PHOTOACCLIMATION OF PHOTOSYNTHESIS IRRADIANCE RESPONSE CURVES AND PHOTOSYNTHETIC PIGMENTS IN MICROALGAE AND CYANOBACTERIA1

The photosynthesis‐irradiance response (PE) curve, in which mass‐specific photosynthetic rates are plotted versus irradiance, is commonly used to characterize photoacclimation. The interpretation of PE curves depends critically on the currency in which mass is expressed. Normalizing the light‐limited rate to chl a yields the chl a‐specific initial slope (α^chl). This is proportional to the light absorption coefficient (a^chl), the proportionality factor being the photon efficiency of photosynthesis (φ_m). Thus, α^chl is the product of a^chl and φ_m. In microalgae α^chl typically shows little (<20%) phenotypic variability because declines of φ_m under conditions of high‐light stress are accompanied by increases of a^chl. The variation of α^chl among species is dominated by changes in a^chl due to differences in pigment complement and pigment packaging. In contrast to the microalgae, α^chl declines as irradiance increases in the cyanobacteria where phycobiliproteins dominate light absorption because of plasticity in the phycobiliprotein:chl a ratio. By definition, light‐saturated photosynthesis (P_m) is limited by a factor other than the rate of light absorption. Normalizing P_m to organic carbon concentration to obtain P_m^C allows a direct comparison with growth rates. Within species, P_m^C is independent of growth irradiance. Among species, P_m^C covaries with the resource‐saturated growth rate. The chl a:C ratio is a key physiological variable because the appropriate currencies for normalizing light‐limited and light‐saturated photosynthetic rates are, respectively, chl a and carbon. Typically, chl a:C is reduced to about 40% of its maximum value at an irradiance that supports 50% of the species‐specific maximum growth rate and light‐harvesting accessory pigments show similar or greater declines. In the steady state, this down‐regulation of pigment content prevents microalgae and cyanobacteria from maximizing photosynthetic rates throughout the light‐limited region for growth. The reason for down‐regulation of light harvesting, and therefore loss of potential photosynthetic gain at moderately limiting irradiances, is unknown. However, it is clear that maximizing the rate of photosynthetic carbon assimilation is not the only criterion governing photoacclimation.     

DIFFERENTIAL IMPACTS OF PHOTOACCLIMATION AND THERMAL STRESS ON THE PHOTOBIOLOGY OF FOUR DIFFERENT PHYLOTYPES OF SYMBIODINIUM (PYRRHOPHYTA)1

The capacity for photoacclimation to light at 100 or 600 μmol photons·m^?2·s^?1 and the subsequent response to thermal stress was examined in four genetically distinct cultures of symbiotic dinoflagellates in the genus Symbiodinium with the ITS2 designations A1, A1.1, B1, and F2. While all algal types showed typical signs of photoacclimation to high light via a reduction in chl a, there was a differential response in cellular growth, photosystem II (PSII) activity, and the chl a‐specific absorption coefficient between cultures. When maintained at 32°C for up to 10 days, significant variation in the susceptibility to thermal stress was observed in the rate of loss in PSII activity and electron transport, PSII reaction center degradation, and cellular growth. The order of thermal tolerance did not change between the two light levels. However, as expected, loss in photosynthetic function was exacerbated in the thermally sensitive phylotypes (B1 and A1.1) when acclimated to the higher light intensity. There was no consistent relationship between thermal tolerance and changes in light energy dissipation via non‐photochemical pathways. Phylotypes F2 and A1 showed a high degree of thermal tolerance, yet the cellular responses to light and temperature were markedly different between these algae. The F2 isolate showed the greatest capacity for photoacclimation and growth at high light and temperature, while the A1 isolate appeared to adjust to thermal stress by a slight decline in PSII activity and a significant decline in growth, possibly at the expense of increased photosystem and cellular repair rates.     

COMBINED EFFECTS OF TEMPERATURE AND IRON ON THE GROWTH AND PHYSIOLOGY OF THE MARINE DIATOM PHAEODACTYLUM TRICORNUTUM (BACILLARIOPHYCEAE)

Phaeodactylum tricornutum Bohlin (Bacillariophyceae) was maintained in exponential growth under Fe‐replete and stressed conditions over a range of temperatures from 5 to 30° C. The maximum growth rate (GR) was observed at 20° C (optimal temperature) for Fe‐replete and ‐stressed cells. There was a gradual decrease in the GR decreasing temperatures below the optimum temperature; however, the growth rate dropped sharply as temperature increased above the optimum temperature. Fe‐stressed cells grew at half the growth rate of Fe‐replete cells at 20° C, whereas this difference became larger at lower temperatures. The change in metabolic activities showed a similar pattern to the change in growth rate temperature aside from their optimum temperature. Nitrate reductase activity (NRA) and respiratory electron transport system activity (ETS) per cell were maximal between 15 and 20° C, whereas cell‐specific photosynthetic rate (P_cell) was maximal at 20° C for Fe‐replete cells. These metabolic activities were influenced by Fe deficiency, which is consistent with the theoretical prediction that these activities should have an Fe dependency. The degree of influence of Fe deficiency, however, was different for the four metabolic activities studied: NRA > P_cell > ETS = GR. NRA in Fe‐stressed cells was only 10% of that in Fe‐replete cells at the same temperature. These results suggest that cells would have different Fe requirements for each metabolic pathway or that the priority of Fe supply to each metabolic reaction is related to Fe nutrition. In contrast, the order of influence of decreasing the temperature from the optimum temperature was ETS > P_cell > NRA > GR. For NRA, the observed temperature dependency could not be accounted for by the temperature dependency of the enzyme reaction rate itself that was almost constant with temperature, suggesting that production of the enzyme would be temperature dependent. For ETS, both the enzyme reactivity and the amount of enzyme accounted for the dependency. This is the first report to demonstrate the combined effects of Fe and temperature on three important metabolic activities (NRA, P_cell, and ETS) and to determine which activity is affected the most by a shortage of Fe. Cellular composition was also influenced by Fe deficiency, showing lower chl a content in the Fe‐stressed cells. Chl a per cell volume decreased by 30% as temperature decreased from 20 to 10° C under Fe‐replete conditions, but chl a decreased by 50% from Fe‐replete to Fe‐stressed conditions.     

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

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

BIO-OPTICAL CHARACTERISTICS AND PHOTOADAPTIVE RESPONSES IN THE TOXIC AND BLOOM-FORMING DINOFLAGELLATES GYRODINIUM AUREOLUM,GYMNODINIUM GALATHEANUM,AND TWO STRAINS OF PROROCENTRUM MINIMUM1

Photoadaptive responses in the toxic and bloom-forming dinoflagellates Gyrodinium aureolum Hulbert, Gymnodinium galatheanum Braarud, and two strains of Prorocentrum minimum (Pavillard)Schiller were evaluated with respect to pigment composition, light-harvesting characteristics, carbon and nitrogen contents, and growth rates in shade- and light-adapted cells. The two former species were grown at scalar irradiances of 30 and 170 μmol · m ^?2 at a 12-h daylength at 20° C. The two strains of P. minimum were grown at 35 and 500 μmol. m^?2· s^?1 at a 2-h daylength at 20° C. For the first time, chlorophyll (chl) c₃, characteristic of several bloom-forming prymnesiophytes, was detected in G. aureolum and G. galatheanum. Photoadaptional status affected the pigment composition strongly, and the interpretation of the variation depended on whether the pigment composition was normalized per cell, carbon, or chl a. Species-specific and photoadaptional differences in chl a-specific absorption (°a_c, 400–700 nm) and chl a-normalized fluorescence excitation spectra of photosystem II fluorescence with or without addition of DCMU (°F and °F_DCMU 400–700 nm) were evident. Gyrodinium aureolum and G. galatheanum exhibited in vivo spectral characteristics similar to chl c₃-containing prymnesiophytes in accordance with their similar pigmentation. Prorocentrum minimum had in vivo absorption and fluorescence characteristics typical for peridinin-containing dinoflagellates. Species-specific differences in in vivo absorption were also observed as a function of package effect vs. growth irradiance. This effect could be explained by differences in intracellular pigment content, cell size/shape, and chloroplast morphology/numbers. Light- and shade-adapted cells of P. minimum contained 43 and 17% of photoprotective carotenoids (diadino + diatoxanthin) relative to chl a, respectively. The photoprotective function of these carotenoids was clearly observed as a reduction in °F and °F ^DCMU at 400–540 nm compared to °ac in light-adapted cells of P. minimum. Spectrally weighted light absorption (normalized to chl a and carbon, 400–700 nm) varied with species and growth conditions. The use of quantum-corrected and normalized fluorescence excitation spectra with or without DCMU-treated cells to estimate photosynthetically usable light is discussed. The usefulness of in vitro absorption and fluorescence excitation spectra for estimation of the degradation status of chl a and the ratio of chl a to total pigments is also discussed.     

PHOTOACCLIMATION IN THE PHOTOTROPHIC MARINE CILIATE MESODINIUM RUBRUM (CILIOPHORA)1

Mesodinium rubrum (=Myrionecta rubra), a marine ciliate, acquires plastids, mitochondria, and nuclei from cryptophyte algae. Using a strain of M. rubrum isolated from McMurdo Sound, Antarctica, we investigated the photoacclimation potential of this trophically unique organism at a range of low irradiance levels. The compensation growth irradiance for M. rubrum was 0.5 μmol quanta · m⁻² · s⁻¹, and growth rate saturated at ∼20 μmol quanta · m⁻² · s⁻¹. The strain displayed trends in photosynthetic efficiency and pigment content characteristic of marine phototrophs. Maximum chl a–specific photosynthetic rates were an order of magnitude slower than temperate strains, while growth rates were half as large, suggesting that a thermal limit to enzyme kinetics produces a fundamental limit to cell function. M. rubrum acclimates to light‐ and temperature‐limited polar conditions and closely regulates photosynthesis in its cryptophyte organelles. By acquiring and maintaining physiologically viable, plastic plastids, M. rubrum establishes a selective advantage over purely heterotrophic ciliates but reduces competition with other phototrophs by exploiting a very low‐light niche.     

Effects of temperature,irradiance, and daylength on the marine diatom leptocylindrus danicus cleve. I. Photosynthesis and cellular composition

Rates of ¹⁴C uptake and cellular composition of C, N, and Chl a in the marine diatom Leptocylindrus danicus Cleve were measured in axenic batch culture under 49 combinations of temperature (5, 10, 15, 20 °C), daylength (15: 9, 12: 12, 9: 15 LD), and irradiance (at least four irradiances per daylength). ¹⁴C uptake exhibited a temperature-dependent daylength effect. Similar P-I curves characterized cells grown under 15: 9 and 12: 12 LD; P_max values were 17.2, 11.2, 4.3, and 1.8 pg C. pg Chl a^?1. h^?1 at 20, 15, 10, and 5°C, respectively. Under 9:15 LD at 20 and 15°C, the lightsaturated photosynthetic rate was ≈50% that in cells grown under longer daylengths. ¹⁴C uptake was independent of daylength at 10 and 5°C. The initial slope, a, of cells grown under long daylengths increased by five-fold between 5 and 20 °C. α values of cells grown under 9: 15 LD at 15 and 20 °C were depressed relative to longer daylengths. Chl a was inversely related to irradiance, and increased with temperature from 10 to 20 °C, whereas cell carbon and nitrogen showed a similar temperature dependence but was not influenced by irradiance or daylength. The C : N ratio and cell volume were independent of temperature, irradiance, and daylength. Both the C : Chl a and N : Chl a ratios increased with irradiance by greater amounts at lower temperatures.     

PIGMENT TURNOVER IN THE MARINE DIATOM THALASSIOSIRA WEISSFLOGII. I. THE 14CO2-LABELING KINETICS OF CHLOROPHYLL a1

The turnover of chlorophyll a (chl a) was investigated in the diatom Thalassiosira weissflogii (Grunow) Fryxell and Hasle using a new method based on the incorporation of ¹⁴C into chl a. The alga was maintained in its exponential growth phase under continuous light; ¹⁴C was supplied as bicarbonate. The time course of label accumulation into the tetrapyrrole ring and the phytol side chain was determined for time periods equivalent to 1–2 cell doublings. The labeling kinetics of the tetrapyrrole ring and the phytol side chain were described satisfactorily by a simple precursor-pigment model with two free parameters, the precursor turnover rate and the pigment turnover rate, both having dimensions of time^?1. The model was fit to the experimental data to determine the values of these two free parameters. The turnover rates of the tetrapyrrole ring and the phytol side chain were not significantly different, ranging from 0.01 to 0.1 per day. These rates are equivalent to turnover times ranging from days to weeks. Growth rate-normalized turnover rates did not vary with irradiance (7.5–825 μE · m^?2· s^?1). The precursor turnover rates of the tetrapyrrole ring and the phytol side chain differed by an order of magnitude. These results indicate that chl a is not degraded significantly in cultures of T. weissflogii grown under continuous light. Neither irradiance nor growth rate affected growth rate-normalized chlorophyll turnover rates. Our results are inconsistent with the hypothesis that steady-state cellular concentrations of chl a are maintained by a dynamic equilibrium between rates of synthesis and degradation.     

THE INFLUENCE OF SALINITY AND TEMPERATURE COVARIATION ON THE PHOTOPHYSIOLOGICAL CHARACTERISTICS OF ANTARCTIC SEA ICE MICROALGAE1

The responses of sea ice microalgae to variation in ambient irradiance (0 to 150 μE · m^?2· s^?1), temperature (–6° to + 6° C), and salinity (0 to 100 ppt) were tested to determine whether these variables act independently or in concert to influence rates of microalgal photosynthesis. The photosynthetic efficiency and maximum photosynthetic rate for sea ice microalgae increased as a function of incubation temperature between -6° and + 6° C. Furthermore, photosynthetic efficiency, maximum photosynthetic rate, and quantum yield were greatest at salinities between SO and 50 ppt. In contrast, the mean specific absorption coefficients were lowest near seawater salinities, and the saturating irradiance, I_s, appeared to be inversely proportional to salinity. Results also suggest that the effects of salinity on the growth of sea ice microalgae are independent of those elicited by temperature or light, and that the functional relationship between salinity and light or temperature is multiplicative. This information is essential to the proper formulation of algorithms used to describe algal growth in environments where light, temperature, and salinity are changing simultaneously, such as within sea ice or within the water column at the marginal ice edge zone.     

Discovery of a resting stage in the harmful,brown‐tide‐causing pelagophyte,Aureoumbra lagunensis: a mechanism potentially facilitating recurrent blooms and geographic expansion

To date, the life stages of pelagophytes have been poorly described. This study describes the ability of Aureoumbra lagunensis to enter a resting stage in response to environmental stressors including high temperature, nutrient depletion, and darkness as well as their ability to revert from resting cells back to vegetative cells after exposure to optimal light, temperature, and nutrient conditions. Resting cells became round in shape and larger in size, filled with red accumulation bodies, had smaller and fewer plastids, more vacuolar space, contained lower concentrations of chl a and RNA, displayed reduced photosynthetic efficiency, and lower respiration rates relative to vegetative cells. Analysis of vegetative and resting cells using Raman microspectrometry indicated resting cells were enriched in sterols within red accumulation bodies and were depleted in pigments relative to vegetative cells. Upon reverting to vegetative cells, cells increased their chl a content, photosynthetic efficiency, respiration rate, and growth rate and lost accumulation bodies as they became smaller. The time required for resting cells to resume vegetative growth was proportional to both the duration and temperature of dark storage, possibly due to higher metabolic demands on stored energy (sterols) reserves during longer period of storage and/or storage at higher temperature (20°C vs. 10°C). Resting cells kept in the dark at 10°C for 7 months readily reverted back to vegetative cells when transferred to optimal conditions. Thus, the ability of Aureoumbra to form a resting stage likely enables them to form annual blooms within subtropic ecosystems, resist temperature extremes, and may facilitate geographic expansion via anthropogenic transport.     

PHOTOSYNTHETIC RESPONSES TO LIGHT AND TEMPERATURE OF THE HETEROMORPHIC MARINE ALGA MASTOCARPUS PAPILLATUS (RHODOPHYTA)1

Physiological differentiation of the heteromorphic life-history phases of the red alga Mastocarpus papillatus (C. Agardh) Kützing was assessed. Photosynthetic responses to light and temperature of the erect, foliose gametophyte were compared to those of the crustose tetrasporophyte. Plants of both phases were collected from four locations on the Pacific coast of Baja California, Mexico, and California, USA, between 32–4l° N latitude. Within each location, the chlorophyll-specific, light-saturated photosynthetic rates of gametophytes were generally five times greater than those of tetrasporophytes. Initial slopes of photosynthesis: irradiance curves were greater for gametophytes than for tetrasporophytes. The crust and the blade from each location were similar with respect to dark respiration rates. For tetrasporophytes from all locations, the photosynthetic temperature optima were between 12–15° C. The photosynthetic temperature optima for gametophytes ranged from 15–17° C for plants from Trinidad Head (41° N) to 22–25° C for plants from Punta Descanso (32° N). Both gametophytes and tetrasporophytes from the northernmost location had significantly higher photosynthetic rates than the same phase from the other three locations. The photosynthetic responses to light support models for the life history in which life history phases have different ecological roles. The gametophyte, thought to be specialized for rapid growth and competition, may allocate more resources to photosynthetic machinery, hence the higher maximum photosynthetic rates. The tetrasporophyte, thought to be specialized for resistance to herbivores, may allocate more resources to structural or chemical defenses in preference to photosynthetic machinery. Consequently, the tetrasporophyte has lower photosynthetic capacity.     

EFFECTS OF LIGHT AND TEMPERATURE ON NET PHOTOSYNTHESIS AND DARK RESPIRATION OF GAMETOPHYTES AND EMBRYONIC SPOROPHYTES OF MACROCYSTIS PYRIFERA1

The rates of net photosynthesis as a function of irradiance and temperature were determined for gametophytes and embryonic sporophytes of the kelp, Macrocystis pyrifera (L.) C. Ag. Gametophytes exhibited higher net photosynthetic rates based on oxygen and pH measurements than their derived embryonic sporophytes, but reached light saturation at comparable irradiance levels. The net photosynthesis of gametophytes reached a maximum of 66.4 mg O₂ g dry wt^?1 h^?1 (86.5 mg CO₂ g dry wt^?1 h^?1), a value approximately seven times the rate reported previously for the adult sporophyte blades. Gametophytes were light saturated at 70 μE m^?2 s^?1 and exhibited a significant decline in photosynthetic performance at irradiances 140 μE m^?1 s^?1. Embryonic sporophytes revealed a maximum photosynthetic capacity of 20.6 mg O₂ g dry wt^?1 h^?1 (25.3 mg CO₂ g dry wt^?1 h^?1), a rate about twice that reported for adult sporophyte blades. Embryonic sporophytes also became light saturated at 70 μE m^?2 s^?1, but unlike their parental gametophytes, failed to exhibit lesser photosynthetic rates at the highest irradiance levels studied; light compensation occurred at 2.8 μE m^?2 s^?1. Light-saturated net photosynthetic rates of gametophytes and embryonic sporophytes varied significantly with temperature. Gametophytes exhibited maximal photosynthesis at 15° to 20° C, whereas embryonic sporophytes maintained comparable rates between 10° and 20° C. Both gametophytes and embryonic sporophytes declined in photosynthetic capacity at 30° C. Dark respiration of gametophytes was uniform from 10° to 25° C, but increased six-fold at 30° C; the rates for embryonic sporophytes were comparable over the entire range of temperatures examined. The broader light and temperature tolerances of the embryonic sporophytes suggest that this stage in the life history of M. pyrifera is well suited for the subtidal benthic environment and for the conditions in the upper levels of the water column.     

EFFECTS OF TEMPERATURE ON GROWTH,LIGHT ABSORPTION,AND QUANTUM YIELD IN DUNALIELLA TERTIOLECTA (CHLOROPHYCEAE)1

The effects of growth temperature on the marine chlorophyte Dunaliella tertiolecta Butcher were studied to provide a more mechanistic understanding of the role of environmental factors in regulating bio-optical properties of phytoplankton. Specific attention was focused on quantities that are relevant for modeling of growth and photosynthesis. Characteristics including chlorophyll a (chl z)-specific light absorption (a*_ph(λ)), C:chl a ratio, and quantum yield for growth (φ_μ) varied as functions of temperature under conditions of excess light and nutrients. As temperature increased over the range examined (12°-28°C), intracellular concentrations of chl a increased by a factor of 2 and a*_ph(λ) values decreased by more than 50% at blue to green wavelengths. The lower values of a*_ph(λ) were due to both a decrease in the abundance of accessory pigments relative to chl a and an increase in pigment package effects arising from higher intracellular pigment concentrations. Intracellular pigment concentration increased as a consequence of higher cellular pigment quotas combined with lower cell volume. At high growth temperatures, slightly more light was absorbed on a per-cell-C basis, but the dramatic increases in growth rate from μ= 0.5 d^?1 at 12° C to μ= 2.2 d^?1 at 28°C were primarily due to an increase in φ_μ (0.015–0.041 mol C (mol quanta)^?1). By comparison with previous work on this species, we conclude the effects of temperature on a*_ph(λ) and φ_μ are comparable to those observed for light and nutrient limitation. Patterns of variability in a*_ph(λ)and φ_μ as a function of growth rate at different temperatures are similar to those previously documented for this species grown at the same irradiance but under a range of nitrogen-limited conditions. These results are discussed in the context of implications for bio-optical modeling of aquatic primary production by phytoplankton.     

PHOTOSYNTHESIS-IRRADIANCE RELATIONSHIPS IN PHYTOPLANKTON FROM THE PHYSICALLY STABLE WATER COLUMN OF A PERENNIALLY ICE-COVERED LAKE (LAKE BONNEY,ANTARCTICA)1

The perennially ice-covered lakes of Antarctica have hydrodynamically stable water columns with a number of vertically distinct phytoplankton populations. We examined the photosynthesis-irradiance characteristics of phytoplankton from four depths of Lake Bonney to determine their physiological condition relative to vertical gradients in irradiance and temperature. All populations studied showed evidence of extreme shade adaptation, including low I_k values (15–45 μE · m^?2· s^?1) and extremely low maximal photosynthetic rates (P^B_m less than 0.3 μg C ·μg chl a^?1· h^?1). Photosynthetic rates were controlled by temperature as well as light variations with depth. Lake Bonney has an inverted temperature profile within the trophogenic zone that increased from 0° C at the ice-water interface to 6° C from 10 to 18 m. Deeper phytoplankton (10 m and 17 m) were found to have photosynthetic capacities (P^B_m) and efficiences (α) three to five times higher than those at the ice-water interface. However, Q₁₀ values were only ca. 2 for P^B_m (no temperature dependence was evident for α), suggesting that a simple temperature response cannot explain all the differences between populations. Lake Bonney phytoplankton (primarily cryptophytes and chlorophytes) had photosynthetic characteristics similar to diatoms from other physically stable environments (e.g. sea ice, benthos) and may be ecologically analogous to multiple deep chlorophyll maxima.     

CHARACTERIZATION OF PSYCHROPHILIC OSCILLATORIANS (CYANOBACTERIA) FROM ANTARCTIC MELTWATER PONDS

Oscillatorian cyanobacteria dominate benthic microbial mat communities in many polar freshwater ecosystems. Capable of growth at low temperatures, all benthic polar oscillatorians characterized to date are psychrotolerant (growth optima > 15° C) as opposed to psychrophilic (growth optima ≤ 15° C). Here, psychrophilic oscillatorians isolated from meltwater ponds on Antarctica's McMurdo Ice Shelf are described. Growth and photosynthetic rates were investigated at multiple temperatures, and compared with those of a psychrotolerant isolate from the same region. Two isolates showed a growth maximum at 8° C, with rates of 0.12 and 0.08 doublings·d ^{? 1}, respectively. Neither displayed detectable growth at 24° C. The psychrotolerant isolate showed almost imperceptible growth at 4° C and a rate of 0.9 doublings·d ^{? 1} at its optimal temperature of ～23° C. In both photosynthesis versus irradiance and photosynthesis versus temperature experiments, exponentially growing cultures were acclimated for 14 days at 3, 8, 12, 20, and 24° C under saturating light intensity, and [¹⁴C] photoincorporation rates were measured. Psychrophilic isolates acclimated at 8° C showed greatest photosynthetic rates; those acclimated at 3° C were capable of active photosynthesis, but photoincorporation was not detected in cells acclimated at 20 and 24° C, because these isolates were not viable after 14 days at those temperatures. The psychrotolerant isolate, conversely, displayed maximum photosynthetic rates at 24° C, though photoincorporation was actively occurring at 3° C. Within acclimation temperature treatments, short‐term photosynthetic rates increased with increasing incubation temperature for both psychrophilic and psychrotolerant isolates. These results indicate the importance of temperature acclimation before assays when determining optimal physiological temperatures. All isolates displayed photosynthetic saturation at low light levels (<128 μmol·m ^{? 2}·s ^{? 1}) but were not photoinhibited at the highest light treatment (233 μmol·m ^{? 2}·s ^{? 1}). Field studies examining the impact of temperature on photosynthetic responses of intact benthic mats, under natural solar irradiance, showed the mat communities to be actively photosynthesizing from 2 to 20° C, with maximum photoincorporation at 20° C, as well as capable of a rapid response to an increase in temperature. The rarity of psychrophilic cyanobacteria, relative to psychrotolerant strains, may be due to their extremely slow growth rates and inability to take advantage of occasional excursions to higher temperatures. We suggest an evolutionary scenario in which psychrophilic strains, or their most recent common ancestor, lost the ability to grow at higher temperatures while maintaining a broad tolerance for fluctuations in other physical and chemical parameters that define shallow meltwater Antarctic ecosystems.     

BROAD THERMAL TOLERANCE OF THE SYMBIOTIC DINOFLAGELLATE SYMBIODINIUM MUSCATINEI (DINOPHYTA) IN THE SEA ANEMONE ANTHOPLEURA ELEGANTISSIMA (CNIDARIA) FROM NORTHERN LATITUDES1

The sea anemone Anthopleura elegantissima (Brandt) hosts two species of symbiotic dinoflagellates, known as zooxanthellae, which coexist within the host at southern latitudes only. One of these species, Symbiodinium muscatinei LaJeunesse et Trench, has a broad latitudinal distribution, occurring in intertidal anemones from Washington state to Southern California. To investigate whether high thermal tolerance contributes to the ability of S. muscatinei to inhabit anemones from northern and southern regions, the upper thermal tolerance limit for photosynthesis of symbionts in northern (48°24′ N) populations of A. elegantissima was determined by subjecting anemones to a gradual increase in temperature from 12°C to 30°C over a 10‐week period. Light‐saturated photosynthetic rates of isolated zooxanthellae were the same over the range of 12°C–24°C and declined significantly at 26°C, which is 14°C and 5°C above average summertime seawater temperatures in northern Puget Sound and Southern California, respectively. At 28°C, zooxanthellae isolated from the anemones, and those expelled by their hosts, exhibited extremely low rates of photosynthesis and highly reduced chl content. The photosynthetic rates and chl content of expelled zooxanthellae were lower than those of retained zooxanthellae. The high thermal tolerance of S. muscatinei isolated from northern populations of anemones supports the broad latitudinal distribution of this symbiont, allowing it to coexist with S. californium (#383, Banaszak et al. 1993 ) in southern populations of anemones.     

VARIABILITY IN THE ECOPHYSIOLOGY OF HALIMEDA SPP. (CHLOROPHYTA,BRYOPSIDALES) ON CONCH REEF,FLORIDA KEYS,USA1

The photosynthetic performance, pigmentation, and growth of a Halimeda community were studied over a depth gradient on Conch Reef, Florida Keys, USA during summer–fall periods of 5 consecutive years. The physiology and growth of H. tuna (Ellis & Solander) Lamouroux and H. opuntia (L.) Lamouroux on this algal dominated reef were highly variable. Maximum rate of net photosynthesis (P_max), respiration rate, and quantum efficiency (α) did not differ between populations of either species at 7 versus 21 m, even though the 21‐m site received a 66% lower photon flux density (PFD). Physiological parameters, as well as levels of photosynthetic pigments, varied temporally. P_max, saturation irradiance, compensation irradiance, and growth were greatest in summer months, whereas α, chl a, chl b, and carotenoid concentrations were elevated each fall. Halimeda tuna growth rates were higher at 7 m compared with 21 m for only two of five growth trials. This may have arisen from variability in light and nutrient availability. Individuals growing at 7 m received a 29% greater PFD in August 2001 than in 1999. In August 1999 and 2001 seawater temperatures were uniform over the 14‐m gradient, whereas in August 2000 cold water regularly intruded upon the 21‐m but not the 7‐m site. These results illustrate the potentially dynamic relationship between nutrients, irradiance, and algal productivity. This suggests the necessity of long‐term monitoring over spatial and temporal gradients to accurately characterize factors that impact productivity.     

The pigment composition of Phaeocystis antarctica (Haptophyceae) under various conditions of light,temperature, salinity,and iron

The pigment composition of Phaeocystis antarctica was monitored under various conditions of light, temperature, salinity, and iron. 19′‐Hexanoyloxyfucoxanthin (Hex‐fuco) always constituted the major light‐harvesting pigment, with remarkably stable ratios of Hex‐fuco‐to‐chl a under the various environmental conditions. Increased pigment‐to‐chl a ratios at low irradiance confirmed the light‐harvesting function of Fucoxanthin (Fuco), 19′‐Hexanoyloxy‐4‐ketofucoxanthin (Hex‐kfuco), 19′‐butanoyloxyfucoxanthin (But‐fuco), and chl c2 and c3. Increased pigment‐to‐chl a ratios at high irradiance, low iron concentrations, and to a lesser extent at high salinity confirmed the photoprotective function of diadinoxanthin, diatoxanthin, and ß,ß‐carotene. Pigment ratios were not always according to expectations. The consistent increase in But‐fuco/chl at high temperature, high salinity, and low iron suggests a role in photoprotection rather than in light harvesting. Low Hex‐kfuco/chl ratios at high salinity were consistent with a role as light harvester, but the high ratios at high temperature were not, leaving the function of Hex‐kfuco enigmatic. Dedicated experiments were performed to test whether or not the light‐harvesting pigment Fuco could be converted into its structural relative Hex‐fuco, and vice versa, in response to exposure to light shifts. Rapid conversions could not be confirmed, but long‐term conversions cannot be excluded. New pigment ratios are proposed for chemotaxonomic applications. The ratios will improve pigment‐based diagnosis of algal species in waters dominated by P. antarctica.     

EFFECTS OF LONG TERM EXPOSURE TO LOW TEMPERATURE ON THE PHOTOSYNTHETIC APPARATUS OF DUNALIELLA TERTIOLECTA (CHLOROPHYCEAE)1

The effects of long term exposure to suboptimal growth temperature on the photosynthetic apparatus of Dunaliella tertiolecta Butcher were investigated using carbon fixation rate versus irradiance curves and the variable fluorescence induction method. Carbon fixation rates per unite chlorophyll a at saturating (p^B_m) and subsaturating (α^B) irradiances were 55% and 39% lower, respectively, at 12° C than at 20° C. Chlorophyll a quotas and the spectrally averaged in vivo absorption cross section normalized to chlorophyll a (a^*) were not significantly different at these two temperatures. Analysis of the fluorescence kinetics revealed 1) no significant variations of the amount of PSII photoactive reaction centers per unit chlorophyll a, 2) a 14% decrease of the PSII quantum yield(+) and 3) a 29% decrease of the energy transfer efficiency between the light harvesting chlorophyll a pigment bed and the PSII reaction centers. The decrease in energy transfer efficiency between the antennae and the PSII reaction centers at 12° C was interpreted as a mechanism to avoid photoinhibition.