Silicon and the ecology of marine plankton diatoms. II. Silicate-uptake kinetics in five diatom species

The variation of the rate of silicate uptake with varying silicate concentration in the medium was investigated in short-term experiments with the following marine diatom species:Skeletonema costatum, Thalassiosira pseudonana, T. decipiens, Ditylum brightwellii, andLicmophora sp. The uptake conformed to Michaelis-Menten kinetics only after a correction had been made for reactive silicate that apparently could not be utilized by the diatoms. The magnitude of this correction was in the range of 0.3 to 1.3 g-at Si/l. Mean values of the half-saturation constant of silicate uptake were calculated for the different species. The lowest value was found inS. costatum (0.80 g-at Si/l) and the highest inT. decipiens (3.37 g-at Si/l). Growth limitation by low silicate concentrations could be a cause of species succession in marine plankton-diatom blooms.     

Growth and competition of the marine diatoms Phaeodactylum tricornutum and Thalassiosira pseudonana. II. Light limitation

The marine diatoms Phaeodactylum tricornutum (Bohlin) and Thalassiosira pseudonana (Hasle and Heimdal) were grown under both continous illumination and a 14 h light: 10 h dark cycle at light intensities ranging from 1.53×10^-4 to 2.95×10^-1 ly min^-1. Under both photoperiods, T. pseudonana exhibited higher division rates than P. tricornutum at high light intensities, but the reverse was true at all light intensities <3×10^-3 ly min^-1. Comparison of these results with available data on light-limited growth of other planktonic algae suggests that P. tricornutum may be unusually efficient at maintaining its cell division rate at low light intensity. This efficiency may contribute substantially to its success in turbid, nutrient-enriched mass algal culture systems, the only environments in which it is known to attain great numbers.Contribution No. 4086 from the Woods Hole Oceanographic Institution.     

Marine diatoms grown in chemostats under silicate or ammonium limitation. IV. Transient response of Chaetoceros debilis,Skeletonema costatum,and Thalassiosira gravida to a single addition of the limiting nutrient

The kinetic response of ammonium- or silicate-limited and ammonium- or silicatestarved populations of Chaetoceros debilis, Skeletonema costatum, and Thalassiosira gravida was determined by a single addition of the limiting nutrient to a steady-state culture and subsequent monitoring of the nutrient disappearance of the limiting and non-limiting nutrients at frequent time intervals. The kinetic response of nonlimited (nutrient) populations of these three species was also determined. Three distinct modes of the uptake of the limiting nutrient were observed for ammonium-or silicate-limited populations of these three species, surge uptake (V _s ), internally (cellular) controlled uptake (V _i ), and externally (ambient limiting nutrient concentration) controlled uptake (V _e ). Non-limited populations did not exhibit the three distinct segments of uptake, V _s , V _i and V _e . Estimates of the maximal uptake rate (V _max) and the Michaelis constant (K _s ) were obtained from nutrient-limited populations during the V _e segment of the uptake curve. Pooled values of V _e for the three ammonium-limited populations yielded V _max and K _s estimates of 0.16 h^-1 and 0.5 g-at NH₄–N l^-1. Kinetic data derived from the V _e segment of the uptake curve for silicate-limited populations yielded different values of V _max and K _s for each of the three species. In a number of parameters that were measured, T. gravida was clearly different from C. debilis and S. costatum and its recovery from nutrient starvation was the slowest. Recovery of all species from silicate limitation or starvation was slower than from ammonium limitation or starvation. Ammonium-starved populations maintained a maximal uptake rate at a substrate concentration an order of magnitude lower (0.1 g-at NH₄–N l^-1) than that observed for NH₄-limited populations (1.0 g-at NH₄–N l^-1). Adaptation to the severity of the nutrient limitation occurred as changes in the magnitude of cellular characteristics, such as short-term uptake potential (V _s ) and affinity for the substrate (K _s ). The consequence of these results are discussed in terms of another possible mechanism to explain changes in species composition and succession in nutrient-depleted environments.Contribution No. 944 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA.     

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.     

Feeding deterrence properties of apo-fucoxanthinoids from marine diatoms. II. Physiology of production of apo-fucoxanthinoids by the marine diatoms Phaeodactylum tricornutum and Thalassiosira pseudonana,and their feeding deterrent effects on the copepod Tigriopus californicus

The marine diatoms Phaeodactylum tricornutum and Thalassiosira pseudonana have been shown to produce apo-fucoxanthinoid compounds which act as feeding deterrents against the harpacticoid copepod Tigriopus californicus. The amounts and types of apo-fucoxanthinoids produced were species specific. Th. pseudonana produced small quantities of apo-12-fucoxanthinal and apo-13-fucoxanthinone only during senescence, while P. tricornutum produced much greater quantities of these two compounds during both log and senescence phases, in addition to producing a third compound, apo-10-fucoxanthinal, only during senescence. For both species, production of apo-fucoxanthinoids increased as the cells entered senescence phase due to phosphate limitation. The amounts of apo-fucoxanthinoids necessary to reduce feeding in T. californicus by 50% ranged from 2.22 to 20.2 ppm. This range was approximately 1000 times lower than the total apo-fucoxanthinoid concentration in P. tricornutum. The amounts of apo-fucoxanthinoids necessary to cause a 50% mortality in a population of T. californicus ranged from 36.8 to 76.7 ppm. Thus, these compounds are present in concentrations which may have ecological significance in the control of bloom formation and grazing. The production of apo-fucoxanthinoids may be a phenomenon common to many diatoms, particularly as they enter senescence due to nutrient limitation.     

Marine diatoms grown in chemostats under silicate or ammonium limitation. I. Cellular chemical composition and steady-state growth kinetics of Skeletonema costatum

Skeletonema costatum was grown in chemostats under ammonium or silicate limitation to examine its growth kinetics and changes in cellular chemical composition at different steady-state growth rates. When the relationship between the effluent limiting substrate concentration and steady-state growth rates was examined, deviations from the simple hyperbolic form of the Monod growth equation were noted at low and high dilution rates. The data from the plot of growth rate and substrate concentration were divided into 4 regions and the relationship of these region to cell quota is discussed. Two physiological states were identified. All populations grown at D<0.05 h^-1, regardless of the size of the cells or the magnitude of Q, exhibited a maximal growth rate of approximately 0.05 h^-1, while populations grown at higher dilution rates (D>0.06 h^-1 to 0.14 h^-1). The maximal value of growth rate is obtained only in cultures grown at very high dilution rates where nutrient shift-up appears to occur, the cell quota approaches a maximum and the heterogeneous cell population becomes more homogeneous.Contribution No. 881 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA. This paper represents a portion of two dissertations submitted by P.J.H. and H.L.C. to the Department of Oceanography, University of Washington, Seattle, in partial fulfillment of the requirements for the Ph.D. degree.     

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.     

Effect of UV-B radiation on the marine diatoms Lauderia annulata and Thalassiorsira rotula grown in different salinities

The marine diatoms Lauderia annulata Cleve and Thalassiosira rotula Meunier were grown at different salinities (20, 35 and 45) and exposed to different levels of midultraviolet, UV-B) 439, 717 and 1230 J m^-2 d^-1, weighted) for 2 d. A low UV-B dose (439 J m^-2 d^-1) usually caused a slight increase in biomass production (dry weight) compared to non-UV-B irradiated cells. Enhanced UV-B radiation (717 J m^-2 d^-1) depressed protein and pigment content (chlorophyll a, chlorophyll c ₁+c₂ and carotenoids), especially in algae grown at 20 or 35 salt concentration of the nutrient solution. The effect of UV-B radiation (717 J m^-2 d^-1) on the pattern and concentration of amino acids was species-dependent. Aspartic acid was reduced in all tested diatoms. A drastic increase in glutamine and a reduction in glutamic acid pools could be observed in L. annulata samples, but no significant variation of the impact of UV-B was found in dependence on the salt concentration of the nutrient medium. T. rotula cells grown at 35 S showed an increase of glutamic acid and a decrease of glutamine levels after UV-B radiation. The results are discussed in relation to the impact of UV-B upon carbon and nitrogen metabolism.     

Study of the nutrient and plankton dynamics in Lake Tanganyika using a reduced-gravity model

An eco-hydrodynamic (ECOH) model is proposed for Lake Tanganyika to study the plankton productivity. The hydrodynamic sub-model solves the non-linear, reduced-gravity equations in which wind is the dominant forcing. The ecological sub-model for the epilimnion comprises nutrients, primary production, phytoplankton biomass and zooplankton biomass. In the absence of significant terrestrial input of nutrients, the nutrient loss is compensated for by seasonal, wind-driven, turbulent entrainment of nutrient-rich hypolimnion water into the epilimnion, which gives rise to high plankton productivity twice in the year, during the transition between two seasons. Model simulations predict well the seasonal contrasts of the measured physical and ecological parameters. Numerical tests indicate that the half saturation constant for grazing by zooplankton and the fish predation rate on zooplankton affect the zooplankton biomass measurably more than that of phytoplankton biomass. This work has implications for the application of this model to predict the climatological biological productivity of Lake Tanganyika.     

Determination of a limiting nutrient regulating algal biomass using in situ experiments of nutrient enrichment bioassay (NEB) and empirical relations of nutrients and chlorophyll-a

Long-term nutrient contents and nutrient ratios indicated that phosphorus was a potential limiting element for algal growth. In situ experiments of nutrient enrichment bioassay supported the evidence of P-limitation. However, regression analyses of log10-transformed chlorophyll-a (CHL) against TP (R2 values < 0.25) showed that seasonal CHL was not closely related to flux of phosphorus during all seasons. Also, two dimensional graphical approach of Trophic State Index (TSI) showed that most values of TSI (CHL) -TSI (TP) and TSI (CHL) -TSI (SD) were less than zero, indicating factors other than phosphorus limited algal biomass (CHL -TP < 0), and that non-algal particles dominated light attenuation (CHL -SD < 0). The weak empirical relations and trophic deviations were explained well by the experiment of NEB-II that was conduced during a period of high inorganic turbidity. Overall results suggest that phosphorus is the primary element regulating the system productivity, but the system also were highly influenced by rapid flushing and high inorganic turbidity.     

Autecology and clonal variability of the marine centric diatom Thalassiosira rotula (Bacillariophyceae) in response to light,temperature and salinity

The influence of 49 combinations of salinity (10–40 S, at 5 S intervals) and temperature (0°–30°C, at 5C° intervals) on the maximum daily division rate (K) and 18 combinations of light intensity (six levels) and temperature (5°, 15°, and 25°C) on photosynthesis, cell division, and chlorophyll a was examined using two clones of Thalassiosira rotula Meunier isolated from the upwelling area of Baja California (clone C8) and from Narragansett Bay, Rhode Islands (clone A8). Physiological differences appear to characterize these to clones with regard to their temperature tolerance (C8 5°–30°C, A8 0°–25°C), maximum growth rate (C8 K=2.9, A8 K=2.4), chlorophyll a content, and in the rates of growth and photosynthesis in response to light intensity and temperature. Optimum salinity for both clones (25–30 S) was generally independent of temperature, while chlorophyll a content decreased with temperature. T. rotula is a cosmopolitan paractic species; experimental studies indicate that it is eurythermal and moderately euryhaline. Comparison of five additional Narragansett Bay isolates of T. rotula reveal minimal spacial or temporal variability in genetically determined physiological characteristics within this local population.     

Tisbe (Copepoda: Harpacticoida) species from the Lagoon of Venice. I. Seasonal fluctuations and ecology

Distribution and abundance of Tisbe species were studied throughout a period of 3 years at 3 stations in the Lagoon of Venice (Italy). At 2 other stations samples were taken occasionally, as also at the Lido station (open sea). Twelve species may be considered as common inhabitants of the lagoon, and 3 species appear to be occasional transients, compared to the 9 species found at the Lido. A comparison of the physical data shows that the stations were similar in temperature and salinity, but differed primarily in pH values, sediment characteristics, algal substratum and faunal benthic community. The fact that remarkable differences in species composition and distribution of Tisbe could be observed between the various parts of the lagoon and the lagoon and the open sea, indicates that temperature and salinity are not so important for species diversity as are biotic factors. A particularly strong difference existed between samples taken at the bottom and from pilings, which constitute a very special biotope for the benthic communities in the lagoon and seem to be a preferred habitat for T. lagunaris and T. cucumariae. The data suggest that there is a certain interaction between T. holothuriae and the other species. Whenever T. holothuriae was abundant, the other species were rare, but T. clodiensis, T. dobzhanskii (Stations 1 and 2) and Tisbe sp. (Station 3) attained high relative abundance whenever T. holothuriae decreased considerably in number. T. holothuriae represents certainly the hardiest species of the lagoon, displaying a great ability to cope with stress conditions, even those present in the interior lagoon where strong fluctuations in salinity, temperature, pH and oxygen prevail.     

Light absorption by a marine diatom: experimental observations and theoretical calculations of the package effect in a small Thalassiosira species

The relationship between in vivo light absorption efficiency of whole cells and in vitro absorption efficiency of algal pigments has been examined experimentally in the marine diatom Thalassiosira sp. In vitro absorption spectra were obtained for cells disrupted by either ultrasonic treatment or high-pressure shearing stress in a low-temperature (-40°C) pressure cell. A dimensionless measure of the magnitude of the package effect (Q _a ^*), calculated from the ratio of whole-cell to disrupted-cell absorption, ranged from about 0.5 at the blue absorption peak of chlorophyll a (λ=435 nm) to 0.7 at the red chlorophyll a peak (λ=670 nm) to 1.0 at the absorption minimum (λ=600 nm). Cell diameter was found to be an inappropriate measure of size for assessing the magnitude of the package effect. Instead, the effective optical diameter for calculation of intracellular self-shading was found to be less than the cell diameter. This observation is consistent with the fact that most algal pigments are contained within chloroplasts, and that chloroplast volume is necessarily smaller than cell volume.     

The development of a natural plankton population in an outdoor tank with nutrient-poor sea water. I. Phytoplankton succession

Due to complex hydrodynamic and biological inhomogeneities, the phytoplankton species succession cannot be satisfactorily observed, apart from the seasonal blooms which occur in temperate waters. Large flexible plastic tanks have proved to be useful for such observations. In 1972, for 28 days, a phytoplankton succession in nutrient-poor water in the outer harbor of Helgoland was observed in a flexible plastic tank (3 m³). During this period, 3 phytoplankton biomass maxima were formed with many significant correlations. In the first 7 days the ammonia concentration decreased from over 6 to 2 mol 1^-1 for 14 days. The nitrate concentration remained in the range of 3–6 mol 1^-1 and then fell abruptly to 0–2 mol 1^-1. The phosphate concentration was about 0.1 mol 1^-1. Lauderia borealis dominated the first period, and its increase was significantly correlated with the decrease in ammonia. The diatom was succeeded by two dinoflagellates, Dinophysis acuminata and Prorocentrum micans. The last period of the experiment was characterized by a stronger development of Rhizosolenia species. The rapid recovery of the crop in the nutrient-poor water points to intensive remineralization processes. The irregular occurrence of ammonia near the surface was correlated with the appearance of Noctiluca miliaris at this depth. It is expected that repetitions of this type of experiment will permit further explanations of statistical correlations which are not yet clear. As a first step, in order to test hypotheses, a correlation analysis was employed to eliminate the statistically non-significant correlations.This work was supported by the Sonderforschungsbereich 94, Hamburg, of the Deutsche Forschungsgemeinschaft.     

Role of the fish Astyanax aeneus (Characidae) as a keystone nutrient recycler in low-nutrient neotropical streams

Nutrient recycling by animals is a potentially important biogeochemical process in both terrestrial and aquatic ecosystems. Stoichiometric traits of individual species may result in some taxa playing disproportionately important roles in the recycling of nutrients relative to their biomass, acting as keystone nutrient recyclers. We examined factors controlling the relative contribution of 12 Neotropical fish species to nutrient recycling in four streams spanning a range of phosphorus (P) levels. In high-P conditions (135 microg/L soluble reactive phosphorus, SRP), most species fed on P-enriched diets and P excretion rates were high across species. In low-P conditions (3 microg/L SRP), aquatic food resources were depleted in P, and species with higher body P content showed low rates of P recycling. However, fishes that were subsidized by terrestrial inputs were decoupled from aquatic P availability and therefore excreted P at disproportionately high rates. One of these species, Astyanax aeneus (Characidae), represented 12% of the total population and 18% of the total biomass of the fish assemblage in our focal low-P study stream but had P excretion rates > 10-fold higher than other abundant fishes. As a result, we estimated that P excretion by A. aeneus accounted for 90% of the P recycled by this fish assemblage and also supplied approximately 90% of the stream P demand in this P-limited ecosystem. Nitrogen excretion rates showed little variation among species, and the contribution of a given species to ecosystem N recycling was largely dependent upon the total biomass of that species. Because of the high variability in P excretion rates among fish species, ecosystem-level P recycling could be particularly sensitive to changes in fish community structure in P-limited systems.     

Influence of pore size of plankton nets and towing speed on the sampling performance of two high-speed samplers (Delfino I and II) and its consequences for the assessment of plankton populations

The sampling performance of two high-speed samplers, (Delfino I and II), fitted with a calibrated flow meter, was studied in the open Ligurian Sea using a battery comprising 6 such Delfinos linked together. Since, in plankton counts, only those organisms should be counted which are definitely retained in the nets, several techniques for the separation of the organisms which pass through the pore of a net of a given pore size and those which are safely retained were tested. Attempts to separate these two fractions by filtration of the fixed sample through nets of different pore sizes were not successful, as living organisms were able to pass through pore sizes which retained dead organisms. However, optical sizing under the dissecting microscope during counting gave reproducible results. Using this counting technique, the influence of the pore size of plankton nets on the reproducibility of sampling of natural populations was studied. The results obtained show that live plankton passes through the net pores at a size at which fixed plankton is retained. Hence, the minimum retention size for living and dead plankton is different. By comparing samples taken with nets of different pore sizes, and counting the organisms caught according to different sizes, it was possible to determine the minimum retention size of live plankton organisms for several different net pore-sizes. The minimum retention size is, therefore, the smallest pore size at which the organisms of a certain width cannot escape through the pores of the net and are, thus, quantitatively retained in the net. By applying the criterion of minimum retention size, the influence of speed towing on the number of plankton organisms caught was studied. It was shown that the number of organisms which are safely retained, i.e., do not escape through the pores, increased with speed, reaching a plateau at 5 to 7 knots.A contribution under the Association Contract CNEN-EURATOM. Contribution No. 668 of the Euratom Biology Division.     

Salinity and nutrient effects on the induction of the galactose permease system in a psychrophilic marine vibrio

An Antarctic psychrophilic marine Vibrio species was isolated having the inducible ability to accumulate nonmetabolizable thiomethyl-14C--galactopyranoside (14C-TMG) through a galactose permease system. Induction of 14C-TMG uptake was found to have a salinity requirement which was higher than that required for uptake. At the optimum salinity, galactose and fucose were the primary inducers. Lactose produced a comparable induction but only at higher concentrations, whereas glucose did not cause induction. The initial rate of 14C-TMG uptake exhibited saturation kinetics with an apparent Km value of 4.8 x 10^–6M. An amino acid, in addition to the inducer, was required for induction which could not be replaced by glycerol or galactose. Evidence is presented which indicates that the uptake of 14C-TMG is energy-dependent and that nutrient availability is more important than salinity for induction and uptake under conditions which would normally be found in the oceanic environment.Technical Paper No. 4911, Oregon Agricultural Experiment Station.     

Female habitat choice as a determinant of the reproductive success of the territorial male marine iguana (Amblyrhynchus cristatus)

Summary The reproductive behaviour of marine iguanas (Amblyrhynchus cristatus) was studied to answer the following questions: Which physical factors determine the number of females found on a male's territory? Do males select territories according to preferences for physical conditions or according to the number of females there? What causes the strong correlation between the number of females on a territory and the number of copulations achieved there? Females preferred as their resting places territories with the following characteristics, in order of importance: High number of shady/shelter places, low temperature of rocks at noon, proximity to the sea, sun-basking places in the afternoon. Other factors measured (probability of being wetted by spray, sun-basking places in the morning and at noon) showed no significant correlation with the number of females. Males, however, preferred territories further from the sea (within about 7 m), with high rock temperatures and a high number of basking places in the morning (in order of importance). Males primarily followed their site preferences and not female density in their choice of a territory. The site choice of females determined the number of copulations achieved on a territory, as females copulated preferentially on their resting places. A shift of females from one territory to another within one colony between two succeeding years was correlated with the change in relative weight of territory owners. This result was interpreted as an indication of female choice. Site preferences of males and females are discussed in relation to the animals' physiological requirements. Possible influences of ecological factors on the evolution of this mating system are considered.     

Nitrate reductase and urease enzyme activity in the marine diatom<Emphasis Type="Italic"> Thalassiosira weissflogii</Emphasis> (Bacillariophyceae): interactions among nitrogen substrates

The dissolved nitrogen pool in aquatic systems is comprised of many different nitrogen forms, both inorganic and organic. Interaction among these nitrogen forms at the level of uptake and enzyme activity is, with the exception of NH₄⁺ and NO₃^–, not completely understood. Nitrate reductase (NR) and urease (UA) activities in the marine diatom Thalassiosira weissflogii (Grunow) Fryxell et Hasle were measured in NO₃^–, NH₄⁺, and urea-sufficient cultures before and after challenge additions of NH₄⁺, NO₃^–, and urea in a factorial design. NR and UA were constitutively expressed during growth on NO₃^–, NH₄⁺, and urea. Growth on NH₄⁺ or urea resulted in NR activities that were <10% of the activity observed in the NO₃^–-grown culture, while growth on NO₃^– resulted in UA values that were ~35% of the activities during growth on either NH₄⁺ or urea. The addition of NH₄⁺ or urea to NO₃^–-grown cultures resulted in an immediate decrease in cellular NO₃^– uptake rate, which was not mirrored by an immediate repression of in vitro NR activity; however, the diel peak in NR was suppressed in these challenge experiments. The addition of NO₃^– or NH₄⁺ to urea-grown cultures resulted in non-significant decreases in the urea uptake rate. UA was not impacted by NO₃^– addition, but NH₄⁺ addition significantly decreased UA throughout the experiment. These studies demonstrate that the uptake and assimilation of NO₃^– and urea may not be subject to the same internal feedback mechanism when challenged with other nitrogen substrates.Communicated by J.P. Grassle, New Brunswick     

Oxygen as a limiting factor in phototaxis and in intraclonal spacing of the sea anemone Anthopleura elegantissima

Phototaxis in Anthopleura elegantissima, a sea anemone symbiotic with zooxanthellae, was investigated with special reference to oxygen as a possible controlling factor. Under high oxygen concentrations in seawater, movement towards light was not observed for symbiotic anamones as it was under normal oxygen concentrations. Both aposymbiotic and symbiotic anemones demonstrated movement towards high oxygen concentrations in seawater. Oxygen is, therefore, implicated as a controlling factor in phototaxis. Under laboratory conditions, increased intraclonal spacing occurred with low oxygen concentrations in seawater. In the field, individuals in symbiotic clones were spaced significantly closer than in aposymbiotic clones. Since intraclonal spacing is controlled by oxygen in the laboratory, spacing may also be affected in the field by oxygen; symbiotic clones may be spaced closer because they have better oxygen availability than do aposymbiotic clones.