temperature, and body size on the growth rate of Calanus pacificus and Pseudocalanus sp.

Changes in dry weight and in weight-specific growth rates were measured for copepodite stages of Calanus pacificus Brodsky and Pseudocalanus sp. cultured under various combinations of phytoplankton concentration and temperature. Mean dry weight of early copepodites was relatively unaffected by either food concentration or temperature, but mean dry weight of late stages increased hyperbolically with food concentration and was inversely related to temperature. The food concentration at which maximum body weight was attained increased with increasing temperature and body size, and it was considerably higher for C. pacificus than for Pseudocalanus sp. This suggests that final body size of small species of copepods may be determined primarily by temperature, whereas final body size of large species may be more dependent on food concentration than on temperature. Individual body weight increased sigmoidally with age. The weight-specific growth rate increased hyperbolically with food concentration. The maximum growth rate decreased logarithmically with a linear increase in body weight, and the slope of the lines was proportional to temperature. The critical food concentration for growth increased with body size proportionally more at high than at low temperature, and it was considerably higher for C. pacificus than for Pseudocalanus sp. Because of these interactions, early copepodites optimized growth at high temperature, even at low food concentrations, but under similar food conditions late stages attained higher growth at low temperature. The same growth patterns were found for both species, but the rates were significantly higher for the larger species, C. pacificus, than for the smaller one, Pseudocalanus sp. On the basis of findings in this study and of analyses of relationships between the maximum growth rate, body size, and temperature from other studies it is postulated (1) that the extrapolation of growth rates from one species to another on the basis of similarity in body size is not justified, even for taxonomically related species; (2) that the allometric model is inadequate for describing the relationship between the maximum weight-specific growth rate and body size at the intraspecific level; (3) that the body-size dependence of this rate is strongly influenced by temperature; and (4) that species of zooplankton seem to be geographically and vertically distributed, in relation to body size and food availability, to optimize growth rates at various stages of their life cycles.Contribution No. 1127 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Interactions of inorganic nitrogen in the uptake and assimilation by marine phytoplankton

The effect of ambient ammonium concentration on the nitrate uptake rate of marine phytoplankton was investigated. These studies consisted of laboratory experiments using unialgal species and field experiments using natural phytoplankton communities. In laboratory experiments, ammonium suppressed the uptake rates of nitrate and nitrite. Approximately 30 min were required for ammonium to exhibit its fully inhibitory effect on nitrate uptake. At high ammonium concentration (>3 g-at/l), a residual nitrate uptake rate of approximately 0.006 h^-1 was observed. When the ambient ammonium concentration was reduced to a value less than 1 g-at/l, the suppressed nitrate uptake rate subsequently attained a value comparable to that observed before the addition of ammonium. A range of 25 to 60% reduction in the nitrate uptake rate of natural phytoplankton communities was observed at ambient ammonium concentrations of 1.0 g-at/l. A mechanism is proposed for the suppression of nitrate uptake rate by ammonium through feedback control of the nitrate permease system and/or the nitrate reductase enzyme system. The feedback control is postulated to be regulated by the level of total amino acids in the cell.Contribution No. 936 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA. This paper represents a portion of a dissertation submitted to the Department of Oceanography, University of Washington, Seattle, in partial fulfillment of the requirements for the Ph.D. degree.     

Heterotrophic nutrition of the marine pennate diatom Nitzschia angularis var. affinis

The nutritional pattern for heterotrophic growth of Nitzschia angularis var. affinis (Grun.) Perag. is more complex than for other diatom species studied previously. This species grew slowly in the dark in the presence of single amino acids, either glutamate or alanine; other amino acids when supplied singly were not used as substrates. Carbon from glutamate was converted to cell carbon with an efficiency of 43%. Glutamine was inhibitory both in the light and in the dark, and aspartate inhibited heterotrophic growth on glutamate. Glucose and tryptone supplied singly did not support heterotrophic growth, but when combined, together they allowed for rapid growth of N. angularis (generation time of 16 h). Glucose in combination with glutamate, alanine, aspartate, or asparagine (but not with any other amino acids) also supported growth in the dark, at a rate considerably more rapid than with glutamate alone. In the presence of excess glucose and limiting concentrations of glutamate, approximately 50% of the cell carbon for heterotrophic growth came from glucose, while in combination with tryptone about 25% of the cell carbon came from glucose. Amino acids were taken up by cells grown either photoautrophically or in the dark in the presence or absence of organic substrates; uptake rates were some-what higher for dark-grown than for light-grown cells. Glucose was taken up only by dark-grown cells; induction of a glucose uptake system in the dark required the presence of glutamate but not of glucose. The rates of uptake of glutamate and glucose by cells incubated in the dark with glutamate were sufficiently high to account for the observed rates of growth on these substrates in the dark. The uptake systems of N. angularis have relatively high affinities for glucose (K _s =0.03 mM) and glutamate (K _s =0.02 mM).Contribution No. 890 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA.     

Growth and grazing loss rates in single-celledPhaeocystis sp. (Prymnesiophyceae)

Growth and grazing loss rates of naturalPhaeocystis sp. single cells were measured using a seawater dilution technique. Measurements were performed during an intensePhaeocystis sp. bloom in the North Sea between 19 April and 5 May 1988. Experimental results yielded rapid carbon turnover rates. Population growth rates varied from 0.033 to 0.098 h^–1, grazing loss rates from 0.037 to 0.174 h^–1. From measured growth rates, average doubling rages of 1.3 doublings d^–1 were calculated. The growth rates would have resulted in maximum carbon production rates of 146 mg C m^–3 d^–1. Grazing rates increased in the course of the bloom and exceeded growth rates at the end. Grazing loss was caused primarily by microzooplankton feeding. Ciliates and heterotrophic dinoflagellates were identified as the major potential consumers of single cells ofPhaeocystis sp. at the beginning of the bloom. The grazing impact of larger microzooplankton species appeared to increase during the progressing bloom.     

Ammonia excretion by zooplankton and its significance to primary productivity during summer

Excretion rates of ammonia have been determined for zooplankton off the coasts of Washington and Oregon (USA). Rates varied from 0.16 to 0.60 g-at NH ₄ ⁺ -N/mg dry weight/day for most planktonic animals, and from 0.02 to 0.06 for jellyfishes. Ammonia concentration in seawater was low in offshore regions. Ammonia released by zooplankton was studied in relation to primary productivity during summer. It was found that, in the Columbia River plume offshore, excreted ammonia contributed about 90% of the total nitrogen requirements of observed production rates. The ammonia-N contribution was 36% in oceanic waters, and was relatively unimportant in the inshore region. The significance of eddy diffusivity in offshore waters and upwelling in inshore waters is also discussed.Contribution No. 747 from the Department of Oceanography, University of Washington, Seattle, USA.     

Development of rapid ammonium uptake during starvation of batch and chemostat cultures of the marine diatom Thalassiosira pseudonana

Cell nitrogen quotas and uptake rates following ammonium additions were measured during ammonium-limited growth transients obtained by starving batch and chemostat cultures of Thalassiosira pseudonana (Clone 3 H). During starvation, cell quotas decreased by more than 50% in batch cultures. In chemostat cultures, the drop in cell quota during starvation decreased with dilution rate, from more than 50% at 1.45 d^-1, to less than 10% at 0.22 d^-1. Minimal levels of 3 to 4×10^-2 pg-at. N cell^-1 were reached after 24 h starvation in both batch and chemostat cultures. Uptake rates over the first minute of perturbation experiments were 3 times the long-term (10 to 30 min) rates. In batch cultures, specific uptake rates increased from 4 d^-1 to 20 d^-1 after 24 h starvation. Uptake rates per cell were independent of starvation time and dilution rate in chemostat cultures, but lower in non-starved batch cultures. The implications of these data for models of phytoplankton growth are discussed: the data support models which predict a depression in average growth rates when diatoms encounter microscale patches in oligotrophic environments.     

Effects of environmental factors and body size on rates of oxygen consumption in Archaeomysis grebnitzkii and Neomysis awatschensis (Crustacea: Mysidae)

The rates of oxygen consumption in relation to oxygen tension, temperature, salinity and body size were determined for Archaeomysis grebnitzkii Czerniavsky and Neomysis awatschensis (Brandt). Oxygen uptake was regulated by both species down to an oxygen tension of about 50 mm Hg (30% of saturation value); below this level, it was related to the oxygen tension in the medium. Oxygen tension below 20 mm Hg was lethal. A statistically significant interaction was noted for the effects of species, temperature, and salinity. In general, oxygen-uptake rate decreased with decreasing salinity in both species. The effect of size is expressed as a power function of body weight. The regression coefficients of oxygen consumption on body weight, 0.70 for A. grebnitzkii and 0.62 for N. awatschensis, were found to be significantly different from each other.Contribution No. 726 from the Department of Oceanography, University of Washington, Seattle, USA.     

Physioecology of zooplankton. IV. Effects of phytoplankton concentration,temperature, and body size on the net production efficiency ofCalanus pacificus

Weight-specific rates of individual production, total metabolic expenditure and assimilation, and net production efficiencies were estimated forCalanus pacificus Brodsky of selected body weights cultured at various phytoplankton concentrations and temperatures. The weight-specific rate of individual production increased hyperbolically with food concentration, and the maximum rate of individual production decreased logarithmically with a linear increase in body weight propotionally more at high than at low temperature. The weight-specific rate of total metabolic expenditure decreased logarithmically with increasing body weight and was unaffected by changes in food concentration. The effects of food concentration and temperature on the weight-specific rate of assimilation were similar to those on the rate of individual production, but the effect of body size differed considerably. The diversity in the temperature and body-size dependence of the maximum weight-specific rates of various physiological processes suggest (1) that, except for the metabolic rate, the allometric model (log-log relation) is inadequate for describing relationships between maximum rates of physiological processes and body size within species, and (2) that the common assumption that temperature affects the rates of various physiological processes in similar ways is not justified. Net production efficiency increased hyperbolically with food concentration, and the maximum production efficiency first increased slightly and then decreased gradually with increasing body weight. Small copepods attained higher efficiency at high temperature but larger ones did so at low temperature. The critical food concentrations for production efficiency and for the rate of individual production increased with increasing temperature and body size. Because of the effects of interactions among critical food concentration, temperature, and body size on the rates of growth and individual production and on net production efficiency, early development stages ofC. pacificus optimized growth and food conversion efficiency at high temperature, but late stages, particularly at low food concentrations, grew best and transformed food more efficiently at low temperature.Contribution No. 1130 From the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Demographic and temporal structure of an allele frequency cline in the mussel Mytilus edulis

Feeding rates, patterns of prey selection, and starvation tolerance were investigated for adult males and females of the cyclopoid copepod Corycaeus anglicus collected from the waters of Friday Harbor, Washington, USA. Selection by C. anglicus was determined largely by prey body-size, but was also affected by species and developmental stage. Small developmental stages of all prey species were fed upon at relatively low rates. The small calanoid species Acartia clausii was increasingly vulnerable to predation by C. anglicus as it progressed through successive developmental stages. Larger prey species, Pseudocalanus sp. and Calanus pacificus, were more vulnerable in intermediate stages, the C3 and N6 stages, respectively. Larger and smaller prey were characteristically attacked at different sites on their bodies; however, attack sites fell within a similar range of body widths, 130 to 170 m. Males of Corycaeus anglicus killed a maximum of 1.4 prey d^-1 when feeding on the optimally-sized adult females of Acartia clausii, which are approximately equivalent to its own body length. Males fed at approximately double the rates of females. Despite its small size and apparent lack of metabolic stores, this cyclopoid is highly tolerant of starvation conditions. Median survival time without food is at least 2 wk for both males and females. In its predatory behavior, C. anglicus employs an ambush-type strategy and seems to be adapted for infrequent encounters with relatively large prey.Contribution No. 1412 from the School of Oceanography, University of Washington, Seattle     

Marine diatoms grown in chemostats under silicate or ammonium limitation. III. Cellular chemical composition and morphology of Chaetoceros debilis,Skeletonema costatum,and Thalassiosira gravida

Three marine diatoms, Skeletonema costatum, Chaetoceros debilis, and Thalassiosira gravida were grown under no limitation and ammonium or silicate limitation or starvation. Changes in cell morphology were documented with photomicrographs of ammonium and silicate-limited and non-limited cells, and correlated with observed changes in chemical composition. Cultures grown under silicate starvation or limitation showed an increase in particulate carbon, nitrogen and phosporus and chlorophyll a per unit cell volume compared to non-limited cells; particulate silica per cell volume decreased. Si-starved cells were different from Si-limited cells in that the former contained more particulate carbon and silica per cell volume. The most sensitive indicator of silicate limitation or starvation was the ratio C:Si, being 3 to 5 times higher than the values for non-limited cells. The ratios Si:chlorophyll a and S:P were lower and N:Si was higher than non-limited cells by a factor of 2 to 3. The other ratios, C:N, C:P, C:chlorophyll a, N:chlorophyll a, P:chlorophyll a and N:P were considered not to be sensitive indicators of silicate limitation or starvation. Chlorophyll a, and particulate nitrogen per unit cell volume decreased under ammonium limitation and starvation. NH₄-starved cells contained more chlorophyll a, carbon, nitrogen, silica, and phosphorus per cell volume than NH₄-limited cells. N:Si was the most sensitive ratio to ammonium limitation or starvation, being 2 to 3 times lower than non-limited cells. Si:chlorophyll a, P:chlorophyll a and N:P were less sensitive, while the ratios C:N, C:chlorophyll a, N:chlorophyll a, C:Si, C:P and Si:P were the least sensitive. Limited cells had less of the limiting nutrient per unit cell volume than starved cells and more of the non-limiting nutrients (i.e., silica and phosphorus for NH₄-limited cells). This suggests that nutrient-limited cells rather than nutrient-starved cells should be used along with non-limited cells to measure the full range of potential change in cellular chemical composition for one species under nutrient limitation.Contribution No. 943 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA.