Seasonal patterns of photosynthetic response and acclimation to elevated carbon dioxide in field-grown strawberry

Strawberry (Fragaria × ananassa) plants were grown in field plots at the current ambient [CO₂], and at ambient + 300 and ambient + 600 μmol mol⁻¹ [CO₂]. Approximately weekly measurements were made of single leaf gas exchange of upper canopy leaves from early spring through fall of two years, in order to determine the temperature dependence of the stimulation of photosynthesis by elevated [CO₂], whether growth at elevated [CO₂] resulted in acclimation of photosynthesis, and whether any photosynthetic acclimation was reduced when fruiting created additional demand for the products of photosynthesis. Stimulation of photosynthetic CO₂ assimilation by short-term increases in [CO₂] increased strongly with measurement temperature. The stimulation exceeded that predicted from the kinetic characteristics of ribulose-1,5-bisphosphate carboxylase at all temperatures. Acclimation of photosynthesis to growth at elevated [CO₂] was evident from early spring through summer, including the fruiting period in early summer, with lower rates under standard measurement conditions in plants grown at elevated [CO₂]. The degree of acclimation increased with growth [CO₂]. However, there were no significant differences between [CO₂] treatments in total nitrogen per leaf area, and photosynthetic acclimation was reversed one day after switching the [CO₂] treatments. Tests showed that acclimation did not result from a limitation of photosynthesis by triose phosphate utilization rate at elevated [CO₂]. Photosynthetic acclimation was not evident during dry periods in midsummer, when the elevated [CO₂] treatments conserved soil water and photosynthesis declined more at ambient than at elevated [CO₂]. Acclimation was also not evident during the fall, when plants were vegetative, despite wet conditions and continued higher leaf starch content at elevated [CO₂]. Stomatal conductance responded little to short-term changes in [CO₂] except during drought, and changed in parallel with photosynthetic acclimation through the seasons in response to the long-term [CO₂] treatments. The data do not support the hypothesis that source-sink balance controls the seasonal occurrence of photosynthetic acclimation to elevated [CO₂] in this species. This revised version was published online in June 2006 with corrections to the Cover Date.     

The temporal and species dynamics of photosynthetic acclimation in flag leaves of rice (Oryza sativa) and wheat (Triticum aestivum) under elevated carbon dioxide

In this study, we tested for the temporal occurrence of photosynthetic acclimation to elevated [CO₂] in the flag leaf of two important cereal crops, rice and wheat. In order to characterize the temporal onset of acclimation and the basis for any observed decline in photosynthetic rate, we characterized net photosynthesis, g_s, g_m, C_i/C_a, C_i/C_c, V_cmax, J_max, cell wall thickness, content of Rubisco, cytochrome (Cyt) f, N, chlorophyll and carbohydrate, mRNA expression for rbcL and petA, activity for Rubisco, sucrose phosphate synthase (SPS) and sucrose synthase (SS) at full flag expansion, mid‐anthesis and the late grain‐filling stage. No acclimation was observed for either crop at full flag leaf expansion. However, at the mid‐anthesis stage, photosynthetic acclimation in rice was associated with RuBP carboxylation and regeneration limitations, while wheat only had the carboxylation limitation. By grain maturation, the decline of Rubisco content and activity had contributed to RuBP carboxylation limitation of photosynthesis in both crops at elevated [CO₂]; however, the sharp decrease of Rubisco enzyme activity played a more important role in wheat. Although an increase in non‐structural carbohydrates did occur during these later stages, it was not consistently associated with changes in SPS and SS or photosynthetic acclimation. Rather, over time elevated [CO₂] appeared to enhance the rate of N degradation and senescence so that by late‐grain fill, photosynthetic acclimation to elevated [CO₂] in the flag leaf of either species was complete. These data suggest that the basis for photosynthetic acclimation with elevated [CO₂] may be more closely associated with enhanced rates of senescence, and, as a consequence, may be temporally dynamic, with significant species variation.     

Effect of panicle removal on photosynthetic acclimation under elevated CO<Subscript>2</Subscript> in rice

To examine the role of sink size on photosynthetic acclimation under elevated atmospheric CO₂ concentrations ([CO₂]), we tested the effects of panicle-removal (PR) treatment on photosynthesis in rice (Oryza sativa L.). Rice was grown at two [CO₂] levels (ambient and ambient + 200 μmol mol⁻¹) throughout the growing season, and at full-heading stage, at half the plants, a sink-limitation treatment was imposed by the removal of the panicles. The PR treatment alleviated the reduction of green leaf area, the contents of chlorophyll (Chl) and Rubisco after the full-heading stage, suggesting delay of senescence. Nonetheless, elevated [CO₂] decreased photosynthesis (measured at current [CO₂]) of plants exposed to the PR treatment. No significant [CO₂] × PR interaction on photosynthesis was observed. The decrease of photosynthesis by elevated [CO₂] of plants was associated with decreased leaf Rubisco content and N content. Leaf glucose content was increased by the PR treatment and also by elevated [CO₂]. In conclusion, a sink-limitation in rice improved N status in the leaves, but this did not prevent the photosynthetic down-regulation under elevated [CO₂].     

Integration of photosynthetic acclimation to CO2 at the whole-plant level

Primary events in photosynthetic (PS) acclimation to elevated CO₂ concentration ([CO₂]) occur at the molecular level in leaf mesophyll cells, but final growth response to [CO₂] involves acclimation responses associated with photosynthate partitioning among plant organs in relation to resources limiting growth. Source–sink interactions, particularly with regard to carbon (C) and nitrogen (N), are key determinants of PS acclimation to elevated [CO₂] at the whole-plant level. In the long term, PS and growth response to [CO₂] are dependent on genotypic and environmental factors affecting the plant's ability to develop new sinks for C, and acquire adequate N and other resources to support an enhanced growth potential. Growth at elevated [CO₂] usually increases N use efficiency because PS rates can be maintained at levels comparable to those observed at ambient [CO₂] with less N investment in PS enzymes. A frequent acclimation response, particularly under N-limited conditions, is for the accumulation of leaf carbohydrates at elevated [CO₂] to lead to repression of genes associated with the production of PS enzymes. The hypothesis that this is an adaptive response, leading to a diversion of N to plant organs where it is of greatest benefit in terms of competitive ability and reproductive fitness, needs to be more rigorously tested. The biological control mechanisms which plants have evolved to acclimate to shifts in source–sink balance caused by elevated [CO₂] are complex, and will only be fully elucidated by probing at all scales along the hierarchy from molecular to ecosystem. Use of environmental manipulations and genotypic comparisons will facilitate the testing of specific hypotheses. Improving our ability to predict PS acclimation to [CO₂] will require the integration of results from laboratory studies using simple model systems with results from whole-plant studies that include measurements of processes operating at several scales. Abbreviations: CAM, crassulacean acid metabolism; FACE, Free-Air CO₂ Enrichment; Pi, inorganic phosphate; LAR, leaf area ratio (m² g^-1); LWR, leaf weight ratio (g g^-1); NAR, net assimilation rate (g m^-2 d^{- 1}); PS, photosynthetic; RGR, relative growth rate (g g^-1 d^-1); R:S, root/shoot ratio; rubisco, ribulose bisphosphate carboxylase/oxygenase; RuBP, ribulose bisphosphate; SLA, specific leaf area (m² g^-1); SPS, sucrose phosphate synthase; WUE, water use efficiency (g biomass g H₂O^-1).     

Effects of fully open-air [CO2] elevation on leaf ultrastructure, photosynthesis, and yield of two soybean cultivars

The objective of this study was to investigate the effect of elevated (550 ± 17 ??mol mol^?1) CO₂ concentration ([CO₂]) on leaf ultrastructure, leaf photosynthesis and seed yield of two soybean cultivars [Glycine max (L.) Merr. cv. Zhonghuang 13 and cv. Zhonghuang 35] at the Free-Air Carbon dioxide Enrichment (FACE) experimental facility in North China. Photosynthetic acclimation occurred in soybean plants exposed to long-term elevated [CO₂] and varied with cultivars and developmental stages. Photosynthetic acclimation occurred at the beginning bloom (R1) stage for both cultivars, but at the beginning seed (R5) stage only for Zhonghuang 13. No photosynthetic acclimation occurred at the beginning pod (R3) stage for either cultivar. Elevated [CO₂] increased the number and size of starch grains in chloroplasts of the two cultivars. Soybean leaf senescence was accelerated under elevated [CO₂], determined by unclear chloroplast membrane and blurred grana layer at the beginning bloom (R1) stage. The different photosynthesis response to elevated [CO₂] between cultivars at the beginning seed (R5) contributed to the yield difference under elevated [CO₂]. Elevated [CO₂] significantly increased the yield of Zhonghuang 35 by 26% with the increased pod number of 31%, but not for Zhonghuang 13 without changes of pod number. We conclude that the occurrence of photosynthetic acclimation at the beginning seed (R5) stage for Zhonghuang 13 restricted the development of extra C sink under elevated [CO₂], thereby limiting the response to elevated [CO₂] for the seed yield of this cultivar.     

Elevated CO2 and limited nitrogen nutrition can restrict excitation energy dissipation in photosystem II of Japanese white birch (Betula platyphylla var. japonica) leaves

Elevated atmospheric CO₂ concentration [CO₂] and different levels of nitrogen (N) nutrition can influence the amount of excess excitation energy in photosystem (PS) II and related photosynthetic properties. The interactive effect of two [CO₂] levels (ambient: 360 µM M⁻¹ and elevated: 720 µM M⁻¹) and two N levels (high: 700 mg N plant⁻¹ and low: 100 mg N plant⁻¹) on these properties was examined in seedlings of Japanese white birch (Betula platyphylla var. japonica) using simultaneous measurements of gas exchange and chlorophyll fluorescence. Photosynthetic acclimation to elevated [CO₂], as indicated by a decline in carboxylation efficiency (CE), was observed in plants grown at elevated [CO₂] especially under low N. Elevated [CO₂] resulted in a decrease in area-based leaf N content (N_area) irrespective of N treatment. The adverse effect of elevated [CO₂] and low N on CE may have been exacerbated by a greater accumulation of leaf sugar and starch contents in these plants leading to a lower electron transport rate (ETR). While these plants also showed higher non-photochemical quenching (Nq_P) that could offset the reduction in energy dissipation through ETR to some extent, they still have a higher risk of photoinhibition from excessive excitation energy in PSII as indicated by a decrease in photochemical quenching (q_P). However, chronic photoinhibition was not observed in plant grown at elevated [CO₂] and low N because they showed no difference in F_v/F_m (the maximum photochemical efficiency of PSII) from those grown at ambient [CO₂] and low N after an overnight dark adaptation. High levels of Nq_P in plants grown at elevated [CO₂] and low N reflect a near saturation of thermal energy dissipation. This impaired capacity of photoprotection would render these plants more vulnerable to photoinhibition in the event of additional environmental stresses such as drought, low or high temperature.     

Photosynthesis of Helianthus annuus does not acclimate toelevated CO2 regardless of N supply

A number of studies have shown that relatively long-term exposure to elevated levels of CO₂ can lead to the downward acclimation of photosynthesis. Although the exact mechanisms are not clearly understood, it has been suggested that such a downward adjustment may be more common under limited N availability. Here we examined the effect of N supply on the photosynthetic acclimation response of Helianthus annuus L. cv. Teddy Bear plants to elevated CO₂ at three growth stages – 18, 38 and 56 d after emergence corresponding to vegetative, pre-flowering and flowering stages. Plants were grown at CO₂ partial pressures of 37 or 70 Pa, and supplied with 0.5, 2.5 or 5 mol·m^–3 N. After 18 d of treatment, photosynthetic capacity of H. annuus as evaluated by parameters derived from the A-C_i data (Rubisco carboxylation capacity, V_c,max; electron transport capacity, J_max; and capacity for triose phosphate utilization, TPU) showed no acclimation to elevated CO₂. The leaf nitrogen concentration, [N], and total non-structural carbohydrates, [TNC], were also comparable between ambient- and elevated-CO₂-grown plants. However, all these photosynthetic parameters as well as leaf [N], but not [TNC], significantly increased in response to N supply. Similarly, after 38 and 56 d of exposure to CO₂ treatments, photosynthetic capacities, foliar [N] and [TNC] did not significantly differ between ambient- and elevated-CO₂ plants. These results suggest that H. annuus plants maintained their photosynthetic capacity during long-term exposure to elevated CO₂ because of their capacity to maintain leaf N-status. It is further suggested that plant capacity to maintain the balance between C and N acquisition rather than simply N-supply level, may determine whether photosynthetic acclimation in response to elevated CO₂ occurs or not.     

Acclimation of peanut (Arachis hypogaea L.) leaf photosynthesis to elevated growth CO2 and temperature

Peanut (Arachis hypogaea L. cv. Florunner) was grown from seed sowing to plant maturity under two daytime CO₂ concentrations ([CO₂]) of 360 μmol mol⁻¹ (ambient) and 720 μmol mol⁻¹ (elevated) and at two temperatures of 1.5 and 6.0 °C above ambient temperature. The objectives were to characterize peanut leaf photosynthesis responses to long-term elevated growth [CO₂] and temperature, and to assess whether elevated [CO₂] regulated peanut leaf photosynthetic capacity, in terms of activity and protein content of ribulose bisphosphate carboxylase-oxygenase (Rubisco), Rubisco photosynthetic efficiency, and carbohydrate metabolism. At both growth temperatures, leaves of plants grown under elevated [CO₂] had higher midday photosynthetic CO₂ exchange rate (CER), lower transpiration and stomatal conductance and higher water-use efficiency, compared to those of plants grown at ambient [CO₂]. Both activity and protein content of Rubisco, expressed on a leaf area basis, were reduced at elevated growth [CO₂]. Declines in Rubisco under elevated growth [CO₂] were 27–30% for initial activity, 5–12% for total activity, and 9–20% for protein content. Although Rubisco protein content and activity were down-regulated by elevated [CO₂], Rubisco photosynthetic efficiency, the ratio of midday light-saturated CER to Rubisco initial or total activity, of the elevated-[CO₂] plants was 1.3- to 1.9-fold greater than that of the ambient-[CO₂] plants at both growth temperatures. Leaf soluble sugars and starch of plants grown at elevated [CO₂] were 1.3- and 2-fold higher, respectively, than those of plants grown at ambient [CO₂]. Under elevated [CO₂], leaf soluble sugars and starch, however, were not affected by high growth temperature. In contrast, high temperature reduced leaf soluble sugars and starch of the ambient-[CO₂] plants. Activity of sucrose-P synthase, but not adenosine 5′-diphosphoglucose pyrophosphorylase, was up-regulated under elevated growth [CO₂]. Thus, in the absence of other environmental stresses, peanut leaf photosynthesis would perform well under rising atmospheric [CO₂] and temperature as predicted for this century.     

Physiological acclimation dampens initial effects of elevated temperature and atmospheric CO2 concentration in mature boreal Norway spruce

Physiological processes of terrestrial plants regulate the land–atmosphere exchange of carbon, water, and energy, yet few studies have explored the acclimation responses of mature boreal conifer trees to climate change. Here we explored the acclimation responses of photosynthesis, respiration, and stomatal conductance to elevated temperature and/or CO₂ concentration ([CO₂]) in a 3‐year field experiment with mature boreal Norway spruce. We found that elevated [CO₂] decreased photosynthetic carboxylation capacity (?23% at 25 °C) and increased shoot respiration (+64% at 15 °C), while warming had no significant effects. Shoot respiration, but not photosynthetic capacity, exhibited seasonal acclimation. Stomatal conductance at light saturation and a vapour pressure deficit of 1 kPa was unaffected by elevated [CO₂] but significantly decreased (?27%) by warming, and the ratio of intercellular to ambient [CO₂] was enhanced (+17%) by elevated [CO₂] and decreased (?12%) by warming. Many of these responses differ from those typically observed in temperate tree species. Our results show that long‐term physiological acclimation dampens the initial stimulation of plant net carbon assimilation to elevated [CO₂], and of plant water use to warming. Models that do not account for these responses may thus overestimate the impacts of climate change on future boreal vegetation–atmosphere interactions.     

Stomatal acclimation to increased CO2 concentration in a Florida scrub oak species Quercus myrtifolia Willd

Native scrub‐oak communities in Florida were exposed for three seasons in open top chambers to present atmospheric [CO₂] (approx. 350 μmol mol^?1) and to high [CO₂] (increased by 350 μmol mol^?1). Stomatal and photosynthetic acclimation to high [CO₂] of the dominant species Quercus myrtifolia was examined by leaf gas exchange of excised shoots. Stomatal conductance (g_s) was approximately 40% lower in the high‐ compared to low‐[CO₂]‐grown plants when measured at their respective growth concentrations. Reciprocal measurements of g_s in both high‐ and low‐[CO₂]‐grown plants showed that there was negative acclimation in the high‐[CO₂]‐grown plants (9–16% reduction in g_s when measured at 700 μmol mol^?1), but these were small compared to those for net CO₂ assimilation rate (A, 21–36%). Stomatal acclimation was more clearly evident in the curve of stomatal response to intercellular [CO₂] (c_i) which showed a reduction in stomatal sensitivity at low c_i in the high‐[CO₂]‐grown plants. Stomatal density showed no change in response to growth in high growth [CO₂]. Long‐term stomatal and photosynthetic acclimation to growth in high [CO₂] did not markedly change the 2·5‐ to 3‐fold increase in gas‐exchange‐derived water use efficiency caused by high [CO₂].     

Does long-term cultivation of saplings under elevated CO2 concentration influence their photosynthetic response to temperature?

Background and Aims Plants growing under elevated atmospheric CO₂ concentrations often have reduced stomatal conductance and subsequently increased leaf temperature. This study therefore tested the hypothesis that under long-term elevated CO₂ the temperature optima of photosynthetic processes will shift towards higher temperatures and the thermostability of the photosynthetic apparatus will increase.Methods The hypothesis was tested for saplings of broadleaved Fagus sylvatica and coniferous Picea abies exposed for 4–5 years to either ambient (AC; 385 µmol mol⁻¹) or elevated (EC; 700 µmol mol⁻¹) CO₂ concentrations. Temperature response curves of photosynthetic processes were determined by gas-exchange and chlorophyll fluorescence techniques.Key Results Initial assumptions of reduced light-saturated stomatal conductance and increased leaf temperatures for EC plants were confirmed. Temperature response curves revealed stimulation of light-saturated rates of CO₂ assimilation (A_max) and a decline in photorespiration (R_L) as a result of EC within a wide temperature range. However, these effects were negligible or reduced at low and high temperatures. Higher temperature optima (T_opt) of A_max, Rubisco carboxylation rates (V_Cmax) and R_L were found for EC saplings compared with AC saplings. However, the shifts in T_opt of A_max were instantaneous, and disappeared when measured at identical CO₂ concentrations. Higher values of T_opt at elevated CO₂ were attributed particularly to reduced photorespiration and prevailing limitation of photosynthesis by ribulose-1,5-bisphosphate (RuBP) regeneration. Temperature response curves of fluorescence parameters suggested a negligible effect of EC on enhancement of thermostability of photosystem II photochemistry.Conclusions Elevated CO₂ instantaneously increases temperature optima of A_max due to reduced photorespiration and limitation of photosynthesis by RuBP regeneration. However, this increase disappears when plants are exposed to identical CO₂ concentrations. In addition, increased heat-stress tolerance of primary photochemistry in plants grown at elevated CO₂ is unlikely. The hypothesis that long-term cultivation at elevated CO₂ leads to acclimation of photosynthesis to higher temperatures is therefore rejected. Nevertheless, incorporating acclimation mechanisms into models simulating carbon flux between the atmosphere and vegetation is necessary.     

Pea plant responsiveness under elevated [CO2] is conditioned by the N source (N2 fixation versus NO3− fertilization)

The main goal of this study was to test the effect of [CO₂] on C and N management in different plant organs (shoots, roots and nodules) and its implication in the responsiveness of exclusively N₂-fixing and NO₃⁻-fed plants. For this purpose, exclusively N₂-fixing and NO₃⁻-fed (10 mM) pea (Pisum sativum L.) plants were exposed to elevated [CO₂] (1000 μmol mol⁻¹ versus 360 μmol mol⁻¹ CO₂). Gas exchange analyses, together with carbohydrate, nitrogen, total soluble proteins and amino acids were determined in leaves, roots and nodules. The data obtained revealed that although exposure to elevated [CO₂] increased total dry mass (DM) in both N treatments, photosynthetic activity was down-regulated in NO₃⁻-fed plants, whereas N₂-fixing plants were capable of maintaining enhanced photosynthetic rates under elevated [CO₂]. In the case of N₂-fixing plants, the enhanced C sink strength of nodules enabled the avoidance of harmful leaf carbohydrate build up. On the other hand, in NO₃⁻-fed plants, elevated [CO₂] caused a large increase in sucrose and starch. The increase in root DM did not contribute to stimulation of C sinks in these plants. Although N₂ fixation matched plant N requirements with the consequent increase in photosynthetic rates, in NO₃⁻-fed plants, exposure to elevated [CO₂] negatively affected N assimilation with the consequent photosynthetic down-regulation.     

Direct and acclimatory responses of stomatal conductance to elevated carbon dioxide in four herbaceous crop species in the field

In order to separate the net effect of growth at elevated [CO₂] on stomatal conductance (g_s) into direct and acclimatory responses, mid‐day values of g_s were measured for plants grown in field plots in open‐topped chambers at the current ambient [CO₂], which averaged 350 μmol mol^?1 in the daytime, and at ambient + 350 μmol mol^?1[CO₂] for winter wheat, winter barley, potato and sorghum. The acclimatory response was determined by comparing g_s measured at 700 μmol mol^?1[CO₂] for plants grown at the two [CO₂]. The direct effect of increasing [CO₂] from 350 to 700 μmol mol^?1 was determined for plants grown at the lower concentration. Photosynthetic rates were measured concurrently with g_s. For all species, growth at the higher [CO₂] significantly reduced g_s measured at 700 μmol mol^?1[CO₂]. The reduction in g_s caused by growth at the higher [CO₂] was larger for all species on days with low leaf to air water vapour pressure difference for a given temperature, which coincided with highest conductances and also the smallest direct effects of increased [CO₂] on conductance. For barley, there was no other evidence for stomatal acclimation, despite consistent down‐regulation of photosynthetic rate in plants grown at the higher [CO₂]. In wheat and potato, in addition to the vapour pressure difference interaction, the magnitude of stomatal acclimation varied directly in proportion to the magnitude of down‐regulation of photosynthetic rate through the season. In sorghum, g_s consistently exhibited acclimation, but there was no down‐regulation of photosynthetic rate. In none of the species except barley was the direct effect the larger component of the net reduction in g_s when averaged over measurement dates. The net effect of growth at elevated [CO₂] on mid‐day g_s resulted from unique combinations of direct and acclimatory responses in the various species.     

Interaction of water supply and N in wheat

The purpose of this study was to investigate effects of N nutrition and water stress on stomatal behavior and CO₂ exchange rate in wheat (Triticum aestivum L. cv Olaf). Wheat plants were grown hydroponically with high (100 milligrams per liter) and low (10 milligrams per liter) N. When plants were 38 days old, a 24-day water stress cycle was begun. A gradual increase in nutrient solution osmotic pressure from 0.03 to 1.95 mega Pascals was achieved by incremental additions of PEG-6,000. Plants in both N treatments adjusted osmotically, although leaf water potential was consistently lower and relative water content greater for low N plants in the first half of the stress cycle. Leaf conductance of high N plants appeared greater than that of low N plants at high water potentials, but showed greater sensitivity to reductions in water potential as indicated by earlier stomatal closure during the stress cycle. The apparent greater stomatal sensitivity of high N plants was associated with a curvilinear relationship between leaf conductance and leaf water potential; low N plants exhibited more of a threshold response. Trends in [CO₂]_INT throughout the stress cycle indicated nonstomatal effects of water stress on CO₂ exchange rate were greater in high N plants. Although estimates of [CO₂]_INT were generally lower in high N plants, they were relatively insensitive to leaf water potential-induced changes in leaf conductance. In contrast, [CO₂]_INT of low N plants dropped concomitantly with leaf conductance at low leaf water potentials. Oxygen response of CO₂ exchange rate for both treatments was affected less by reductions in water potential than was CO₂ exchange rate at 2.5% O₂, suggesting that CO₂ assimilation capacity of the leaves was affected more by reductions in leaf water potential than were processes related to photorespiration.     

Effect of nitrogen application and elevated CO2 on photosynthetic gas exchange and electron transport in wheat leaves

Nitrogen (N) availability is a critical factor affecting photosynthetic acclimation of C₃ plants under elevated atmospheric CO₂ concentration ([CO₂]_e). However, current understanding of N effects on photosynthetic electron transport rate and partitioning, as well as its impact on photosynthesis under [CO₂]_e, is inadequate. Using controlled environment open-top chambers, wheat (Triticum aestivum L.) was grown at two N levels (0 and 200 mg(N) kg^?1 soil) and two atmospheric CO₂ concentrations of 400 ([CO₂]_a) and 760 μmol mol^?1([CO₂]_e) during 2009 and 2010. Under [CO₂]_e high N availability increased stomatal conductance and transpiration rate, reduced limitations on the activity of triose phosphate isomerase, a Calvin cycle enzyme, and increased the rate of net photosynthesis (P _N). Considering photosynthetic electron transport rate and partitioning aspects, we suggest that greater N availability increased P _N under [CO₂]_e due to four following reasons: (1) higher N availability enhanced foliar N and chlorophyll concentrations, and the actual photochemical efficiency of photosystem (PS) II reaction centers under irradiance increased, (2) increase of total electron transport rate and proportion of open PSII reaction centers, (3) enhancement of the electron transport rate of the photochemical and carboxylation processes, and (4) reduced limitations of the Calvin cycle enzymes on the photosynthetic electron transport rate. Consequently, sufficient N improved light energy utilization in wheat flag leaves under [CO₂]_e, thus benefiting to photosynthetic assimilation.     

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。     

Elevated [CO2] and increased N supply reduce leaf disease and related photosynthetic impacts on Solidago rigida

To evaluate whether leaf spot disease and related effects on photosynthesis are influenced by increased nitrogen (N) input and elevated atmospheric CO₂ concentration ([CO₂]), we examined disease incidence and photosynthetic rate of Solidago rigida grown in monoculture under ambient or elevated (560 μmol mol⁻¹) [CO₂] and ambient or elevated (+4 g N m⁻² year⁻¹) N conditions in a field experiment in Minnesota, USA. Disease incidence was lower in plots with either elevated [CO₂] or enriched N (−57 and −37%, respectively) than in plots with ambient conditions. Elevated [CO₂] had no significant effect on total plant biomass, or on photosynthetic rate, but reduced tissue%N by 13%. In contrast, N fertilization increased both biomass and total plant N by 70%, and as a consequence tissue%N was unaffected and photosynthetic rate was lower on N fertilized plants than on unfertilized plants. Regardless of treatment, photosynthetic rate was reduced on leaves with disease symptoms. On average across all treatments, asymptomatic leaf tissue on diseased leaves had 53% lower photosynthetic rate than non-diseased leaves, indicating that the negative effect from the disease extended beyond the visual lesion area. Our results show that, in this instance, indirect effects from elevated [CO₂], i.e., lower disease incidence, had a stronger effect on realized photosynthetic rate than the direct effect of higher [CO₂].     

Profiles of isoprene emission and photosynthetic parameters in hybrid poplars exposed to free‐air CO2 enrichment†

Poplar (Populus × euroamericana) saplings were grown in the field to study the changes of photosynthesis and isoprene emission with leaf ontogeny in response to free air carbon dioxide enrichment (FACE) and soil nutrient availability. Plants growing in elevated [CO₂] produced more leaves than those in ambient [CO₂]. The rate of leaf expansion was measured by comparing leaves along the plant profile. Leaf expansion and nitrogen concentration per unit of leaf area was similar between nutrient treatment, and this led to similar source–sink functional balance. Consequently, soil nutrient availability did not cause downward acclimation of photosynthetic capacity in elevated [CO₂] and did not affect isoprene synthesis. Photosynthesis assessed in growth [CO₂] was higher in plants growing in elevated than in ambient [CO₂]. After normalizing for the different number of leaves over the profile, maximal photosynthesis was reached and started to decline earlier in elevated than in ambient [CO₂]. This may indicate a [CO₂]‐driven acceleration of leaf maturity and senescence. Isoprene emission was adversely affected by elevated [CO₂]. When measured on the different leaves of the profile, isoprene peak emission was higher and was reached earlier in ambient than in elevated [CO₂]. However, a larger number of leaves was emitting isoprene in plant growing in elevated [CO₂]. When integrating over the plant profile, emissions in the two [CO₂] levels were not different. Normalization as for photosynthesis showed that profiles of isoprene emission were remarkably similar in the two [CO₂] levels, with peak emissions at the centre of the profile. Only the rate of increase of the emission of young leaves may have been faster in elevated than in ambient [CO₂]. Our results indicate that elevated [CO₂] may overall have a limited effect on isoprene emission from young seedlings and that plants generally regulate the emission to reach the maximum at the centre of the leaf profile, irrespective of the total leaf number. In comparison with leaf expansion and photosynthesis, isoprene showed marked and repeatable differences among leaves of the profile and may therefore be a useful trait to accurately monitor changes of leaf ontogeny as a consequence of elevated [CO₂].