Acclimation to high irradiance in temperate deciduous trees in the field: changes in xanthophyll cycle pool size and in photosynthetic capacity along a canopy light gradient

To test the hypothesis that in temperate deciduous trees acclimation to potentially damaging high irradiances occurs via long-term adjustments in foliar photosynthetic capacity, and short-term changes in xanthophyll cycle pool size in response to weather fluctuations, nitrogen concentration and pigment composition were examined along a canopy light gradient in three species –Betula pendula, Populus tremula and Tilia cordata (from most shade intolerant to tolerant), and foliage photosynthetic potentials in P. tremula and T. cordata. Integrated quantum flux density (Q_i) incident on leaves was estimated with a method combining hemispherical photography and light measurements with quantum sensors made over the growing season. Long- and short-term light indices – average total seasonal daily integrated quantum flux density (T_s, mol m^–2 d^–1) and that of the 3 d preceding foliage sampling (T_3d) – were calculated for each sampled leaf. In addition to total integrated quantum flux density, the part of Q_i attributable to direct flux was also computed. Strong linear relationships between the capacity for photosynthetic electron transport per area (J^a_max), estimated from in situ measurements of effective quantum yield of photosystem II (PS II), and Q_i averaged over the season and over the preceding 3 d were found for all studied species. However, the major determinant of J^a_max, the product of electron transport capacity per leaf dry mass (J^m_max) and leaf dry mass per area (M_A), was M_A rather than J^m_max, which was relatively constant along the light gradient. There was evidence that J^a_max is more tightly related to T_s, which characterizes the light climate during foliar development, than to short-term integrated light, possibly because there is little flexibility in adjustments in M_A after the completion of foliar growth. Leaf chlorophyll concentrations and the investment of leaf nitrogen in chlorophyll (Chl/N) were negatively related to Q_i– an investment pattern which improves light harvesting in low light. Xanthophyll cycle pool size (VAZ, violaxanthin + antheraxanthin + zeaxanthin) either expressed per unit chlorophyll (VAZ/Chl) or as a fraction of total carotenoids (VAZ/Car) increased with increasing Q_i in all species. However, contrary to J^a_max, it tended to correlate more strongly with short-term than with long-term average integrated light. There were few interspecific differences in J^a_max, Chl/N, VAZ/Chl and VAZ/Car when the variability in light level incident to the leaves was accounted for, indicating that the foliage of both shade-intolerant and -tolerant temperate tree species possesses considerable phenotypic flexibility. Collectively these results support the view that rapid adjustment of the xanthophyll cycle pool size provides an important means for acclimation to light fluctuations in a time scale of days, during which the potential for photosynthetic quenching of excitation energy is not likely to change appreciably.     

CYANOBACTERIAL ACCLIMATION TO RAPIDLY FLUCTUATING LIGHT IS CONSTRAINED BY INORGANIC CARBON STATUS1

Acclimation to rapidly fluctuating light, simulating shallow aquatic habitats, is altered depending on inorganic carbon (C_i) availability. Under steady light of 50 μmol photons·m^?2·s^?1, the growth rate of Synechococcus elongatus PCC7942 was similar in cells grown in high C_i (4 mM) and low C_i (0.02 mM), with induced carbon concentrating mechanisms compensating for low C_i. Growth under fluctuating light of a 1‐s period averaging 50 μmol photons·m^?2·s^?1 caused a drop in growth rate of 28%±6% in high C_i cells and 38%±8% in low C_i cells. In high C_i cells under fluctuating light, the PSI/PSII ratio increased, the PSII absorption cross‐section decreased, and the PSII turnover rate increased in a pattern similar to high‐light acclimation. In low C_i cells under fluctuating light, the PSI/PSII ratio decreased, the PSII absorption cross‐section decreased, and the PSII turnover remained slow. Electron transport rate was similar in high and low C_i cells but in both was lower under fluctuating than under steady light. After acclimation to a 1‐s period fluctuating light, electron transport rate decreased under steady or long‐period fluctuating light. We hypothesize that high C_i cells acclimated to exploit the bright phases of the fluctuating light, whereas low C_i cells enlarged their PSII pool to integrate the fluctuating light and dampen the variation of the electron flux into a rate‐restricted C_i pool. Light response curves measured under steady light, widely used to predict photosynthetic rates, do not properly predict photosynthetic rates achieved under fluctuating light, and exploitation of fluctuating light is altered by C_i status.     

The photosynthetic characteristics of papyrus in a tropical swamp

Summary Photosynthesis and transpiration was measured in the large emergent C₄ sedge Cyperus papyrus (papyrus) which occupies wide areas of wetland on the African continent. The maximum observed value of net assimilation was 35 mol CO₂ m^-2 s^-1 at full sunlight but light saturation of photosynthesis did not occur. The quantum yield of photosynthesis obtained from the initial slope of the light response curves (0.06 mol mol^-1 incident light) was relatively high and close to previously recorded values for some C₄ grasses. Measurements made over two days showed that stomatal conductance was sensitive to the ambient air vapour pressure deficit (VPD) and was consistently lower on the day when VPD's were higher. There was, however, no marked midday closure of the stomata. Photosynthesis was also reduced on the day when VPD's were higher. The relationship between net photosynthesis and stomatal conductance was close to linear over the range of measurement conditions, with the result that intercellular CO₂ concentrations (C _i ) did not vary markedly. There was some evidence that C _i decreased at high VPD's. The regulation of stomatal movement in papyrus appears to minimise excessive water loss while not severely limiting photosynthesis. The significance of this strategy for a wetland species with plentiful supplies of water is discussed.     

The effect of light quality on the induction of efficient photosynthesis under low CO2 conditions in Chlamydomonas reinhardtii and Chlorella pyrenoidosa

The effect of blue and red light on the adaptation to low CO₂ conditions was studied in high-CO₂ grown cultures of Chlorella Pyrenoidosa (82T) and Chlamydomonas reinhardtii(137⁺) by measuring O₂ exchange under various inorganic carbon (Cⁱ) concentrations. At equal photosynthetic photon flux density (PPFD), blue light was more favourable for adaptation in both species, compared to red light. The difference in photosynthetic oxygen evolution between cells adapted to low C_iunder blue and red light was more pronounced when oxygen evolution was measured under low C_i compared to high C_i conditions. The effect of light quality on adaptation remained for several hours. The different effects caused by blue and red light was observed in C. pyrenoidosa over a wide range of PPFD with increasing differences at increasing PPFD. The maximal difference was obtained at a PPFD above 1 500 μmol m^?2s^?1. We found no difference in the extracellular carbonic anhydrase activity between blue- and red light adapted cells. The light quality effect recorded under C_i-limiting conditions in C. reinhardtii cells adapted to air, was only 37% less when instead of pure blue light red light containing 12.5% of blue light (similar PPFD as blue light) was used during adaptation to low carbon. This indicates that in addition to affecting photosynthesis, blue light affected a sensory system involved in algal adaptation to low C_i conditions. Since the affinity for C_i of C. Pyrenoidosa and C. reinhardtii cells adapted to air under blue light was higher than that of cells adapted under red light, we suggest that induction of some component(s) of the C_i accumulating mechanism is regulated by the light quality.     

Phosphorus recycling in photorespiration maintains high photosynthetic capacity in woody species

Leaf photosynthetic CO₂ responses can provide insight into how major nutrients, such as phosphorus (P), constrain leaf CO₂ assimilation rates (A_net). However, triose‐phosphate limitations are rarely employed in the classic photosynthesis model and it is uncertain as to what extent these limitations occur in field situations. In contrast to predictions from biochemical theory of photosynthesis, we found consistent evidence in the field of lower A_net in high [CO₂] and low [O₂] than at ambient [O₂]. For 10 species of trees and shrubs across a range of soil P availability in Australia, none of them showed a positive response of A_net at saturating [CO₂] (i.e. A_max) to 2 kPa O₂. Three species showed >20% reductions in A_max in low [O₂], a phenomenon potentially explained by orthophosphate (P_i) savings during photorespiration. These species, with largest photosynthetic capacity and P_i > 2 mmol P m^?2, rely the most on additional P_i made available from photorespiration rather than species growing in P‐impoverished soils. The results suggest that rarely used adjustments to a biochemical photosynthesis model are useful for predicting A_max and give insight into the biochemical limitations of photosynthesis rates at a range of leaf P concentrations. Phosphate limitations to photosynthetic capacity are likely more common in the field than previously considered.     

Photosynthetic characteristics of dwarf and fringe Rhizophora mangle L. in a Belizean mangrove

Twin Cays (Belize) is a highly oligotrophic mangrove archipelago dominated by Rhizophora mangle L. Ocean‐fringing trees are 3–7 m tall with a leaf area index (LAI) of 2.3, whereas in the interior, dwarf zone, trees are 1.5 m or less, and the LAI is 0.7. P‐fertilization of dwarf trees dramatically increases growth. As a partial explanation of these characteristics, it was hypothesized that differences in stature and growth rates would reflect differences in leaf photosynthetic capacity, as determined by the photochemical and biochemical characteristics at the chloroplast level. Gas exchange and chlorophyll fluorescence were used to compare photosynthesis of dwarf, fringe and fertilized trees. Regardless of zonation or treatment, net CO₂ exchange (A) and photosynthetic electron transport were light saturated at less than 500 µmol photons m^?2 s^?1, and low‐light quantum efficiencies were typical for healthy C₃ plants. On the other hand, light‐saturated A was linearly related to stomatal conductance (g_s), with seasonal, zonal and treatment differences in photosynthesis corresponding linearly to differences in the mean g_s. Overall, photosynthetic capacity appeared to be co‐regulated with stomatal conductance, minimizing the variability of C_i at ambient CO₂ (and hence, C_i/C_a). Based on the results of in vitro assays, regulation of photosynthesis in R. mangle appeared to be accomplished, at least in part, by regulation of Rubisco activity.     

高大气CO2浓度下氮素对小麦叶片光能利用的影响

关于氮素对高大气CO₂浓度下C₃植物光合作用适应现象的调节机理已有较为深入的研究, 但对其光合作用适应现象的光合能量转化和分配机制缺乏系统分析。该文以大气CO₂浓度和施氮量为处理手段, 通过测定小麦(Triticum aestivum)抽穗期叶片的光合作用-胞间CO₂浓度响应曲线以及荧光动力学参数来测算光合电子传递速率和分配去向, 研究了长期高大气CO₂浓度下小麦叶片光合电子传递和分配对施氮量的响应。结果表明, 与正常大气CO₂浓度处理相比, 高大气CO₂浓度下小麦叶片较多的激发能以热量的形式耗散, 增施氮素可使更多的激发能向光化学反应方向的分配, 降低光合能量的热耗散速率; 大气CO₂浓度升高后小麦叶片光化学淬灭系数无明显变化, 高氮叶片的非光化学猝灭降低而低氮叶片明显升高, 施氮促进PSII反应中心的开放比例, 降低光能的热耗散; 高大气CO₂浓度下高氮叶片通过PSII反应中心的光合电子传递速率(J_F)较高, 而且参与光呼吸的非环式电子流速率(J₀)显著降低, 较正常大气CO₂浓度处理的高氮叶片下降了88.40%, 光合速率增加46.47%; 高大气CO₂浓度下小麦叶片J_F-J₀升高而J₀/J_F显著下降, 光呼吸耗能被抑制, 更多的光合电子分配至光合还原过程。因此, 大气CO₂浓度增高条件下, 小麦叶片激发能的热耗散速率增加, 但增施氮素后小麦叶片PSII反应中心开放比例提高, 光化学速率增加, 进入PSII反应中心的电子流速率明显升高, 光呼吸作用被抑制, 光合电子较多地进入光化学过程, 这可能是高氮条件下光合作用适应性下调被缓解的一个原因。     

No photosynthetic down-regulation in sweetgum trees (Liquidambar styraciflua L.) after three years of CO2 enrichment at the Duke Forest FACE experiment

Photosynthetic capacity and leaf properties of sun and shade leaves of overstorey sweetgum trees (Liquidambar styraciflua L.) were compared over the first 3 years of growth in ambient or ambient + 200 μL L^?1 CO₂ at the Duke Forest Free Air CO₂ Enrichment (FACE) experiment. We were interested in whether photosynthetic down‐regulation to CO₂ occurred in sweetgum trees growing in a forest ecosystem, whether shade leaves down‐regulated to a greater extent than sun leaves, and if there was a seasonal component to photosynthetic down‐regulation. During June and September of each year, we measured net photosynthesis (A) versus the calculated intercellular CO₂ concentration (C_i) in situ and analysed these response curves using a biochemical model that described the limitations imposed by the amount and activity of ribulose‐1,5‐bisphosphate carboxylase/oxygenase (Vc_max) and by the rate of ribulose‐1,5‐bisphosphate (RuBP) regeneration mediated by electron transport (J_max). There was no evidence of photosynthetic down‐regulation to CO₂ in either sun or shade leaves of sweetgum trees over the 3 years of measurements. Elevated CO₂ did not significantly affect Vc_max or J_max. The ratio of Vc_max to J_max was relatively constant, averaging 2·12, and was not affected by CO₂ treatment, position in the canopy, or measurement period. Furthermore, CO₂ enrichment did not affect leaf nitrogen per unit leaf area (N_a), chlorophyll or total non‐structural carbohydrates of sun or shade leaves. We did, however, find a strong relationship between N_a and the modelled components of photosynthetic capacity, Vc_max and J_max. Our data over the first 3 years of this experiment corroborate observations that trees rooted in the ground may not exhibit symptoms of photosynthetic down‐regulation as quickly as tree seedlings growing in pots. There was a strong sustained enhancement of photosynthesis by CO₂ enrichment whereby light‐saturated net photosynthesis of sun leaves was stimulated by 63% and light‐saturated net photosynthesis of shade leaves was stimulated by 48% when averaged over the 3 years. This study suggests that this CO₂ enhancement of photosynthesis will be sustained in the Duke Forest FACE experiment as long as soil N availability keeps pace with photosynthetic and growth processes.     

Photosynthetic acclimation in trees to rising atmospheric CO2: A broader perspective

Analysis of leaf-level photosynthetic responses of 39 tree species grown in elevated concentrations of atmospheric CO₂ indicated an average photosynthetic enhancement of 44% when measured at the growth [CO₂]. When photosynthesis was measured at a common ambient [CO₂], photosynthesis of plants grown at elevated [CO₂] was reduced, on average, 21% relative to ambient-grown trees, but variability was high. The evidence linking photosynthetic acclimation in trees with changes at the biochemical level is examined, along with anatomical and morphological changes in trees that impact leaf- and canopy-level photosynthetic response to CO₂ enrichment. Nutrient limitations and variations in sink strength appear to influence photosynthetic acclimation, but the evidence in trees for one predominant factor controlling acclimation is lacking. Regardless of the mechanisms that underlie photosynthetic acclimation, it is doubtful that this response will be complete. A new focus on adjustments to rising [CO₂] at canopy, stand, and forest scales is needed to predict ecosystem response to a changing environment.Abbreviations A/C_i photosynthesis as a function of internal [CO₂] - J_max maximum rate of electron transport - Rubisco ribulose-1,5-bisphosphate carboxylase/oxygenase - Vc_max maximum rate of carboxylation The U.S. Government right to retain a non-exclusive, royalty free licence in and to any copyright is acknowledged.     

The quantum efficiency of photosynthesis in macroalgae and submerged angiosperms

Summary Photon absorption and photosynthesis under conditions of light limitation were determined in six temperate marine macroalgae and eight submerged angiosperms. Photon absorption and photosynthetic efficiency based on incident light increased in proportion to chlorophyll density per area and approached saturation at the highest densities (300 mg chlorophyll m^–2) encountered. Absorption and photosynthetic efficiency were higher in brown and red algae than in green algae and angiosperms for the same chlorophyll density because of absorption by accessory pigments. Among thin macroalgae and submerged angiosperms chlorophyll variations directly influence light absorption and photosynthesis, whereas terrestrial leaves have chlorophyll in excess and thus there is only a minor influence of pigment variability on light-limited photosynthesis. The quantum efficiency of photosynthesis averaged 0.062±0.019 (±SD) mol O₂ mol^–1 photons absorbed for macroalgae and, significantly less, 0.049±0.016 mol O₂ mol^–1 photons for submerged angiosperms. Of the measurements 80% were between 0.037 and 0.079 mol O₂ mol^–1 photons. The results are lower than values given in the literature for unicellular algae and terrestrial C₃ species at around 0.1 mol O₂ mol^–1 photons, but resemble values for other marine macroalgae and terrestrial C₄ species. The reason for these differences remains unknown, but may be sought for in differential operation of cyclic photophosphorylation and photorespiration.     

Scaling of photosynthetic production of aquatic macrophytes – a review

Most studies on photosynthetic production of aquatic macrophytes have been made on detached leaves and algal thalli. This may have given the false impression that production is often saturated by light and that inorganic carbon and nutrients are more important limiting factors. However, studies on the more relevant ecological scale of macrophyte communities lead to a completely different perception because community production is light limited due to intense self‐shading. Relatively high irradiances are needed for photosynthesis to balance respiratory costs and not even maximum irradiances at noon during summer saturate photosynthetic production. The fundamental importance of light is confirmed by the close coupling to community light absorptance of both maximum production in high‐light environments and efficiency of light use in low‐light environments. The upper boundaries of light‐limited and light‐saturated production are distinctly and linearly related to community absorptance. Moreover, higher diversity in the community has a positive and stabilizing influence on light absorptance and production because different species supplement each other temporally and spatially. Close predictions of actual production rates in macroalgal communities throughout the year are possible solely from determinations of incident and absorbed irradiance. Along with the increasing regulating role of light from leaves or thalli to entire communities the importance of temperature, inorganic carbon and other resources decreases. Thus, ten‐fold rise of CO₂ relative to atmospheric saturation does not enhance maximum production in dense communities of efficient HCO₃^?‐users and only doubles production of pure CO₂‐users. A challenge therefore exists establishing the importance of light for photosynthetic production of macrophytes from individuals to communities and re‐evaluating the postulated main importance of inorganic carbon and temperature in the scenarios of globally rising CO₂ and temperature.     

Light respiration by subtropical seaweeds

Here, we report the first‐ever measurements of light CO₂ respiration rate (CRR) by seaweeds. We measured the influence of temperature (15–25°C) and light (irradiance from 60 to 670 μmol · m^?2 · s^?1) on the light CCR of two subtropical seaweed species, and measured the CRR of seven different seaweed species under the same light (150 μmol · m^?2 · s^?1) and temperature (25°C). There was little effect of irradiance on light CRR, but there was an effect of temperature. Across the seven species light CRR was similar to OCR (oxygen consumption rate in the dark), with the exception of a single species. The outlier species was a coralline alga, and the higher light CRR was probably driven by calcification. CRR could be estimated from OCR, as well as carbon photosynthetic rates from oxygen photosynthetic rates, which suggests that previous studies have probably provided good estimations of gross photosynthesis for seaweeds.     

Do the capacity and kinetics for modification of xanthophyll cycle pool size depend on growth irradiance in temperate trees?

The present study investigated the interaction of growth irradiance (Q_int) with leaf capacity for and kinetics of adjustment of the pool size of xanthophyll cycle carotenoids (sum of violaxanthin, antheraxanthin and zeaxanthin; VAZ) and photosynthetic electron transport rate (J_max) after changes in leaf light environment. Individual leaves of lower‐canopy/lower photosynthetic capacity species Tilia cordata Mill. and upper canopy/higher photosynthetic capacity species Populus tremula L. were either illuminated by additional light of 500–800 µmol m^?2 s^?1 for 12 h photoperiod or enclosed in shade bags. The extra irradiance increased the total amount of light intercepted by two‐fold for the upper and 10–15‐fold for the lower canopy leaves, whereas the shade bags transmitted 45% of incident irradiance. In control leaves, VAZ/area, VAZ/Chl and J_max were positively associated with leaf growth irradiance (Q_int). After 11 d extra illumination, VAZ/Chl increased in all cases due to a strong reduction in foliar chlorophyll, but VAZ/area increased in the upper canopy leaves of both species, and remained constant or decreased in the lower canopy leaves of T. cordata. The slope for VAZ/area changes with cumulative extra irradiance was positively associated with Q_int only in T. cordata, but not in P. tremula. Nevertheless, all leaves of P. tremula increased VAZ/area more than the most responsive leaves of T. cordata. Shading reduced VAZ content only in P. tremula, but not in T. cordata, again demonstrating that P. tremula is a more responsive species. Compatible with the hypothesis of the role of VAZ in photoprotection, the rates of photosynthetic electron transport declined less in P. tremula than in T. cordata after the extra irradiance treatment. However, foliar chlorophyll contents of the exposed leaves declined significantly more in the upper canopy of P. tremula, which is not consistent with the suggestion that the leaves with the highest VAZ content are more resistant to photoinhibition. This study demonstrates that previous leaf light environment may significantly affect the adaptation capacity of foliage to altered light environment, and also that species differences in photosynthetic capacity and acclimation potentials importantly alter this interaction.     

Water availability affects seasonal CO2‐induced photosynthetic enhancement in herbaceous species in a periodically dry woodland

Elevated atmospheric CO₂ (eCO₂) is expected to reduce the impacts of drought and increase photosynthetic rates via two key mechanisms: first, through decreased stomatal conductance (g_s) and increased soil water content (V_SWC) and second, through increased leaf internal CO₂ (C_i) and decreased stomatal limitations (S_lim). It is unclear if such findings from temperate grassland studies similarly pertain to warmer ecosystems with periodic water deficits. We tested these mechanisms in three important C₃ herbaceous species in a periodically dry Eucalyptus woodland and investigated how eCO₂‐induced photosynthetic enhancement varied with seasonal water availability, over a 3 year period. Leaf photosynthesis increased by 10%–50% with a 150 μmol mol^?1 increase in atmospheric CO₂ across seasons. This eCO₂‐induced increase in photosynthesis was a function of seasonal water availability, given by recent precipitation and mean daily V_SWC. The highest photosynthetic enhancement by eCO₂ (>30%) was observed during the most water‐limited period, for example, with V_SWC <0.07 in this sandy surface soil. Under eCO₂ there was neither a significant decrease in g_s in the three herbaceous species, nor increases in V_SWC, indicating no “water‐savings effect” of eCO₂. Periods of low V_SWC showed lower g_s (less than ≈ 0.12 mol m^?2 s^?1), higher relative S_lim (>30%) and decreased C_i under the ambient CO₂ concentration (aCO₂), with leaf photosynthesis strongly carboxylation‐limited. The alleviation of S_lim by eCO₂ was facilitated by increasing C_i, thus yielding a larger photosynthetic enhancement during dry periods. We demonstrated that water availability, but not eCO₂, controls g_s and hence the magnitude of photosynthetic enhancement in the understory herbaceous plants. Thus, eCO₂ has the potential to alter vegetation functioning in a periodically dry woodland understory through changes in stomatal limitation to photosynthesis, not by the “water‐savings effect” usually invoked in grasslands.     

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.     

Mitochondrial uncoupling protein silence is compromised in photosynthesis and redox poise

Mitochondrial uncoupling proteins play important roles in generation of metabolic thermogenesis, response to stress situations, and regulation of energy metabolism. We demonstrated here that the absence of LeUCP in tomato inhibited photosynthesis using virus-induced gene-silencing approach. A significant decrease in the rate of CO₂ assimilation in LeUCP-silencing plants was observed over a range of different light intensities. Absence of LeUCP resulted in lower net photosynthetic rate, light-saturated rate of the CO₂ assimilation (A _sat), maximum carboxylation rates (V _cmax) and maximum RuBP regeneration rate (J _max). Activities of ribulose-1,5-bisphosphate carboxylase/oxygenase Rubisco and stromal fructose-1,6-bisphosphatase and genes expression levels encoded Calvin cycle enzymes of LeUCP gene-silencing plants were inhibited. However, silencing of LeUCP gene had no effect on F _v/F _m, but decreased photochemical quenching and electron transport rate. Meanwhile, non-photochemical quenching and Je (PSII), the distribution of Je (PCR) and Je (PCO), the content of AsA, NAD, and the ratios of NAD⁺/NADH, AsA/DHA were significantly reduced with increased reactive oxygen species while GSH and GSSG were unaltered. Taken together, these results strongly suggest that LeUCP gene in tomato leaves is crucial in maintaining the redox poise of the mitochondrial electron transport chain to facilitate photosynthetic metabolism.     

Changes in needle nitrogen partitioning and photosynthesis during 80 years of tree ontogeny in Picea abies

Needle nitrogen partitioning and photosynthesis of Norway spruce were studied in a forest chronosequence in Järvselja Experimental Forest, Estonia. Current- and previous-year shoots were sampled from upper and lower canopy positions in four stands, ranging in age from 13 to 82 years. A/c _i curves were determined to obtain maximum carboxylation rate (V _cmax) and maximum rate of electron transport (J _max), whereas needle nitrogen partitioning into carboxylation (P _R), bioenergetics associated with electron transport (P _B) and thylakoid light harvesting components (P _L) was calculated from the values of V _cmax, J _max and leaf chlorophyll concentration. The greatest changes in studied needle characteristics took place between tree ages of 13 and 26 years, and this pattern was independent of needle age and canopy position. Needle mass per projected area (LMA) was lowest in the 13-year-old stand and mass-based nitrogen concentration (N^M) was generally highest in that stand. The values of LMA were significantly higher and those of N^M lower in the 26-year-old stand. Mass-based V _cmax and J _max were highest in the 13-year-old stand. Area-based photosynthetic capacity was independent of tree age. The proportion of photosynthetic nitrogen (P _R, P _B and P _L) was highest and that of non-photosynthetic nitrogen lowest in the 13-year-old stand. Current-year needles had lower LMA and P _L, but higher photosynthetic capacity compared to 1-year-old foliage. Needles from lower canopy positions exhibited lower LMA, area-based nitrogen concentration and photosynthetic capacity than needles from upper canopy. The period of substantial reductions in needle photosynthetic capacity and changes in nitrogen partitioning coincides with the onset of reproductive phase during tree ontogeny.     

Light dependency of the photosynthetic recovery of Nostoc flagelliforme

PS II photochemical efficiency (F_v/F_m) of Nostoc flagelliforme was examined after rewetting in order to investigate the light-dependency of its photosynthetic recovery. F_v/F_m was not detected in the dark, but was immediately recognized in the light. Different levels of light irradiation (4, 40 and 400 μmol photon m² s^-1) displayed different effects on the recovery process of photosynthesis. The intermediate level led to the best recovery of photochemical efficiency; the low light required longer and the high light inhibited the extent of the recovered efficiency. It was concluded that the photosynthetic recovery of N. flagelliforme is both light-dependent and influenced by photon flux density. This revised version was published online in June 2006 with corrections to the Cover Date.     

Steady-state stomatal responses of C3 and C4 species to blue light fraction: Interactions with CO2 concentration

Blue light induced stomatal opening has been studied by applying a short pulse (~5 to 60 s) of blue light to a background of saturating photosynthetic red photons, but little is known about steady-state stomatal responses. Here we report stomatal responses to blue light at high and low CO₂ concentrations. Steady-state stomatal conductance (g_s) of C₃ plants increased asymptotically with increasing blue light to a maximum at 20% blue (120 μmol m⁻² s⁻¹). This response was consistent from 200 to 800 μmol mol⁻¹ atmospheric CO₂ (C_a). In contrast, blue light induced only a transient stomatal opening (~5 min) in C₄ species above a C_a of 400 μmol mol⁻¹. Steady-state g_s of C₄ plants generally decreased with increasing blue intensity. The net photosynthetic rate of all species decreased above 20% blue because blue photons have lower quantum yield (moles carbon fixed per mole photons absorbed) than red photons. Our findings indicate that photosynthesis, rather than a blue light signal, plays a dominant role in stomatal regulation in C₄ species. Additionally, we found that blue light affected only stomata on the illuminated side of the leaf. Contrary to widely held belief, the blue light-induced stomatal opening minimally enhanced photosynthesis and consistently decreased water use efficiency.     

In situ temperature relationships of biochemical and stomatal controls of photosynthesis in four lowland tropical tree species

Net photosynthetic carbon uptake of Panamanian lowland tropical forest species is typically optimal at 30–32 °C. The processes responsible for the decrease in photosynthesis at higher temperatures are not fully understood for tropical trees. We determined temperature responses of maximum rates of RuBP‐carboxylation (V_CMax) and RuBP‐regeneration (J_Max), stomatal conductance (G_s), and respiration in the light (R_Light) in situ for 4 lowland tropical tree species in Panama. G_s had the lowest temperature optimum (T_Opt), similar to that of net photosynthesis, and photosynthesis became increasingly limited by stomatal conductance as temperature increased. J_Max peaked at 34–37 °C and V_CMax ~2 °C above that, except in the late‐successional species Calophyllum longifolium, in which both peaked at ~33 °C. R_Light significantly increased with increasing temperature, but simulations with a photosynthesis model indicated that this had only a small effect on net photosynthesis. We found no evidence for Rubisco‐activase limitation of photosynthesis. T_Opt of V_CMax and J_Max fell within the observed in situ leaf temperature range, but our study nonetheless suggests that net photosynthesis of tropical trees is more strongly influenced by the indirect effects of high temperature—for example, through elevated vapour pressure deficit and resulting decreases in stomatal conductance—than by direct temperature effects on photosynthetic biochemistry and respiration.