Photosynthetic and optical properties of the marine chlorophyte Dunaliella tertiolecta grown under fluctuating light caused by surface-wave focusing

Photosynthetic and optical properties of the marine chlorophyte Dunaliella tertiolecta Butcher were studied in response to irradiance fluctuations caused by surface-wave focusing. The experimental conditions simulated the prominent features of the light field (high average irradiance, spectral composition and statistical properties) in the uppermost few meters of the water column under sunny surface conditions. The properties of algae grown under high-frequency fluctuations were compared with control cells grown under constant light at the same average irradiance (800 mol quantam^-2s^-1). No significant differences were found for a number of parameters, including growth rate, cellular chlorophyll a and pigment ratios, photosynthetic unit size and density of Photosystem I reaction centers, the rate of photosynthesis at the growth irradiance, dark respiration, and in vivo fluorescence of chlorophyll a per cell. Photosynthetic parameters were not affected by whether the incident light for oxygen exchange measurements was fluctuating or constant. This was the case whether the cells had been previously acclimated to either fluctuating or constant irradiance. Such a photosynthetic response indicates that cells are accomplishing a time integration of the fluctuating light. In addition, although D. tertiolecta is capable of dramatically changing its optical properties in response to low or high growth irradiance levels, the refractive index of the cells, the efficiency factors for light absorption and scattering by individual cells, and chlorophyll-specific absorption and scattering coefficients of cell suspensions, were all very similar under high irradiance, whether or not wave focusing was present.Contribution to the program of GIROQ (Groupe Interuniversitaire de Recherches Océanographiques du Québec)     

Light quality and concentration of proteins,RNA, DNA and photosynthetic pigments in two species of marine plankton algae

Chlorophyll, protein, DNA and RNA concentrations in cultures ofCyclotella nana (Hustedt) andDunaliella tertiolecta (Butcher) were higher in blue light and lower in green light than in white light of the same intensity. Total carotenoid concentrations were highest in green light. Total pigment concentrations were highest in cells grown in blue light, lowest in those from green light. The relative changes in pigment concentrations associated with differences in spectral composition of the radiation are comparable to those often observed in phytoplankton from various depths in stratified natural water. Light adaptation in algae as a response to light quality is suggested. The chemical composition of plankton algae can be expected to vary considerably with depth.     

Kinetic evidence for common photosynthetic step in diverse seaweeds

The turnover time of photosynthetic oxygen production was the same, near 0.5 msec, for the seaweeds Ulva lactuca, Codium fragile, Porphyryaumbilicalis, Chondrus crispus, Champia parvula and Fucus vesiculosis. This turnover time did not change for sun- and shade-adapted U. lactuca and P. umbilicalis. The similarity of these turnover times to those of other algae, higher plants, and cyanobacteria is strong evidence for similarity in the kinetics, and thus for a universal mechanism of photosynthesis. Evolution in the light-gathering apparatus responsible for the color of seaweeds has occurred without variation in the mechanism of photosynthesis. The size of the Emerson-Arnold photosynthetic unit (the ratio of chlorophyll to oxygen formed in a single tumover light flash) in the green seaweeds is about the same as that in algae, i.e., 2000. It is about half this number for the red and brown seaweeds. The different accessory pigments in the latter two groups compensate for the smaller amount of chlorophyll. The size of the unit was independent of sun- and shade-adaptation in U. lactuca, but did increase in shade-adapted P. umbilicalis.     

Adaptation of light-harvesting pigments to downwelling light and the consequent photosynthetic performance of the eulittoral rockweeds Ascophyllum nodosum and Fucus vesiculosus

We compared the effect of habitat and water depth on the light-harvesting pigment content for Ascophyllum nodosum and Fucus vesiculosus at two near-shore stations in Long Island Sound (USA). Excised pieces of seaweeds were attached at depth intervals to a vertically buoyed line, and left in situ for 7 days. For comparison, fronds were collected from sun and shade habitats in the littoral zone. The three major antenna (light-harvesting) pigments increased in concentration with depth or shade. Chlorophyll c to a ratios remained stable at about 0.2. Fucoxanthin to chlorophyll a ratios decreased by 20 to 30% with depth or shade. Although pigment composition for the two rockweed species was equivalent, the maximum photosynthetic performance of F. vesiculosus exceeded that of A. nodosum by a factor of 2, while the compensation depths for 4 m-adapted A. nodosum and F. vesiculosus under natural limiting light conditions were equivalent. Plants held at 4 m had higher photosynthetic rates compared with plants held at 0 m, no matter the depth of measurement. Indirect evidence indicates that the enhanced photosynthesis of 4 m-adapted plants is due not only to higher concentrations of antenna pigments but to other physiological factors as well. We conclude that the clearly delineated vertical distribution of these two canopy species, the F. vesiculosus zone over the A. nodosum zone, is not determined by light quantity or quality, but by biotic factors as evidenced by the experiments of Menge which are cited herein.     

Ecotypic differentiation in the marine diatom Skeletonema costatum: influence of light intensity on the photosynthetic apparatus

Three genetically distinct clones of Skeletonema costatum (Grev.) Cleve were grown at 20°C under high (274 E m^-2 s^-1) and low (27 E m^-2 s^-1) light conditions and their photoadaptive photosynthetic responses compared. When all three clones were grown under low light, pigment analyses and fluorescence excitation spectra demonstrated that the accessory pigments, chlorophyll c and fucoxanthin, became more important in light-harvesting compared to chlorophyll a. Photosynthetic unit sizes increased for Photosystems I and II in low light, but photosynthesis vs irradiance characteristics were not reliable predictors of photosynthetic unit features. Fluorescence excitation spectra and photosynthesis vs irradiance (P-I) relationships indicated that changes in energy transfer occurred independent of changes in pigment content. Large increases in accessory pigment content were not accompanied by large increases in excitation from these pigments. Changes in energy transfer properties were as important as changes in PSU size in governing the photoadaptive responses of S. costatum. When the three clones were grown under identical conditions, each had a separate and distinct pattern of photoadaptation. Significant differences among clones were found for pigment ratios, photosynthetic unit sizes for Photosystems I and II and efficiency of energy transfer between pigments. These strikingly different photoadaptive strategies among clones may partially account for the great ecological success of the diatom species. This is the first quantitative investigation of the importance of both chlorophyll c and fucoxanthin to the adaptive responses of diatoms to light intensity, and represents the most complete characterization of the photoadaptive responses of a single species of marine phytoplankter to differences in light environment.     

Adaptation of solitary corals and their zooxanthellae to low light and UV radiation

Adaptation of solitary corals, Fungia repanda and F. echinata, and their zooxanthellae to low light and ultraviolet light B (UV-B) was studied with respect to changes in their protein contents, photosynthetic pigment contents and the photosynthesis-irradiance (P-I) curves. The corals were collected from 1 to 50 m depths in the Republic of Belau (Paulau) in 1990 and 1991. The chlorophyll a content in a unit surface area of the coral did not change significantly with the depth of the habitat, whereas cellular chlorophyll a in the algae increased with the depth. Zooxanthellae density and protein content in a unit surface area of Fungia spp. decreased with the depth. Photosynthetic parameters normalized by a unit surface area of the Fungia spp., maximum gross photosynthetic rate (P _gmax area^-1) and dark respiration rate (R area^-1), were negatively correlated with the depth, while initial slope of the P-I curve () did not show significant correlation with the depth. Compensation light intensity (Ic) decreased with the depth. In isolated zooxanthellae, P _max chl a ^-1, and R chl a ^-1 decreased with the depth, while _{chl a} was constant. P _gmax cell^-1 and R cell^-1 did not change significantly but _cell increased with the depth. Ic decreased with the depth as in the intact corals. Reduction of protein content in a unit area of the coral from deeper habitat implies decrease of host animal tissues. Reduction of Ic can be explained by decrease of R area^-1, which may be due to the diminution of animal tissues. The photoadaptational response to low light intensity of intact Fungia spp. was found to be a combination of the photoadaptation of symbiotic algae and the decrease of host animal tissue. In order to study their adaptation to ultraviolet (UV) radiation, P-I curves of Fungia spp. and isolated zooxanthellae were analyzed before and after UV-B irradiation. 1 h UV-B irradiation showed no effect on the photosynthetic rate of the shallow water (1 m) corals, while it inhibited the photosynthesis of the deep water (30 m) corals and zooxanthellae isolated from both shallow and deep water corals. These results indicate that the host, Fungia spp., in shallow water have protective mechanism for intense UV-B in their habitat. These photoadaptational mechanisms seem to allow the Fungia spp. to have wide vertical distribution where light intensity spans more than two orders of magnitude.     

Effects of wave-induced lightflecks in the intertidal zone on photosynthesis in the macroalgae Postelsia palmaeformis and Hedophyllum sessile (Phaeophyceae)

Intertidal algae are exposed to a highly variable photic regime because of crashing waves. We measured photosynthetic rate of whole fronds of the seaweeds Postelsia palmaeformis (Ruprecht) (at Bodega Marine Laboratory, 1991) and Hedophyllum sessile (Setchell) (Phaeophyceae) (at Friday Harbor Laboratories, 1990) in flashing and steady-state photosynthetically active radiation (PAR) of equal irradiance, using a recirculating metabolism chamber designed to minimize the thickness of diffusional boundary layers on the surfaces of algal tissues. The dimensionless ratio of photosynthetic rate under flecking PAR (P_fleck) and the sum of photosynthetic rate under steady state PAR (P_s) of high and low irradiance was computed for lightfleck periods from 0.2 to 100 s. P_fleck:P_s is a measure of the light-flash utilization-efficiency in flashing light, and was greater than unity at periods between lighflecks of 2 to 30 s, with a peak at 10 s. We used a novel optical fiber irradiance meter to measure PAR incident on fronds of P. palmaeformis as they were washed about by waves breaking in the intertidal zone, and compared the light records to that obtained by a stationary sensor under the canopy. PAR flashing period was closely correlated with the period of breaking waves in stands of P. palmaeformis. We estimated the seasonal spectrum of period between light flashes in stands of this species by examining the spectral density of ocean waves at Bodega Marine Laboratory, Bodega Bay, California. The wave spectrum peaks at a period (10 s) where light-flashes may theoretically enhance the light-flash utilization-efficiency of seaweeds. We calculate that the enhanced light-flash utilization-efficiency wrought by wave-induced light-flashes may contribute to significant gains in primary productivity of these macroalgae under some conditions.     

Floristic and physiological differences between the shallow and the deep nanophytoplankton community in the euphotic zone of the open tropical Atlantic revealed by HPLC analysis of pigments

Suspended matter sampled in 1982 in the North Equatorial Current, in the open Atlantic to the west of West Africa, was analyzed by high performance liquid chromatography. The pigment fingerprint of samples taken in the surface mixed layer was dominated by zeaxanthin and chlorophyll a, in agreement with observed dominance of coccoid cyanobacteria. Near the bottom of the euphotic zone the fingerprint was more complicated, with a sharp transition at the depth of the deep chlorophyll maximum layer to dominance of chlorophyll b, 19-hexanoyloxyfucoxanthin and an unknown fucoxanthin derivative in the lower part of this layer; this fingerprint suggests dominance of eukaryotes (green algae, Prymnesiophyceae and Chrysophyceae) at depth. Up to 90% of the chl a was contained in particles smaller than 8 m, and in the surface mixed layer even more than 50% in particles smaller than 1 m. The high concentration of zeaxanthin relative to chl a near the surface suggests adaptation of the cyanobacteria to exposure to high irradiance. Evidence of this adaptation was the very high specific phytoplankton growth rate between sunrise and sunset (=0.16 h^-1), measured by recording ¹⁴C incorporation into organic carbon and into chl a carbon after isolation of the latter by HPLC. The high concentration of chl b relative to chl a at depth was possibly caused by shade-adapted green algae containing more chl b than chl a. The specific growth rate of the deep shade community was low (<0.04 h^-1), yet net primary production, calculated on the basis of chl a increase during incubation, was greatest at depth.     

Interplay of host morphology and symbiont microhabitat in coral aggregations

The Belizean reef coral Agaricia tenuifolia Dana forms aggregations in which rows of thin, upright blades line up behind each other. On average, the spacing between blades increases with depth and hence with decreasing ambient irradiance. We designed and built a small, inexpensive light meter and used it to quantify the effect of branch spacing on light levels within colonies at varying distances from branch tips. Concurrently, we measured photosynthetic pigment concentrations and population densities of symbiotic dinoflagellates (zooxanthellae) extracted from coral branches of colonies with tight (≤3 cm) vs wide (≥6 cm) branch spacing, collected at 15 to 17 m and from colonies with tight branch spacing collected at 1 to 2 m. Light levels decreased significantly with tighter branch spacing and with distance from the branch tips. Total cellular pigment concentrations (chlorophylls a, c ₂ and peridinin) as well as chlorophyll a:c ₂ and chlorophyll a: peridinin ratios all increased significantly with distance from the branch tip, indicating very localized differences in photoacclimation within individual branches. Zooxanthellae from colonies with widely-spaced branches displayed significantly lower chlorophyll a:c ₂ and chlorophyll a:peridinin ratios, and were present at significantly higher population densities than those from colonies with tightly-spaced branches collected at the same depth (15 m). Tightly-spaced colonies collected from shallow environments (1 to 2 m) displayed pigment ratios similar to those from widely-spaced colonies from deeper water (15 m), but maintained zooxanthellae populations at levels similar to those in tightly-branched colonies from deeper water. Thus, variation in colony morphology (branch spacing and distance from branch tip) can affect symbiont physiology in a manner comparable to an increase of over 15 m of water depth. These results show that a host's morphology can strongly determine the microhabitat of its symbionts over very small spatial scales, and that zooxanthellae can in turn display steep gradients in concordance with these altered physical conditions. Received: 12 June 1997 / Accepted: 24 June 1997     

Phytoplankton photosynthetic adaptation to high frequency light fluctuations simulating those induced by sea surface waves

In order to test the ability of phytoplankton to adapt to the high frequency light fluctuations induced by sea surface waves, the green alga Dunaliella tertiolecta was grown under both steady and fluctuating (0.1, 1.0 and 10 Hz) illuminations. The latter conditions reproduced those fluctuations experienced by phytoplankton in the upper photic layer. For each culture, photosynthesis versus irradiance were measured under four incubation frequencies (steady, 0.1, 1.0 and 10 Hz fluctuating illuminations). Results indicated that growth rates were similar for algae grown under steady light and 10 Hz fluctuating light (0.26–0.33 d^–1). Cells grown at 0.1 and 1.0 Hz showed lower growth rates (0.17–0.26 d^–1). Chlorophyll a and b were significantly higher under 0.1 and 10 Hz frequencies than under steady illumination; at 1.0 Hz, there were no significant differences with steady light. No changes in carotenoids were evidenced at any frequency tested. Photosynthetic measurements showed that algae grown under steady illumination had higher photosynthetic efficiency and capacity when incubated under steady and 0.1 Hz fluctuating light. Photosynthetic characteristics of algae grown under 0.1 Hz illumination did not show any clear responses to fluctuating light. Algae grown under 1.0 or 10 Hz had higher photosynthetic efficiency and capacity than those grown under steady illumination, when incubated under 1.0 and 10 Hz light. This suggests that microalgae grown under high frequency illumination (1.0 and 10 Hz) can adapt their photosynthetic characteristics to the rapidly fluctuating light regime experienced during growth, and that algae grown under steady conditions respond better to steady or slowly fluctuating (0.1 Hz) light. Such an adaptation provides a means of probing the photosynthetic responses of phytoplankton to vertical mixing.Contribution to the program of GIROQ (Groupe interuniversitaire de recherches océanographiques du Québec)     

Adaptation strategies of the corallimorpharian <Emphasis Type="Italic">Rhodactis rhodostoma</Emphasis> to irradiance and temperature

Corallimorpharians may dominate some habitats on coral reefs and compete with stony corals for access to light, yet little is known concerning their photosynthetic traits. At Eilat in the northern Red Sea, we observed that the abundance of individuals of the corallimorpharian Rhodactis rhodostoma decreased significantly with depth on the reef slope. Field and laboratory experiments revealed that they employ several mechanisms of photoadaptation to high irradiance on the shallow reef flat. Their endosymbiotic microalgae (zooxanthellae) varied significantly in both abundance and chlorophyll content with level of irradiance. Use of a diving pulse amplitude modulated fluorometer revealed that the zooxanthellae of R. rhodostoma effectively disperse excess light energy by expressing significantly higher values of non-photochemical quenching and maximum excitation pressure on photosystem II when experimentally exposed to high light (HL) versus low light (LL). Host corallimorpharian tissues mediated this response by shielding the algal symbionts from high irradiance. The endoderm of host tentacles thickened significantly and microalgal cells were located further from the mesoglea in HL than in LL. The clades of zooxanthellae hosted by the corallimorpharians also varied with depth. In shallow water, all sampled individuals hosted clade C zooxanthellae, while in deep water the majority hosted clade D. The photosynthetic output of individuals of R. rhodostoma was less affected by HL than was that of a stony coral examined. When exposed to both high temperature (HT) and HL, individuals of R. rhodostoma reduced their maximum quantum yield, but not when exposed to HL at low temperature (LT). In contrast, colonies of the scleractinian coral Favia favus reduced their photosynthetic output when exposed to HL in both temperature regimes. After 2 weeks of HT stress, R. rhodostoma polyps appeared to bleach completely but re-established their zooxanthella populations upon return to ambient temperature. We conclude that mechanisms of photoadaptation to high irradiance employed by both the endosymbiotic zooxanthellae and host corallimorpharians may explain in part the abundance of R. rhodostoma on some shallow reef flats. The ability to survive for weeks at HT while bleached also may allow corallimorpharians to repopulate shallow reef areas where scleractinians have been killed by thermal stress. B. Kuguru and G. Winters contributed equally to this work.     

Aspects of water column primary productivity in the Chukchi Sea during summer

Measurements of primary productivity, chlorophyll a, incident solar radiation, phosphate-P, silicate-Si, nitrate-N, nitrite-N, ammonium-N, temperature and salinity were made in the Marginal Ice Zone of the Chukchi Sea in summer 1974. Low to moderate levels of primary productivity (0.07 to 0.97 g C m^-2 half-day^-1) were observed; primary productivity exceeded 3 g C m^-2 half-day^-1 at two stations. Surface primary productivity was nitrogen-limited at most stations. Mean chlorophyll a concentration in the photic zone varied from 0.4 to 17.8 mg m^-3. Higher concentrations and significant subsurface accumulation of chlorophyll a, reaching 40 mg m^-3, were observed in July at stations near the ice-edge than those in open water. No chlorophyll maximum was noted in September, when values ranged from 0.4 to 2.2 mg m^-3. It is postulated that the contribution of sea-ice algae to the total chlorophyll content can be substantial, but that the stay of these cells in the water column may not be long. Non-linear regression estimates from solar radiation and chlorophyll-specific primary productivity data showed a maximal photosynthetic rate of 18 mg C mg chlorophyll a ^-1 half-day^-1, an optimal light intensity of 54 langleys half-day^-1, and markedly reduced primary productivity at moderately higher light intensities. These features indicate that phytoplankton was shade-adapted.     

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.     

Photosynthetic characteristics of Prochloron sp./ascidian symbioses

The prokaryotic green alga Prochloron sp. (Prochlorophyta) is found in symbiotic association with colonial didemnid ascidians that inhabit warm tropical waters in a broad range of light environments. We sought to determine the light-adaptation features of this alga in relation to the natural light environments in which the symbioses are found, and to characterize the temperature sensitivity of photosynthesis and respiration of Prochloron sp. in order to assess its physiological role in the productivity and distribution of the symbiosis. Colonies of the host ascidian Lissoclinum patella were collected from exposed and shaded habitats in a shallow lagoon in Palau, West Caroline Islands, during February and March, 1983. Some colonies from the two light habitats were maintained under conditions of high light (2 200 E m^–2 s^–1) and low light (400 E m^–2 s^–1) in running seawater tanks. The environments were characterized in terms of daily light quantum fluxes, daily periods of light-saturated photosynthesis (H_sat), and photon flux density levels. Prochloron sp. cells were isolated from the hosts and examined for their photosynthesis vs irradiance relationships, respiration, pigment content and photosynthetic unit features. In addition, daily P:R ratios, photosynthetic quotients, carbon balances and photosynthetic carbon release were also characterized. It was found that Prochloron sp. cells from low-light colonies possessed lower chlorophyll a/b ratios, larger photosynthetic units sizes based on both reaction I and reaction II, similar numbers of reaction center I and reaction center II per cell, lower respiration levels, and lower P_max values than cells from high-light colonies. Cells isolated from low-light colonies showed photoinhibition of P_max at photon flux densities above 800 E m^–2 s^–1. However, because the host tissue attenuates about 60 to 80% of the incident irradiance, it is unlikely that these cells are normally photoinhibited in hospite. Collectively, the light-adaptation features of Prochloron sp. were more similar to those of eukaryotic algae and vascular plant chloroplasts than to those of cyanobacteria, and the responses were more sensitive to the daily flux of photosynthetic quantum than to photon flux density per se. Calculation of daily minimum carbon balances indicated that, though high-light cells had daily P:R ratios of 1.0 compared to 4.6 for low-light cells, the cells from the two different light environments showed nearly identical daily carbon gains. Cells isolated from high-light colonies released between 15 and 20% of their photosynthetically-fixed carbon, levels sufficient to be important in the nutrition of the host. Q₁₀ responses of photosynthesis and respiration in Prochloron sp. cells exposed briefly (15–45 min) to temperatures between 15° and 45°C revealed a discontinuity in the photosynthetic response at the ambient growth temperatures. The photosynthetic rates were found to be more than twice as sensitive to temperatures below ambient (Q₁₀=3.47) than to temperatures above ambient (Q₁₀=1.47). The Q₁₀ for respiration was constant (Q₁₀=1.66) over the temperature range examined. It appears that the photosynthetic temperature sensitivity of Prochloron sp. may restrict its distribution to warmer tropical waters. The ecological implications of these findings are discussed in relation to published data on other symbiotic systems and free-living algae.     

Nutrient enrichment of seagrass beds in a rhode island coastal lagoon

Seagrass and algal beds showed a variety of reponses when the water column was treated with low level additions of ammonium, nitrate and phosphate. The nutrients were added separately to 3 uniform seagrass beds of a temperature coastal lagoon during 1979 and 1980. (1) Ammonium caused the production of dense mats of free-floating green algae Enteromorpha plumosa and Ulva lactuca. It also stimulated growth in both the leaf and root-rhizome fractions of Zostera marina. This growth response in Z. marina was greater in the area where current reached 12 cm · s^-1 than in the area with little or no current. The concentration of nitrogen in the tissue did not change. In contrast, where current was lacking, Z. marina growth increase with ammonium was small, but the concentration of nitrogen in the tissue doubled over that in control plots. The growth of Ruppia maritima was inversely related to the growth of green algae in the same plots. The red alga Gracilaria tikvahiae did not grow better in ammonium, but its tissue reddened. (2) Nitrate additions enhanced the growth of the green seaweeds Enteromorpha spp. and U. lactuca, but not Z. marina or R. maritima. G. tikvahiae, when fertilized in isolation from other plants, showed a marginal response to this nutrient, and the tissue always reddened. (3) Phosphate enhanced growth in Z. marina and R. maritima exposed to moderate current. G. tikvahiae growing alone showed a small growth response to phosphate. The phosphate made no difference in the growth of the green seaweeds. (4) None of the nutrient supplements noticeably altered the species composition of either epiphytic or planktonic algae associated with the beds, although we did detect small increases in their numbers. The rapid and dense growth of green algae in nitrogen-enriched water probably limited growth of adjacent seagrasses and red algae. Because these seaweeds did not use the phosphate, it became available to other plant components. The overall floral response to nutrient addition in seagrass communities depends, therefore, upon the particular nutrient supplied, the ability of alternate species in the area to compete for that nutrient and the velocity of current in the specific area.     

Pigment content and photosynthetic rate of the fronds of Macrocystis pyrifera

The Macrocystis pyrifera (L.) C. Ag. frond is here described in terms of chlorophyll a, fucoxanthin, chlorophyll c and photosynthetic rate. Pigment concentrations increased back from the apical meristem reaching a maximum after 2 to 3 m. Pigment concentrations were then generally constant throughout most of the length of the frond, finally decreasing again in the oldest parts of the frond with the exception of the sporophylls. Pigment ratios remained relatively constant throughout. Maximum net photosynthetic rates on a given frond showed a decrease with tissue age on both an area basis (1040 down to 463 nmol O₂ cm^-2 h^-1) and on a chlorophyll a basis, which was shown as half-saturation constants (quantum irradiance) which dropped on an area basis from 85 mol m^-2 sec^-1 at 4.5 m above the holdfast to 26 mol m^-2 sec^-1 at 15.5 m. Young sporophytes transplanted from the sea floor to the surface (12 m) tended to decrease pigment content, while those transplanted to the bottom tended to increase all pigments, but especially fucoxanthin. Photosynthetic rates, however, changed little on a unit area basis. The results of these data are considered in the light of recent work on photosynthetic units, tissue age effects and general adaptations of the M. pyrifera frond to its light environment.     

Light absorption and utilization among hermatypic corals: a study in Jamaica,West Indies

The chlorophyll specific absorption coefficient ( _c) was measured for zooxanthellae from six hermatypic coral species obtained, where possible, from four depths (1, 10, 30, 50 m) on reef sites near Discovery Bay, Jamaica in February and March 1983. Measurements of photosynthetic rates versus irradiance, as well as cellular and areal chlorophyll a, were also performed on these colonies or sister colonies. Together the data were used to compare minimum quantum requirements (1/Φ m) among species and depths and to assess the importance of light utilization to the growth and depth distribution of these corals. Our data suggest that, although _c was found to decrease with depth, interspecific differences in _c do not occur for zooxanthellae from the corals investigated. Minimum quantum requirements (1/Φ m) decreased significantly with depth, thereby reflecting an increase in photosynthetic light utilization efficiency with decreasing irradiance. Interspecific differences in 1/Φ m determinations were suggested but not statistically conclusive. We conclude that interspecific differences in gross photosynthesis, and perhaps growth and depth distribution, are primarily attributable to differences in the light utilization capacity of the whole coral, as reflected by the product of _c ^′ and chlorophyll per unit surface area, and in-situ quantum efficiencies. This research was performed under the auspices of the US Department of Energy under Contract No. DE-AC02-76CH00016     

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.     

Photoadaptation of zooxanthellae in the sponge Cliona vastifica from the Red Sea, as measured in situ

In the Red Sea, the zooxanthellate sponge Cliona vastifica (Hancock) is mainly present at >15 m depth or in shaded areas. To test whether its scarcity in unshaded areas of shallower waters is linked to the functional inefficiency of its photosymbionts at high irradiances, sponges were transferred from 30 m to a six times higher light regime at 12 m depth, and then returned to their original location. During this time, photosynthetic responses to irradiance were measured as rapid light curves (RLCs) in situ by pulse amplitude modulated (PAM) fluorometry using a portable underwater device, and samples were taken for microscopic determinations of zooxanthellar abundance. The zooxanthellae harboured by this sponge adapted to the higher irradiance at 12 m by increasing both their light saturation points and relative photosynthetic electron transport rates (ETRs). The ETRs at light saturation increased almost fourfold within 15–20 days of transfer to the shallower water, and decreased back to almost their original values after the sponges were returned to 30 m depth. This, as well as the fact that the photosynthetic light responses within an individual sponge were in accordance with the irradiance incident to specific surfaces, shows that these photosymbionts are highly adaptable to various irradiances. There was no significant change in the number of zooxanthellae per sponge area throughout these experiments, and the different photosynthetic responses were likely due to adaptations of the photosynthetic apparatus within each zooxanthella. In conclusion, it seems that parameters other than the hypothesised inability of the photosymbionts to adapt adequately to high light conditions are the cause of C. vastifica's rareness in unshaded shallow areas of the Red Sea. Received: 25 April 2000 / Accepted: 13 October 2000     

Vertical heterogeneity in physiological characteristics of Ulva spp. mats

In eutrophic areas, green macroalgae are frequently and for long periods arranged in mats, resulting in a steep light gradient. This study investigates the effect of this gradient on physiological characteristics [tissue nitrogen content, maximal photosynthetic efficiency (F_v/F_m), glutathione levels and redox ratio, absorbance and absorption spectra] of the green macroalga Ulva spp. Mats were sampled during the build-up (June), stationary (July), and decomposing (September) phases of a macroalgal bloom in the Veerse Meer, a eutrophic brackish (salinity 15–20 psu) lake in the southwest Netherlands. Water samples were taken for nutrient analyses. At all three sampling dates, the mats were composed almost entirely of Ulva spp.; in September the mats were in decay and covered with silt and epiphytes. In June and July, total dissolved inorganic nitrogen concentration (DIN) of the water within the mat was significantly higher than outside the mat. Pronounced vertical differences were found in tissue N, F_v/F_m values, total glutathione levels, glutathione redox ratios, and absorbance. In June and July, tissue N decreased from over 2.2% dry weight (DW; N-sufficient) in the bottom layers to around 1% DW (minimum level for survival) in the top layers. Wide-band absorption increased with depth in the mat and throughout the season, probably due to higher Chl a and b and lutein contents. The shape of the absorption spectrum was similar for all layers. The absorption of the silt/epiphyte film on the top Ulva layer was highest; its absorption spectrum (high absorption in the 500–560 nm range) indicates that the film on the top layers of the macroalgal mats mainly consisted of diatoms. In June, F_v/F_m and the glutathione redox ratio of the algae increased with depth in the layer, while total glutathione decreased. Low F_v/F_m values in the bottom and middle layers in September reflect the bad condition of the algae; the mats were largely decaying. It is concluded that multiple growth-limiting gradients occur in macroalgal mats: upper layers suffer from nitrogen limitation and photoinhibition while bottom layers are light limited. The algae in the mat acclimatize to low light conditions by increasing their absorption through increased pigment contents and by higher photosynthetic efficiency during the build-up and stationary period. This study qualifies the glutathione redox ratio as a promising candidate for stress indicator in macroalgae and provides suggestions for its further development.Communicated by S.A. Poulet, Roscoff