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

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

STRATEGIES AND KINETICS OF PHOTOACCLIMATION IN THREE ANTARCTIC NANOPHYTOFLAGELLATES1

Three Antarctic nanophytoflagellates (two cryptophyte species and a Pyramimonas sp.) were compared for their capacity to phiotoacclimate and for their kinetic responses in changing photic environments. Division rate, cell size cellular fluorescence, and chlorophyll a content were measured steady and transient states of semi-continuous cultures maintain at 1.0° C. Of all parameters tested, cell size was most affected by irradiance. Acclimation kinetics were modeled using a first-order equation. Rates of change in cell size following shifts in irradiance were comparable with rates of change in chemical composition reported for temperate algae. Response rates of cellular in vivo red and orange fluorescence were lower. In many cases, however, responses could not be described by the first-order kinetic model. Division rates remained high for approximately 3 days following a shift down in irradiance, after which new division rates were established. The nanoflagellates studied here appear to respond to small irradiance perturbations at low rates. However, they may fail to adapt and abrupt changes in photon flux density (PFD). When shade-adapted (25 μmol, m^?2, m^?2, s^?1) cells were exposed to high PFD (400 μmol, m^?2, s^?1) for 1–3 days, cell were incapable of readapting division rate and pigment content to the initial irradiance condition (25 μmol, m^?2, s^?1) for about 1 month following the shift-down step. The ecological role of the kinetics of photoacclimation in nanophytoflagellate growth performance in Antarctic ecosystems is discussed.     

NONEQUILIBRIUM RATES OF PHOTOSYNTHESIS AND RESPIRATION UNDER DYNAMIC LIGHT SUPPLY1

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

Photosynthetic performance of freshwater Rhodophyta in response to temperature, irradiance, pH and diurnal rhythm

Responses of net photosynthetic rates to temperature, irradiance, pH/inorganic carbon and diurnal rhythm were analyzed in 15 populations of eight freshwater red algal species in culture and natural conditions. Photosynthetic rates were determined by oxygen concentration using the light and dark bottles technique. Parameters derived from the photosynthesis–irradiance curves indicated adaptation to low irradiance for all freshwater red algae tested, confirming that they tend to occur under low light regimes. Some degree of photo‐inhibition (β= ‐0.33–0.01 mg O₂ g^?1 DW h^?1 (μmol photons m^?2 s^?1)^?1) was found for all species/populations analyzed, whereas light compensation points (Ic) were very low (≤ 2 μmol photons m‐ photons s^?1) for most algae tested. Saturation points were low for all algae tested (Ik = 6–54 μmol photons m^?2 s^?1; Is = 20–170 umol photons m^?2 s^?1). Rates of net photosynthesis and dark respiration responded to the variation in temperature. Optimum temperature values for net photosynthesis were variable among species and populations so that best performances were observed under distinct temperature conditions (10, 15, 20 or 25°C). Rates of dark respiration exhibited an increasing trend with temperature, with highest values under 20–25°C. Results from pH experiments showed best photosynthetic performances under pH 8.5 or 6.5 for all but one species, indicating higher affinity for inorganic carbon as bicarbonate or indistinct use of bicarbonate and free carbon dioxide. Diurnal changes in photosynthetic rates revealed a general pattern for all algae tested, which was characterized by two relatively clear peaks, with some variations around it: a first (higher) during the morning (07.00–11.00 hours.) and a second (lower) in the afternoon (14.00–18.00 hours). Comparative data between the ‘Chantransia’ stage and the respective gametophyte for one Batrachospermum population revealed higher values (ca 2‐times) in the latter, much lower than previously reported. The physiological role of the ‘Chantransia’ stage needs to be better analyzed.     

Changes in the condition of photosynthetic apparatus of a diatom alga <Emphasis Type="Italic">Thallassiosira weisflogii</Emphasis> during photoadaptation and photodamage

Two populations of a diatom alga Thallassiosira weisflogii were grown at photon flux densities (PFD) of 0.8 and 8 μmol/(m² s). For both diatom populations, the recovery of chlorophyll fluorescence parameters (F ₀, F _m, F _v/F _m, and NPQ) was monitored after nondestructive irradiation by visible light at PFD of 40 μmol/(m² s) and after high-intensity irradiation by visible light (1000–4000 μmol/(m² s)). The exposure of diatoms to PFD of 40 μmol/(m² s)—higher than PFD used for algal growth but still nondamaging to photosynthetic apparatus—induced nonphotochemical quenching (NPQ), which was stronger in algae grown at higher PFD (8 μmol/(m² s)) than in algae grown at low light. After irradiation with high-intensity light, the recovery of chlorophyll fluorescence parameters was more pronounced in algae grown at elevated PFD level. During short-term irradiation of diatoms with high-intensity visible light (1000 μmol/(m² s)), a stronger NPQ was observed in the culture adapted to high irradiance. After the treatment of algae with dithiothreitol (an inhibitor of carotenoid deepoxidase in the diadinoxanthin cycle) or NH₄Cl (an agent abolishing the proton gradient at thylakoid membranes), a short exposure of algae to PFD of 40 μmol/(m² s) induced hardly any nonphotochemical quenching. The results indicate the dominant contribution of xanthophyll cycle carotenoids to energy-dependent quenching.     

Changes in photon flux can induce stomatal patchiness

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

SEASONAL VARIATIONS OF PHOTOSYNTHETIC PIGMENTS, TOTAL C, N, AND P CONTENT, AND PHOTOSYNTHESIS IN PHYLLARIOPSIS PURPURASCENS: (PHAEOPHYTA) FROM THE STRAIT OF GIBRALTAR

Photosynthetic pigments, C, N, and P tissue composition, and photosynthetic rate were measured from April to October in the brown alga Phyllariopsis purpurascens (C. Agardh) Henry et South (Laminariales, Phaeophyta) growing at a 30-m depth in the Strait of Gibraltar. Ir-radiance reaching the population ranged from 13.5 to 27.5 mol.m^-2.mo^-1. The available light for this species, expressed as a percentage of the irradiance above the water, was 1.8%. Dissolved inorganic nitrogen forms, NO3-and NH₄+, were constant from April to October, whereas phosphate was depleted in August. Chlorophyll a decreased from 520.0 ± 165.0 to 199.6 ± 159.9 μg.g^-1 dry weight; in contrast, chlorophyll c and carotenoids did not change until September but increased threefold in October. C:N and N:P ratios changed in the same way and in the same range. They were constant until July but increased from 15–17 up to 42 (C:N) and from 14 to 40 (N:P) in October, suggesting a severe P limitation of growth of this species. The dark respiration rate and the light compensation point were constant from April to October (0.5 ± 0.1 μmol O₂. m^-2.s^-1 and 6.5 ± 0.2 μmol.m^-2. s^-1, respectively), whereas the maximum rate of apparent photosynthesis, light onset saturation parameter, and half saturation constant for light were maximum in April to May (3.7 μmol O₂. m^-2.s^-1and 40 and 41.5 μmol.m^-2. s^-1, respectively) and October (3.6 μmol O₂. m^-2.s^-1 and 50 and 53.7 μmol.m^-2. s^-1, respectively). They were minimum in August (1.2 μmol O₂.m^-2.s^-1 and 11.3 and 12 μmol.m^-2.s^-1, respectively). These minimum figures yielded a negative carbon budget in August and 0 in September, whereas it was positive the rest of the year. Photosynthetic efficiency, estimated by the ratio between maximum apparent photosynthesis and light half saturation constant, showed a strong agreement with productivity measured by means of an independent method. These results indicate that lamina expansion in this species is controlled by photosynthetic efficiency.     

Photosynthetic characteristics of charophytes from tropical lotic ecosystems

Responses of photosynthetic rates, determined by oxygen evolution using the light and dark bottles technique, to different temperatures, irradiances, pH, and diurnal rhythm were analyzed under laboratory conditions in four charophyte species (Chara braunii Gmelin, C. guairensis R. Bicudo, Nitella subglomerata A. Braun and Nitella sp.) from lotic habitats in southeastern Brazil. Parameters derived from the photosynthesis versus irradiance curves indicated affinity to low irradiances for all algae tested. Some degree of photoinhibition, [β= ‐(0.30–0.13) mg O2 g^?1 dry weight Ir¹ (μmol photons m^?2 s^?1)^?1], low light compensation points (I_c= 4–20 μmol photons m^?2 s^?1) were found for all species analyzed, as well as low values of light saturation parameter (I_k) and saturation (I_s) 29–130 and 92–169 μmol photons m^?2 s^?1, respectively. Photoacclimation was observed in two populations of N. subglomerata collected from sites with different irradiances, consisting of variations in photosynthetic parameters (higher values of a, and lower of I_k and maximum photosynthetic rate, P_max, in the population under lower irradiance). The highest photosynthetic rates for Chara species were observed at 10–15°C, while for Nitella the highest photosynthetic rate was observed at 20–25°C, despite the lack of significant differences among most levels tested. Rates of dark respiration significantly increase with temperature, with the highest values at 25°C. The results from pH experiments showed highest photosynthetic rates under pH 4.0 for all algae, suggesting higher affinity for inorganic carbon in the form of carbon dioxide, except in one population of N. subglomerata, with similar rates under the three levels, suggesting indistinct use of bicarbonate and carbon dioxide. Diurnal changes in photosynthetic rates revealed a general pattern for most algae tested, which was characterized by two peaks: the first (higher) during the morning (07.00–11.00) and the second (lower) in the afternoon (14.00–17.00). This suggests an endogenous rhythm determining the daily variations in photosynthetic rates.     

PHOTOINHIBITION OF MECHANICALLY STIMULABLE BIOLUMINESCENCE IN THE AUTOTROPHIC DINOFLAGELLATE CERATIUM FUSUS (PYRROPHYTA)1

Ceratium fusus (Ehrenb.) Dujardin was exposed to light of different wavelengths and photon flux densities (PFDs) to examine their effects on mechanically stimulable bioluminescence (MSL). Photoinhibition of MSL was proportional to the logarithm of PFD. Exposure to I μmol photons·m^?2s^?1 of broadband blue light (ca. 400–500 nm) produced near-complete photoinhibition (≥90% reduction in MSL) with a threshold at ca. 0.01 μmol photons·m^?2·s^?1. The threshold of photoinhibition was ca. an order of magnitude greater for both broadband green (ca. 500–580 nm) and red light (ca. 660–700 nm). Exposure to narrow spectral bands (ca. 10 nm half bandwidth) from 400 and 700 nm at a PFD of 0.1 μmol photons·m^?2·s^?1 produced a maximal response of photoinhibition in the blue wavelengths (peak ca. 490 nm). A photoinhibition response (≥ 10%) in the green (ca. 500–540 nm) and red wavelengths (ca. 680 nm) occurred only at higher PFDs (1 and 10 μmol photons·m^?2·s^?1). The spectral response is similar to that reported for Gonyaulax polyedra Stein and Pyrocystis lunula Schütt and unlike that of Alexandrium tamarense (Lebour) Balech et Tangen. The dinoflagellate's own bioluminescence is two orders of magnitude too low to result in self-photoinhibition. The quantitative relationships developed in the laboratory predict photoinhibition of bioluminescence in populations of C. fusus in the North Atlantic Ocean.     

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

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

Heterogeneity of Light Availability and its Effects on Simulated Carbon Gain of Tree Leaves in a Small Gap and the Understory in a Tropical Rain Forest1

To clarify the small-scale heterogeneity of light regimes in a rain forest, photosynthetic photon flux density (PFD) was measured at 1-min intervals during six days at 12 microsites in each of two plots, a small gap and an understory in Pasoh Forest Reserve, Peninsular Malaysia. Frequency distribution of microsite PFD was unimodal with the peak value between 16 and 32 μmol/m²/sec in the small gap, but between 8 and 16 μmol/m²/sec in the understory. In the small gap, PFD was more variable among microsites; total daily PFD and daily sunfleck PFD exceeding 10 μmol/ m²/sec tended to be higher (P <0.05; t-test) compared to those in the understory. Sunfleck PFD exceeding 50 μmol/ m²/sec, however, showed no difference between the two plots. Diffuse PFD transmittance, defined as the ratio of PFD in the forest to that measured at 43 m above ground during the periods 0800-0810 and 1750-1800 h, was significantly higher in the small gap than in the understory plot. Diffuse PFD transmittance was also positively correlated with microsite total daily PFD. To examine the effects of the subtle heterogeneity of light regimes on leaf carbon gain, we simulated carbon gain by sun and shade leaves in a typical shade-tolerant species, Brosimum aticastrum Sw. (Moraceae). Despite the similarity in total daily PFD, total daily carbon gain was considerably higher in the gap than in the understory for both sun and shade leaves. This study suggests that frequency distribution of PFD is critical in describing microsite PFD regimes and determining leaf carbon gain in the tropical forest floor.     

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

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

Annual carbon fixation in terrestrial populations of Nostoc commune (Cyanobacteria) from an Antarctic dry valley is driven by temperature regime

Nostoc commune Vaucher (a cyanobacterium) is a very conspicuous terrestrial primary producer in Victoria Land, continental Antarctica. Because polar ecosystems are considered to be especially sensitive to environmental changes, understanding the environmental constraints on net carbon (C) fixation by N. commune is necessary to determine the effects of environmental changes on the ecological functioning of ice‐free areas of the continent. A model describing net C fixation in terrestrial populations of N. commune in an Antarctic dry valley was constructed using field and laboratory measurements in which N. commune colonies were exposed to different combinations of incident irradiance (400–700 nm), temperature, and degree of desiccation. For desiccated N. commune mats with water content ≤ 30% saturation, net C fixation was highly variable between replicates and could not be modelled. However, for colonies at > 30% saturation, rates of net C fixation and dark respiration depended strongly on irradiance and temperature. Net C fixation reached a maximum rate of 21.6 μg C m^{− 2} s^{− 1} at irradiance of approximately 250 μmol m^{− 2} s^{− 1} and the optimum temperature of 20.5 °C. Agreement between predicted short‐term net C fixation and field and laboratory measurements allowed estimation of total seasonal fixation, using previously published environmental data. Annual net C fixation was estimated in the range 14.5–21.0 g C fixed m^{− 2}Nostoc mat, depending on year/season. Estimates for different seasons correlated with thermal time (accumulated hours above 0 °C during the year) rather than irradiance, in contrast to communities in local lacustrine environments, where irradiance is the main driver of primary productivity. In the terrestrial habitat, N. commune appears to compromise between an ability to capitalize on short periods of higher temperature and efficient utilization of lower irradiance at low temperature. The relationship between thermal time and net annual C fixation by N. commune is strongly linear.     

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

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

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

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

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

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

EFFECT OF SEASONAL SEA ICE BREAKOUT ON THE PHOTOSYNTHESIS OF BENTHIC DIATOM MATS AT CASEY,ANTARCTICA1

Photosynthesis of marine benthic diatom mats was examined before and after sea ice breakout at a coastal site in eastern Antarctica (Casey). Before ice breakout the maximum under‐ice irradiance was between 2.5 and 8.2 μmol photons·m^?2·s^?1 and the benthic microalgal community was characterized by low E_k (12.1–32.3 μmol photons·m^?2·s^?1), low relETR_max (9.2–32.9), and high alpha (0.69–1.1). After breakout, 20 days later, the maximum irradiance had increased to between 293 and 840 μmol photons·m^?2·s^?1, E_k had increased by more than an order of magnitude (to 301–395 μmol photons·m^?2·s^?1), relETR_max had increased by more than five times (to 104–251), and alpha decreased by approximately 50% (to 0.42–0.68). During the same time interval the species composition of the mats changed, with a decline in the abundance of Trachyneis aspera (Karsten) Hustedt, Gyrosigma subsalsum Van Heurck, and Thalassiosira gracilis (Karsten) Hustedt and an increase in the abundance of Navicula glaciei Van Heurck. The benthic microalgal mats at Casey showed that species composition and photophysiology changed in response to the sudden natural increase in irradiance. This occurred through both succession shifts in the species composition of the mats and also an ability of individual cells to photoacclimate to the higher irradiances.     

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

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

PHOTOSYNTHESIS IN AN EXTREME SHADE ENVIRONMENT: BENTHIC MICROBIAL MATS FROM LAKE HOARE, A PERMANENTLY ICE-COVERED ANTARCTIC LAKE

We investigated the composition of benthic microbial mats in permanently ice-covered Lake Hoare, Antarctica, and their irradiance vs. photosynthetic oxygen exchange relationships. Mats could be subdivided into three distinct depth zones: a seasonally ice-free “moat” zone and two under-ice zones. The upper under-ice zone extended from below the 3.5 m thick ice to approximately 13 m and the lower from below 13 m to 22 m. Moat mats were acclimated to the high irradiance they experienced during summer. They contained photoprotective pigments, predominantly those characteristic of cyanobacteria, and had high compensation and saturating irradiances (E_c and E_k) of 75 and 130 μmol photons·m⁻²·s⁻¹, respectively. The moat mats used light inefficiently. The upper under-ice community contained both cyanobacteria and diatoms. Within this zone, biomass (as pigments) increased with increasing depth, reaching a maximum at 10 m. Phycoerythrin was abundant in this zone, with shade acclimation and efficiency of utilization of incident light increasing with depth to a maximum of 0.06 mol C fixed·mol⁻¹ incident photons under light-limiting conditions. Precipitation of inorganic carbon as calcite was associated with this community, representing up to 50% of the carbon sequestered into the sediment. The lower under-ice zone was characterized by a decline in pigment concentrations with depth and an increasing prevalence of diatoms. Photosynthesis in this community was highly shade acclimated and efficient, with E_c and E_k below 0.5 μmol·m⁻²·s⁻¹ and 2 μmol·m⁻²·s⁻¹, respectively, and maximum yields of 0.04 mol C fixed·mol⁻¹ incident quanta. Carbon uptake in situ by both under-ice and moat mats was estimated at up to 100 and 140 mg·m⁻²·day⁻¹, based on the photosynthesis–irradiance curves, incident irradiance, and light attenuation by ice and the water column.     

Effect of CO2 concentration on the PIC/POC ratio in the coccolithophore Emiliania huxleyi grown under light-limiting conditions and different daylengths

We compared the effect of CO₂ concentration ([CO₂], ranging from ∼5 to ∼34 μmol l⁻¹) at four different photon flux densities (PFD=15, 30, 80 and 150 μmol m⁻² s⁻¹) and two light/dark (L/D) cycles (16/8 and 24/0 h) on the coccolithophore Emiliania huxleyi. With increasing [CO₂], a decrease in the particulate inorganic carbon to particulate organic carbon (PIC/POC) ratio was observed at all light intensities and L/D cycles tested. The individual response in cellular PIC and POC to [CO₂] depended strongly on the PFD. POC production increased with rising [CO₂], irrespective of the light intensity, and PIC production decreased with increasing [CO₂] at a PFD of 150 μmol m⁻² s⁻¹, whereas below this light level it was unaffected by [CO₂]. Cell growth rate decreased with decreasing PFD, but was largely independent of ambient [CO₂]. The diurnal variation in PIC and POC content, monitored over a 38-h period (16/8 h L/D, PFD=150 μmol m⁻² s⁻¹), exceeded the difference in carbon content between cells grown at high (∼29 μmol l⁻¹) and low (∼4 μmol l⁻¹) [CO₂]. However, consistent with the results described above, cellular POC content was higher and PIC content lower at high [CO₂], compared to the values at low [CO₂], and the offset was observed throughout the day. It is suggested that the observed sensitivity of POC production for ambient [CO₂] may be of importance in regulating species-specific primary production and species composition.