Interaction Effects of Light, Temperature and Nutrient Limitations (N, P and Si) on Growth, Stoichiometry and Photosynthetic Parameters of the Cold-Water Diatom Chaetoceros wighamii

Light (20-450 μmol photons m^-2 s^-1), temperature (3-11°C) and inorganic nutrient composition (nutrient replete and N, P and Si limitation) were manipulated to study their combined influence on growth, stoichiometry (C:N:P:Chl a) and primary production of the cold water diatom Chaetoceros wighamii. During exponential growth, the maximum growth rate (~0.8 d^-1) was observed at high temperture and light; at 3°C the growth rate was ~30% lower under similar light conditions. The interaction effect of light and temperature were clearly visible from growth and cellular stoichiometry. The average C:N:P molar ratio was 80:13:1 during exponential growth, but the range, due to different light acclimation, was widest at the lowest temperature, reaching very low C:P (~50) and N:P ratios (~8) at low light and temperature. The C:Chl a ratio had also a wider range at the lowest temperature during exponential growth, ranging 16-48 (weight ratio) at 3°C compared with 17-33 at 11°C. During exponential growth, there was no clear trend in the Chl a normalized, initial slope (α*) of the photosynthesis-irradiance (PE) curve, but the maximum photosynthetic production (P_m) was highest for cultures acclimated to the highest light and temperature. During the stationary growth phase, the stoichiometric relationship depended on the limiting nutrient, but with generally increasing C:N:P ratio. The average photosynthetic quotient (PQ) during exponential growth was 1.26 but decreased to <1 under nutrient and light limitation, probably due to photorespiration. The results clearly demonstrate that there are interaction effects between light, temperature and nutrient limitation, and the data suggests greater variability of key parameters at low temperature. Understanding these dynamics will be important for improving models of aquatic primary production and biogeochemical cycles in a warming climate.

Effects of N supply on the accumulation of photosynthetic pigments and photoprotectors in Gracilaria tenuistipitata (Rhodophyta) cultured under UV radiation

We have studied the effects of nitrate supply under photosynthetic active radiation (PAR) plus ultraviolet radiation (UVR) exposure on photosynthetic pigments (chlorophyll a and carotenoids), photoprotective UV screen mycosporine-like amino acids (MAAs), and photosynthetic parameters, including the maximum quantum yield (F(v)/F(m)) and electron transport rate (ETR) on the red agarophyte Gracilaria tenuistipitata. Apical tips of G. tenuistipitata were cultivated under ten different concentrations of NO(3)(-) for 7 days. It has been shown that G. tenuistipitata cultured under laboratory conditions has the ability to accumulate high amounts of MAAs following a nitrate concentration-dependent manner under PAR+UVR. Two MAAs were identified, shinorine and porphyra-334. The relative concentration of the first increased under high concentrations of nitrate, while the second one decreased. The presence of antheraxanthin is reported for the first time in this macro-algae, which also contains zeaxanthin, lutein, and beta-carotene. The accumulation of pigments, photoprotective compounds, and photosynthetic parameters of G. tenuistipitata is directly related to N availability. All variables decreased under low N supplies and reached constant maximum values with supplements higher than 0.5 mM NO(3)(-). Our results suggest a high potential to acclimation and photoprotection against stress factors (including high PAR and UVR) directly related to N availability for G. tenuistipitata.

Evidence of higher photosynthetic plasticity in the early successional Guazuma ulmifolia Lam. compared to the late successional Hymenaea courbaril L. grown in contrasting light environments

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Photosynthetic characteristics of a tropical population of Nitella cernua (Characeae, Chlorophyta)

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Light-related photosynthetic characteristics of freshwater rhodophytes

Photosynthetic characteristics in response to irradiance were analysed in 21 field and culture populations of thirteen freshwater red algal species applying two distinct techniques (chlorophyll fluorescence and oxygen evolution). Photosynthesis-irradiance (PI) curves indicated adaptations to low irradiances in all species analysed, essentially characterized by occurrence of photoinhibition, low values of the saturation parameter (I-k < 225 mu mol m(-2) s(-1)) and compensation irradiance (I-c < 20 mu mol m(-2) s(-1)) and relatively high values of the effective quantum yield of photosystem II (Delta F/F'(m) >= 45). These characteristics have been reported in freshwater red algae and were confirmed from data based on the two techniques, indicating they are typically shade-adapted plants. on the other hand, some species (e.g. Batrachospermum delicatulum) can tolerate high irradiances (up to 2400 mu mol m(-2) s(-1)), suggesting they have mechanisms that enable them to avoid photodarnage of the photosynthetic apparatus. One of these mechanisms is the increase in dissipation of excessive energy captured by reaction centres after exposure to continuous irradiance, as reflected by the non-photochemical quenching fluorescence parameter in dark/light induction curves. Photo-inhibition occurred in all algae tested by both techniques. Light acclimation was evident particularly in field populations, as revealed by lower values of the saturation parameter (Ik) and the compensation irradiance (I-c) and higher values of Delta F/F'(m) in algae under low irradiances (shaded or heavily shaded stream segments), and vice-versa. Forms living within the boundary layer (e.g. crusts), in a region of reduced current velocity, tended to be more shade-adapted than semi-erect plants (e.g. non-mucilaginous or mucilaginous filaments), as indicated by highest values of photosynthetic efficiency (alpha = 0.31) and effective quantum yield (Delta F/F'(m) = 0.88) under natural conditions. Higher photo- synthetic rates (P-max) for the same species or population were observed under culture than field conditions when measured with the oxygen evolution technique, whereas the opposite trend was observed using chlorophyll fluorescence. (c) 2005 Elsevier B.V. All rights reserved.

Light-related photosynthetic characteristics of lotic macroalgae

Photosynthetic characteristics in response to irradiance were analysed in 42 populations of 33 macroalgal species by two distinct techniques (chlorophyll fluorescence and oxygen evolution). Photosynthesis-irradiance (PI) curves based on the two techniques indicated adaptations to low irradiance reflected by low saturation values, high to moderate values of photosynthetic efficiency (alpha) and photoinhibition (beta), for Bacillariophyta and Rhodophyta, which suggests they are typically shade-adapted algae. In contrast, most species of Chlorophyta were reported as sun adapted algae, characterized by high values of I-k and low of alpha, and lack of or low photoinhibition. Cyanophyta and Xanthophyta were intermediate groups in terms of light adaptations. Photoinhibition was observed in variable degrees in all algal groups, under field and laboratory conditions, which confirms that it is not artificially induced by experimental conditions, but is rather a common and natural phenomenon of the lotic macroalgae. Low values of compensation irradiance (I-c) were found, which indicate that these algae can keep an autotrophic metabolism even under very low irradiances. High ratios (>2) of photosynthesis/respiration were found in most algae, which indicates a considerable net gain. These two physiological characteristics suggest that macroalgae may be important primary producers in lotic ecosystems. Saturation parameters (I-k and I-s) occurred in a relatively narrow range of irradiances (100-400 mumol photons m(-2) s(-1)), with some exceptions (higher in some filamentous green algae or lower in red algae). These parameters were way below the irradiances measured at collecting sites for most algae, which means that most of the available light energy was not photochemically converted via photosynthesis. Acclimation to ambient PAR was observed, as revealed by lower values of I-k and I-c and higher values of alpha and quantum yield in algae from shaded streams, and vice versa. Forms living within the boundary layer (crusts) showed responses of shade-adapted species and had the highest values of P-max, alpha and quantum yield, whereas the opposite trend was observed in gelatinous forms (colonies and. laments). These results suggests adaptation to the light regime rather than functional attributes related to the growth form.

Acclimation to short-term low temperatures in two Eucalyptus globulus clones with contrasting drought resistance

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)

Photosynthetic responses to temperature in tropical lotic macroalgae

A comparative analysis of the photosynthetic responses to temperature (10-30°C) was carried out under short-term laboratory conditions by chlorophyll fluorescence and oxygen (O2) evolution. Ten lotic macroalgal species from southeastern Brazil (20°11-20°48′S, 49°18-49°41′W) were tested, including Bacillariophyta, Chlorophyta, Cyanophyta, Rhodophyta and Xanthophyta. Temperature had significant effects on electron transport rate (ETR) only for three species (Terpsinoe musica, Bacillariophyta; Cladophora glomerata, Chlorophyta; and C. coeruleus, Rhodophyta), with highest values at 25-30°C, whereas the remaining species had no significant responses. It also had similar effects on non-photochemical quenching and ETR. Differences in net photosynthesis/dark respiration ratios at distinct temperatures were found, with an increasing trend of respiration with higher temperatures. This implies in a decreasing balance between net primary production and temperature, representing more critical conditions toward higher temperatures for most species. In contrast, high net photosynthesis and photosynthesis/dark respiration ratios at high and wide ranges of temperature were found in three species of green algae, suggesting that these algae can be important primary producers in lotic ecosystems, particularly in tropical regions. Optimal photosynthetic rates were observed under similar environmental temperatures for five species (two rhodophytes, two chlorophytes and one diatom) considering both techniques, suggesting acclimation to their respective ambient temperatures. C. coeruleus was the only species with peaks of ETR and O 2 evolution under similar field-measured temperatures. All species kept values of ETR and net photosynthesis close to the optimum under a broad range of temperatures. Increased non-photochemical quenching, as a measure of thermal dissipation of excess energy, toward higher temperatures was observed in some species, as well as positive correlation of non-photochemical quenching with ETR, and were interpreted as two mechanisms of adaptation of the photosynthetic apparatus to temperature changes. Different optimal temperatures were found for individual species by each technique, generally under lower temperatures by O2 evolution, indicating dependence on distinct factors: increases in temperature generally induced higher ETR due to increased enzymatic activity, whereas increments of enzymatic activity were compensated by increased respiration and photorespiration leading to decreases in net photosynthesis.

INSTANTANEOUS PHOTOSYNTHETIC RESPONSE TO TEMPERATURE OF MATURE FOREST CANOPIES AND EXPERIMENTALLY WARMED SEEDLINGS

Tropical trees have been shown to be more susceptible to warming compared to temperate species, and have shown growth and photosynthetic declines at elevated temperatures as little as 3oC above ambient. However, regional and global vegetation models lack the data needed to accurately represent physiological response to increased temperatures in tropical forests. We compared the instantaneous photosynthetic responses to elevated temperatures of four mature tropical rainforest tree species in Puerto Rico and the temperate broadleaf species sugar maple (Acer saccharum) in Michigan. Contrary to expectations, leaves in the upper canopy of both temperate and tropical forests had temperature optima that are already exceeded by mean daily leaf temperatures. This indicates that tropical and temperate forests are already seeing photosynthesis decline at mid-day temperature. This decline may worsen as air temperatures rise with climate change if trees are unable to acclimate, increasing the likelihood that forests may shift from carbon sinks to sources. A secondary study was conducted on experimentally warmed sugar maple seedlings to determine if photosynthesis had been able to acclimate to +5oC air temperature over four years. Species abundance models had predicted a decline of sugar maple within the Upper Peninsula of Michigan over the next 100 years, due to elevated temperature and altered precipitation. Instantaneous photosynthetic temperature response curves on both control and heated seedlings showed that the differences between treatments were not statistically significant, though there was a 16% increase in temperature optima and a 3% increase in maximum rates of photosynthesis in warmed plots. Though evidence of acclimation was not significant, the seedlings did not fare poorly as the models suggest.

Growth at moderately elevated temperature alters the physiological response of the photosynthetic apparatus to heat stress in pea (Pisum sativum L.) leaves

The impact of heat stress on the functioning of the photosynthetic apparatus was examined in pea (Pisum sativum L.) plants grown at control (25 °C; 25 °C-plants) or moderately elevated temperature (35 °C; 35 °C-plants). In both types of plants net photosynthesis (Pn) decreased with increasing leaf temperature (LT) and was more than 80% reduced at 45 °C as compared to 25 °C. In the 25 °C-plants, LTs higher than 40 °C could result in a complete suppression of Pn. Short-term acclimation to heat stress did not alter the temperature response of Pn. Chlorophyll a fluorescence measurements revealed that photosynthetic electron transport (PET) started to decrease when LT increased above 35 °C and that growth at 35 °C improved the thermal stability of the thylakoid membranes. In the 25 °C-plants, but not in the 35 °C-plants, the maximum quantum yield of the photosystem II primary photochemistry, as judged by measuring the Fv/Fm ratio, decreased significantly at LTs higher than 38 °C. A post-illumination heat-induced reduction of the plastoquinone pool was observed in the 25 °C-plants, but not in the 35 °C-plants. Inhibition of Pn by heat stress correlated with a reduction of the activation state of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco). Western-blot analysis of Rubisco activase showed that heat stress resulted in a redistribution of activase polypeptides from the soluble to the insoluble fraction of extracts. Heat-dependent inhibition of Pn and PET could be reduced by increasing the intercellular CO2 concentration, but much more effectively so in the 35 °C-plants than in the 25 °C-plants. The 35 °C-plants recovered more efficiently from heat-dependent inhibition of Pn than the 25 °C-plants. The results show that growth at moderately high temperature hardly diminished inhibition of Pn by heat stress that originated from a reversible heat-dependent reduction of the Rubisco activation state. However, by improving the thermal stability of the thylakoid membranes it allowed the photosynthetic apparatus to preserve its functional potential at high LTs, thus minimizing the after-effects of heat stress.

Increased CO2 modifies the carbon balance and the photosynthetic yield of two common Arctic brown seaweeds: Desmarestia aculeata and Alaria esculenta

Ocean acidification affects with special intensity Arctic ecosystems, being marine photosynthetic organisms a primary target, although the consequences of this process in the carbon fluxes of Arctic algae are still unknown. The alteration of the cellular carbon balance due to physiological acclimation to an increased CO2 concentration (1300 ppm) in the common Arctic brown seaweeds Desmarestia aculeata and Alaria esculenta from Kongsfjorden (Svalbard) was analysed. Growth rate of D. aculeata was negatively affected by CO2 enrichment, while A. esculenta was positively affected, as a result of a different reorganization of the cellular carbon budget in both species. Desmarestia aculeata showed increased respiration, enhanced accumulation of storage biomolecules and elevated release of dissolved organic carbon, whereas A. esculenta showed decreased respiration and lower accumulation of storage biomolecules. Gross photosynthesis (measured both as O2 evolution and 14C fixation) was not affected in any of them, suggesting that photosynthesis was already saturated at normal CO2 conditions and did not participate in the acclimation response. However, electron transport rate changed in both species in opposite directions, indicating different energy requirements between treatments and species specificity. High CO2 levels also affected the N-metabolism, and 13C isotopic discrimination values from algal tissue pointed to a deactivation of carbon concentrating mechanisms. Since increased CO2 has the potential to modify physiological mechanisms in different ways in the species studied, it is expected that this may lead to changes in the Arctic seaweed community, which may propagate to the rest of the food web.

Strong shift from HCO3- to CO2 uptake in Emiliania huxleyi with acidification: new approach unravels acclimation versus short-term pH effects

Effects of ocean acidification on Emiliania huxleyi strain RCC 1216 (calcifying, diploid life-cycle stage) and RCC 1217 (non-calcifying, haploid life-cycle stage) were investigated by measuring growth, elemental composition, and production rates under different pCO2 levels (380 and 950 µatm). In these differently acclimated cells, the photosynthetic carbon source was assessed by a (14)C disequilibrium assay, conducted over a range of ecologically relevant pH values (7.9-8.7). In agreement with previous studies, we observed decreased calcification and stimulated biomass production in diploid cells under high pCO2, but no CO2-dependent changes in biomass production for haploid cells. In both life-cycle stages, the relative contributions of CO2 and HCO3 (-) uptake depended strongly on the assay pH. At pH values =< 8.1, cells preferentially used CO2 (>= 90 % CO2), whereas at pH values >= 8.3, cells progressively increased the fraction of HCO3 (-) uptake (~45 % CO2 at pH 8.7 in diploid cells; ~55 % CO2 at pH 8.5 in haploid cells). In contrast to the short-term effect of the assay pH, the pCO2 acclimation history had no significant effect on the carbon uptake behavior. A numerical sensitivity study confirmed that the pH-modification in the (14)C disequilibrium method yields reliable results, provided that model parameters (e.g., pH, temperature) are kept within typical measurement uncertainties. Our results demonstrate a high plasticity of E. huxleyi to rapidly adjust carbon acquisition to the external carbon supply and/or pH, and provide an explanation for the paradoxical observation of high CO2 sensitivity despite the apparently high HCO3 (-) usage seen in previous studies.

Acclimation of Arabidopsis Leaves Developing at Low Temperatures. Increasing Cytoplasmic Volume Accompanies Increased Activities of Enzymes in the Calvin Cycle and in the Sucrose-Biosynthesis Pathway1

Photosynthetic and metabolic acclimation to low growth temperatures were studied in Arabidopsis (Heynh.). Plants were grown at 23°C and then shifted to 5°C. We compared the leaves shifted to 5°C for 10 d and the new leaves developed at 5°C with the control leaves on plants that had been left at 23°C. Leaf development at 5°C resulted in the recovery of photosynthesis to rates comparable with those achieved by control leaves at 23°C. There was a shift in the partitioning of carbon from starch and toward sucrose (Suc) in leaves that developed at 5°C. The recovery of photosynthetic capacity and the redirection of carbon to Suc in these leaves were associated with coordinated increases in the activity of several Calvin-cycle enzymes, even larger increases in the activity of key enzymes for Suc biosynthesis, and an increase in the phosphate available for metabolism. Development of leaves at 5°C also led to an increase in cytoplasmic volume and a decrease in vacuolar volume, which may provide an important mechanism for increasing the enzymes and metabolites in cold-acclimated leaves. Understanding the mechanisms underlying such structural changes during leaf development in the cold could result in novel approaches to increasing plant yield.

Acclimation of Photosynthesis to Elevated CO2 under Low-Nitrogen Nutrition Is Affected by the Capacity for Assimilate Utilization. Perennial Ryegrass under Free-Air CO2 Enrichment1

Acclimation of photosynthesis to elevated CO2 has previously been shown to be more pronounced when N supply is poor. Is this a direct effect of N or an indirect effect of N by limiting the development of sinks for photoassimilate? This question was tested by growing a perennial ryegrass (Lolium perenne) in the field under elevated (60 Pa) and current (36 Pa) partial pressures of CO2 (pCO2) at low and high levels of N fertilization. Cutting of this herbage crop at 4- to 8-week intervals removed about 80% of the canopy, therefore decreasing the ratio of photosynthetic area to sinks for photoassimilate. Leaf photosynthesis, in vivo carboxylation capacity, carbohydrate, N, ribulose-1,5-bisphosphate carboxylase/oxygenase, sedoheptulose-1,7-bisphosphatase, and chloroplastic fructose-1,6-bisphosphatase levels were determined for mature lamina during two consecutive summers. Just before the cut, when the canopy was relatively large, growth at elevated pCO2 and low N resulted in significant decreases in carboxylation capacity and the amount of ribulose-1,5-bisphosphate carboxylase/oxygenase protein. In high N there were no significant decreases in carboxylation capacity or proteins, but chloroplastic fructose-1,6-bisphosphatase protein levels increased significantly. Elevated pCO2 resulted in a marked and significant increase in leaf carbohydrate content at low N, but had no effect at high N. This acclimation at low N was absent after the harvest, when the canopy size was small. These results suggest that acclimation under low N is caused by limitation of sink development rather than being a direct effect of N supply on photosynthesis.

The Regulation of Photosynthetic Electron Transport during Nutrient Deprivation in Chlamydomonas reinhardtii1

The light-saturated rate of photosynthetic O2 evolution in Chlamydomonas reinhardtii declined by approximately 75% on a per-cell basis after 4 d of P starvation or 1 d of S starvation. Quantitation of the partial reactions of photosynthetic electron transport demonstrated that the light-saturated rate of photosystem (PS) I activity was unaffected by P or S limitation, whereas light-saturated PSII activity was reduced by more than 50%. This decline in PSII activity correlated with a decline in both the maximal quantum efficiency of PSII and the accumulation of the secondary quinone electron acceptor of PSII nonreducing centers (PSII centers capable of performing a charge separation but unable to reduce the plastoquinone pool). In addition to a decline in the light-saturated rate of O2 evolution, there was reduced efficiency of excitation energy transfer to the reaction centers of PSII (because of dissipation of absorbed light energy as heat and because of a transition to state 2). These findings establish a common suite of alterations in photosynthetic electron transport that results in decreased linear electron flow when C. reinhardtii is limited for either P or S. It was interesting that the decline in the maximum quantum efficiency of PSII and the accumulation of the secondary quinone electron acceptor of PSII nonreducing centers were regulated specifically during S-limited growth by the SacI gene product, which was previously shown to be critical for the acclimation of C. reinhardtii to S limitation (J.P. Davies, F.H. Yildiz, and A.R. Grossman [1996] EMBO J 15: 2150–2159).