Light climate and plankton in the deep chlorophyll maxima in North Patagonian Andean lakes

The light climate at the deep chlorophyll maxima (DCM) was analysedin a set of lakes of the North Patagonian Andean region. Apparentand inherent optical properties in relation to the chlorophylla vertical distribution were investigated in seven lakes includingdeep (Z_max > 90 m) and shallow (Z_max < 12 m) ones. Samplingwas carried out during the thermal stratification period (summer)of the deep lakes since in the shallower lakes no stable thermalstratification was detected. The large deep lakes presentedvery low diffuse attenuation coefficients of photosyntheticallyactive radiation (K_d PAR), and a DCM situated at 0.98–2%of surface PAR irradiance, coinciding with the maximum abundanceof the mixotrophic ciliate Ophrydium naumanni and autotrophicpicoplankton. Both fractions seemed to be favoured by dim lightconditions of particular wavelengths, since at these DCM layersmainly green and blue wavelengths prevailed (<600 nm). Incontrast, shallow lakes showed higher K_d PAR values, with ahigher concentration of dissolved yellow substances, which causedsubstantial differences in the spectral quality that may havecontributed to explain the absence of this ciliate populationin these lakes.     

Symbiotic association of the ciliate Ophrydium naumanni with Chlorella causing a deep chlorophyll a maximum in an oligotrophic South Andes lake

Vertical profiles of temperature, light and chlorophyll a concentration were examined in Lake Moreno Oeste, an oligotrophic South Andean lake (Argentina), during the warmest period of the year (November-April), when thermal stratification is characteristic. Concurrent samples for the enumeration of phytoplankton and green ciliates were taken, and the different contribution of these fractions to total chlorophyll a concentration was analysed. The development of a distinctive deep chlorophyll maximum was observed during summer months. The deep chlorophyll maximum was situated near the limit of the euphotic zone and just below the upper limit of the metalimnion. The results showed that the green ciliate Ophrydium naumanni with endosymbiotic Chlorella dominated the metalimnion causing the deep chlorophyll maximum. Additional laboratory experiments revealed a strong dependence of O.naumanni on light. Therefore, the symbiotic association appears to be an effective exploitation of the water column in poor-nutrient-high-light ecosystems like large Andean lakes.      

Feeding of Boeckella gracilipes (Copepoda, Calanoida) on ciliates and phytoflagellates in an ultraoligotrophic Andean lake

The calanoid copepod, Boeckella gracilipes, is the dominantcrustacean zooplankton in South Andean deep ultra-oligotrophiclakes. Combining field and experimental data we explored thefeeding of the copepod and its access to the mixotrophic ciliate,Ophrydium naumanni, in Lake Moreno Oeste (Patagonia, Argentina).Phytoplankton was dominated by nanoflagellates throughout thewater column. Ophrydium naumanni, which accumulates much ofthe chlorophyll a, as do copepodites and adults of B. gracilipes,has a deep distribution during the day, with maximal abundancesaround 30 m depth. Mouth-part morphology analysis of B. gracilipesindicated that the copepod has an omnivorous diet. Laboratoryexperiments showed that B. gracilipes could access O. naumannionly when it is offered as a single food item. However, whennatural phytoplankton and ciliate assemblages (including O.naumanni)are offered, B. gracilipes did not eat Ophrydium and preyedon the oligotrich, Strombidium viride, and phytoflagellateslike Chrysochromulina parva. The range of ingested sizes wasbroad (3.9–33 µm of equivalent spherical diameter)but all selected particles were motile ones with distinctivemovements, which would enhance the copepod particle detection.     

Growth rates of phytoplankton under fluctuating light

• 1 The effect of light fluctuations on the growth rates of four species of freshwater phytoplankton was investigated. Experimental light regimes included constant irradiance and fluctuations of a step function form, with equal proportion of high (maximum of 240 µmol photons m^‐2 s^‐1) and low light (minimum of 5 µmol photons m^‐2 s^‐1) (or dark) in a period. Fluctuations of 1, 8 and 24‐h periods were imposed over several average irradiances (25, 50, 100 and 120 µmol photons m^‐2 s^‐1).
• 2 Growth rate responses to fluctuations were species‐specific and depended on both the average irradiance and the period of fluctuations. Fluctuations at low average irradiances slightly increased growth rate of the diatom Nitzschia sp. and depressed growth of the cyanobacterium Phormidium luridum and the green alga Sphaerocystis schroeteri compared to a constant irradiance.
• 3 Fluctuations at higher average irradiance did not have a significant effect on the growth rates of Nitzschia sp. and Sphaerocystis schroeteri (fluctuations around saturating irradiances) and slightly increased the growth rates of the cyanobacteria Anabaena flos‐aquae and Phormidium luridum (when irradiance fluctuated between limiting and inhibiting levels).
• 4 In general, the effect of fluctuations tended to be greater when irradiance fluctuated between limiting and saturating or inhibiting levels of a species growth‐irradiance curve compared to fluctuations within a single region of the curve.
• 5 The growth rates of species under fluctuating light could not always be predicted from their growth‐irradiance curves obtained under constant irradiance. When fluctuations occur between limiting and saturating or inhibiting irradiances for the alga and when the period of fluctuations is long (greater than 8 h), steady‐state growth‐irradiance curves may be insufficient to predict growth rates adequately. Consequently, additional data on physiological acclimation, such as changes in photosynthetic parameters, may be required for predictions under non‐constant light supply in comparison to constant conditions.

     

Mixotrophic ciliates in an Andean lake: dependence on light and prey of an Ophrydium naumanni population

1. Planktonic ciliates were examined during a spring–summer period (November 1998–April 1999) in the ultraoligotrophic Lake Moreno Oeste (41°5' S and 71°33' W, 758 m a.s.l), which belongs to the Nahuel Huapi System (Patagonia, Argentina). The lake is deep ( Z _max=90 m) and warm monomictic.
2. Sampling was performed at a mid-lake station, where vertical profiles of temperature and light were measured in situ , and samples for bacteria and ciliates enumeration were taken throughout the water column.
3. The peritrich Ophrydium naumanni , a freshwater pelagic ciliate with endosymbiotic Chlorella , was the dominant ciliate in the lake.
4.  Ophrydium naumanni and autotrophic picoplankton exhibit a clear coincidence in their vertical distribution ( P  < 0.05), preferring levels at or near the 1% of surface photosynthetically active radiation (PAR) irradiance. Both may have the same light requirements, or the coincidence may reflect a trophic relationship.
5. Dependence on light and prey by O. naumanni were studied using field experiments, in which we analysed ciliate grazing on bacteria, and in laboratory experiments, in which we compared particle uptake under dark and light conditions.
6.  Ophrydium naumanni was able to ingest particles [latex microspheres and fluorescently labelled bacteria (FLB)] in field and laboratory experiment, indicating that it has the potential to affect bacteria population of Lake Moreno Oeste.
7. Ciliate particle ingestion was observed to be dependent on light availability because under dark conditions, the ingestion was lowered ( P  < 0.05).     

The influence of nitrogen on rain forest dipterocarp seedlings exposed to a large increase in irradiance

Dipterocarps dominate the canopy of lowland tropical rain forest in South‐east Asia. Seedlings of these species form diverse assemblages on the forest floor where low irradiance severely limits their growth. Further growth depends largely upon the increased irradiance that can occur with the creation of canopy gaps. However, the response of dipterocarp seedlings to increased irradiance and their subsequent establishment in the canopy may be influenced by the availability of other resources, such as nutrient availability. We investigated the influence of nitrogen supply on aspects of the photosynthetic physiology and growth of seedlings of four dipterocarp species (Shorea leprosula, Shorea johorensis, Shorea oleosa and Dryobalanops lanceolata) growing under low irradiance, during transfer from low to high irradiance, and during subsequent growth at high irradiance. All four species increased growth and photosynthetic capacity in response to N‐supply at high irradiances but not at low irradiance approximating that which can be expected to occur in the forest understorey. When seedlings grown at low irradiances and varying N‐supply were exposed to a large increase in irradiance, all species showed some degree of initial photodamage (measured through chlorophyll fluorescence), the extent of which was similar between species but differed markedly depending on the pre‐exposure growth irradiance and N‐supply. Greater photodamage occurred in seedlings grown at lower compared with higher N‐supply and irradiance. Despite these initial difference in the extent of this photodamage, all seedlings demonstrated a similar capacity to recover from damage. However, the alterations in the photosynthetic physiology of leaves during this recovery differed between species and depended on N‐supply. Under high N‐supply all species apart from S. oleosa increased photosynthetic capacity per unit chlorophyll following exposure to high irradiance by increasing photosynthetic capacity per unit leaf area while, under low N‐supply, an increase in photosynthetic capacity per unit leaf only occurred in D. lanceolata. Our results suggest that variations in N‐availability may have a much greater impact on the relative competitiveness of dipterocarp seedlings during the regenerative phase following canopy gap formation than physiological differences between seedlings. Our results demonstrate a potentially significant role for N‐availability in the regeneration dynamics and distribution of canopy‐dominating dipterocarp species.     

THE EFFECT OF PHOTON IRRADIANCE ON THE BEHAVIORAL ECOLOGY AND POTENTIAL NICHE SEPARATION OF FRESHWATER PHYTOPLANKTONIC FLAGELLATES1

Photoresponsive behavior in phytoplanktonic flagellates potentially controls depth regulation, vertical migration, and the accumulation of cells in preferred conditions and hence has major implications for photosynthesis, growth, and species competition. This study used microscale laboratory chambers with cell track analysis and mesoscale lake studies to investigate the responses of five phylogenetically contrasting freshwater flagellates to gradients of light. Laboratory results demonstrated that these species differed in their light preferences despite being grown at the same photon irradiance. Preferred photon irradiances were 20–120, 20, 65–120, 4–20, and>200 μmol photons·m^?2·s^?1, respectively, for Ceratium furcoides (Levander) Langhans, Chlamydomonas moewusii Gerloff, Dinobryon sertularia Ehrenberg, Euglena gracilis Klebs, and Plagioselmis nannoplanctica (Skuja) Novarino. Analysis of the response of individual cells showed that in all species, photoresponsive preference was the result of positive and negative phototaxis combined with step‐up and step‐down photophobic reactions. There was no evidence for photokinesis or cell memory. Only in C. furcoides did the preferred photon irradiance alter with growth conditions. In C. furcoides, D. sertularia, and P. nannoplanctica, irradiance preference matched the optimal irradiance for growth, whereas in the two remaining species a lower than optimal irradiance was preferred, suggesting that light may be used as an indicator of other ecological conditions. Mesoscale experiments in a lake demonstrated that the laboratory microscale measurements provide information relevant to understanding ecological distributions. Behavioral responses to light contribute to the delineation of vertical niche separation and provide a method for predicting the spatial and temporal distribution of flagellates.     

Effects of temperature, irradiance and photoperiod on growth and pigment content in some freshwater red algae in culture

The responses of relative growth rate (% day‐1) and pigment content (chlorophyll a, phycocyanin and phycoerythrin) to temperature, irradiance and photoperiod were analyzed in culture in seven freshwater red algae: Audouinella hermannii (Roth) Duby, Audouinella pygmaea (Kützing) Weber‐van Bosse, Batrachospermum ambiguum Montagne, Batrachospermum delicatulum (Skuja) Necchi et Entwisle,‘Chantransia’ stages of B. delicatulum and Batrachospermum macrosporum Montagne and Compsopogon coeruleus (C. Agardh) Montagne. Experimental conditions included temperatures of 10, 15, 20 and 25°C and low and high irradiances (65 and 300 μmol photons m^?2 s^?1, respectively). Long and short day lengths (16:8 and 8:16 LD cycles) were also applied at the two irradiances. Growth effects of temperature and irradiance were evident in most algae tested, and there were significant interactions among treatments. Most freshwater red algae had the best growth under low irradiance, confirming the preference of freshwater red algae for low light regimens. In general there was highest growth rate in long days and low irradiance. Growth optima in relation to temperature were species‐specific and also varied between low and high irradiances for the same alga. The most significant differences in pigment content were related to temperature, whereas few significant differences could be attributed to variation in irradiance and photoperiod or interactions among the three parameters. The responses were species‐specific and also differed for pigments in distinct temperatures, irradiances and photoperiods in the same alga. Phycocyanin was generally more concentrated than phycoerythrin and phycobiliproteins were more concentrated than chlorophyll a. The highest total pigment contents were found in two species typical of shaded habitats: A. hermannii and C. coeruleus. The expected inverse relationship of pigment with irradiance was observed only in C. coeruleus. In general, the most favorable conditions for growth were not coincident with those with highest pigment contents.     

Ecophysiology of photosynthesis in bryophytes: major roles for oxygen photoreduction and non‐photochemical quenching?

CO₂ fixation in mosses saturates at moderate irradiances. Relative electron transport rate (RETR) inferred from chlorophyll fluorescence saturates at similar irradiance in shade species (e.g. Plagiomnium undulatum, Trichocolea tomentella), but many species of unshaded habitats (e.g. Andreaea rothii, Schistidium apocarpum, Sphagnum spp. and Frullania dilatata) show non‐saturating RETR at high irradiance, with high non‐photochemical quenching (NPQ). In P. undulatum and S. apocarpum, experiments in different gas mixtures showed O₂ and CO₂ as interchangeable electron sinks. Nitrogen + saturating CO₂ gave high RETR and depressed NPQ. In S. apocarpum, glycolaldehyde (inhibiting photosynthesis and photorespiration) depressed RETR in air more at low than at high irradiance; in CO₂‐free air RETR was maintained at all irradiances. Non‐saturating electron flow was not suppressed in ambient CO₂ with 1% O₂. The results indicate high capacity for oxygen photoreduction when CO₂ assimilation is limited. Non‐saturating light‐dependent H₂O₂ production, insensitive to glycolaldehyde, suggests that electron transport is supported by oxygen photoreduction, perhaps via the Mehler‐peroxidase reaction. Consistent with this, mosses were highly tolerant to paraquat, which generates superoxide at photosystem I (PSI). Protection against excess excitation energy in mosses involves high capacity for photosynthetic electron transport to oxygen and high NPQ, activated at high irradiance, alongside high reactive oxygen species (ROS) tolerance.     

EFFECTS OF TEMPERATURE AND IRRADIANCE ON THE SEASONAL VARIATION OF A SPIROGYRA (CHLOROPHYTA) POPULATION IN A MIDWESTERN LAKE (U.S.A.)

Although Spirogyra Link (1820) is a common mat‐forming filamentous alga in fresh waters, little is known of its ecology. A 2‐year field study in Surrey Lake, Indiana, showed that it grew primarily in the spring of each year. The population consisted of four morphologically distinct filamentous forms, each exhibiting its own seasonal distribution. A 45‐μm‐wide filament was present from February to late April or early May, a 70‐μm‐wide form was present from late April to mid‐June, a 100‐μm‐wide form was present from February to mid‐June, and a 130‐μm‐wide form appeared only in February of 1 of 2 study years. The 70‐ and 100‐μm‐wide forms contributed to the peak amount of biomass observed in late May and early June. Multiple regression analysis indicated that the presence of the 45‐, 70‐, and 100‐μm‐wide forms was negatively correlated with temperature. Presence of the 130‐μm‐wide form was negatively correlated with irradiance. Isolates of these filament forms were exposed to temperature (15, 25, and 35° C)/irradiance (0, 60, 200, 400, 900, and 1500 μmol·m^?2·s^?1) combinations in the laboratory. Growth rates of the 45‐μm‐wide form were negative at all irradiances at 35° C, suggesting that this form is susceptible to high water temperatures. However, growth rates of the other forms did not vary at the different temperatures or at irradiances of 60 μmol·m^?2·s^?1 or above. Net photosynthesis was negative at 35° C and 1500 μmol·m^?2·s^?1 for the 100‐ and 130‐μm‐wide forms but positive for the 70‐μm‐wide form. All forms lost mat cohesiveness in the dark, and the 100‐ and 130‐μm‐wide forms lost mat cohesiveness under high irradiances and temperature. Thus, the morphological forms differed in their responses to irradiance and temperature. We hypothesize that the rapid disappearance of Spirogyra populations in the field is due to loss of mat cohesiveness under conditions of reduced net photosynthesis, for example, at no to low light for all forms or at high light and high temperatures for the 100‐ and 130‐μm‐wide forms. Low light conditions can occur in the interior of mats as they grow and thicken or under shade produced by other algae.     

ADAPTATION OF CERATIUM FURCA AND GONYAULAX POLYEDRA (DINOPHYCEACE) TO DIFFERENT TEMPERATURES AND IRRADIANCES: GROWTH RATES AND CELL VOLUMES1

Growth rates and cell volumes of Ceratium furca Ehrenberg and Gonyaulax polyedra Stein were determined during the log phase of growth in cultures which had been extensively adapted to one of three temperatures and five irradiances. At each temperature, curves for the growth rate vs. irradiance for both species had light-limited and light-saturated regions. Three properties of these curves characterized the response of each species to temperature: the light-saturated growth rate, the irradiance at which growth became light-saturated and the compensation irradiance for growth. For both species, the first two properties generally decreased with declining growth temperature, while the compensation irradiance declined for Ceratium but had a V-shaped response pattern for Gonyaulax. The light-saturated growth rates were generally higher for Ceratium than for Gonyaulax, while the irradiance at which growth became saturated and the compensation irradiance were lower for Ceratium. The changes in cell volume associated with the irradiance and temperature of growth were very different for Ceratium and Gonyaulax. The cell size of Gonyaulax increased as irradiance and temperature decreased, while cell volumes of Ceratium did not change with temperature but were smallest at the highest and lowest growth irradiances. In general, the growth rate patterns were similar for Ceratium and Gonyaulax, while those for cell size were different. The maximum growth rate, the irradiance at which growth became saturated, the compensation irradiance, and the cell volume all showed that Ceratium grew at the same rate or faster than Gonyaulax over the entire range of irradiances and temperatures examined.     

UV-B irradiance gradient affects photosynthesis and pigments but not food quality of periphyton

• 1 This laboratory study examined the effect of a gradient of UV‐B radiation (280–320 nm) on photosynthesis and food quality of periphyton, the trophic base of many freshwater benthic communities. Four irradiances of UV‐B (0, 0.6, 1.2, and 2.3 W m^‐2) were delivered by UV‐B lamps (313 nm peak irradiance) over a 13‐day period in the first experiment and over a 4‐h period in the second experiment. These irradiances were roughly equivalent to 0, 1, 2, and 4 times the ambient biologically effective (DNA) midsummer, midday UV‐B irradiance in Tennessee.
• 2 Rates of photosynthesis and photosynthetic pigments were significantly reduced by irradiances greater than ambient during the 13‐day experiment, suggesting that food supply rates to grazers would be depressed by increases in current UV‐B levels. Effects on community structure were minor, but mean diatom cell size decreased at higher UV‐B irradiances.
• 3 Irradiated periphyton was fed in surplus to juvenile snails (Physella gyrina) in the first experiment as a bioassay for food quality. Snail growth was the same on all four diets, suggesting that UV‐B did not affect food quality. Nitrogen and phosphorus content of the periphyton were not affected by UV‐B, either.
• 4 Photosynthesis by low‐biomass periphyton in the second experiment was significantly depressed by irradiances above ambient after only 4 h. Photosynthesis by the high biomass periphyton was not significantly affected by UV‐B, suggesting that self‐shading reduced UV‐B effects.

     

Ontogenetic Shifts in the Ability of the Cladoceran,Moina macrocopa Straus and Ceriodaphnia cornuta Sars to Utilize Ciliated Protists as Food Source

The ontogenetic diet shifts and age specific ability of the two cladoceran species Moina macrocopa and Ceriodaphnia cornuta to derive energy from ciliated protists have been investigated in laboratory. The postembryonic developmental rates and life table demography (longevity, age and size at first reproduction, fecundity and intrinsic rate of natural increase) of the cladocerans have been elucidated on algae (Chlorella vulgaris) and the ciliated protists (Tetrahymena pyriformis, Colpoda (c.f.) steini) as food. For either of the cladoceran, the somatic growth rate and average body size at first reproduction were higher with algal diet. During initial stages of development (0–5 days), either cladoceran realized higher rate of somatic growth on algal diet, subsequently ciliated protists supported significantly higher growth rate than the alga. Algal and ciliate diets did not differ in maximum body size (C. cornuta: 539–554 μm; M. macrocopa: 1274.8–1309 μm) reached by either of the cladocerans. The maximum body sizes were larger than size at first reproduction with either of the ciliated protists, however, with algal diet the maximum body sizes did not differ from the size at first reproduction in each case. In case of C. cornuta the generation time (20.5 ± 0.3 days on ciliate; 15.6 ± 0.17 days on algal diet), reproductive rates (net reproductive rate: 20.05 ± 3.2 on ciliate; 15.5 ± 1.2 on algal diet), and average life expectancy at hatching (27 ± 0.8 days on ciliate; 22.7 ± 0.71 days on alga) were higher, whereas the size at first reproduction (482 μm on ciliate; 521 μm on alga) was smaller with the ciliate than with an algal diet. The algal and the ciliate diets did not differ in survival (life expectancy at hatching: 9.2 ± 0.7 days) and fecundity (NRR: 23.6 ± 2.4) for M. macrocopa. The two ciliates used in the experiment did not differ in their performance as food source for either cladoceran species. Our results suggest that both the cladoceran species are able to utilize smaller ciliate (e.g., T. pyriformis, C. (c.f.) steini) as food; however with differential ability to derive energy from the ciliate diet and this ability is size and age structured in both cases. (© 2008 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Macrozooplankton and the persistence of the deep chlorophyll maximum in a stratified lake

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Photoheterotrophy and light-dependent uptake of organic and organic nitrogenous compounds by Planktothrix rubescens under low irradiance

1. Planktothrix rubescens is the dominant photoautotrophic organism in Lake Zürich, a prealpine, deep, mesotrophic freshwater lake with an oxic hypolimnion. Over long periods of the year, P. rubescens accumulates at the metalimnion and growth occurs in situ at irradiance near the photosynthesis compensation point. Experiments were conducted to evaluate the contribution of photoheterotrophy, heterotrophy and light‐dependent uptake of nitrogenous organic compounds to the carbon and nitrogen budget of this cyanobacterium under conditions of restricted availability of light quanta. 2. We used both purified natural populations of P. rubescens from the depth of 9 m and an axenic culture grown under low irradiance at 11 μmol m^?2 s^?1 on a light : dark cycle (10 : 14 h) to determine the uptake rates of various amino acids, urea, glucose, fructose, acetate and inorganic carbon. The components were added to artificial lake water in low amounts that simulated the naturally occurring potential concentrations. 3. The uptake rates of acetate and amino acids (glycine, serine, glutamate and aspartate) were strongly enhanced at low irradiance as compared with the dark. However, no difference was observed in the uptake of arginine, which was taken up at high rates under both treatments. The uptake rates of glucose, fructose and urea were very low under all conditions. Similar results were obtained for both axenic P. rubescens and for purified natural populations of P. rubescens that were separated from bacterioplankton and other phytoplankton. 4. Metalimnetic P. rubescens that was stratified at low irradiance for weeks exhibited much higher uptake rates than filaments that were entrained in the deepening surface mixed layer and experienced higher irradiance. The added organic compounds contributed up to 62% to the total carbon uptake of metalimnetic P. rubescens. On the basis of a molar C : N ratio of 4.9, the nitrogen uptake as organic compounds satisfied up to 84% of the nitrogen demand. 5. The experiments indicate that photoheterotrophy and light‐dependent uptake of nitrogenous organic compounds may contribute significantly to the carbon and nitrogen budget of filaments at low irradiance typical for growth of P. rubescens in the metalimnion and at the bottom of the surface mixed layer.     

IMPORTANCE OF MACRO‐ VERSUS MICROSTRUCTURE IN MODULATING LIGHT LEVELS INSIDE CORAL COLONIES1

Adjusting the light exposure and capture of their symbiotic photosynthetic dinoflagellates (genus Symbiodinium Freud.) is central to the success of reef‐building corals (order Scleractinia) across high spatio‐temporal variation in the light environment of coral reefs. We tested the hypothesis that optical properties of tissues in some coral species can provide light management at the tissue scale comparable to light modulation by colony architecture in other species. We compared within‐tissue scalar irradiance in two coral species from the same light habitat but with contrasting colony growth forms: branching Stylophora pistillata and massive Lobophyllia corymbosa. Scalar irradiance at the level of the symbionts (2 mm into the coral tissues) were <10% of ambient irradiance and nearly identical for the two species, despite substantially different light environments at the tissue surface. In S. pistillata, light attenuation (90% relative to ambient) was observed predominantly at the colony level as a result of branch‐to‐branch self‐shading, while in L. corymbosa, near‐complete light attenuation (97% relative to ambient) was occurring due to tissue optical properties. The latter could be explained partly by differences in photosynthetic pigment content in the symbiont cells and pigmentation in the coral host tissue. Our results demonstrate that different strategies of light modulation at colony, polyp, and cellular levels by contrasting morphologies are equally effective in achieving favorable irradiances at the level of coral photosymbionts.     

PRIMARY PRODUCTION OF CRUSTOSE CORALLINE RED ALGAE IN A HIGH ARCTIC FJORD1

Crustose coralline algae occupied ～1%–2% (occasionally up to 7%) of the sea floor within their depth range of 15–50 m, and they were the dominant encrusting organisms and macroalgae beyond 20 m depth in Young Sound, NE Greenland. In the laboratory, oxygen microelectrodes were used to measure net photosynthesis (P) versus downwelling irradiance (E_d) and season for the two dominant corallines [Phymatolithon foecundum (Kjellman) Düwel et Wegeberg 1996 and Phymatolithon tenue (Rosenvinge) Düwel et Wegeberg 1996] representing> 90% of coralline cover. Differences in P‐E_d curves between the two species, the ice‐covered and open‐water seasons, or between specimens from 17 and 36 m depth were insignificant. The corallines were low light adapted, with compensation irradiances (E_c) averaging 0.7–1.8 μmol photons·m ^{? 2}·s ^{? 1} and light adaptation (E_k) indices averaging 7–17 μmol photons·m ^{? 2}·s ^{? 1}. Slight photoinhibition was evident in most plants at irradiances up to 160 μmol photons·m ^{? 2}·s ^{? 1}. Photosynthetic capacity (P_m) was low, averaging 43–67 mmol O₂·m ^{? 2} thallus·d ^{? 1} (～250–400 g C·m ^{? 2} thallus·yr ^{? 1}). Dark respiration rates averaged ～5 mmol O₂·m ^{? 2} thallus·d ^{? 1}. In ice covered periods, E_d at 20 m depth averaged ～1 μmol photons·m ^{? 2}·s ^{? 1}, with daily maxima of 2–3 μmol photons·m ^{? 2}·s ^{? 1}. During the open water season, E_d at 20 m depth averaged ～7 μmol photons·m ^{? 2}·s ^{? 1} with daily maxima of ～30 μmol photons·m ^{? 2}·s ^{? 1}. Significant net primary production of corallines was apparently limited to the 2–3 months with open water, and the small contribution of corallines to primary production seems due to low P_m values, low in situ irradiance, and their relatively low abundance in Young Sound.     

Effects of riparian leaf dynamics on periphyton photosynthesis and light utilisation efficiency

1. Streambed light regimes change dramatically when riparian trees gain leaves in spring and lose them in autumn. This study examined the effect of these changes on periphyton photosynthetic characteristics, primary production, and light utilisation efficiency in two eastern Tennessee streams. 2. Photosynthesis–irradiance responses were measured at intervals covering leaf emergence and abscission in spring and autumn. Photosynthetic efficiency (α^chl) increased with declining streambed irradiances during spring leaf emergence, but returned to pre‐emergence values after autumn leaf fall. The onset of photosaturation (I_k) displayed the opposite pattern, decreasing during leaf emergence and increasing after leaf fall. Both α^chl and I_k were closely associated (P < 0.01) with daily integrated streambed irradiance, as were periphyton carotenoids. Internal shading by photoprotective carotenoids is hypothesised to account for lower α^chl when streambed irradiances are high. 3. An in situ shading experiment confirmed that the temporal changes observed in periphyton photosynthetic characteristics and carotenoids were primarily the result of changing light levels and not other environmental factors (e.g. nutrients, temperature). 4. Daily chlorophyll‐specific primary production (PP^chl) was calculated with P–I models and recorded streambed irradiances. In both streams, PP^chl was the highest in early spring when trees were leafless, and then declined markedly as leaves emerged, reaching a minimum in summer. PP^chl increased after leaf abscission, but was still lower than it was in early spring, when the sun was higher and daylength was longer. A hyperbolic tangent equation fit to PP^chl and daily integrated irradiance (r²=0. 85) suggested that primary production was light saturated at 4–8 mol m^–2 d^–1. 5. Light utilisation efficiency (Ψ) increased 10‐fold during leaf emergence. Photosaturation at high irradiances and photoacclimation at lower irradiances were responsible for a negative hyperbolic relationship between Ψ and daily integrated irradiance.     

Abundance and primary production of filamentous green algae Zygogonium ericetorum in an extremely acid (pH 2.9) mining lake and its impact on alkalinity generation

1. In extremely acid mining lakes, benthic filamentous green algae (Zygnemataceae, Chlorophyta) thrive as effective competitors for limited carbon (C). These algae could supply C for microbial‐mediated benthic alkalinity generation. However, biomass, productivity and impact of the acidobiontic filamentous green algae at pH ≤3 have not previously been determined. 2. Periphytic filamentous green algae was mapped by harvesting their biomass from 85 1 × 1 m quadrats in mining lake Grünewalder Lauch. Zygogonium ericetorum colonised water depths between 1.6 and 10.5 m covering 88% of total area. Biomass peaked at 5–6 m depth. Total Zygogonium biomass amounted to 72.2 t dry weight for the whole lake (0.94 km²), which corresponds to 16.1 t C and the accumulation of primary production from 2.2 years. 3. Growth of Zygogonium is moderately N, C and extremely P deficient, and seriously stressed by high rates of Fe deposition during summer. Consequently, net primary production (NPP) of Zygogonium, calculated from measured photosynthesis versus irradiance characteristics and calculated underwater irradiance (0.13 g C m^?2 year^?1) and in situ oxygen measurements (7.8 g C m^?2 year^?1), corresponds to only 0.3% and 18.1% of pelagic NPP. 4. Neither pelagic nor benthic Zygogonium primary production can supply enough C for efficient acidity removal. However, at rates of benthic NPP in summer of 21.4 mg C m^?2 day^?1, Zygogonium contributed 26% of the C equivalents to remove acidity associated with ferric iron, contributing at least seasonally to efficient alkalinity generation.     

Phenology,irradiance and temperature characteristics of a freshwater red alga,Nemalionopsis tortuosa (Thoreales), from Kagoshima,southern Japan

Phenology, irradiance and temperature characteristics of a freshwater benthic red alga, Nemalionopsis tortuosa Yoneda et Yagi (Thoreales), were examined from Kagoshima Prefecture, southern Japan for the conservation of this endemic and endangered species. Field surveys confirmed that algae occurred in shaded habitats from winter to early summer, and disappeared during August through November. A net photosynthesis–irradiance (P–E) model revealed that net photosynthetic rate quickly increased and saturated at low irradiances, where the saturating irradiance (E_k) and compensation irradiance (E_c) were 10 (8–12, 95% credible interval (CRI)) and 8 (6–10, 95% CRI) μmol photon m^?2 s^?1, respectively. Gross photosynthesis and dark respiration was determined over a range of temperatures (8–36°C) by dissolved oxygen measurements, and revealed that the maximum gross photosynthetic rate was highest at 29.5 (27.4–32.0, 95%CRI) °C. Dark respiration also increased linearly when temperature increased from 8°C to 36°C, indicating that the increase in dark respiration at higher temperature most likely caused decreases in net photosynthesis. The maximum quantum yield (Fv/Fm) that was determined using a pulse amplitude modulated‐chlorophyll fluorometer (Imaging‐PAM) was estimated to be 0.51 (0.50–0.52, 95%CRI) and occurred at an optimal temperature of 21.7 (20.1–23.4, 95%CRI) °C. This species can be considered well‐adapted to the relatively low natural irradiance and temperature conditions of the shaded habitat examined in this study. Our findings can be applied to aid in the creation of a nature‐reserve to protect this species.