Effect of nitrogen supply on photosynthesis and carbonic anhydrase activity in the green seaweed Ulva rigida (Chlorophyta)

Photosynthesis rate and carbonic anhydrase (CA) activity have been studied in the green seaweed Ulva rigida C. Agardh (Chlorophyta) grown in seawater (SW) and SW supplemented with 40 M NH₄Cl (N-SW). Higher growth and maximal O₂ evolution rates were observed in N-SW- than in SW-grown sea-weeds. Western blot analysis of the total homogenates probed with antibodies raised against small subunits of ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBisCO) showed crossreaction with a 15 kdalton polypeptide in both SW- and N-SW-grown plants, although the band was more intense in N-SW-grown plants. Carbonic anhydrase activities in the total homogenate and in the soluble protein fraction were higher in N-SW-grown plants. Although the pellets from both plants showed a considerable CA activity, the activity of CA in the thylakoid membranes was undetectable. The low nitrogen concentration is a major environmental factor that affects the level of RuBisCO and CA, and therefore CO₂ assimilation in U. rigida.     

Ultrastructure of tetrasporogenesis in the red alga Osmundea spectabilis var. spectabilis (Rhodomelaceae: Ceramiales: Rhodophyta)

Tetrasporogenesis in Osmundea spectabilis (Postels and Ruprecht) K.W. Nam var. spectabilis proceeds through three developmental stages. A uninucleate tetraspore mother cell with synaptonemal complexes depicted during early prophase of meiosis I comprises the youngest stage. Subsequently, the nuclear envelope obtains a highly irregular profile and is surrounded by perinuclear endoplasmic reticulum (ER) and numerous mitochondria. Proplastids with peripheral thylakoid and numerous straight-profiled dictyosomes are present. Dictyosome vesicles discharge their contents, initiating tetraspore wall formation. A four-nucleate tetraspore mother cell is formed prior to tetrahedral cleavage. During the second stage of tetraspore formation the nuclei are very irregular in outline and move toward the central region where they fuse. The formation of fibrous vacuoles originates from the fibrous vacuole associated organelles (FVAOs). Thylakoid formation from concentric lamellar bodies (CLBs) seems to occur in plastids. Extremely hypertrophied dictyosomes with two tightly compressed and densely laminated mid-regions produce vesicles which contribute to enlargement of the fibrous vacuoles. During the second stage of tetraspore development striated vesicles are formed and starch granule formation increases. Mature tetraspores are characterized by numerous cored vesicles and abundant starch granules. Plastids have a fully developed system of internal thylakoids, and cleavage is usually complete at this stage. Mature tetraspores are surrounded by a bi-layered wall and the tetrasporangial wall. Unbranched, tubular structures connecting the plasmalemma with the peripheral sheets of ER are possibly involved in wall deformation. A distinct blebbing nuclear envelope appears to sequester vesicles into the cytoplasm.     

Interactions between light and temperature on the physiological ecology ofGracilaria tikvahiae (Gigartinales: Rhodophyta)

Main effects and interactions of light and temperature on rates of growth (), net photosynthesis (P_s), and dark respiration (R) of the red seaweedGracilaria tikvahiae were investigated in outdoor, nutrient-replete continuous-flow seawater culture chambers. Below 15°C,G. tikvahiae did not grow and between 15° and 30°C, both main effects and interactions of light and temperature on and P_s were significant, which explains the occurrence of this alga as a summer annual in its northern range. Temperature interacted with light (I) through its influence on the vs I and P_s vs I curves. The initial slope of the vs I curve, , the light saturation intensity, I_s, and maximum growth rate, _max, were all significantly lowerat 15°C compared to 20°, 25°, or 30°C. Maximum values of _max, the P_s:R ratio and the net photosynthesis:gross photosynthesis ratio (P_s:P_g) all occurred at 25°C, suggesting that this is the best temperature for growth ofG. tikvahiae. Values for P_max increased up to 30°C, indicating that the temperature for maximum growth and net photosynthesis are not the same forG. tikvahiae. Significant photoinhibition of growth and photosynthesis at full incident sunlight (I₀) occurred at 15°C but not at 20°, 25°, or 30°C. Steele's equation fit the 15°C vs I data best, whereas the hyperbolic tangent function fit the 20°, 25°, and 30°C data best. Main effects and interactionof light intensity and temperature on rates of R were also significant (P<0.001). R was highly intercorrelated with and P_s (0.86r0.94), indicating that R inG. tikvahiae is primarily regulated by growth rate and not temperatureper se. Environmental factors that regulate growth, such as light intensity, exert a great influence on R inG. tikvahiae.     

Interactions between light and temperature on the physiological ecology ofGracilaria tikvahiae (Gigartinales: Rhodophyta)

The biology and ecology of the Antarctic scallop Adamussium colbecki has been investigated on the west side of McMurdo Sound, Antarctica, principally at Explorers Cove, during the austral summer of 1981–1982. This conspicuous benthic invertebrate exhibits highest densities, 85 individuals m^-2, in shallow (4 to 6 m) water. Densities decrease to 20 m^-2 at 30 m. Biomass levels are highest in shallow water, 1 600 g wet wt m^-2. Provisional growth information suggests that 8 cm individuals are about 12 yr old. Mortality is apparently caused chiefly by a hyposaline lens of seawater, which forms under the sea ice during the summer melt; predators do not appear to be important. High biomass levels and a short generation time suggest that A. colbecki is an important species in a very productive community on the west side of McMurdo Sound. The shallow benthos of Explorers Cove is an important exception to the generalization that Antarctic ecosystems are dominated by indigestible filter-feeders.     

High-temperature tolerance of photosynthesis in the red alga Chondrus crispus

Chondrus crispus (Stackhouse) is a perennial red seaweed, common in intertidal and shallow sublittoral communities throughout the North Atlantic Ocean. In the intertidal zone, C. crispus may experience rapid temperature changes of 10 to 20C° during a single immerison-emerision cycle, and may be exposed to temperatures that exceed the thermal limits for long-term survival. C. crispus collected year-round at Long Cove Point, Chamberlain, Maine, USA, during 1989 and 1990, underwent phenotypic acclimation to growth temperature in the laboratory. This phenotypic acclimation enhanced its ability to withstand brief exposure to extreme temperature. Plants grown at summer seawater temperature (20°C) were able to maintain constant rates of lightsaturated photosynthesis at 30°C for 9 h. In contrast, light-saturated photosynthetic rates of plants grown at winter seawater temperature (5°C) declined rapidly following exposure to 30°C, reached 20 to 25% of initial values within 10 min, and then remained constant at this level for 9 h. The degree of inhibition of photosynthesis at 30°C was also dependent upon light intensity. Inhibition was greatest in plants exposed to 30°C in darkness or high light (600 mol photons m^-2s^-1) than in plants maintained under moderate light levels (70 to 100 mol photons m^-2s^-1). Photosynthesis of 20°C-acclimated plants was inhibited by exposure to 30°C in darkness or high light, but the degree of inhibition was less than that exhibited by 5°C-grown plants. Not only was light-saturated photosynthesis of 20°C plants less severely inhibited by exposure to 30°C than that of 5°C plants, but the former also recovered faster when they were returned to growth conditions. The mechanistic basis of this acclimation to growth temperature is not clear. Our results indicate that there were no differences between 5 and 20°C-grown plants in the thermal stability of respiration, electron transport associated with Photosystems I or II, Rubisco or energy transfer between the phycobilisomes and Photosystem II. Overall, our results suggest that phenotypic acclimation to seawater temperature allows plants to tolerate higher temperatures, and may play an important role in the success of C. crispus in the intertidal environment.     

Effects of the environment on developmental rates of sporelings of two Hypnea species (Rhodophyta: Gigartinales)

Growth of sporelings of Hypnea cervicornis J. Agardh and H. chordacea Kuetzing in response to gradients of selected environmental factors has been investigated. The sporelings of H. cervicornis tolerated higher levels of the tested parameters (e.g. salinity and temperature) than those of H. chordacea. In all the treatments, the sporelings of H. cervicornis grew significantly faster than those of H. chordacea. The ecological and agronomic implications of these findings are discussed.Extracted from a Ph. D. dissertation submitted to the Graduate Division of the University of Hawaii in Fall, 1974.     

Ultrastructure of carposporophyte development in the red algaCaulacanthus ustulatus (Gigartinales: Caulacanthaceae)

The ultrastructure of carposporophyte development is described for the red algaCaulacanthus ustulatus (Turner) Kützing collected at Epanomi (Gulf of Thessaloniki, Greece) and studied in 1988/1989. Following presumed fertilization the diploid nucleus is transferred to the auxiliary cell which expands and cuts off several multinucleate gonimoblast initials. The gonimoblast initials are distributed around the periphery of the cystocarpic cavity and divide further to produce generative gonimoblast cells. The latter cleave repeatedly to form clusters of carpospores. Thus, carposporophyte development proceeds inwardly. The first formed gonimoblast cell is transformed into a storage gonimoblast cell containing large quantities of starch granules and presumably functioning to supply nutrients. A fusion cell is never formed. Gonimoblast initials contain typical red algal proplastids and numerous sacs and/or vesicles originating from cytoplasmic concentric membranes.Please address all correspondence and requests for reprints to Dr S. G. Delivopoulos     

Spore germination and early stages of development in Hypnea musciformis (Rhodophyta,Gigartinales)

The early stages of development of the carpospores and tetraspores of Hypnea muscilormis (Wulfen) Lamouroux have been investigated. Both types of spores germinated immediately after liberation. The spores segmented repeatedly into, 2, 3, 4 cells, etc. until a multicellular ball of cells was produced. The germination pattern was thus of the discal type, i.e., Typus discalis mediatus (Inoh, 1947) or the Dumontia-type in the terminology of Chemin (1937). Subsequently, a relatively expansive attachment holdfast was produced from each sporeling. This was followed by the development of up to 4 or more shoot axes arising from the basal disc. These findings are discussed with reference to an earlier study by the senior author on the developmental biology of other species of Hypnea Lamouroux.     

Effect of freezing on photosynthesis of intertidal macroalgae: relative tolerance of Chondrus crispus and Mastocarpus stellatus (Rhodophyta)

The effect of freezing on photosynthetic metabolism was studied in the red algae, Chondrus crispus and Mastocarpus stellatus. Plants of both species were collected from the intertidal at Chamberlain or Kresge Point, Maine, USA (43°56N, 69°54W) between February and March 1987. Photosynthetic rates were measured immediately after freezing at-20°C and following recovery periods in seawater. Photosynthesis in C. crispus declined rapidly following freezing, falling to 70% of control values within 1 h and 30% after 3 h exposure. Minimum photosynthetic rates (7 to 9% of controls) occurred following freezing exposures of 12 h or more. Full photosynthetic recovery in C. crispus after 3 h at-20°C required 48 h. Photosynthesis in C. crispus did not fully recover in plants frozen for 6 h or more. In contrast, photosynthesis in M. stellatus was relatively unaffected by freezing exposures of <12 h. Twelve hours or more at-20°C reduced photosynthesis to 55% of controls. Photosynthesis in M. stellatus fully recovered from 24 h at-20°C within 24 h. In both species the reduction of photosynthesis by freezing was associated with damage to the plasma membrane and reduced efficiency of energy transfer from phycobilisomes to chlorophyll a, but did not appear to involve ribulose-1,5-bisphosphate carboxylase oxygenase activity. The freezing tolerance of C. crispus and M. stellatus positively correlates with their respective intertidal distributions, suggesting that freezing may be involved in controlling the distributions of these species on the shore.     

Alkaline phosphatase and carbonic anhydrase activity associated with arm regeneration in the seastarAsterias forbesi

A sharp increase in alkaline phosphatase (AP) activity occurred 7 d after arm autotomy inAsterias forbesi. This activity remained elevated at 20 d post-autotomy, during which time active ossicle-calcification was occurring. By 60 d post-autotomy, AP activity had returned to the level of non-regenerating individuals. Conversely, carbonic anhydrase (CA) activity exhibited no initial change in response to arm autotomy. By Day 60 a 50% increase in CA activity was observed. AP is localized in the outer membranes and associated microvilli of the coelomic epithelium and is not associated directly with the tissues involved in calcification. CA activity was highest in the tissues containing the ossicles, but was also present in the non-calcifying tube feet. These data suggest that inA. forbesi AP is involved in cell division and differentiation associated with wound-healing and the initiation of the regeneration process, and that CA may facilitate the ongoing maturation process, of newly formed ossicles.     

Chemically mediated interactions between the red algaPlocamium hamatum (Rhodophyta) and the octocoralSinularia cruciata (Alcyonacea)

Interactions between the red algaPlocamium hamatum J. Agardh (Rhodophyta) and other benthic organisms including the alcyonacean soft coralSinularia cruciata (Tixier-Durivault) were investigated on an inshore fringing reef environment in whichP. hamatum was the dominant large fleshy alga. Field observations of sessile reef organisms including octocorals and sponges living in close proximity toP. hamatum revealed that varying degrees of tissue necrosis were suffered by the invertebrates when in physical contact with the alga. In order to establish whether the chemical constituents of the alga, especially chloromertensene, played a role in this necrosis, manipulative field experiments were carried out in the Pelorus Channel, Palm Island group (18°34S; 146°29E), North Queensland, Australia, in November and December 1988. The first experiment involved the relocation of healthy plants and soft corals into contact and non-contact situations on a mesh grid. In all cases of contact betweenP. hamatum andS. cruciata, the soft coral suffered tissue necrosis (n=6,p=0.0022). The second experiment had the same design, but involved the use of artificial plants both uncoated and coated with natural levels of chloromertensene, in contact withS. cruciata. In all cases of contact with coated treatments, necrosis was observed inS. cruciata (n=4,p=0.025). In cases where uncoated artificial fronds were placed in contact with soft corals,S. cruciata showed minor abrasion effects, but no appreciable necrosis. Coated treatments were not fouled by epiphytes during the experiment and were not consumed by predators. Uncoated treatments were rapidly reduced in size by predation and any remaining material was biofouled. These experiments thus demonstrated that the deleterious effects observed in soft corals in the field were caused by contact with the algaP. hamatum, that these effects were indeed chemically mediated by chloromertensene, and that physical contact without chemical intervention caused no such deleterious effects. This is the first experimental evidence which conclusively demonstrates allelopathy between an alga and other marine organisms and identifies the compound responsible for the observed allelopathic effects.     

Effect of carbon dioxide concentration on calcification in the red coralline alga Bossiella orbigniana

The relationship between various experimental concentrations of CO₂ and calcification in Bossiella orbigniana (Decaisne) was studied by measuring Ca-45 incorporation into the crystalline matrix. Air containing CO₂ at partial pressures (P_{CO 2}) of 0.04 to 5.5% was bubbled through synthetic seawater in incubation vessels. The resultant pH values in the presence of plants ranged from 6.5 to 8.7. The maximum calcification rate appears to lie between 0.11 and 1.05% P_{CO 2}. The data suggest that calcification is controlled by a biological process that may be sensitive to pH and/or to the relative bicarbonate concentration. The data also suggest that a severalfold increase in CO₂ over the present atmospheric level might lead to increased calcification in this marine alga.     

Stress tolerance of photosynthesis in sporelings of the red alga Grateloupia doryphora compared to that of Stage III thalli

More experimental evidence is needed to understand the role of propagules in macroalgal biology. There are no reports in the literature on the comparative physiology (e.g. photosynthesis) of sporelings and adults. In this paper we report on the␣variation␣in␣photosynthetic parameters (maximum photosynthesis, P _max , and efficiency, alpha, and dark respiration (R _d ) of cultivated young sporelings of the red alga␣Grateloupia␣doryphora (Montagne) Howe under normal conditions and after a short-term incubation at different salinities and temperatures. The results are compared to those␣for␣adult Stage III thalli obtained in laboratory culture from the same population of sporelings. The pigment composition of sporelings (more chlorophyll a and less phycoerthryn and phycocyanin than adults) promotes a better photosynthetic performance (higher P _max and alpha and lower R _d ) under chlorophyll a excitation. The younger sporelings were also more tolerant to variations in salinity and temperature than Stage III, in which the highest variation in maximum photosynthesis and dark respiration was observed. Received: 21 October 1996 / Accepted: 5 February 1997     

Physiological role of branchial carbonic anhydrase in the shore crabCarcinus maenas

Excretion of total CO₂ and uptake of sodium and chloride ions across the branchial epithelium of the posterior gills of the shore crabCarcinus maenas, collected from Kiel Bay (Baltic Sea) in 1989, were measured using isolated perfused gill preparations. Total CO₂ effluxes depended on the HCO ₃ ^- concentration of the internal perfusate in a saturable mode and were inhibited by internally and externally applied acetazolamide at 10^–4 M. Potential differences between hemolymph space and medium did not change significantly during experimental treatments. Neither a bicarbonate gradient (6 mM) directed from the internal perfusate to external bath solution nor symmetrically applied 10^–4 M acetazolamide significantly influenced the influxes of Na⁺ and Cl^–. Results confirmed the role of carbonic anhydrase in CO₂ excretion but called into question the assumed functioning of the enzyme in branchial ion transport processes.     

Photosynthesis and growth of red and green morphotypes of Kappaphycus alvarezii (Rhodophyta) from the Philippines

The effect of photosynthetic available radiation (PAR) levels, light quality, ultraviolet (UV) radiation, and temperature on photosynthesis, growth, and chlorophyll fluorescence was evaluated in red and green morphotypes of the rhodophyte Kappaphycus alvarezii (Doty) Doty under controlled conditions. Chlorophyll a and phycoerythrin (PE) levels were similar in the red and green morphotypes cultured under the same conditions, but phycocyanin (PC) and allophycocyanin (APC) levels were 2-fold greater in the green than in the red morphotype. Pigment characterization indicated that the overexpression of PC and APC masked the red pigmentation in the green morphotype. Maximum photosynthesis and photosynthetic efficiency were similar between the two morphotypes assayed at a wide temperature range, which was reflected in the similar growth rates observed in outdoor culture systems. In the green morphotype, photosynthetic efficiency increased 2-fold relative to the red morphotype when assayed with red light (λ > 600 nm), indicating that photosynthetic characteristics are modified as a result of pigment variation in these morphotypes. Such increase in photosynthetic efficiency in the green morphotype, however, did not result in greater growth rates when cultured under white light. Short exposure to high levels of solar radiation (UV-A + UV-B + PAR), and filtered solar radiation (UV-A + PAR or PAR) decreased effective quantum yield (ΔF/F _m′) in both morphotypes. The reduction of ΔF/F _m′ values in the red and green morphotypes was accounted for by high levels of PAR and not by the UV-A + UV-B + PAR and UV-A + PAR treatments. Photoinhibition caused by UV-A, UV-B, or PAR was completely reversed within 30 h after incubations. Recovery rates from photoinhibition, however, were significantly reduced in the green morphotype when incubated with UV-B radiation. The results here suggest that the overexpression of pigments do not necessarily increase photosynthesis and growth in these morphotypes. Received: 19 June 2000 / Accepted: 28 November 2000     

Reproduction in Amphiura filiformis (Echinodermata: Ophiuroidea): Seasonality in gonad development

The burrowing ophiuroid Amphiura filiformis (O. F. Müller) (Echinodermata: Ophiuroidea) colonises an extensive area and is a numerically dominant member of the macrofauna in Galway Bay on the west coast of Ireland. There, it has a discrete, relatively short annual breeding period, from June to September, with peak activity in the month of August. The breeding period approximates in timing those of other European populations. The oocytes of a particular year are laid down in the autumn of the previous year; they begin to develop in spring, with the period of fastest growth in May/June. Gonad growth corresponds quite closely to the annual rise in water temperature, with spawning taking place during the warmest months of the year. The species is polytelic, and it is suggested that the long-lived Galway Bay individuals may breed every year for several years. The relationship between spawning and population dynamics is discussed in the light of current, available literature concerning A. filiformis from other parts of Europe. The role of so-called ultimate factors in the process of ophiuroid maturation and spawning is discussed.     

Large-scale estimate of somatic growth in Amphiura filiformis (Echinodermata: Ophiuroidea)

Quantitative data on size structure of the ophiuroid Amphiura filiformis (O.F. Müller) from 35 benthic stations in the Kattegat sampled twice with a 143 d interval, June and October 1991, were used to estimate somatic growth. The material was objectively divided into cohorts, and cohorts, from the two occasions were paired to give estimates of growth. The growth constant, K, in the Von Bertalanffy equation, was estimated from a Ford-Walford plot to 0.54 yr^-1. Results were in agreement with previous estimates from a few single sites, and suggest that the main part of dise growth occurs within the first 5 to 7 yr of living. Size specific growth in oral width was density independent despite high densities of A. filiformis (>3000 ind m^-2) and high total benthic biomass (up to 1000 g wet wt m^-2) in some areas. Growth was uncorrelated with plant pigment concentrations in the sediment and showed weak positive correlation with sediment carbon and nitrogen, as well as water depth. Growth was higher in fine sediments. This is the first attempt to estimate growth in this important species over a large area and to relate growth in the field to environmental factors.     

Reproductive biology of the red alga Polysiphonia lanosa (Ceramiales) in the Bay of Fundy, Canada

The purpose of this investigation was to establish reproductive phenology of the dioecious marine floridean red alga Polysiphonia lanosa (L.) Tandy. Specifically, the distribution and frequencies of male and female plants, timing of their sexual maturity, fecundity and success of fertilization were investigated. The presence of cystocarps was taken as an indication of syngamy. Populations of P. lanosa in the Bay of Fundy reproduce in spring, summer and autumn. Both male and female filaments were present in equal abundance at all sampling locations, grew in close proximity, and occurred with equal abundance at all elevations above chart datum. During the growing season most of the mature plants contained reproductive organs with apparently functional gametangia containing gametes the structure and behavior of which conformed to those reported in the literature for other species of the genus. Male and female plants of P. lanosa matured synchronously during late May to early July at all collecting sites. The ratio of spermatia to carpogonia calculated for the sampled population was high, 3000–4800 spermatia:1 carpogonium. At the end of the growing season (late October) the fertilization success was at its maximum of 50 to 90% and at its minimum in July at 24%. The three free-living life-history phases (females, males and tetrasporophytes) were abundant and occurred with equal frequency, indicating that a complete life cycle was common at the studied location. Received: 10 December 1996 / Accepted: 16 January 1997     

Growth and production of the red alga Gelidium sesquipedale off the Basque coast (northern Spain)

Several aspects of the growth of the agarophyte Gelidium sesquipedale Clemente (Thuret) were studied in sublittoral populations of the Basque coast (northern Spain) from 1981 to 1986. The standing stock of the populations varied greatly with season. During two consecutive years, 60 to 70% of summer biomass had been lost by the end of the unfavourable period of year to growth (autumn and winter). The decline in biomass was mainly due to partial losses of branches, with more branches being lost from larger fronds. Yearly variations in the standing stock were also recorded. Production estimates obtained by a destructive sampling method were not satisfactory because of the spatial heterogeneity of the populations at the three locations studied. Indirect estimation of production by measuring the growth of marked fronds in a control quadrat gave more reliable estimates. Production values of 622±124, 735±147 and 615±123 g dry wt m^-2 yr^-1 were obtained at the three locations, respectively. Frond elongation rates of 7.5, 6.1, 7.5 and 7.9 cm yr^-1, respectively, were recorded during four consecutive years. In winter elongation rates of fronds were rather low. Elongation did not seem to be affected by frond length, the self-shading effect of the canopy, or changes in depth. Cuts resulting from grazing were frequent on fronds through the year, decreasing significantly with increasing depth. Sheared fronds regenerated promptly, independent of season or height at which the cut was made. For commercial purposes the best season for harvesting is summer, which is the period of maximum standing stock, infertility, and calm warm waters that provide easier working conditions for divers.