In situ seagrass photosynthesis measured using a submersible, pulse-amplitude modulated fluorometer

Assessments of photosynthetic activity in marine plants can now be made in situ using a newly developed, submersible, pulse-amplitude modulated (PAM) fluorometer: Diving-PAM. PAM fluorometry provides a measure of chlorophyll a fluorescence using rapid-light curves in which the electron-transport rate can be determined for plants exposed to ambient light conditions. This technique was used to compare the photosynthetic responses of seagrasses near Rottnest Island, Western Australia. Several fluorescence parameters were measured as a function of time of day and water depth; electron-transport rate (ETR), quantum yield, photochemical quenching and non-photochemical quenching and Photosystem II (PSII) photochemical efficiency (F _v :F _m ratio) were measured. Results indicate that recent light-history plays a crucial role in seagrass photosynthetic responses. Maximum ETR of Posidonia australis, Amphibolis antarctica and Halophila ovalis is influenced by the irradiance during the diurnal cycle, with low rates at dawn and dusk (<10 μmol electron m⁻² s⁻¹), highest rates in late morning (40 to 60 μmol electron m⁻² s⁻¹) and a mid-day depression. Maximum ETR and PSII photochemical efficiency varied widely between seagrass species and were not correlated. A comparison of photochemical to non-photochemical quenching indicated that seagrasses in shallow water receiving high light have a high capacity for non-photochemical quenching (e.g. light protection) compared to seagrasses in deep water. These results indicate that in situ measurements of photosynthesis will provide new insights into the mechanisms and adaptive responses of marine plants. Received: 26 May 1997 / Accepted: 27 May 1998     

In situ measurements of photosynthetic irradiance responses of two Red Sea sponges growing under dim light conditions

Photosynthetic responses to irradiance by the photosymbionts of the two Red Sea sponges Theonella swinhoei (Gray) and Clionavastifica (Hancock) growing under dim light conditions were measured in situ (in September 1997) using a newly developed underwater pulse amplitude modulated (PAM) fluorometer. Relative rates of photosynthetic electron transport (ETR) were calculated as the effective quantum yield of photosystem II (Y ) multiplied with the photosynthetic photon flux (PPF). Photosynthesis versus irradiance (P-I ) curves, obtained within minutes, showed that individual specimens of both sponges, growing under very low light conditions, feature lower light saturation points as well as lower maximal ETRs than individuals growing under higher light. Evaluations of such curves using low irradiances of the actinic light source (20 to 130 μmol photons m⁻² s⁻¹) showed a general decrease in Y, with a shoulder from the lowest irradiance applied till 20 to 30 μmol photons m⁻² s⁻¹. Point measurements yielded ETRs close to what could be estimated from the P-I curves. These point measurements also revealed good correlations between the diurnally changing ambient irradiances (1 to 50 μmol photons m⁻² s⁻¹) and average ETR values for both species. Further analysis showed that although Y values varied considerably between the different point measurements, they did not decrease significantly with light under these very low irradiances. Therefore, PPF rather than Y seems to determine the in situ diel photosynthetic performance at the low ambient irradiances experienced by these sponges. Received: 22 November 1997 / Accepted: 8 April 1998     

Photosynthetic activity of the non-dormant <Emphasis Type="Italic">Posidonia oceanica</Emphasis> seed

The photosynthetic adaptive features of non-dormant seeds in Posidonia oceanica were studied in order to evaluate the effects of light on germination success. Transmission electron micrographs showed the presence of chloroplasts in the epidermal cells, close to the nucleus at the periphery of the cytoplasm. The well-developed thylakoid membranes and the presence of starch granules indicated that the chloroplasts were photosynthetically active. The relationship between photosynthesis versus irradiance in P. oceanica seeds incubated at 15 and 21°C was analysed. The net photosynthesis in the non-dormant seed of P. oceanica was positive and compensated its respiration demand (90 μmol quanta m⁻² s⁻¹) at both temperatures. Net photosynthesis was negative at the other irradiance values. To test the effects of light on germination success, seeds were placed both in dark and light conditions. Germination success was significantly higher in light rather than in dark condition. The characteristics observed in the photosynthesis in P. oceanica seed could be a mechanism to guarantee seedling survival in temperate waters, demonstrating though the specialized nature of this species.     

Effects of light exposure on the retention of kleptoplastic photosynthetic activity in the sacoglossan mollusc <Emphasis Type="Italic">Elysia viridis</Emphasis>

The effects of light exposure on the photosynthetic activity of kleptoplasts were studied in the sacoglossan mollusc Elysia viridis. The photosynthetic activity of ingested chloroplasts was assessed in vivo by non-destructively measuring photophysiological parameters using pulse amplitude modulation (PAM) fluorometry. Animals kept under starvation were exposed to two contrasting light conditions, 30 μmol photons m⁻² s⁻¹ (low light, LL), and 140 μmol photons m⁻² s⁻¹ (high light, HL), and changes in photosynthetic activity were monitored by measuring the maximum quantum yield of photosystem II (PSII), F _v/F _m, the minimum fluorescence, F _o, related to chlorophyll a content, and by measuring rapid light-response curves (RLC) of relative electron transport rate (rETR). RLCs were characterised by the initial slope of the curve, α_RLC, related to efficiency of light capture, and the maximum rETR level, rETR_m,RLC, determined by the carbon-fixation metabolism. Starvation induced the decrease of all photophysiological parameters. However, the retention of photosynthetic activity (number of days for F _v/F _m > 0), as well as the rate and the patterns of its decrease over time, varied markedly with light exposure. Under HL conditions, a rapid, exponential decrease was observed for F _v/F _m, α_RLC and rETR_m,RLC, F _o not showing any consistent trend of variation, and retention times ranged between 6 and 15 days. These results suggested that the retention of chloroplast functionality is limited by photoinactivation of PSII reaction center protein D1. In contrast, under LL conditions, a slower decrease in all parameters was found, with retention times varying from 15 to 57 days. F _v/F _m, α_RLC and rETR_m,RLC exhibited a bi-phasic pattern composed by a long phase of slow decrease in values followed by a rapid decline, whilst F _o decayed exponentially. These results were interpreted as resulting from lower rates of D1 photoinactivation under low light and from the gradual decrease in carbon provided by photosynthesis due to reduction of functional photosynthetic units.     

Effect of acclimatization to low temperature and reduced light on the response of reef corals to elevated temperature

This study tested the effects of acclimatization on the response of corals to elevated temperature, using juvenile massive Porites spp. and branching P. irregularis from Moorea (W149°50′, S17°30′). During April and May 2006, corals were acclimatized for 15 days to cool (25.7°C) or ambient (27.7°C) temperature, under shaded (352 μmol photons m⁻² s⁻¹) or ambient (554 μmol photons m⁻² s⁻¹) natural light, and then incubated for 7 days at ambient or high temperature (31.1°C), under ambient light (659 μmol photons m⁻² s⁻¹). The response to acclimatization was assessed as biomass, maximum dark-adapted quantum yield of PSII (F _v/F _m), and growth, and the effect of the subsequent treatment was assessed as F _v/F _m and growth. Relative to the controls (i.e., ambient temperature/ambient light), massive Porites spp. responded to acclimatization through increases in biomass under ambient temperature/shade, and low temperature/ambient light, whereas P. irregularis responded through reduced growth under ambient temperature/shade, and low temperature/ambient light. Acclimatization affected the response to thermal stress for massive Porites spp. (but not P. irregularis), with an interaction between the acclimatization and subsequent treatments for growth. This interaction resulted from a lessening of the negative effects of high temperature after acclimatizing to ambient temperature/shade, but an accentuation of the effect after acclimatizing to low temperature/shade. It is possible that changes in biomass for massive Porites spp. are important in modulating the response to high temperature, with the taxonomic variation in this effect potentially resulting from differences in morphology. These results demonstrate that corals can acclimatize during short exposures to downward excursions in temperature and light, which subsequently affects their response to thermal stress. Moreover, even con-generic taxa differ in this capacity, which could affect coral community structure. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Microsensor studies of photosynthesis and respiration in the symbiotic foraminifer Orbulina universa

Oxygen and pH microelectrodes were used to investigate the microenvironment of the planktonic foraminifer Orbulina universa and its dinoflagellate endosymbionts. A diffusive boundary layer surrounds the foraminiferal shell and limits the O₂ and proton transport from the shell to the ambient seawater and vice versa. Due to symbiont photosynthesis, high O₂ concentrations of up to 206% air saturation and a pH of up to 8.8, i.e. 0.5 pH units above ambient seawater, were measured at the shell surface of the foraminifer at saturating irradiances. The respiration of the host–symbiont system in darkness decreased the O₂ concentration at the shell surface to <70% of the oxygen content in the surrounding air-saturated water. The pH at the shell surface dropped to 7.9 in darkness. We measured a mean gross photosynthetic rate of 8.5 ± 4.0 nmol O₂ h⁻¹ foraminifer⁻¹. The net photosynthesis averaged 5.3 ± 2.7 nmol O₂ h⁻¹. In the light, the calculated respiration rates reached 3.9 ± 1.9 nmol O₂h⁻¹, whereas the dark respiration rates were significantly lower (1.7 ± 0.7 nmol O₂ h⁻¹). Experimental light–dark cycles demonstrated a very dynamic response of the symbionts to changing light conditions. Gross photosynthesis versus scalar irradiance curves (P vs E _o curves) showed light saturation irradiances (E _k) of 75 and 137 μmol photons m⁻² s⁻¹ in two O. universa specimens, respectively. No inhibition of photosynthesis was observed at irradiance levels up to 700 μmol photons m⁻² s⁻¹. The light compensation point of the symbiotic association was 50 μmol photons m⁻² s⁻¹. Radial profile measurements of scalar irradiance (E _o) inside the foraminifera showed a slight increase at the shell surface up to 105% of the incident irradiance (E _d). Received: 26 January 1998 / Accepted: 11 April 1998     

Growth and grazing responses of Chrysochromulina ericina (Prymnesiophyceae): the role of irradiance, prey concentration and pH

The effect of irradiance, prey concentration and pH on the growth and grazing responses of the mixotrophic prymnesiophyte Chrysochromulina ericina under N-and P-replete conditions was studied using the pedinophyte Marsupiomonas pelliculata as prey. The two organisms were inoculated in monocultures and in mixed cultures at different predator: prey ratios at three irradiances and allowed to grow for 4–7 days. All cultures were non-axenic. Algal densities and pH were monitored throughout the experiments and growth and grazing rates were measured. An increase in growth of C. ericina cultures at irradiances of 25 and 70 μmol photons m⁻² s⁻¹ was observed after the addition of prey, while growth of C. ericina cultures at the high irradiance (150 μmol photons m⁻² s⁻¹) was unaffected by the addition of prey. However, although the growth of C. ericina increased at low irradiance (25 μmol photons m⁻² s⁻¹), it did not reach the same level as monocultures at the high irradiance (150 μmol photons m⁻² s⁻¹), suggesting that phagotrophy can only partly replace photosynthesis in C. ericina. Maximum growth rates of C. ericina at irradiances of 25 and 70 μmol photons m⁻² s⁻¹ were obtained at concentrations of > 0.15–0.3×10⁵ M. pelliculata ml⁻¹, corresponding to 50–100 μg C 1⁻¹. Ingestion of M. pelliculata cells by C. ericina did not generally follow Michaelis—Menten kinetics. Deviation from the expected saturation kinetics was especially pronounced at irradiances of 70 and 150 μmol photons m⁻² s⁻¹. At these irradiances ingestion of M. pelliculata cells by C. ericina decreased at high concentrations of M. pelliculata, indicating an increased uptake of bacterial prey in these cultures. The growth rate of C. ericina was affected in both monocultures and in mixed cultures when pH increased above 8.6, and growth stopped around pH 9. The prey alga M. pelliculata tolerated high pH better and, consequently, took over in the mixed cultures when pH exceeded 9. The ecological significance of mixotrophy in the genus Chrysochromulina is discussed. Published online: 4 July 2002     

Photosynthetic utilisation of carbon and light by two tropical seagrass species as measured in situ

In situ measurements of seagrass photosynthesis in relation to inorganic carbon (Ci) availability, increased pH and an inhibitor of extracellular carbonic anhydrase were made using an underwater pulse amplitude modulated (PAM) fluorometer. By combining the instrument with a specially designed Perspex chamber, we were able to alter the water surrounding a leaf without removing it from the growing plant. Responses to Ci within the chamber showed that subtidal plants of the seagrasses Cymodocea serrulata and Halophila ovalis had photosynthetic rates that were limited by the ambient Ci concentration depending on the irradiance that was available during short-term photosynthesis–irradiance trials. Relative electron transport rates (RETRs) at light saturation (up to 500 μ mol photons m⁻² s⁻¹) increased by 66–100% when the Ci concentration was increased from ca. 2.2 to 6.2 mM. On the other hand, intertidal plants of the same species exhibited a much lesser limitation of photosynthesis by Ci at any irradiance (up to 1500 μ mol photons m⁻² s⁻¹). Both species were able to use HCO⁻ ₃ efficiently, and there was stronger evidence for direct uptake of HCO⁻ ₃ rather than extracellular dehydration of HCO⁻ ₃ to CO₂ prior to Ci uptake. Subtidally, H. ovalis and C. serrulata grew to 10 and 12 m, respectively, where ambient irradiances were approximately 16 and 11% of those at the surface. Maximum RETRs (at light saturation) were lower for these deep-growing plants than for the intertidally growing ones. For both species, the onset of light saturation of photosynthesis (E _k) occurred at approximately 100 μ mol photons m⁻² s⁻¹ for the deep water populations, which was four and two times lower than for the shallow populations of C. serrulata and H. ovalis, respectively. This, and the differences in maximal photosynthetic rates (RETR _max), reflects an acclimation of the deep-growing populations to the lower light environment. The results presented here show that photosynthesis, as measured in situ, was limited by the availability of Ci for the deeper growing plants in Zanzibar, while the intertidally growing plants photosynthesised at close to Ci saturation. The latter result is contrary to previous conclusions regarding Ci limitations for these intertidal plants, and, in general, our findings highlight the need for performing similar experiments in situ rather than under laboratory conditions. Received: 4 April 2000 / Accepted: 31 August 2000     

Distribution and maximum growth depth of Fucus vesiculosus along the Gulf of Finland

 A survey of the distribution and maximum depth of a continuous Fucus vesiculosus belt was carried out in the Gulf of Finland in 1991. F. vesiculosus is widely distributed throughout the Gulf of Finland, including the vicinity of Vyborg Bay, Russia in the east. The maximum growth depth of F. vesiculosus in the Gulf of Finland reflects two different patterns according to the exposure to wave action. The most robust and continuous F. vesiculosus belt is observed on exposed shores, where the maximum growth depth is 5 to 6 m, with the optimum at 2 to 3 m. On moderately exposed shores the maximum growth depth is 3 m, with an optimum growth depth of <2 m. The maximum growth depth also varies geographically, with a decreasing trend towards the east. Maximum growth depth of F. vesiculosus correlates with light intensity. The compensation point for F. vesiculosus photosynthesis is about 25 μmol m⁻² s⁻¹, and photosynthesis is saturated at a light intensity of 300 μmol m⁻² s⁻¹. Vertical irradiance attenuation measurements in situ in summer revealed that for F. vesiculosus photosynthesis the quantity of light is optimal (200 to 300 μmol m⁻² s⁻¹) at <3 m depth. At depths >5 m the quantity of light is near or below the photosynthesis compensation point and insufficient for growth. These depth limits of light penetration coincide with measured growth depths of F. vesiculosus in the Gulf of Finland. Received: 7 May 1999 / Accepted: 18 November 1999     

Irradiance and temperature regulation of oxygenic photosynthesis and O2 consumption in a hypersaline cyanobacterial mat (Solar Lake, Egypt)

 Short-term effects of temperature and irradiance on oxygenic photosynthesis and O₂ consumption in a hypersaline cyanobacterial mat were investigated with O₂ microsensors in a laboratory. The effect of temperature on O₂ fluxes across the mat–water interface was studied in the dark and at a saturating high surface irradiance (2162 μmol photons m⁻² s⁻¹) in the temperature range from 15 to 45 °C. Areal rates of dark O₂ consumption increased almost linearly with temperature. The apparent activation energy of 18 kJ mol⁻¹ and the corresponding Q ₁₀ value (25 to 35 °C) of 1.3 indicated a relative low temperature dependence of dark O₂ consumption due to mass transfer limitations imposed by the diffusive boundary layer at all temperatures. Areal rates of net photosynthesis increased with temperature up to 40 °C and exhibited a Q ₁₀ value (20 to 30 °C) of 2.8. Both O₂ dynamics and rates of gross photosynthesis at the mat surface increased with temperature up to 40 °C, with the most pronounced increase of gross photosynthesis at the mat surface between 25 and 35 °C (Q ₁₀ of 3.1). In another mat sample, measurements at increasing surface irradiances (0 to 2319 μmol photons m⁻² s⁻¹) were performed at 25, 33 (the in situ temperature) and 40 °C. At all temperatures, areal rates of gross photosynthesis saturated with no significant reduction due to photoinhibition at high irradiances. The initial slope and the onset of saturation (E _k = 148 to 185 μmol photons m⁻² s⁻¹) estimated from P versus E _d curves showed no clear trend with temperature, while maximal photosynthesis increased with temperature. Gross photosynthesis was stimulated by temperature at each irradiance except at the lowest irradiance of 54 μmol photons m⁻² s⁻¹, where oxygenic gross photosynthesis and also the thickness of the photic zone was significantly reduced at 40 °C. The compensation irradiance increased with temperature, from 32 μmol photons m⁻² s⁻¹ at 25 °C to 77 μmol photons m⁻² s⁻¹ at 40 °C, due to increased rates of O₂ consumption relative to gross photosynthesis. Areal rates of O₂ consumption in the illuminated mat were higher than dark O₂ consumption at corresponding temperatures, due to an increasing O₂ consumption in the photic zone with increasing irradiance. Both light and temperature enhanced the internal O₂ cycling within hypersaline cyanobacterial mats. Received: 30 November 1999 / Accepted: 11 April 2000     

Light requirements for growth of six shade-acclimated Mediterranean macroalgae

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

Photoacclimation in phytoplankton: implications for biomass estimates, pigment functionality and chemotaxonomy

Chl a and C-normalized pigment ratios were studied in two dinophytes (Prorocentrum minimum and Karlodinium micrum), three haptophytes (Chrysochromulina leadbeateri, Prymnesium parvum cf. patelliferum, Phaeocystis globosa), two prasinophytes (Pseudoscourfieldia marina, Bathycoccus prasinos) and the raphidophyte Heterosigma akashiwo, in low (LL, 35 μmol photons m⁻² s⁻¹) and high light (HL, 500 μmol photons m⁻² s⁻¹). Pigment ratios in LL and HL were compared against a general rule of photoacclimation: LL versus HL ratios ≥1 are typical for light-harvesting pigments (LHP) and <1 for photoprotective carotenoids. Peridinin, prasinoxanthin, gyroxanthin-diester and 19′-butanoyloxy-fucoxanthin were stable chemotaxonomic markers with less than 25% variation between LL versus HL Chl a–normalized ratios. As expected, Chls exhibited LL/HL to Chl a ratios >1 with some exceptions such as Chl c ₃ in P. globosa and MV Chl c ₃ in C. leadbeateri. LL/HL to Chl a ratios of photosynthetic carotenoids were close to 1, except Hex-fuco in P. globosa (four-fold higher Chl a ratio in HL vs LL). Although pigment ratios in P. globosa clearly responded to the light conditions the diadinoxanthin-diatoxanthin cycle remained almost unaltered at HL. Total averaged pigment and LHP to C ratios were significantly higher in LL versus HL, reflecting the photoacclimation status of the studied species. By contrast, the same Chl a-normalized ratios were weakly affected by the light intensity due to co-variation with Chl a. Based on our data, we suggest that the interpretation of PPC and LHP are highly dependent on biomass normalization (Chl a vs. C).     

Diurnal cycle and kinetics of ammonium assimilation in the green alga <Emphasis Type="Italic">Ulva pertusa</Emphasis>

The kinetics of ammonium assimilation was investigated in Ulva pertusa (Chlorophyceae, Ulvales) from northeastern New Zealand. Ammonium assimilation exhibited Michaelis–Menten kinetics with a maximum rate of assimilation (V _max) of 54 ± 5 μmol g⁻¹ dry weight h⁻¹ and half-saturation constant (K _m) of 23 ± 8 μM. In contrast, values for ammonium uptake were considerably higher with a V _max of 316 ± 59 μmol g⁻¹ dry weight h⁻¹ and K _m of 135 ± 46 μM. At environmentally relevant ammonium concentrations (5 μM), assimilation accounted for most (70%) of the ammonium taken up. Darkness decreased the maximum rate of ammonium assimilation by 83%. We investigated the hypothesis that rates of biosynthetic processes are greater in the early part of the day in Ulva. Consistent with this hypothesis, the maximum rate of ammonium assimilation in U. pertusa peaked in the morning and coincided with low levels of the photosynthetic product sucrose, which peaked in the afternoon. There was a diurnal cycle in the rate of ammonium uptake and assimilation in light and dark, but the amplitude was much greater for assimilation than uptake. Moreover, our data suggest that net ammonium assimilation only occurs during the day in U. pertusa. We suggest that two major roles for diurnal cycles are minimisation of interspecific competition for resources and metabolic costs.     

Phytoplankton community structure and primary production in small intertidal estuarine-bay ecosystem (eastern English Channel,France)

From May 2002 to October 2003, a fortnightly sampling programme was conducted in a restricted macrotidal ecosystem in the English Channel, the Baie des Veys (France). Three sets of data were obtained: (1) physico-chemical parameters, (2) phytoplankton community structure illustrated by species composition, biovolume and diversity, and (3) primary production and photosynthetic parameters via P versus E curves. The aim of this study was to investigate the temporal variations of primary production and photosynthetic parameters in this bay and to highlight the potential links with phytoplankton community structure. The highest level of daily depth-integrated primary production Pz (0.02–1.43 g C m⁻² d⁻¹) and the highest maximum photosynthetic rate P ^B _max (0.39–8.48 mg C mg chl a ⁻¹ h⁻¹) and maximum light utilization coefficient α^B [0.002–0.119 mg C mg chl a ⁻¹ h⁻¹ (μmol photons m⁻² s⁻¹)] were measured from July to September. Species succession was determined based on biomass data obtained from cell density and biovolume measurements. The bay was dominated by 11 diatoms throughout the year. However, a Phaeocystis globosa bloom (up to 25 mg chl a m⁻³, 2.5 × 10⁶ cells l⁻¹) was observed each year during the spring diatom bloom, but timing and intensity varied interannually. Annual variation of primary production was due to nutrient limitation, light climate and water temperature. The seasonal pattern of microalgal succession, with regular changes in composition, biovolume and diversity, influenced the physico-chemical and biological characteristics of the environment (especially nutrient stocks in the bay) and thus primary production. Consequently, investigation of phytoplankton community structure is important for developing the understanding of ecosystem functioning, as it plays a major role in the dynamics of primary production.     

Photoinhibition of photosynthesis in a sun and a shade species of the red algal genus Porphyra

Gametophytes of two species of Porphyra collected around San Juan Island, Washington in 1986 and acclimated to low light conditions in culture showed different resistances to photoinhibition of photosynthesis. The intertidal species P. perforata J. Agardh exhibited photoinhibition at one-third the rate exhibited by the subtidal species P. nereocystis Anderson following treatments at 2000 mol photons m^-2 s^-1 under conditions of full hydration and optimal temperature. The greater resistance of P. perforata to photoinhibition could not be attributed to reduced photosynthetic pigment concentration, higher photosynthetic capacity, avoidance of light by chloroplast movement or to enhanced rates of photorespiration. Total carotenoid concentrations were similar in the two species. It is probable that the mechanisms of this resistance are operating at the level of the thylakoid membranes. Resistance to photoinhibition represents an adaptation of photosynthesis in P. perforata which may contribute to its persistance in the extreme environment of its intertidal habitat.     

Microsensor studies of photosynthesis and respiration in larger symbiotic foraminifera. I The physico-chemical microenvironment of Marginopora vertebralis, Amphistegina lobifera and Amphisorus hemprichii

 The physico-chemical microenvironment of larger benthic foraminifera was studied with microsensors for O₂, CO₂, pH, Ca²⁺ and scalar irradiance. Under saturating light conditions, the photosynthetic activity of the endosymbiotic algae increased the O₂ up to 183% air saturation and a pH of up to 8.6 was measured at the foraminiferal shell surface. The photosynthetic CO₂ fixation decreased the CO₂ at the shell down to 4.7 μM. In the dark, the respiration of host and symbionts decreased the O₂ level to 91% air saturation and the CO₂ concentration reached up to 12 μM. pH was lowered relative to the ambient seawater pH of 8.2. The endosymbionts responded immediately to changing light conditions, resulting in dynamic changes of O₂, CO₂ and pH at the foraminiferal shell surface during experimentally imposed light–dark cycles. The dynamic concentration changes demonstrated for the first time a fast exchange of metabolic gases through the perforate, hyaline shell of Amphistegina lobifera. A diffusive boundary layer (DBL) limited the solute exchange between the foraminifera and the surrounding water. The DBL reached a thickness of 400–700 μm in stagnant water and was reduced to 100–300 μm under flow conditions. Gross photosynthesis rates were significantly higher under flow conditions (4.7 nmol O₂ cm⁻³ s⁻¹) than in stagnant water (1.6 nmol O₂ cm ⁻³ s⁻¹), whereas net photosynthesis rates were unaffected by flow conditions. The Ca²⁺ microprofiles demonstrated a spatial variation in sites of calcium uptake over the foraminiferal shells. Ca²⁺ gradients at the shell surface showed total Ca²⁺ uptake rates of 0.6 to 4.2 nmol cm⁻² h⁻¹ in A. lobifera and 1.7 to 3.6 nmol cm⁻² h⁻¹ in Marginopora vertebralis. The scattering and reflection of the foraminiferal calcite shell increased the scalar irradiance at the surface up to 205% of the incident irradiance. Transmittance measurements across the calcite shell suggest that the symbionts are shielded from higher light levels, receiving approximately 30% of the incident light for photosynthesis. Received: 6 July 1999 / Accepted: 28 April 2000     

Spring sea ice photosynthesis,primary productivity and biomass distribution in eastern Antarctica, 2002–2004

While it is known that Antarctic sea ice biomass and productivity are highly variable over small spatial and temporal scales, there have been very few measurements from eastern Antarctic. Here we attempt to quantify the biomass and productivity and relate patterns of variability to sea ice latitude ice thickness and vertical distribution. Sea ice algal biomass in spring in 2002, 2003 and 2004 was low, in the range 0.01–8.41 mg Chl a m⁻², with a mean and standard deviation of 2.08 ± 1.74 mg Chl a m⁻² (n = 199). An increased concentration of algae at the bottom of the ice was most pronounced in thicker ice. There was little evidence to suggest that there was a gradient of biomass distribution with latitude. Maximum in situ production in 2002 was approximately 2.6 mg C m⁻² h⁻¹ with assimilation numbers of 0.73 mg C (mg Chl a)⁻¹ h⁻¹. Assimilation numbers determined by the ¹⁴C incubations in 2002 varied between 0.031 and 0.457 mg C (mg Chl a)⁻¹ h⁻¹. Maximum fluorescence quantum yields of the incubated ice samples in 2002 were 0.470 ± 0.041 with E _k indices between 19 and 44 μmol photons m⁻² s⁻¹. These findings are consistent with the shade-adapted character of ice algal communities. In 2004 maximum in situ production was 5.9 mg C m⁻² h⁻¹ with an assimilation number of 5.4 mg C (mg Chl a)⁻¹ h⁻¹. Sea ice biomass increased with ice thickness but showed no correlation with latitude or the time the ice was collected. Forty-four percent of the biomass was located in bottom communities and these were more commonly found in thicker ice. Surface communities were uncommon.     

A method for the assessment of light-induced oxidative stress in embryos of fucoid algae via confocal laserscan microscopy

A method was developed for measurement of active oxygen production in embryonic stages of the brown seaweed Fucus spiralis, using the label CM-DCFH-DA. Active oxygen species convert the label into the green fluorescent CM-DCF (exc/em 488/530 nm) that is detected via confocal laserscan microscopy and quantitative image analysis. Loading of the label did not harm the embryos; loading efficiency was age-independent, and the esterases needed for conversion to CM-DCFH were not inhibited by the effective UV dose (2 W m⁻²) applied in the experiments. After correction for daily variation of the laser power, and calibration with DCF standard solutions, this automated analysis of confocal images rendered active oxygen concentrations in fucoid embryos (μM DCF). An experiment was designed for the assessment of active oxygen production following irradiance stress in the light-sensitive embryos. Dim-light-acclimated, 1-, 2- and 4-day-old embryos were transferred for 60 min to defined high-light conditions (4π-irradiance 300 μmol photons m⁻² s⁻¹), optionally without UV radiation, including UVA, or including UVA plus UVB. PSII yield measurements (PAM fluorometer) were carried out in order to assess the degree of photoinhibition under these light conditions. The imposed light stress initially caused a rapid decrease of the PSII yields (Φ_P). With increasing embryo age, minimum Φ_P values attained under light stress remained higher. Consequently, electron transport rates (ETR) would increase with embryo age, i.e., with the development of their photosynthetic apparatus. Active oxygen production increased with ETR, and when UVB was included, relatively greater amounts of active oxygen were produced. A slow, second-phase decrease of Φ_P under light stress that was proportional to active oxygen production indicated that some photooxidative damage was caused, in particular during UVB exposure. Recovery from light stress was a rapid process in the absence of UVB; in such cases Φ_P was almost restored to the initial values within 60 min. The relative state of recovery of Φ_P was correlated with both the effective UV dose and active oxygen production rate (DCF). Recovery was slowest in embryos exposed for 60 min to an experimental UVB dose, which was representative of a situation at low tide, on a sunny day. The results suggest that active oxygen may cause an in situ inhibition of growth of the earliest life stages of F. spiralis. Received: 26 January 2000 / Accepted: 4 September 2000     

Root respiration and oxygen flux in salt marsh grasses from different elevational zones

Plants growing in waterlogged environments are subjected to low oxygen levels around submerged tissues. While internal oxygen transport has been postulated as an important factor governing flooding tolerance, respiration rates and abilities to take up oxygen under hypoxic conditions have been largely ignored in plant studies. In this study, physiological characteristics related to internal oxygen transport, respiration, and oxygen affinity were studied in low intertidal marsh species (Spartina alterniflora and S. anglica) and middle to high intertidal species (S. densiflora, S. patens, S. foliosa, a S. alterniflora × S. foliosa hybrid, S. spartinae, and Distichlis spicata). These marsh plants were compared to the inland species S. pectinata and the crop species rice (Oryza sativa), corn (Zea mays), and oat (Avena sativa). Plants were grown in a greenhouse under simulated estuarine conditions. The low marsh species S. anglica was found to transport oxygen internally at rates up to 2.2 μmol O₂ g fresh root weight⁻¹ h⁻¹. In contrast, marsh species from higher zones and crop species were found to transport significantly less oxygen internally, although rice plants were able to transport 1.4 μmol g⁻¹ h⁻¹. Under hypoxic conditions, low marsh species were better able to remove dissolved oxygen from the medium compared to higher marsh species and crops. The oxygen concentration at which respiration rates declined due to limited oxygen (P _crit) was significantly lower in low marsh species compared to inland and crop species; P _crit ranged from <4 μM O₂ in the low marsh species S. anglica up to 20 μM in the inland species corn. Flooding-sensitive crop species had significantly higher aerobic respiration rates compared to flooding-tolerant species in this study. Crop species took up 3.6–6.7 μmol O₂ g⁻¹ h⁻¹ while all but one marsh species took up <3.5 μmol O₂ g⁻¹ h⁻¹. We conclude that oxygen transport, aerobic demand, and oxygen affinity all play important and interrelated roles in flood tolerance and salt marsh zonation.     

Photobiology of endolithic microorganisms in living coral skeletons: 1. Pigmentation, spectral reflectance and variable chlorophyll fluorescence analysis of endoliths in the massive corals Cyphastrea serailia, Porites lutea and Goniastrea australensis

We used microscopy, reflectance spectroscopy, pigment analysis, and photosynthesis-irradiance curves measured with variable fluorescence techniques to characterise the endolithic communities of phototrophic microorganisms in the skeleton of three massive corals from a shallow reef flat. Microscopic observations and reflectance spectra showed the presence of up to four distinct bands of photosynthetic microorganisms at different depths within the coral skeleton. Endolithic communities closer to the coral surface exhibited higher photosynthetic electron transport rates and a green zone dominated by Ostreobium quekettii nearest the surface had the greatest chlorophyll pigment concentration. However, Ostreobium was also present and photosynthetically active in the colourless band between the coral tissue and the green band. The spectral properties and pigment density of the endolithic bands were also found to closely correlate to photosynthetic rates as assessed by fluorometry. All endolithic communities were extremely shade-adapted, and photosynthesis was saturated at irradiances <7 μmol photons m⁻²s⁻¹.