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     

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     

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     

A comparison of two cartographic exposure methods using Fucus vesiculosus as an indicator

Morphological variation and vertical distribution of Fucus vesiculosus were quantified at several sites in the Finnish archipelago (Baltic Sea). F. vesiculosus samples were obtained from skerries at geographical distances of 1 km or more (large scale) and at intervals of ca 100 m around a single island (small scale). The results were examined in relation to wave exposure, calculated by Baardseth and effective fetch cartographic methods. Despite the fact that the exposure indices were calculated differently they correlated strongly. Vegetative morphological characteristics of F. vesiculosus illustrate the morphological differences both within and between exposure gradients. The tallest and widest F. vesiculosus plants were found at the sheltered end of the large-scale exposure gradient. Those from equally sheltered sites of the island were smaller in all respects. Thus, the trend from small narrow plants to large wide sheltered plants was expressed differently over the different geographical scales. Consequently localities with similar exposure indices may have morphologically different F. vesiculosus populations. Shores with similar cartographic exposure indices can be different in nature. Underwater topography and shore locations, either close to the mainland or at the outermost sites of the archipelago, affect the exposure. Although a sheltered shore is indicated, the sublittoral zone may be quite exposed to the movements of water. In contrast, in an open shore environment underwater rocks, boulders and shallow water areas can provide sheltered habitats. The depth range of the F. vesiculosus belt exhibited two distinctive patterns. At sheltered sites, around islands in the outermost reaches of the archipelago F. vesiculosus can grow to a maximum depth of 5 m. In exposed habitats the belt becomes narrower, reaching a maximum depth of 3 m. Closer to the mainland F. vesiculosus is found at exposed sites to a maximum depth of 5 m; the depth range at sheltered sites is narrower, only reaching depths of 2 m or less. In conclusion, the changes in plant morphology and in the vertical belt distribution are similar to each other along both gradients at the exposed ends of the wave action spectrum; however, the two gradients diverge at the sheltered ends of the spectrum. Received: 10 August 1998 / Accepted: 11 January 1999     

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     

Photosynthetic performance of giant clams, Tridacna maxima and T. squamosa, Red Sea

Two species of giant clams, Tridacna maxima and T. squamosa, coexist in the Red Sea, but exhibit distinctly different depth distributions: T. maxima mostly occurs in shallow waters (reef flat and edge), while T. squamosa may occur down to the lower fore-reef slope. Giant clams have been described as mixotrophic, capable of both filter-feeding and photosynthesis due to algal symbionts (zooxanthellae), therefore, observed depth preferences were investigated in relation to possible differences in autotrophy vs. heterotrophy. This study was conducted from April to June 2004, at the reef near the Marine Science Station, Aqaba, Gulf of Aqaba, Red Sea, and in May 2007, at a reef near Dahab, Sinai Peninsula, Egypt. In situ measurements using a submersible pulse amplitude modulated fluorometer (Diving PAM), revealed no significant differences in effective PSII quantum yield (ΔF/Fm′) and relative electron transport rates (ETR) between the two species; but rapid light curves (ETR vs. light, photosynthetically active irradiance, PAR) showed significant differences in maximum photosynthetic rates (ETR_max), with 20% higher values in T. maxima. Chamber incubations displayed higher net and gross oxygen production by T. maxima (88.0 and 120.3 μmol O₂ cm⁻² mantle area day⁻¹) than T. squamosa (56.7 and 84.8 μmol O₂ cm⁻² mantle area day⁻¹); even under shading conditions (simulated depth of 20 m) T. maxima still achieved 93% of the surface gross O₂ production, whereas T. squamosa reached only 44%. A correlation was found between ETR and net photosynthesis measured as oxygen production (T. maxima: R ² = 0.53; T. squamosa: R ² = 0.61). Calculated compensation depth (CD) (gross photosynthesis equals respiration) in T. maxima (16 m) matches the maximum depth of occurrence in this study (17 m). By contrast, the CD of T. squamosa (9 m) was much shallower than the maximum vertical range (42 m). Findings suggest T. maxima is a strict functional photoautotroph limited by light, whereas T. squamosa is a mixotroph whose photoautotrophic range is extended by heterotrophy. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Fitting fertilisation kinetics models for free-spawning marine invertebrates

 To determine how fertilisation varied with sperm concentration for two species of scallop, Chlamys (Equichlamys) bifrons (Lamarck) and C. asperrima (Lamarck), we performed a simple series of sperm dilution experiments, and measured egg size and sperm swimming speeds. C. bifrons eggs were much larger (average diam=116.5 μm), and sperm swimming speeds faster (209.8 μm s⁻¹), than C. asperrima (71.2 μm, 166.0 μm s⁻¹). In both species, maximum fertilisation occurred at an ambient sperm concentration of around 100 sperm μl⁻¹; the maximum proportion of eggs fertilised was less than 0.70 in the C. bifrons experiments, but nearer 1.0 with C. asperrima. At high sperm concentrations (>100 sperm μl⁻¹), fertilisation decreased (presumably due to polyspermy) with increasing sperm concentration, but decreased more rapidly in C. bifrons than C. asperrima. A polyspermy-adjusted fertilisation kinetics model could be fitted to the experimental data, but unique parameter estimates could not be determined. Received: 7 October 1999 / Accepted: 8 July 2000     

Symbiotic anemones can grow when starved: nitrogen budget for Anemonia viridis in ammonium-supplemented seawater

The ability of endosymbioses between anthozoans and dinoflagellate algae (zooxanthellae) to retain excretory nitrogen and take up ammonium from seawater has been well documented. However, the quantitative importance of these processes to the nitrogen budget of such symbioses is poorly understood. When starved symbiotic Anemonia viridis were incubated in a flow-through system in seawater supplemented with 20 μM ammonium for 91 d under a light regime of 12 h light at 150 μmol photons m⁻² s⁻¹ and 12 h darkness, they showed a mean net growth of 0.197% of their initial weight per day. Control anemones in unsupplemented seawater with an ammonium concentration of <1 μM lost weight by a mean of 0.263% of their initial weight per day. Attempts to construct a nitrogen budget showed that, over a 14 d period, ≃40% of the ammonium taken up could be accounted for by growth of zooxanthellae. It was assumed that the remainder was translocated from zooxanthellae to host. However, since the budget does not balance, only 60% of the growth of host tissue was accounted for by this translocation. The value for host excretory nitrogen which was recycled to the symbionts equalled that taken in by ammonium uptake from the supplemented seawater, indicating the importance of nitrogen retention to the symbiotic association. Received: 23 December 1997 / Accepted: 12 September 1998     

Resource acquisition in two intertidal fucoid seaweeds, Fucus serratus and Himanthalia elongata : seasonal variation and effects of reproductive development

Seasonal variations and the effect of reproductive development on resource acquisition by two intertidal fucoid species, the iteroparous Fucus serratus L. and the semelparous Himanthalia elongata (L.) S. F. Gray were examined. The oxygen-exchange characteristics of vegetative apical tissue of both non-fertile and fertile plants and receptacle tissue were compared at monthly intervals throughout reproductive development. Respiratory rates in non-fertile F. serratus varied seasonally between 1.5 and 8.0 μmol g⁻¹ fresh wt h⁻¹; in fertile plants the receptacle had a significantly lower respiratory rate than the vegetative tissue. The respiratory rate of the vegetative button of fertile H. elongata displayed less seasonal variation and was lower than that of the receptacle, which varied from a maximum of 9.5 μmol g⁻¹ fresh wt h⁻¹ at receptacle initiation in October to a minimum of 2.0 μmol g⁻¹ fresh wt h⁻¹ in February. The maximum photosynthetic rate (P _max) of non-fertile plants of both species did not vary in a distinct seasonal manner (∼60 μmol g⁻¹ fresh wt h⁻¹ for F. serratus and ∼12 μmol g⁻¹ fresh wt h⁻¹ for H. elongata). In fertile plants, the P _max of the receptacle tissue was (∼50% lower in F. serratus, and at its peak three times higher in H. elongata, than that of vegetative tissue. The stable carbon-isotope ratio (δ¹³C) did not differ between different tissue types in F. serratus, but values did vary seasonally, being less negative in the summer than in the winter (−13.5‰ compared to −18‰). The receptacle tissue of H. elongata also displayed a distinct seasonal variation in δ¹³C values (−12‰ in summer, −16‰ in winter), whilst the δ¹³C of the vegetative button did not vary seasonally. The rate of uptake of inorganic nitrogen by the vegetative thallus was lower in H. elongata than in F. serratus. The receptacle tissue of F. serratus had lower uptake rates than the vegetative tissue, whilst the uptake rate by H. elongata receptacle tissue was higher than that of the vegetative button. Received: 14 March 1997 / Accepted: 22 April 1997     

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     

Phytoplankton blooms are strongly impacted by microzooplankton grazing in coastal North Pacific waters

Phytoplankton growth and microzooplankton grazing were measured in two productive coastal regions of the North Pacific: northern Puget Sound and the coastal Gulf of Alaska. Rates of phytoplankton growth (range: 0.09–2.69 day⁻¹) and microzooplankton grazing (range: 0.00–2.10 day⁻¹) varied seasonally, with lowest values in late fall and winter, and highest values in spring and summer. Chlorophyll concentrations also varied widely (0.19–13.65 μg l⁻¹). Large (>8 μm) phytoplankton cells consistently dominated phytoplankton communities under bloom conditions, contributing on average 65% of total chlorophyll biomass when chlorophyll exceeded 2 μg l⁻¹. Microzooplankton grazing was an important loss process affecting phytoplankton, with grazing rates equivalent to nearly two-thirds (64%) of growth rates on average. Both small and large phytoplankton cells were consumed, with the ratio of grazing to growth (g:μ) for the two size classes averaging 0.80 and 0.42, respectively. Perhaps surprisingly, the coupling between microzooplankton grazing and phytoplankton growth was tighter during phytoplankton blooms than during low biomass periods, with g:μ averaging 0.78 during blooms and 0.49 at other times. This tight coupling may be a result of the high potential growth and ingestion rates of protist grazers, some of which feed on bloom-forming diatoms and other large phytoplankton. Large ciliates and Gyrodinium-like dinoflagellates contributed substantially to microzooplankton biomass at diatom bloom stations in the Gulf of Alaska, and microzooplankton biomass overall was strongly correlated with >8 μm chlorophyll concentrations. Because grazing tended to be proportionally greater when phytoplankton biomass was high, the absolute amount of chlorophyll consumed by microzooplankton was often substantial. In nearly two-thirds of the experiments (14/23), more chlorophyll was ingested by microzooplankton than was available for all other biological and physical loss processes combined. Microzooplankton were important intermediaries in the transfer of primary production to higher trophic levels in these coastal marine food webs. Received: 12 November 1999 / Accepted: 4 October 2000     

Effects of ambient ammonia levels on blood ammonia, ammonia excretion and heart and scaphognathite rates of Nephrops norvegicus

During commercial handling of Nephropsnorvegicus (L.) there are a number of situations when the prawns may be exposed to very high ambient ammonia levels. These experiments evaluated the effects of increased levels of ambient total ammonia (TA = NH₃ + NH₄ ⁺) on␣blood ammonia, ammonia efflux rates and on the cardio-ventilatory performance of N. norvegicus. When prawns were taken from <1 to 2000 μmol TA l⁻¹ medium, blood TA concentrations increased rapidly for the first 2 h but tended to drop thereafter. Original blood TA levels were restored 6 h after the prawns were transferred back from seawater containing 2000 to <1 μmol TA l⁻¹. Sudden exposure to 500, 1000, 2000 or 4000 μmol TA l⁻¹ medium induced blood TA concentrations to increase respectively to 50, 30, 33 and 36% of external concentrations (normally, internal TA values are much higher than external levels). Immediately after transfer back to seawater with low ammonia concentration ( <1 μmol TA l⁻¹), excretion rates were higher than those of control prawns, and the absolute amounts of TA excreted were considerably higher than those calculated to have accumulated in the haemolymph. Heart rate (HR) and scaphognathite rate (SR) were not altered when prawns were subjected to sudden alterations in ambient ammonia ( <1 to 2000 to <1 μmol TA l⁻¹). When water ammonia concentrations were altered more gradually, both rates increased, but only at 4000 μmol TA l⁻¹. These results show that N. norvegicus is able to remove ammonia from the haemolymph and/or transform ammonia into some other substance when subjected to increased levels of ambient ammonia. Possible mechanisms involved (e.g. active transport across the gills; storage in some other tissue; glutamate synthe sis) are discussed. Received: 20 May 1996 / Accepted: 1 July 1996     

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     

Growth and productivity of the high Antarctic Bryozoan Melicerita obliqua

We analysed growth of the Antarctic bryozoan Melicerita obliqua (Thornely, 1924) by x-ray photography and stable isotope analysis. M. obliqua colonies form one segment per year, thus attaining maximum length of about 200 mm within 50 years. In the Weddell and Lazarev Seas, annual production/biomass ratio of M. obliqua is 0.1 yr⁻¹, which is in the range of other Antarctic benthic invertebrate populations. Production amounts to 3.34 mg C_orgm⁻² yr⁻¹ and 90.6 mg ash m⁻² yr⁻¹ on the shelf (100 to 600 m water depth), and to 0.13 mg C_orgm⁻² yr⁻¹ and 36.8 mg ash m⁻² yr⁻¹ on the slope (600 to 1250 m water depth). Received: 27 February 1998 / Accepted: 8 May 1998     

Changes in male guppy courting distance in response to a fluctuating light environment

In the guppy (Poecilia reticulata), effective courting by a male requires visual contact with the female. Therefore, environmental light intensity may affect male display behavior, particularly initial courtship distance. We found that male guppies courted at exact and predictable distances from the female given a particular light level, both in field and laboratory studies. In lower light levels (<0.1 μmol m⁻² s⁻¹), for example at dawn, dusk, or under heavy canopy, males court females at closer and less variable distances (<3 cm). At higher light levels, which occur during most of the day and with less canopy cover, males often court from twice or three times further out. Light levels over guppy streams change over relatively short time periods and ranges, correlating with variation in courtship distances. Laboratory manipulations of irradiance confirmed that courtship distance depends on illumination. Hence, courtship distances may be set by the effect of lighting on signal efficiency, minimization of energy or time expenditures, or predation risk. Received: 16 December 1997 / Accepted after revision: 8 August 1998     

Activity and inhibitor sensitivity of ATPases in the hydrothermal vent tubeworm Riftia pachyptila : a comparative approach

Phosphorylated ATPases may be involved in the effective pH regulation seen in the hydrothermal vent tubeworm Riftia pachyptila. R. pachyptila appears not only to have a large concentration of ATPases, but the main function of these ATPases seems to have shifted from other types of transport, such as Na⁺ and K⁺ movement, to the facilitation of H⁺ elimination. Plume and trophosome ATPase activity for R. pachyptila measured 646.2 ± 29.5 and 481.4 ± 32.0 μmol Pi (inorganic phosphate) g⁻¹ wet wth⁻¹, respectively. Plume tissue ATPase activity (both mass-specific and protein-specific) in R. pachyptila was higher (between 7% and 55%) than the activity measured in any tissue for 7 other shallow- and deep-living species, in this study. This supports the hypothesis that R. pachyptila regulates acid/base balance via high concentrations of H⁺-ATPases, including Na⁺/H⁺ and K⁺/H⁺ exchangers and possibly electrogenic H⁺-ATPases, as evidenced by a higher total ATPase concentration (646 μmol Pi g⁻¹ wet wt h⁻¹), lesser Na⁺/K⁺-ATPase activity (13% of the total, as compared to 20−40% found in other animals), and higher H⁺-ATPase activity (226–264 μmol Pi g⁻¹wet wt h⁻¹). Overall, R. pachyptila appears to demonstrate elevated ATPase activity, with a greater fraction of the enzymes devoted to proton elimination, in order to effectively control its extracellular pH in the face of processes acting to acidify the internal environment. Received: 9 May 2000 / Accepted: 4 October 2000     

Differences in chloroplast ultrastructure of Phaeocystis antarctica in low and high light

Phaeocystis antarctica Karsten exhibits optical changes in pigment packaging during acclimation to drastically different light levels. Here, the three-dimensional morphological rearrangements are shown for two light conditions mimicking limiting and saturating conditions for photosynthesis. Cultures of P. antarctica were grown semi-continuously under light-limiting conditions for growth (14 μmol quanta m⁻² s⁻¹) and a light-saturating condition (259 μmol quanta m⁻² s⁻¹) for growth. Increased numbers of thylakoids were observed under the low light treatment. In contrast, there were less amounts of thylakoid stacking in each chloroplast under the high light treatment. Electron microscopic tomographic reconstructions illustrate the complexity of the chloroplast organelle where the thylakoids ‘interact’ with the pyrenoid and the chloroplast membrane. Highly complex characteristics, such as bi- and tri-furcations in the thylakoid stacks, were continuous throughout the chloroplast. Other organelles, such as the Golgi apparatus and dense vesicles that may potentially affect cellular scattering and absorption were also observed in both high and low light. These three dimensional thylakoid arrangements have profound implications for cellular photophysiology. They represent a new view of algal chloroplast structure, and provide a starting point for more accurate optical modeling studies.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Temporal and spatial occurrence of UV-absorbing mycosporine-like amino acids in tissues of the antarctic sea urchin Sterechinusneumayeri during springtime ozone-depletion

From September to November 1991, UV-absorbing mycosporine-like amino acids (MAAs) were monitored in a natural population of the sea urchin Sterechinusneumayeri from a coastal area of Anvers Island (Antarctic Peninsula). MAA concentrations were determined for specific tissues (gonad, digestive tract and body wall) from adults collected at four depths (intertidal, 8, 15 and 24 m). Four MAAs were identified: mycosporine-glycine, shinorine, porphyra-334 and paly-thine. Concentrations of MAAs among replicate individuals varied considerably. Ovaries had high concentrations of MAAs (84 to 1389 μg g⁻¹ dry wt), while testes had non-detectable levels. The relative abundance of specific MAAs in ovaries appeared to be related to the spawning cycle. Digestive-tract samples had MAA concentrations as high as 3000 μg g⁻¹ dry wt, but the mean MAA content in intertidal individuals decreased by 70% over 3 mo during spring. The body walls of sea urchins had very low amounts of MAAs (≤ 0.08 μg g⁻¹ dry wt). There were significant depth differences in the␣total MAA content of the ovary ( p <0.001), ( p <0.015), digestive tract ( p <0.001), and body wall with organisms from the intertidal and 8 m depth having the highest concentrations of MAAs. Biological dosimetry indicated that UV-B (280 to 320 nm) wavelengths penetrated 3 to 7 m below the sea ice during the study period. The total MAA content in ovaries decreased with depth on all sample dates; however, the MAA content of the digestive tract and body wall did not exhibit a consistent pattern of change with depth. The MAA content of tissues did not change significantly with the temporal gradient of light exposure that was established by both ozone depletion and increasing photoperiod, except in the digestive tract sampled from intertidal specimens. Adult urchins are probably well-protected from UV exposure by the water column and a calcareous test; however, the results of this study suggest that, even under ice cover, depth of habitation is a determinant of MAA content in S. neumayeri. Large daily and seasonal fluctuations in the light regime, which are characteristic of Antarctic coastal environments, apparently do not provide reliable cues to elicit a detectable, temporal, biochemical response. Received: 19 February 1997 / Accepted: 26 March 1997     

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     

Effect of temperature fluctuations and food supply on the growth and metabolism of juvenile sea scallops (Placopecten magellanicus)

On the eastern shore of Nova Scotia late summer atmospheric systems cause upwelling of shelf water; the associated temperature variations of 10 °C with a 6 to 8 d period are comparable in magnitude to the seasonal variation. A laboratory study was undertaken to assess the effects of these temperature fluctuations on sea scallop (Placopecten magellanicus) growth and metabolism. In a factorial design, scallops were subjected to constant (10 °C) or a variable (6 to 15 °C) 8 d temperature cycle, and either a low (seston in filtered seawater) or high (seston supplemented with cultured phytoplankton) food diet. During the 48 d experiment scallop mortality was low and growth positive in all treatments. Shell and total tissue growth rate did not differ between temperature treatments, but growth in the high food treatments was 40 to 50% higher than in the low food treatments. However, soft tissue (excluding adductor) growth did show a temperature treatment effect; growth rates were significantly higher in the fluctuating temperature treatment, due in part to greater gonad development. Weight-standardized rates of scallop oxygen consumption (V _sO₂ , μmol O₂ g⁻¹ h⁻¹) were 20 to 25% higher in high food than in low food treatments, consistent with the expected increase in respiration due to the higher growth rates. Scallop metabolism did not acclimate to the fluctuating temperature cycle; V _sO₂ and ammonium excretion (V _sNH⁺ ₄, μmol O₂ g⁻¹ h⁻¹) remained dependent on ambient temperature throughout the experiment. V _sNH⁺ ₄ Q₁₀ (2.77) was higher than V _sO₂ Q₁₀ (2.01) which was reflected in a decrease in the O:N ratio at 15 °C, indicating a shift toward increased protein catabolism and a stressed state. At 10 °C, V _sO₂ and V _sNH⁺ ₄ in the variable temperature treatments were 15 to 18% lower than in the constant temperature treatments, a difference that was not detected in growth measurements. Results demonstrate that the metabolism of Placopecten magellanicus, unlike some bivalve species, is tightly coupled to fluctuations in ambient temperature. Although an absence of compensatory acclimation had a minimal effect on growth in this study, if high temperatures were combined with low food conditions a reduction in scallop production could result. Received: 23 June 1998 / Accepted: 8 February 1999