Effects of simulated current on the growth rate and nitrogen metabolism of Gracilaria tikvahiae (Rhodophyta)

In a series of multifactorial laboratory experiments, Gracilaria tikvahiae apical segments were grown in an apparatus in which they were exposed simultaneously to 3 simulated current speeds (7.5, 15, 22.5 cm s^-1) and a still control, and either 3 ammonium concentrations (<6, 37–39, and 119–136 M) under ample uniform light (ca 200 E m^-2 s^-1) or 3 light intensities (ca 35, 90, and 270 E m^-2 s^-1) with uniform surplus ammonium. Growth rates of apical segments were determined in each experiment as well as nitrogen and carbon composition of tissues and fluxes of NH₄, NO₃/NO₂, and PO₄ in media. In a supplementary series of field experiments, apical segments of G. tikvahiae weresimultaneously exposed to 2 different regimes of water motion in adjacent chambers at several sites characterized by widely different ammonium regimes. The application of simulated current significantly enhanced growth rates in all experiments which utilized recently collected plants. Generally, this enhancement was fully realized at 7.5 cm s^-1, with growth rates tending to plateau above that speed. Growth enhancement by simulated current was independent of ammonium concentration and was considerably reduced at the lowest light intensity. In experiments conducted with plants that had been maintained for several months in aquaria, simulated current failed to enhance growth rates. This suggested that growth responses were at least partly a function of prior conditioning. Growth rates were a direct function of light intensity and an inverse function of ammonium concentration, indicating ammonium inhibition at the higher applied concentrations. Simulated current slightly enhanced rates of ammonium uptake but this did not consistently result in reduced C:N ratios, suggesting that the growth-stimulating effect of relative water motion was attributable to factors other than N uptake. There was evidence of luxury consumption of ammonium. In field experiments, growth rates were not significantly related to exposure to water motion.     

Survival,metabolism and growth of Ulva lactuca under winter conditions: a laboratory study of bottlenecks in the life cycle

Photosynthesis and growth in low light and survival under simulated winter conditions were studied in the freefloating green alga Ulva lactuca L., collected in Roskilde Fjord, Denmark during late autumn and maintained in stock in natural water. It adapts efficiently to low light by increasing chlorophyll concentration and light absorption and continues to grow at the lowest irradiance tested, 0.6 E m^-2 s^-1. This irradiance corresponds to minimum light requirements of deep-living marine macroalgae and phytoplankton growing under ice. The photosynthetic efficiency per unit of incident light is five-fold higher for U. lactuca grown at 1.7 E PAR m^-2 s^-1 as compared with 56.3 E m^-2 s^-1, and the efficiency per unit of light absorbed is twice as high. The maximum photosynthetic efficiency (0.051 mol C E^-1 absorbed) is similar to values for shade-adapted marine phytoplankton. U. lactuca is able to survive for two months in the dark and to resume growth immediately when transferred to light. Exposure to anoxia and sulphide gradually reduces vitality, but does not affect survival over two months. Rigorous deep freezing is detrimental to survival of U. lactuca, while field samples show that more gradual, natural freezing is not. U. lactuca is not easily fitted into one of the traditional strategy concepts. U. lactuca is a very plastic species that combines rapid growth during favourable periods (opportunism) with high survival capacity in the same type of tissue during stress periods (persistence). U. lactuca occupies a niche as a free-floating form in sedimentary coastal areas that are devoid of attached algae.     

Nutrient uptake kinetics and growth of zooxanthellae maintained in laboratory culture

Growth characteristics and nutrient uptake kinetics were determined for zooxanthellae (Gymnodinium microadriaticum) in laboratory culture. The maximum specific growth rate (_max) was 0.35 d^-1 at 27 °C, 12 hL:12 hD cycle, 45 E m^-2 s^-1. Anmmonium and nitrate uptake by G. microadriaticum in distinct growth phases exhibited Michaelis-Menten kinetics. Ammonium half-saturation constants (K_s) ranged from 0.4 to 2.0 M; those for nitrate ranged from 0.5 to 0.8 M. Ammonium maximum specific uptake rates (V_max) (0.75 to 1.74 d^-1) exceeded those for nitrate (0.14 to 0.39 d^-1) and were much greater than the maximum specific growth rate (0.35 d^-1), suggesting that ammonium is the more significant N source for cultured zooxanthellae. Ammonium and nitrate V_max values compare with those reported from freshly isolated zooxanthellae. Light enhanced ammonium and nitrate uptake; ammonium inhibited nitrate uptake which was not reported for freshly isolated zooxanthellae, suggesting that physiological differences exist between the two. Knowledge of growth and nutrient uptake kinetics for cultured zooxanthellae can provide insight into the mechanisms whereby nutrients are taken up in coral-zooxanthelae symbioses.Contribution No. 1515 from the University of Maryland Center for Environmental and Estuarine Studies, Chesapeake Biological Laboratory, Solomons, Maryland 20688-0038, USA     

Light absorption,photosynthesis and growth of Nannochloris atomus in nutrient-saturated cultures

Nannochloris atomus was maintained in exponential growth at photon flux densities (PFD) from 400 to 700 nm, ranging from 10 to 200 mol m^-2 s^-1. Growth was lightsaturated at PFDs in excess of 100 mol m^-2 s^-1, with a mean light-saturated growth rate at 23 °C of 1.5×10^-5s^-1 (1.2 d^-1). The light-limited growth rates extrapolated to a compensation PFD for growth that was not significantly different from zero, although no changes in cell numbers were observed in a single culture incubated at a PFD of 1.0 mol m^-2s^-1. Dark-respiration rates were independent of PFD, averaging 1.7×10^-6 mol O₂ mol^-1 C s^-1 (0.14 mol O₂ mol^-1 C d^-1). The maximum photon (quantum) efficiency of photosynthesis was also independent of PFD, with a mean value of 0.12 mol O₂ mol^-1 photon. The chlorophyll a-specific light absorption cross-section ranged from 3 to 6×10^-3 m² mg^-1 chl a and was lowest at low PFDs due to intracellular self-shading of pigments associated with high cell-chlorophyll a contents. The C:chl a ratio increased from 10 to 40 mg C mg^-1 chl a between PFDs of 14 and 200 mol m^-2 s^-1. These new observations for N. atomus are compared with our previous observations for the diatom Phaeodactylum tricornutum in terms of an energy budget for microalgal growth.     

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.     

Nitrogen fixation by blue-green algae associated with the siphonous green seaweed Codium decorticatum: effects on ammonium uptake

Nitrogen fixation (acetylene reduction) at rates of up to 1.2 g N₂ g dry wt^-1 h^-1 was measured for the siphonous green seaweed Codium decorticatum. No nitrogenase activity was detected in C. isthmocladum. The nitrogenase activity was light sensitive and was inhibited by the addition of DCMU and triphenyl tetrazolium chloride. Additions of glucose did not stimulate nitrogen fixation. Blue-green algae (Calothrix sp., Anabaena sp., and Phormidium sp.) were implicated as the organisms responsible for the nitrogenase activity. They occurred in a reduced microzone within the C. decorticatum thallus where nitrogen fixation was optimized. Nitrogen fixation did not affect the kinetic constants for ammonium uptake in C. decorticatum (K_s=12.0 M, V_max=13.4 mol NH₃ g dry wt^-1 h^-1) determined using the perturbation method. Nevertheless, C. decorticatum thalli which fixed nitrogen had internal dissolved nitrogen concentrations which were over 1.4 times higher than in non-fixing thalli. This suggests that if C. decorticatum does derive part of its nitrogen requirement from the blue-green algae which it harbors, the transfer does not involve competition between this process and the uptake of ambient ammonium.     

The ammonium/methylammonium uptake system of Symbiodinium microadriaticum

The substrate analogue [¹⁴C]-methylammonium was used to study ammonium/methylammonium uptake by Symbiodinium microadriaticum (zooxanthellae). The value of the Michaelis constant (K _m) for the uptake system was approximately 35 M with methylammonium as substrate; ammonium was a competitive inhibitor of methylammonium uptake, and the K _m for ammonium uptake (determined as the inhibition constant, K _i, for methylammonium) was 6.6 M. Methylammonium uptake by zooxanthellae was light-dependent. Methylammonium uptake rates of zooxanthellae which had been freshly isolated from the hermatypic coral Acropora formosa (0.85±0.05x10^-10 mol min^-1 cell^-1) were lower than those of axenic cultures of the zooxanthellae from Montipora verrucosa (Acroporidae) grown under various nitrogen regimes (1.6 to 12x10^-10 mol min^-1 cell^-1). Maximum uptake rates were found for ammonium-starved cultured M. verrucosa zooxanthellae (10.2 to 12x10^-10 mol min^-1 cell^-1); M. verrucosa zooxanthellae growing with ammonium as nitrogen source and zooxanthellae which had been freshly isolated from A. formosa gave similar and considerably lower uptake rates (0.85 to 1.6x10^-1 mol min^-1 cell^-1). These results suggest that either coral tissue contains sufficient ammonium to repress synthesis of the uptake system of the algal symbionts or, alternatively, there are additional barriers to ammonium transport for zooxanthellae in vivo.     

Light requirements for growth of six shade-acclimated Mediterranean macroalgae

Six mediterranean macroalgae were cultivated for more than 2 yr under shade culture conditions, after which light requirements for growth were investigated at 16±2°C. The saturation light levels for growth in the logarithmic phase were related to the bathymetric distribution of the algae on the shore. The eulittoral to supralittoral red alga Bangia atropurpurea was saturated at a photon fluence rate of 71 mol photons m^-2 s^-1, the upper sublittoral to eulittoral brown algae Scytosiphon lomentaria, Colpomenia peregrina and Kuckuckia spinosa and the sublittoral brown alga Stictyosiphon soriferus at 39 to 71 mol photons m^-2 s^-1, and the deep-water alga Choristocarpus tenellus at 19 mol photons m^-2 s^-1. The minimum light requirements for growth of B. atropurpurea and C. tenellus were determined by observing length increase for 56 d under limiting light conditions. The compensation and minimum irradiances required for growth of B. atropurpurea were 0.5 and 1 mol photons m^-2 s^-1 respectively. The corresponding values for C. tenellus were 0.15 to 0.28 and 0.5 mol photons m^-2 s^-1 respectively. C. tenellus was the siowest-growing species tested at saturating light conditions, but it grew faster than B. atropurpurea at 1 mol photons m^-2 s^-1. Both B. atropurpurea and C. tenellus were able to survive 56 d in darkness, but only the latter grew under darkness in the first 14 d.     

Nitrogen uptake and growth of the germlings and mature thalli of Fucus distichus

Fucus distichus L. was collected near Vancouver, Canada, in late fall and early winter, 1981. The effects of the forms of nitrogen (nitrate, ammonium or urea) and periodic exposure to air on growth, rhizoid development and nitrogen uptake in germlings was investigated. Gamete release, fertilization, germination and germling growth had no requirement for a specific form of nitrogen. Periodic exposure to air increased secondary rhizoid development twofold. Nitrate and ammonium uptake rates of the germlings were higher than for the mature thalli (20 to 40 times for nitrate and 8 times for ammonium), while the halfsaturation constant (K _s) values for nitrate were similar (1 to 5 M). The germlings showed saturable uptake kinetics but the mature thalli did not. When germlings were exposed to air it caused a 70% decrease in nitrate uptake, but not change in ammonium uptake. Ammonium uptake in the mature thalli was proportional to the ambient ammonium concentration. Nitrate uptake in the mature thalli appeared to follow saturation kinetics at low nitrate concentrations, but showed a non-saturable component at concentrations greater than 10 M. Presence of ammonium inhibited nitrate uptake by the mature plants but not by the germlings.     

The annual cycle of heterotrophic planktonic ciliates in the waters surrounding the Isles of Shoals,Gulf of Maine: an assessment of their trophic role

Macrofauna living on subtidal rocks reefs in southern California excrete ammonium, a potentially important nutrient for benthic algae. Ammonium excretion rates of eleven macroinvertebrate and five fish taxa were determined from a total of 324 in situ incubations conducted between October 1984 and August 1985 at 14 to 17 m depths off Santa Catalina Island, California. Total ammonium excretion ranged from over 100 mol h^-1 by the kelp bass Paralabrax clathratus to less than 0.1 mol h^-1 by the gastropod Conus californicus. Weight-specific ammonium excretion generally ranged from 0.5 to 4 mol g^-1 h^-1 in invertebrates and from 3 to 7 mol g^-1 h^-1 in fishes. Intraspecific excretion rates varied substantially. Coefficient of variation of excretion rates were higher than reported for laboratory studies and multiple regression indicated that 50 to 90% of the variation in ammonium excretion rates of five species studied in detail could not be explained by the combined variation in dry weight, water temperature, time of day, and incubation dates. The excretion data, along with estimates of population densities and size-frequency distributions, indicate that benthic macrofauna release a total of 25 to 30 mol NH ₄ ⁺ m^-2 h^-1 both day and night. The species that generally make the largest contributions are a gobiid fish (Lythrypnus dalli), followed by three gastropods (Astraea undosa, Tegula eiseni, and T. aureotincta) and a sea urchin (Centrostephanus coronatus). The amount of ammonium excreted by these macrofauna on rocky reefs is insignificant compared to our previously published data on the nighttime excretion of blacksmith (Chromis punctipinnis), a pomacentrid fish that feeds in the water column during the day and shelters on the reef at night. Including blacksmiths, we estimate that the amount released by rocky-reef macrofauna at night is >280 mol m^-2 h^-1, a rate that is similar to that for many other marine communities. Additional studies are required to determine if benthic algae utilize ammonium released by these macrofauna, especially at night.Contribution No. 58 of the Ocean Studies Institute; Contribution No. 123 of the Catalina Marine Science Center     

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.     

Ammonium excretion by gelationous zooplankton and their contribution to the ammonium requirements of microplankton in Chesapeake Bay

Ammonium excretion rates of recently collected specimens of gelatinous zooplankton, the scyphomedusan Chrysaora quinquecirrha DeSor and the etenophore Mnemiopsis leidyi A. Agassiz, were correlated with body mass and water temperature in measurements made from April to October 1989 and 1990. Rates ranged between 3.5 and 5.0 g atoms NH ₄ ⁺ -N (g dry wt)^-1h^-1 for C. quinquecirrha and 3.0 to 4.9 g atoms NH ₄ ⁺ -N (g dry wt)^-1h^-1 for M. leidyi. Excretion rate equations and in situ data on the size distributions and biomasses of gelatinous zooplankters and water temperature were used to estimate the contribution of ammonium by medusae and ctenophores to mesohaline Chesapeake Bay waters on several dates during April to October 1989 and 1990. We then compared the estimated contributions to direct measurements of ¹⁵NH ₄ ⁺ uptake by microplankton. The maximum estimated regeneration by gelatinous zooplankton was 5.8 g atoms NH ₄ ⁺ -N m^-3h^-1 at night in August 1990, when medusae biomass was greatest. This represents about 4% of the ammonium required by the microplankton. During the daytime on all dates, less than 1% of the ammonium required by microplanktion was supplied by gelatinous zooplankton. Therefore, gelatinous zooplankton appear to play a minor role in the ammonium cycle of Chesapeake Bay.     

Nitrogenous nutrition in the euphotic zone of the Central North Pacific Gyre

The uptake rates of the three nitrogen compounds ammonium, nitrate, and urea were measured in the oligotrophic North Central Pacific Gyre in August–September 1985. The measurements were performed by using ¹⁵N-labelled substrates and incubating for short-time periods (3 to 4 h) under simulated in situ conditions. Ambient concentrations of the nitrogenous nutrients were generally below 0.10 mol l^-1. The average total daily nitrogen uptake rate, integrated over the euphotic zone, was 12.5 mmol N m^-2 d^-1. Diel studies in the upper water mass resulted in a calculated phytoplankton growth rate of 1.3 d^-1. Ammonium was the dominating nutrient, accounting for on the average 54% of the total nitrogen uptake, while urea uptake represented 32% and nitrate 14%. Ammonium uptake rates at a coastal station off the Hawaiian Islands were very close to the rates found at the oceanic station. Organisms <3 m dominated the nitrogen assimilation, being responsible for about 75% of the ammonium uptake. The nitrogen uptake rates in this study seem to be higher than those found by earlier investigations in the area, but correlated well with other productivity measurements performed during the same cruise.     

Seasonal photosynthetic characteristics of Ascoseira mirabilis (Ascoseirales,Phaeophyceae) from King George Island,Antarctica

Monthly variation in photosynthesis, dark respiration, chlorophyll a content and carbon: nitrogen (C:N) ratios in different lamina sections of adult plants of Ascoseira mirabilis Skottsberg from King George Island, Antarctica, was investigated between September 1993 and February 1994. Light saturated net photosynthesis (P _max) showed maximum values in September (12 to 25 mol O₂ g^-1 fr wt h^-1), and decreased towards the summer to values ranging between 2.0 and 5.0 mol O₂ g^-1. In the distal section, however, a second optimum occurred in December (25 mol O₂ g^-1 fr wt h^-1). Dark respiration rates were also highest in October and November and decreased strongly in December to February (6.0 and 1.0 mol O₂ g^-1 fr wt h^-1, respectively). Gross photosynthesis exhibited high values between September and December. Concomitant with the seasonal decrease of photosynthetic efficiency () from mean values of 1.2 mol O₂ g^-1 fr wt h^-1 (mol photons cm^-2 s^-1)^-1 in September to 0.3 mol O₂ g^-1 fr wt h^-1 (mol photons cm^-2 s^-1)^-1 in January, the initial light saturating point (I _k) gradually increased from 19 to 60 mol photons m^-2 s^-1. Likewise C:N ratios were low in spring (12 to 13) and increased in summer (20). In general, the photosynthetic parameters P _max, gross photosynthesis, and Chl a concentrations were significantly higher in the distal section of the thallus. In contrast, C:N ratios were lower in the distal section of the lamina. The results show that photosynthesis obviously strongly supports growth of the alga in late winter to spring, as it does in some morphologically related brown algae from temperate and polar regions. The question whether growth is additionally powered     

Photosystem II photoinhibition and altered kinetics of photosynthesis during nutrient-dependent high-light photoadaptation in Gonyaulax polyedra

Gonyaulax poledra Stein was transferred at different cell densities from increasingly nutrient-limited low-light (LL, 80 E m^-2 s^-1) batch-cultures to high-light (HL, 330 E m^-2 s^-1) growth conditions. Several age-dependent differences in HL-adaptation strategies were apparent. Short-term (3h) susceptibility to photosynthetic photoinhibition increased with culture age, with light-limited rates of photosynthesis exhibiting greater photosuppression than light-saturated rates at all stages of growth. These shortterm changes were not accompanied by photobleaching of chlorophyll but were directly related to age-dependent photoinactivation of Photosystem II electron-transport rates. The capacity of electron transport by Photosystem I was only slightly affected. Prolonged exposure of LL log-phase cells to HL conditions did induce photobleaching of chlorophyll associated with increased cell volume, a transient decrease of organic carbon and nitrogen content, enhanced cellular-, carbon-and chlorophyll-based rates of light-saturated photosynthesis (P _max) and suppressed cellular rates of light-limited photosynthesis. As a result, the density of LL log-phase cells doubled and their cellular photosynthetic performance nearly tripled within 1 d of HL exposure while cellular respiratory demands remained unchanged. By contrast, prolonged HL incubation of LL stationary populations induced a transitory burst in cell division and a large reduction in cell volume, leading to a short-term increase in volume-based organic carbon and nitrogen content. Despite reduced cell volume and lowered carbon demand, the cellular-, carbon-and chlorophyll-based rates of P _max in nondividing populations fell by 64, 48 and 27%, respectively, over a 4 d exposure to HL, while light-limited rates were almost fully suppressed within 1 d and chlorophyll a content was reduced by 56%. As a result, the photosynthetic performance of LL-aged cells declined immediately under HL conditions. Addition of inorganic nutrients to LL stationary cultures at the time of HL transfer led to immediate and complete suppression of photosynthesis and cell lysis within 1 d. Addition of nutrients following transfer to HL induced cell responses intermediate to those described for LL log and aged cells exposed to HL. Results support the view that declining nutrient-status impairs HL photoadaptive responses in phytoplankton populations and that the rate and pattern of photoadaptive responses may be used as physiological growth indicators in field studies. The study was conducted from March 1981 to May 1983.     

Effect of feeding regime and irradiance on the photophysiology of the symbiotic sea anemone Aiptasia pulchella

To determine how the animal and algal components of the symbiotic sea anemone Aiptasia pulchella respond to changes in food availability and culture irradiance, sea anemones from a single clone were maintained at four irradiance levels (320, 185, 115, and 45 E m^-2 s^-1) and either starved or fed for 5 wk. Changes in protein biomass of sea anemones maintained under these conditions were not related to the productivity of zooxanthellae, since the protein biomass of fed A. pulchella decreased with increase in irradiance and there was no difference in protein biomass among starved sea anemones at the four irradiance levels. Except for the starved high-light sea anemones, the density of symbiotic zooxanthellae was independent of culture irradiance within both starved and fed. A. pulchella. Starved sea anemones contained over twice the density of zooxanthellae as fed sea anemones. Within both starved and fed individuals, chlorophyll per zooxanthella increased with decreasing culture irradiance while algal size remained constant (in fed sea anemones) at about 8.80 m diameter. Chlorophyll a: c ₂ ratios of zooxanthellae increased with decreasing culture irradiance in zooxanthellae from starved sea anemones but remained constant in zooxanthellae from fed sea anemones. As estimated from mitotic index data, the in situ growth rates of zooxanthellae averaged 0.007 d^-1 and did not vary with irradiance or feeding regime. Photosynthesis-irradiance (P-I) responses of fed A. pulchella indicated an increase in photosynthetic efficiency with decreasing culture irradiance. But there was no consistent pattern in photosynthetic capacity with culture irradiance. Respiration rates of fed sea anemones also did not vary in relation to culture irradiance. The parameter I _k , defined as the irradiance at which light-saturated rates of photosynthesis are first attained, was the only parameter from the P-I curves which increased linearly with increasing culture irradiance. The daily ratio of net photosynthesis to respiration for A. pulchella ranged from 1.6 to 2.8 for sea anemones maintained at the three higher irradiances, but was negative for those maintained at 45 E m^-2 s^-1. Since the final protein biomass was greatest for sea anemones maintained at the lowest irradiance, these results indicate that sea anemone growth cannot be directly related to productivity of zooxanthellae in this symbiotic association.     

Evidence for facultative heterotrophy in cultured zooxanthellae

The locus of symbiotic dinoflagellates within host cells provides a habitat which could potentially be exploited by the alga through heterotrophic uptake of host-derived organic substrates. Using zooxanthellae (Symbiodinium sp.) isolated from the tropical sea anemone Aiptasia pulchella collected from Kaneohe Bay, Hawaii, the effect of various potential organic substrates on growth in vitro was assessed in Erdschreiber seawater medium (ES) supplemented with organic compounds. Zooxanthellae maintained at 5 to 7 E m^-2 s^-1 (below compensation irradiance) grew heterotrophically when supplied with 100 M glycerol, glycolate, acetate, malate, or propionate, and grew in darkness on 100 M propionate. Zooxanthellae exposed to irradiance below compensation were able to utilize carbon sources in the unsupplemented ES medium for slow growth, but generally the growth rate of cultured zooxanthellae was a function of incubation irradiance. Zooxanthellae incubated for 10 wk in unsupplemented ES at 5 to 7 E m^-2 s^-1 were capable of growth at this low irradiance, but were also capable of net photosynthetic oxygen production at higher irradiances. This suggests that zooxanthellae can be photoautotrophic or facultatively heterotrophic. An estimate for the duration of mitosis (t _d ) is made on the basis of growth rate of cultured zooxanthellae in log-phase; this estimate of t _d =4.88 h is less than half the estimated t _d for zooxanthellae in situ.     

Effects of four environmental variables on photosynthesis-irradiance relationships in Antarctic sea-ice microalgae

The effects of temperature, salinity, growth irradiance and diel periodicity of incident irradiance on photosynthesis-irradiance (P-I) relationships were examined in natural populations of sea-ice microalgae from McMurdo Sound in the austral spring of late 1984. Both P _m ^b (photosynthetic rate at optimum irradiance) and ^b (initial slope or P-I curve) were temperature-dependent reaching optimal rates at approximately +6° and +2°C, respectively. P-I relationships showed little difference at 20 and 33 S; however, no measurable photosynthesis by sea-ice microalgae was detected in a 60 S solution of brine collected from the upper layers of congelation ice. Although diel periodicity characteristic of the under-ice light field appeared to have little effect on P-I relationships, changes in growth irradiance had a profound effect. An increase in growth irradiance from 7 E m^-2 s^-1 (ambient) to 35 or 160 E m^-2 s^-1 resulted in a transient three-fold increase in P _m ^b and I _k (index of photoadaptation) during the first four days, followed by a sharp decline. The effects of these environmental factors on ice algal photosynthesis may influence the distribution of microalgae in sea-ice environments.     

Photosynthesis-irradiance responses and photosynthetic periodicity in the sea anemone Aiptasia pulchella and its zooxanthellae

Sea anemones (Aiptasia pulchella) containing zooxanthellae (Symbiodinium microadriaticum) were maintained in a long-term laboratory culture on a 12 h light (100 E m^-2 s^-1):12 h dark cycle. Photosynthetic oxygen production was measured for the symbiotic association and for freshlyisolated zooxanthellae. Light utilization efficiencies () were similar for both sets of zooxanthellae, suggesting negligible shading of zooxanthellae by animal tissue in this association. Whereas freshly-isolated zooxanthellae were photoinhibited at high irradiances (800 to 1 800 E m^-2 s^-1), zooxanthellae in the host continued to function at photosynthetic capacity. Time of day may influence photosynthetic measurements in symbiotic organisms, as it was found that photosynthesis in A. pulchella followed a diel periodicity at both light-saturating (1 200 E m^-2 s^-1) and subsaturating (150 E m^-2 s^-1) irradiances. There was a peak period of photosynthesis between 12.00 and 14.00 hrs. Light stimulated dark respiration rates of A. pulchella. Dark respiration of sea anemones increased somewhat towards the end of the light cycle and was always greater after exposure to high irradiances.     

Production and role of hyaline hairs in Ceramium rubrum

Light intensity within the range of 40 to 210 E · m^-2 · s^-1 had relatively little effect on the abundance, length or distribution of Ceramium rubrum unicellular, hyaline hairs. External NH ₄ ⁺ concentrations less than 0.5 M stimulated hair initiation and growth in apical regions of the thalli. Ammonium concentrations in excess of 20 M inhibited hair formation on all regions of the thalli. Ammonium uptake velocities of plants with hairs were approximately twice those of plants without hairs. These hairs may be adaptive to take advantage of intermittent bursts of nutrients by increasing the plant's surface area and, hence, the number of nutrient uptake sites.Woods Hole Oceanographic Institution Contribution No. 5526