Responses of vegetative gametophytes of Undaria pinnatifida to high irradiance in the process of gametogenesis

The population of Undaria pinnatifida in its ecologic niche sustains itself in high temperature summer in the form of vegetative gametophytes, the haploid stage in its heteromorphic life cycle. Gametogenesis initiates when seawater temperature drops below the threshold levels in autumn in the northern hemisphere. Given that the temperature may fall into the appropriate range for gametogenesis, the level of irradiance determines the final destiny of a gametophytic cell, either undergoing vegetative cell division or initiating gametogenesis. In elucidating how vegetatively propagated gametophytes cope with changes of irradiance in gametogenesis, we carried out a series of culture experiments and found that a direct exposure to irradiance as high as 270 μmol photons m^?2 s^?1 was lethal to dim‐light (7–10 μmol photons m^?2 s^?1) adapted male and female gametophytes. This lethal effect was linearly corelated with the exposure time. However, dim‐light adapted vegetative gametophytes were shown to be able tolerate as high as 420 μmol photons m^?2 s^?1 if the irradiance was steadily increased from dim light levels (7–10 μmol photons m^?2 s^?1) to 90, 180 and finally 420 μmol photons m^?2 s^?1, respectively, at a minimum of 1–3 h intervals. Percentage of female gametophytic cells that turned into oogonia and were eventually fertilized was significantly higher if cultured at higher but not lethal irradiances. Findings of this investigation help to understand the dynamic changes of population size of sporophytic plants under different light climates at different site‐specific ecologic niches. It may help to establish specific technical details of manipulation of light during mass production of seedlings by use of vegetatively propagated gametophytes.     

GROWTH,PHOTOSYNTHESIS AND MAINTENANCE METABOLIC COST IN THE DIATOM PHAEODACTYLUM TRICORNUTUM AT VERY LOW LIGHT LEVELS 1

The compensation point for growth of Phaeodactylum tricornutum Bohlin is less than 1 μmol. m^?2s^?1. Growth at low PFDs (<3.5 μmol. m^?2.s^?1) does not appear to reduce the maximum quantum efficiency of photosynthesis (ø_m) or to greatly inhibit the potential for light-saturated, carbon-specific photosynthesis (P_m^c). The value for ø_m in P. tricornutum is 0.10–0.12 mol O₂-mol photon^?1, independent of acclimation PFD between 0.75 and 200 μmol.m^?2.s^?1 in nutrient-sufficient cultures. P_m^c in cells of P. tricornutum acclimated to PFDs <3.5 μmol m^?2?s^?1 is approximately 50% of the highest value obtained in nutrient-sufficient cultures acclimated to growth-rate-saturating PFDs. In addition, growth at low PFDs does not severely restrict the ability of cells to respond to an increase in light level. Cultures acclimated to growth at lees than 1% of the light-saturated growth rate respond rapidly to a shift-up in PFD after a short initial lag period and achieve exponential growth rates of 1.0 d^?1 (65% of the light- and nutrient-saturated maximum growth rate) at both 40 and 200 μmol.m^?2.s^?1     

Plasticity in relative growth rate and its components following a change in irradiance

A total of 244 plants from two species, Lythrum salicaria and Epilobium glandulosum, were grown individually in hydroponic sand culture from seed for 36 d. Until day 27 all plants experienced an irradiance of 550 μmol m^?2 s^?1 PFD and on day 27 half of the plants were subjected to a neutral shade treatment in which irradiance was reduced to 100 μmol m^?2 s^?1 photon fluy density (PFD). Measures of relative growth rate, net assimilation rate, specific leaf area, biomass partitioning to leaves, roots, structural tissues (i.e. stems, petioles and inflorescences) and tissue density were obtained from intensive harvests three or four times per day. The shade treatment caused an immediate decrease in relative growth rate and net assimilation rate. Within hours the specific leaf area of the shaded plants increased and leaf tissue density decreased, thus partially offsetting the decrease in relative growth rate. Biomass partitioning was not affected.     

LIGHT DEPENDENCE OF GROWTH AND PHOTOSYNTHESIS IN PHAEODACTYLUM TRICORNUTUM (BACILLARIOPHYCEAE)1

Phaeodactylum tricornutum Bohlin was maintained in exponential growth over a range of photon flux densities (PFD) from 7 to 230 μmol·m^?2s^?1. The chlorophyll a-specific light absorption coefficient, maximum quantum yield of photosynthesis, and C:N atom ratio were all independent of the PFD to which cells were acclimated. Carbon- and cell-specific, light-satuated, gross photosynthesis rates and dark respiration rates were largely independent of acclimation PFD. Decreases in the chlorophyll a-specific, gross photosynthesis rate and the carbon: chlorophyll ratio and increases of cell- or carbon-specific absorption coefficients were associated with an increase in cell chlorophyll a in cultures acclimated to low PFDs. The compensation PFD for growth was calculated to be 0.5 μmol·m^?2s^?1. The maintenance metabolic rate (2 × 10^?7s^?1), calculated on the basis of the compensation PFD, is an order of magnitude lower than the measured dark respiration rate(2.7 × 10^?6mol O₂·mol C^?1s^?1). Maintenance of high carbon-specific, light-saturated photosynthesis rates in cells acclimated to low PFDs may allow effective use of short exposures to high PFDs in a temporally variable light environment.     

THE INTERACTION BETWEEN INORGANIC CARBON ACQUISITION AND LIGHT SUPPLY IN PALMARIA PALMATA (RHODOPHYTA)1

The dependence of the carbon concentrating mechanism of Palmaria palmata (L.) Kuntze on the growth light level was examined 1) to determine whether or not there is a threshold photon flux density (PFD) at which the inorganic carbon uptake mechanism can operate and 2) to attempt to quantify the relative energetic costs of acclimation to the two different limiting factors, PFD and dissolved inorganic carbon (DIC) concentration. Plants were grown at six PFDs: 5, 25, 50, 75, 95, and 125 μmol photons. m^?2.s^?1. Growth rates increased with increasing PFD from 5 to 50 μmol photons. m^?2. s^?1 and were light-saturated at 75, 95, and 125 μmol photons. m^?2. s^?1 Values of δ¹³C increased continuously with increasing growth PFD and did not saturate over the range of light levels tested. Time-resolved fluorescence characteristics indicated a progressive photoacclimation below 50 μmol photons. m^?2. s^?1. Analysis of chlorophyll fluorescence induction showed three levels of light use efficirncy associated with growth at 5 or 25, 50, and >75 μmol photons. m^?2. s^?1. The light-haruesting efficiency was inversely proportional to the effectiveness of DIC acquisition in plants grown at the six PFDs. These data were interpreted to indicate that there is a physiological tradeoff between photosynthetic efficiency and bicarbonate use in this species.     

PHOTOSYNTHESIS AND CELL DIVISION BY ANTARCTIC MICROALGAE: COMPARISON OF BENTHIC,PLANKTONIC AND ICE ALGAE1

Irradiance-dependent rates of photosynthesis and cell division of six species of microalgae isolated from the benthos, plankton and sea ice microbial community in McMurdo Sound, Antarctica were compared. Microalgae isolated from different photic environments had distinct photosynthetic and growth characteristics. For benthic and ice algae, photosynthesis saturated at 6 to 20 μE.m^?2.s^?1 and was photoinhibited at 10 to 80 μE.m^?2.s^?1 while for the planktonic algae, saturation irradiances were up to 13 times higher and photoinhibition was not detected. The slope of the light-limited portion of the P-I relationship was up to 50 times greater for the benthic algae than for either the ice or planktonic algae suggesting that benthic algae used the low irradiances more efficiently for carbon uptake. Cell division was dependent on the incubation irradiance for all but one microalga examined. The dependence of division rates on irradiance was however much smaller than for carbon uptake, suggesting that cell division buffers the influence of short term variations of irradiance on cellular metabolism.     

EFFECT OF TEMPERATURE AND IRRADIANCE ON GROWTH OF HAEMATOCOCCUS PLUVIALIS (CHLOROPHYCEAE)1

The growth characteristics of Haematococcus pluvialis Flotow were determined in batch culture. Optimal temperature for growth of the alga was between 25° and 28°C, at which the specific growth rate was 0.054 h^?1. At higher temperatures, no cell division was observed, and cell diameter increased from 5 to 25 μm. The saturated irradiance for growth of the alga was 90 μmol quanta · m^?2·s^?1; under higher irradiances (e.g. 400 μmol quanta·m^?2·s^?1) astaxanthin accumulation was induced. Growth rate, cell cycle, and astaxanthin accumulation were significantly affected by growth conditions. Careful attention should be given to the use of optimal growth conditions when studying these processes.     

SHORT‐TERM EFFECT OF TEMPERATURE ON THE PHOTOKINETICS OF MICROALGAE FROM THE SURFACE LAYERS OF ANTARCTIC PACK ICE1

Microalgae growing within brine channels (85 psu salinity) of the surface ice layers of Antarctic pack ice showed considerable photosynthetic tolerance to the extreme environmental condition. Brine microalgae exposed to temperatures above ?5°C and at irradiances up to 350 μmol photons·m^?2·s^?1 showed no photosynthetic damage or limitations. Photosynthesis was limited (but not photoinhibited) when brine microalgae were exposed to ?10°C, provided the irradiance remained under 50 μmol photons·m^?2·s^?1. The highest level of photosynthetic activity (maximum relative electron transport rate [rETR_max]) in brine microalgae growing within the surface layer of sea ice was at approximately 18 μmol electrons·m^?2·s^?1, which occurred at ?1.8°C. Effective quantum yield of PSII and rETR_max of the halotolerant brine microalgae exhibited a temperature‐dependent pattern, where both parameters were higher at ?1.8°C and lower at ?10°C. Relative ETR_max at temperatures above ?5°C were stable across a wide range of irradiance.     

Changes in photon flux can induce stomatal patchiness

Images of chlorophyll fluorescence were used to detect the occurrence of stomatal patchiness in leaves from eight species under variable photon flux conditions. Pronounced stomatal patchiness was induced within 5–10 min after PFD was changed from intermediate (～450 μmol quanta m^?2 s^?1) to low (～150 μmol quanta m^?2 s^?1) levels. This effect was completely reversible by returning PFD to intermediate levels. The pattern of heterogeneous fluorescence for each leaf was usually similar during repeated applications of medium and low PFD. In three species, stomatal patchiness could only be induced in slightly water-stressed plants. Leaves of more severely water-stressed Xanthium strumarium plants in low air humidity exhibited oscillations in fluorescence that corresponded with oscillatory changes in leaf diffusion conductance for water vapour. Stomatal patchiness was also induced by illuminating dark-adapted leaves with low PFD (below 200–300 μmol quanta m^?2 s^?1). Infiltration of leaves with distilled water showed that heterogeneous chlorophyll fluorescence was caused by changes in stomatal apertures.     

High rates of net ecosystem carbon assimilation by Brachiara pasture in the Brazilian Cerrado

To investigate the consequences of land use on carbon and energy exchanges between the ecosystem and atmosphere, we measured CO₂ and water vapour fluxes over an introduced Brachiara brizantha pasture located in the Cerrado region of Central Brazil. Measurements using eddy covariance technique were carried out in field campaigns during the wet and dry seasons. Midday CO₂ net ecosystem exchange rates during the wet season were ?40 μmol m^?2 s^?1, which is more than twice the rate found in the dry season (?15 μmol m^?2 s^?1). This was observed despite similar magnitudes of irradiance, air and soil temperatures. During the wet season, inferred rates of canopy photosynthesis did not show any tendency to saturate at high solar radiation levels, with rates of around 50 μmol m^?2 s^?1 being observed at the maximum incoming photon flux densities of 2200 μmol m^?2 s^?1. This contrasted strongly to the dry period when light saturation occurred with 1500 μmol m^?2 s^?1 and with maximum canopy photosynthetic rates of only 20 μmol m^?2 s^?1. Both canopy photosynthetic rates and night‐time ecosystem CO₂ efflux rates were much greater than has been observed for cerrado native vegetation in both the wet and dry seasons. Indeed, observed CO₂ exchange rates were also much greater than has previously been reported for C₄ pastures in the tropics. The high rates in the wet season may have been attributable, at least in part, to the pasture not being grazed. Higher than expected net rates of carbon acquisition during the dry season may also have been attributable to some early rain events. Nevertheless, the present study demonstrates that well‐managed, productive tropical pastures can attain ecosystem gas exchange rates equivalent to fertilized C₄ crops growing in the temperate zone.     

Effects of light intensity and quality on the growth rate and photosynthetic pigment content of Spirulina platensis

The quantitative and qualitative effects of light on carotenoid production by Spirulina were studied. Maximum total carotenoid production was measured in cells grown under white light at an irradiance of 432 μmol photon m^?2 s^?1, the onset of light saturation for this organism as determined by growth rates. A true maximum may exist at irradiances above 1500 μmol photon m^?2 s^?1 under white light. Individual carotenoids responded differently to light conditions. Under white light, β-carotene and echinenone were most abundant at the lowest and highest irradiance levels tested. Myxoxanthophyll and lutein/zeaxanthin did not change over the same irradiance range. Under red and blue light, we found decreased values of myxoxanthophyll, while β-carotene increased and lutein/zeaxanthin and echinenone showed little change. In general, maximum carotenoid production requires optimization of the culture conditions that favor growth.     

Effects of irradiance, water temperature and nutrients on the growth of sporelings of the crustose coralline alga Lithophyllum yessoense Foslie (Corallinales, Rhodophyceae)

Lithophyllum yessoense Foslie is a markedly dominant subtidal, crustose coralline alga in south–western Hokkaido, Japan. In this study, the effects of irradiance, water temperature and nutrients (nitrate and phosphate) on the growth of sporelings of the alga were examined. The relative growth rate (RGR) was saturated at 17.6% d^?1 at a high irradiance (240 umol photon m²s^?1). Even at a low irradiance (10.7–49.9 umol photon m^?2s^?1), RGR was 7.1–12.7% d^?1 The survival rate of sporelings was greater than 80% at irradiance above 10.7 μmol photon m^?2s^?1 throughout the culture period. The growth of L. yessoense sporelings was promoted at 15°C and 20°C, but inhibited at 5°C. The half‐saturation constants (Ks) for growth were about 0.5 umol L^?1 and 0.14 umol L^?1 for nitrate and phosphate, respectively. Saturated nitrate and phosphate concentrations for the growth were about 4.0 μmol L^?1 and 0.4 μmol L^?1, respectively, suggesting that L. yessoense is adaptable to a relatively high water temperature, a wide range of irradiance, and low ambient nitrate and phosphate concentrations. The results provide a possible explanation of why L. yessoense is dominant in the environments of south‐western Hokkaido.     

SIMAZINE-INDUCED INHIBITION IN PHOTOACCLIMATED POPULATIONS OF ANABAENA CIRCINALIS (CYANOPHYTA)1

The effects of the triazine herbicide, simazine, on photosynthetic oxygen evolution and growth rate in photoacclimated populations of Anabaena circinalis Rabenhorst were investigated. Chemostat populations were acclimated to photon flux densities (PFDs) of 50, 130, and 230 μmol·m^?2·s^?1 of photosynthetic active radiation (PAR), Decreases in chlorophyll a (Chl a). c-phycocyanin (CPC), and total carotenoid (TCar) contents and CPC: Chl a and CPC: TCar ratios of populations coincided with increasing PFD, Polynomial regression models that characterize inhibition of photosynthesis for populations acclimated to 50 and 130 μmol photons·m^?2·s^?1 PAR were distinct from the model for populations acclimated to 230 μmol photons·m^?2·s^?1 PAR. Simazine concentrations that, depressed oxygen evolution 50% compared to controls decreased with increasing PFD. Increases and decreases in both biomass and growth rate coincided with increasing PFD and simazine concentration, respectively. Simazine concentrations that depressed growth rate 50% compared to controls increased with decreasing PFD. The differences in photosynthetic and growth inhibition among photoacclimated populations indicate that sensitivity to photosystem II inhibitors is affected by alterations in pigment contents.     

Relative enhancement of photosynthesis and growth at elevated CO2 is greater under sunflecks than uniform irradiance in a tropical rain forest tree seedling

The survivorship of dipterocarp seedlings in the deeply shaded understorey of South‐east Asian rain forests is limited by their ability to maintain a positive carbon balance. Photosynthesis during sunflecks is an important component of carbon gain. To investigate the effect of elevated CO₂ upon photosynthesis and growth under sunflecks, seedlings of Shorealeprosula were grown in controlled environment conditions at ambient or elevated CO₂. Equal total daily photon flux density (PFD) (～7·7 mol m^?2 d^?1) was supplied as either uniform irradiance (～170 µmol m^?2 s^?1) or shade/fleck sequences (～30 µmol m^?2 s^?1/～525 µmol m^?2 s^?1). Photosynthesis and growth were enhanced by elevated CO₂ treatments but lower under flecked irradiance treatments. Acclimation of photosynthetic capacity occurred in response to elevated CO₂ but not flecked irradiance. Importantly, the relative enhancement effects of elevated CO₂ were greater under sunflecks (growth 60%, carbon gain 89%) compared with uniform irradiance (growth 25%, carbon gain 59%). This was driven by two factors: (1) greater efficiency of dynamic photosynthesis (photosynthetic induction gain and loss, post‐irradiance gas exchange); and (2) photosynthetic enhancement being greatest at very low PFD. This allowed improved carbon gain during both clusters of lightflecks (73%) and intervening periods of deep shade (99%). The relatively greater enhancement of growth and photosynthesis at elevated CO₂ under sunflecks has important potential consequences for seedling regeneration processes and hence forest structure and composition.     

COMPARATIVE EFFECTS OF LIGHT ON PIGMENTS OF TWO STRAINS OF EMILIANIA HUXLEYI (HAPTOPHYTA)1

Calcifying and a noncalcifying strains of Emiliania huxleyi were cultured in nutrient replete turbidostats under a photon flux density (PFD) gradient from 50 to 600 μmol E·m^?2·s^?1. For both strains, growth was PFD‐saturated at 300 μmol E·m^?2·s^?1. The strains, although with clearly different physiological properties due to the presence or absence of calcification, showed the same trends and magnitude of change in their pigment compliment as a function of PFD. Light‐controlled pigment composition and the trends of change in pigment composition were identical in both strains. Fucoxanthin (Fuco) was the major carotenoid in the calcifying strain, while in the noncalcifying strain this role was assumed by 19′ hexanoyloxyfucoxanthin (19 Hex). The photoprotective pigments and 19 Hex, normalized to chl a, increased with increasing light, while chl a content per cell and chl c's and Fuco, normalized to chl a, decreased with increasing PFD. The sum of all carotenoids normalized to chl a was remarkably similar in all PFDs used. Collectively, our results suggest that 19 Hex was synthesized from Fuco with light as a modulating factor and that the total amount of carotenoids is strain‐specific and synthesized/catabolized in tandem with chl a to a genetically predefined level independent of PFD.     

NONEQUILIBRIUM RATES OF PHOTOSYNTHESIS AND RESPIRATION UNDER DYNAMIC LIGHT SUPPLY1

To identify processes that might account for differences in growth rates of rhodophytes under constant and dynamic light supply, we examined nonequilibrium gas exchange by measuring time courses of photoinduction, loss of photoinduction, and respiration rates immediately after the light–dark transition. Using the rhodophyte species Palmaria palmata (Huds.) Lamour and Lomentaria articulata (Huds.) Lyngb., we compared the effects of growth-saturating constant photon flux density (PFD) (95 μmol photons · m^?2· s^?1) to those of a dynamic light supply modeled on canopy movements in the intertidal zone (25 μmol photons · m^?2· s^?1 background PFD plus light flecks of 350 μmol photons · m^?2· s^?1, 0.1 Hz). The time required for P. palmata and L. articulata to be fully photoinduced was not affected by the dynamics of light supply. L. articulata required only 6 min of illumination with either fluctuating or constant light to be completely induced compared to 20 min for P. palmata. The latter species also lost photoinduction more rapidly than did L. articulata in the dark. There was no significant decline in photoinduction state for either species at the background PFD. The time courses of respiration after illumination with constant and fluctuating light were significantly different for P. palmata but not for L. articulata when the total photon dose was equal. In general, gas exchange of P. palmata appeared to be particularly sensitive to the temporal distribution of light supply whereas that of L. articulata was sensitive to the amplitude of variations, being photoinhibited at high PFD. These results are discussed in terms of the different mechanisms of inorganic carbon acquisition in the two species.     

Chlorophyll fluorescence quenching and violaxanthin deepoxidation of FBPase antisense plants at low light intensities and low temperatures

Genetically modified potato (Solanum tuberosum L. cv. Desiree) and tobacco (Nicotiana tabacum cv. Samsun N.N.) plants were used to analyze the effects exerted by the chloroplastic (cp) fructose- 1,6-bisphosphatase (FBPase) on the regulation of light energy discrimination at the level of photosystem II. The cp-FBPase activity was progressively inhibited by an mRNA antisense to this FBPase. The chlorophyll fluorescence quenching parameters of these transgenic plants were compared to those of wild-type and transgenic plants that were acclimated to low temperatures. In particular various lines of the transgenic potato and tobacco plants were exposed to a temperature treatment of 10 and 20°C for 10 days. Light intensities were kept low to reduce photoinhibition so that we could analyze exclusively the effects of a modification in the carbon fixation cycle on the chlorophyll fluorescence quenching parameters. The photon flux densities (PFDs) employed at the level of the middle leaves of all plants were set to two different values of 10 μmol m^?2 s^?1 and 50 μmol m^?2 s^?1. Subsequent to this 10-day acclimation the chlorophyll-fluorescence parameters of all plants were measured. Photoinhibition as expressed by the F_y/F_m ratio was minor in plants subjected to a PFD of 10 μmol m^?2 s^?1. Higher photon fluence rates of 50 μmol m^?2 s^?1 at temperatures of 10°C gave rise to a significant reduction in the F_y/F_m ratios obtained from the transgenic plants which were characterized by a restriction in cp-FBPase capacity to 20% of normal activity. Furthermore, a progressive inhibition of the cp-FBPase activity induced an amplified nonphotochemical quenching of chlorophyll fluorescence with in the genetically manipulated species (except at 10°C and 50 μmol m^?2 s^?1). The increase in nonphotochemical quenching depended upon light and temperature. Photochemical quenching of light quanta within the antisense plants declined relative to that in the wild type. To further characterize the mechanisms producing higher levels of nonphotochemical fluorescence quenching. we analyzed several of the xanthophyll cycle pigments. The deepoxidation state of the xanthophyll cycle pigments in potato plants increased with attenuating FBPase activities under all conditions. For tobacco plants, this elevation of the deepoxidation state was only observed at a PFD of 50 μmol m^?2 s^?1.     

Effect of irradiance and temperature on the photosynthesis of a cultivated red alga,Pyropia tenera (= Porphyra tenera), at the southern limit of distribution in Japan

The effect of irradiance and temperature on the photosynthesis of the red alga, Pyropia tenera, was determined for maricultured gametophytes and sporophytes collected from a region that is known as one of the southern limits of its distribution in Japan. Macroscopic gametophytes were examined using both pulse‐amplitude modulated fluorometry and/or dissolved oxygen sensors. A model of the net photosynthesis–irradiance (P‐E) relationship of the gametophytes at 12°C revealed that the net photosynthetic rate quickly increased at irradiances below the estimated saturation irradiance of 46 μmol photons m^?2 s^?1, and the compensation irradiance was 9 μmol photons m^?2 s^?1. Gross photosynthesis and dark respiration for the gametophytes were also determined over a range of temperatures (8–34°C), revealing that the gross photosynthetic rates of 46.3 μmol O₂ mg_chl‐a^?1 min^?1 was highest at 9.3 (95% Bayesian credible interval (BCI): 2.3–14.5)°C, and the dark respiration rate increased at a rate of 0.93 μmol O₂ mg_chl‐a^?1 min^?1°C^?1. The measured dark respiration rates ranged from ?0.06 μmol O₂ mg_chl‐a^?1 min^?1 at 6°C to ?25.2 μmol O₂ mg_chl‐a^?1 min^?1 at 34°C. The highest value of the maximum quantum yield (Fv/Fm) for the gametophytes occurred at 22.4 (BCI: 21.5–23.3) °C and was 0.48 (BCI: 0.475–0.486), although those of the sporophyte occurred at 12.9 (BCI: 7.4–15.1) °C and was 0.52 (BCI: 0.506–0.544). This species may be considered well‐adapted to the current range of seawater temperatures in this region. However, since the gametophytes have such a low temperature requirement, they are most likely close to their tolerable temperatures in the natural environment.     

GROWTH AND PHOTOSYNTHESIS OF ULVA ROTUNDATA (CHLOROPHYTA) AS A FUNCTION OF TEMPERATURE AND SQUARE WAVE IRRADIANCE IN INDOOR CULTURE1

Temperature and photon flux density (PFD) vary independently in estuaries, e.g. high PFD may occur at any temperature, so it is necessary to consider synergistic effects of these factors on algal growth. Because natural PFD is highly variable and daylength changes confound seasonal temperature cycles, it is easier to interpret factorial experiments in controlled laboratory conditions. Clonal Ulva rotundata Blid. (Chlorophyta) has been studied extensively in outdoor culture. In this study it was maintained indoors under square wave photoperiods at five PFDs and three temperatures. Growth rate, photqsynthetic light response (P-I) curves, and photosystem II chlorophyll fluorescence properties were measured at the growth temperature following acclimation. Interactions between PFD and growth temperature were strongly indicated in all physiological parameters measured. Greatest PFD response occurred at the highest temperature, and the largest temperature response occurred at the highest PFD. Light-saturated photosynthesis (P_m) dark respiration (R_d), and light-limited quantum yield (Φ_m) were sufficient to describe acclimation status. The light-saturation parameter (I_k) was redundant and potentially misleading. Although U. rotundata exhibits a great amplitude of photoacclimation, it apparently has little capacity for temperature acclimation compared to the kelp, Laminaria saccharina, for which published data indicate similar photosynthetic rates over a broad range of growth temperatures. Diurnal variation of P_m and R_d at a growth PFD of ～ 1700 ± 200 μmol photons · m^?2· s^?1 was similar to the pattern observed previously in outdoor culture, suggesting endogenous control of these parameters. Quantum yield and the ratio of variable to maximum chlorophyll fluorescence (F_v/F_m), which were depressed in midday sunlight exceeding ～ 1500 μmol photons · m^?2· s^?1, were relatively invariant through the day in indoor culture, indicating that these parameters are controlled primarily by instantaneous PFD. Growth and fluorescence data are also presented for some other macroalgae for comparative purposes.     

Photosynthesis and growth of Ulva ohnoi and Ulva pertusa (Ulvophyceae) under high light and high temperature conditions,and implications for green tide in Japan

The macroalga Ulva ohnoi constitutes a considerable fraction of green tides in coastal areas of Japan, but little is known about the physiological characteristics of this species. To investigate the environmental factors that promote the formation of green tides, we tested the responses of U. ohnoi and another common Japanese species, Ulva pertusa, to various levels of irradiance at different water temperatures. Because the two species are morphologically similar, we identified them using the PCR‐restriction fragment length polymorphism method. Under laboratory conditions, we evaluated the photosynthetic, dark respiration, and relative growth rate at a range of water temperatures (5 to 35°C) and photosynthetically active radiation (0 to 1000 μmol photons m^?2 s^?1). The maximum gross photosynthetic rate of U. ohnoi was larger than that of U. pertusa. The dark respiration rates revealed no significant differences among the species and temperature conditions. At 500 μmol photons m^?2 s^?1, the relative growth rate of U. ohnoi was larger than that of U. pertusa in higher temperature and the difference was the largest at 20°C. The estimated compensation irradiance and estimated saturation irradiance of U. ohnoi and U. pertusa ranged from 0.709 to 5.510 and 40.530 to 58.674 μmol photons m^?2 s^?1, which were lower than those in other intertidal green macroalgae, from 6 to 11 and 50 to 82 μmol photons m^?2 s^?1, respectively. Thus, U. ohnoi which exists as free‐floating near the water surface and accumulating inside the green tide can survive extensively in the water column of the intertidal zone, furthermore, the species can maintain rapid growth in this situation. Therefore, as a result of this study, it is suggested that the ecological success of U. ohnoi in shallow waters such as the tidal flats, estuarine, and coasts of the inner bay in comparison with U. pertusa.