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].     

The stomatal CO<Subscript>2</Subscript> proxy does not saturate at high atmospheric CO<Subscript>2</Subscript> concentrations: evidence from stomatal index responses of Araucariaceae conifers

The inverse relationship between the number of stomata on a leaf surface and the atmospheric carbon dioxide concentration ([CO₂]) in which the leaf developed allows plants to optimise water-use efficiency (WUE), but it also permits the use of fossil plants as proxies of palaeoatmospheric [CO₂]. The ancient conifer family Araucariaceae is often represented in fossil floras and may act as a suitable proxy of palaeo-[CO₂], yet little is known regarding the stomatal index (SI) responses of extant Araucariaceae to [CO₂]. Four Araucaria species (Araucaria columnaris, A. heterophylla, A. angustifolia and A. bidwillii) and Agathis australis displayed no significant relationship in SI to [CO₂] below current ambient levels (~380 ppm). However, representatives of the three extant genera within the Araucariaceae (A. bidwillii, A. australis and Wollemia nobilis) all exhibited significant reductions in SI when grown in atmospheres of elevated [CO₂] (1,500 ppm). Stomatal conductance was reduced and WUE increased when grown under elevated [CO₂]. Stomatal pore length did not increase alongside reduced stomatal density (SD) and SI in the three araucariacean conifers when grown at elevated [CO₂]. These pronounced SD and SI reductions occur at higher [CO₂] levels than in other species with more recent evolutionary origins, and may reflect an evolutionary legacy of the Araucariaceae in the high [CO₂] world of the Mesozoic Era. Araucariacean conifers may therefore be suitable stomatal proxies of palaeo-[CO₂] during periods of “greenhouse” climates and high [CO₂] in the Earth’s history.     

Effects of the interaction between vapor-pressure deficit and salinity on growth and photosynthesis of <Emphasis Type="Italic">Cucumis sativus</Emphasis> seedlings under different CO<Subscript>2</Subscript> concentrations

We studied growth and photosynthesis of cucumber (Cucumis sativus) seedlings under two vapor-pressure deficit levels (VPD; 0.4 and 3.0 kPa), two salinity levels (0 mM and 34 mM NaCl), and two CO₂ concentrations ([CO₂]; 400 and 1,000 μmol mol^–1). Relative growth rate (RGR) decreased with increasing VPD, but the causal factor differed between salinity levels and CO₂ concentrations. Under ambient [CO₂], RGR decreased with increasing VPD at low salinity mainly due to decreased leaf area ratio (LAR), and decreased net assimilation rate (NAR) at high salinity. The decrease in intercellular [CO₂] (C_i) with decreasing stomatal conductance caused by high VPD did not significantly limit net photosynthetic rate (P_N) at low salinity, but P_N was potentially limited by C_i at high salinity. At high [CO₂], high VPD reduced LAR, but did not affect NAR. This is because the decrease in C_i occurred where slope of P_N–C_i curve was almost flat.     

Plant responses to elevated CO<Subscript>2</Subscript> concentration at different scales: leaf,whole plant,canopy, and population

Elevated CO₂ enhances photosynthesis and growth of plants, but the enhancement is strongly influenced by the availability of nitrogen. In this article, we summarise our studies on plant responses to elevated CO₂. The photosynthetic capacity of leaves depends not only on leaf nitrogen content but also on nitrogen partitioning within a leaf. In Polygonum cuspidatum, nitrogen partitioning among the photosynthetic components was not influenced by elevated CO₂ but changed between seasons. Since the alteration in nitrogen partitioning resulted in different CO₂-dependence of photosynthetic rates, enhancement of photosynthesis by elevated CO₂ was greater in autumn than in summer. Leaf mass per unit area (LMA) increases in plants grown at elevated CO₂. This increase was considered to have resulted from the accumulation of carbohydrates not used for plant growth. With a sensitive analysis of a growth model, however, we suggested that the increase in LMA is advantageous for growth at elevated CO₂ by compensating for the reduction in leaf nitrogen concentration per unit mass. Enhancement of reproductive yield by elevated CO₂ is often smaller than that expected from vegetative growth. In Xanthium canadense, elevated CO₂ did not increase seed production, though the vegetative growth increased by 53%. As nitrogen concentration of seeds remained constant at different CO₂ levels, we suggest that the availability of nitrogen limited seed production at elevated CO₂ levels. We found that leaf area development of plant canopy was strongly constrained by the availability of nitrogen rather than by CO₂. In a rice field cultivated at free-air CO₂ enrichment, the leaf area index (LAI) increased with an increase in nitrogen availability but did not change with CO₂ elevation. We determined optimal LAI to maximise canopy photosynthesis and demonstrated that enhancement of canopy photosynthesis by elevated CO₂ was larger at high than at low nitrogen availability. We also studied competitive asymmetry among individuals in an even-aged, monospecific stand at elevated CO₂. Light acquisition (acquired light per unit aboveground mass) and utilisation (photosynthesis per unit acquired light) were calculated for each individual in the stand. Elevated CO₂ enhanced photosynthesis and growth of tall dominants, which reduced the light availability for shorter subordinates and consequently increased size inequality in the stand.     

Stomatal conductance and not stomatal density determines the long-term reduction in leaf transpiration of poplar in elevated CO<Subscript>2</Subscript>

Using a free-air CO₂ enrichment (FACE) experiment, poplar trees (Populus × euramericana clone I214) were exposed to either ambient or elevated [CO₂] from planting, for a 5-year period during canopy development, closure, coppice and re-growth. In each year, measurements were taken of stomatal density (SD, number mm⁻²) and stomatal index (SI, the proportion of epidermal cells forming stomata). In year 5, measurements were also taken of leaf stomatal conductance (g _s, μmol m⁻² s⁻¹), photosynthetic CO₂ fixation (A, mmol m⁻² s⁻¹), instantaneous water-use efficiency (A/E) and the ratio of intercellular to atmospheric CO₂ (C_i:C_a). Elevated [CO₂] caused reductions in SI in the first year, and in SD in the first 2 years, when the canopy was largely open. In following years, when the canopy had closed, elevated [CO₂] had no detectable effects on stomatal numbers or index. In contrast, even after 5 years of exposure to elevated [CO₂], g _s was reduced, A/E was stimulated, and C_i:C_a was reduced relative to ambient [CO₂]. These outcomes from the long-term realistic field conditions of this forest FACE experiment suggest that stomatal numbers (SD and SI) had no role in determining the improved instantaneous leaf-level efficiency of water use under elevated [CO₂]. We propose that altered cuticular development during canopy closure may partially explain the changing response of stomata to elevated [CO₂], although the mechanism for this remains obscure.     

Photosynthetic sunfleck utilization potential of understory saplings growing under elevated CO<Subscript>2</Subscript> in FACE

Few studies have evaluated elevated CO₂ responses of trees in variable light despite its prevalence in forest understories and its potential importance for sapling survival. We studied two shade-tolerant species (Acer rubrum, Cornus florida) and two shade-intolerant species (Liquidambar styraciflua, Liriodendron tulipifera) growing in the understory of a Pinus taeda plantation under ambient and ambient+200 ppm CO₂ in a free air carbon enrichment (FACE) experiment. Photosynthetic and stomatal responses to artificial changes in light intensity were measured on saplings to determine rates of induction gain under saturating light and induction loss under shade. We expected that growth in elevated CO₂ would alter photosynthetic responses to variable light in these understory saplings. The results showed that elevated CO₂ caused the expected enhancement in steady-state photosynthesis in both high and low light, but did not affect overall stomatal conductance or rates of induction gain in the four species. Induction loss after relatively short shade periods (<6 min) was slower in trees grown in elevated CO₂ than in trees grown in ambient CO₂ despite similar decreases in stomatal conductance. As a result leaves grown in elevated CO₂ that maintained induction well in shade had higher carbon gain during subsequent light flecks than was expected from steady-state light response measurements. Thus, when frequent sunflecks maintain stomatal conductance and photosynthetic induction during the day, enhancements of long-term carbon gain by elevated CO₂ could be underestimated by steady-state photosynthetic measures. With respect to species differences, both a tolerant, A. rubrum, and an intolerant species, L. tulipifera, showed rapid induction gain, but A. rubrum also lost induction rapidly (c. 12 min) in shade. These results, as well as those from independent studies in the literature, show that induction dynamics are not closely related to species shade tolerance. Therefore, it cannot be concluded that shade-tolerant species necessarily induce faster in the variable light conditions common in understories. Although our study is the first to examine dynamic photosynthetic responses to variable light in contrasting species in elevated CO₂, studies on ecologically diverse species will be required to establish whether shade-tolerant and -intolerant species show different photosynthetic responses in elevated CO₂ during sunflecks. We conclude that elevated CO₂ affects dynamic gas exchange most strongly via photosynthetic enhancement during induction as well as in the steady state. Received: 1 April 1999 / Accepted: 16 August 1999     

The impact of salt stress on the water status of barley plants is partially mitigated by elevated CO2

With the changing climate, plants will be facing increasingly harsh environmental conditions marked by elevated salinity in the soils and elevated concentrations of CO₂ in the atmosphere. These two factors have opposite effects on water status in plants. Therefore, our objective was to determine the interaction between these two factors and to determine whether elevated [CO₂] might alleviate the adverse effects of salt stress on water status in two barley cultivars, Alpha and Iranis, by studying their relative water content and their water potential and its components, transpiration rate, hydraulic conductance, and water use efficiency. Both cultivars maintained their water status under salt stress, increasing water use efficiency and conserving a high relative water content by (1) reducing water potential via passive dehydration and active osmotic adjustment and (2) decreasing transpiration through stomatal closure and reducing hydraulic conductance. Iranis showed a greater capacity to achieve osmotic adjustment than Alpha. Under the combined conditions of salt-stress and elevated [CO₂], both cultivars (1) achieved osmotic adjustment to a greater extent than at ambient [CO₂], likely due to elevated rates of photosynthesis, and (2) decreased passive dehydration by stomatal closure, thereby maintaining a greater turgor potential, relative water content, and water use efficiency. Therefore, we found an interaction between salt stress and elevated [CO₂] with regard to water status in plants and found that elevated [CO₂] is associated with improved water status of salt-stressed barley plants.     

The impact of salt stress on the water status of barley plants is partially mitigated by elevated CO2

With the changing climate, plants will be facing increasingly harsh environmental conditions marked by elevated salinity in the soils and elevated concentrations of CO₂ in the atmosphere. These two factors have opposite effects on water status in plants. Therefore, our objective was to determine the interaction between these two factors and to determine whether elevated [CO₂] might alleviate the adverse effects of salt stress on water status in two barley cultivars, Alpha and Iranis, by studying their relative water content and their water potential and its components, transpiration rate, hydraulic conductance, and water use efficiency. Both cultivars maintained their water status under salt stress, increasing water use efficiency and conserving a high relative water content by (1) reducing water potential via passive dehydration and active osmotic adjustment and (2) decreasing transpiration through stomatal closure and reducing hydraulic conductance. Iranis showed a greater capacity to achieve osmotic adjustment than Alpha. Under the combined conditions of salt-stress and elevated [CO₂], both cultivars (1) achieved osmotic adjustment to a greater extent than at ambient [CO₂], likely due to elevated rates of photosynthesis, and (2) decreased passive dehydration by stomatal closure, thereby maintaining a greater turgor potential, relative water content, and water use efficiency. Therefore, we found an interaction between salt stress and elevated [CO₂] with regard to water status in plants and found that elevated [CO₂] is associated with improved water status of salt-stressed barley plants.     

Effects of salinity on chlorophyll fluorescence and CO<Subscript>2</Subscript> fixation in C<Subscript>4</Subscript> estuarine grasses

The effects of salinity (sea water at 0 ‰ versus 30 ‰) on gross rates of O₂ evolution (J _O2) and net rates of CO₂ uptake (P _N) were measured in the halotolerant estuarine C₄ grasses Spartina patens, S. alterniflora, S. densiflora, and Distichlis spicata in controlled growth environments. Under high irradiance, salinity had no significant effect on the intercellular to ambient CO₂ concentration ratio (C _i/C _a). However, during photosynthesis under limiting irradiance, the maximum quantum efficiency of CO₂ fixation decreased under salinity across species, suggesting there is increased leakage of the CO₂ delivered to the bundle sheath cells by the C₄ pump. Growth under salinity did not affect the maximum intrinsic efficiency of photosystem 2, PS2 (F_V/F_M) in these species, suggesting salinity had no effect on photosynthesis by inactivation of PS2 reaction centers. Under saline conditions and high irradiance, P _N was reduced by 75 % in Spartina patens and S. alterniflora, whereas salinity had no effect on P _N in S. densiflora or D. spicata. This inhibition of P _N in S. patens and S. alterniflora was not due to an effect on stomatal conductance since the ratio of C _i/C _a did not decrease under saline conditions. In growth with and without salt, P _N was saturated at ∼500 μmol(quantum) m⁻² s⁻¹ while J _O2 continued to increase up to full sunlight, indicating that carbon assimilation was not tightly coupled to photochemistry in these halophytic species. This increase in alternative electron flow under high irradiance might be an inherent function in these halophytes for dissipating excess energy.     

Elevated CO<Subscript>2</Subscript> increased photosynthesis and yield without decreasing stomatal conductance in broomcorn millet

Broomcorn millet (Panicum miliaceum L.) is one of the important C₄ crops in the semiarid regions of northern China. It is a close relative of biofuel crop switchgrass. Yet, there is no information on how these crops might respond to a climate change in China. In order to gain insight into such a response, we studied the effect of elevated CO₂ concentration (EC) on broomcorn millet. The changes in leaf photosynthesis, chlorophyll fluorescence, morphological parameters, biomass and yield in response to EC [i.e., + 200 µmol(CO₂) mol^?1] over two years were determined at the open-top chamber (OTC) experimental facility in north China. EC increased net photosynthetic rate, stomatal conductance, intercellular CO₂ concentration, transpiration rate, instantaneous transpiration efficiency, effective quantum yield of PSII photochemistry, and photochemical quenching coefficient of fully expanded flag leaves. Maximal quantum yield of PSII photochemistry declined under EC in 2013, but was not affected in 2014. EC significantly decreased intrinsic efficiency of PSII in 2013, but increased in 2014. Leaf nonphotochemical quenching decreased under EC both in 2013 and 2014. EC significantly enhanced the aboveground biomass and yield by average of 31.4 and 25.5% in both years, respectively. The increased yield of broomcorn millet under EC occurred due to the enhanced number of grains per plant. We concluded that photosynthesis of broomcorn millets was improved through increased stomatal conductance in leaves under EC, which led to an increase in height, stem diameter, aboveground biomass, and yield. This study extends our understanding of the response of this ancient C₄ crop to elevated CO₂ concentration.     

Photosynthetic limitations in olive cultivars with different sensitivity to salt stress

Olive (Olea europea L) is one of the most valuable and widespread fruit trees in the Mediterranean area. To breed olive for resistance to salinity, an environmental constraint typical of the Mediterranean, is an important goal. The photosynthetic limitations associated with salt stress caused by irrigation with saline (200 mm ) water were assessed with simultaneous gas‐exchange and fluorescence field measurements in six olive cultivars. Cultivars were found to possess inherently different photosynthesis when non‐stressed. When exposed to salt stress, cultivars with inherently high photosynthesis showed the highest photosynthetic reductions. There was no relationship between salt accumulation and photosynthesis reduction in either young or old leaves. Thus photosynthetic sensitivity to salt did not depend on salt exclusion or compartmentalization in the old leaves of the olive cultivars investigated. Salt reduced the photochemical efficiency, but this reduction was also not associated with photosynthesis reduction. Salt caused a reduction of stomatal and mesophyll conductance, especially in cultivars with inherently high photosynthesis. Mesophyll conductance was generally strongly associated with photosynthesis, but not in salt‐stressed leaves with a mesophyll conductance higher than 50 mmol m^?2 s^?1. The combined reduction of stomatal and mesophyll conductances in salt‐stressed leaves increased the CO₂ draw‐down between ambient air and the chloroplasts. The CO₂ draw‐down was strongly associated with photosynthesis reduction of salt‐stressed leaves but also with the variable photosynthesis of controls. The relationship between photosynthesis and CO₂ draw‐down remained unchanged in most of the cultivars, suggesting no or small changes in Rubisco activity of salt‐stressed leaves. The present results indicate that the low chloroplast CO₂ concentration set by both low stomatal and mesophyll conductances were the main limitations of photosynthesis in salt‐stressed olive as well as in cultivars with inherently low photosynthesis. It is consequently suggested that, independently of the apparent sensitivity of photosynthesis to salt, this effect may be relieved if conductances to CO₂ diffusion are restored.     

The effects of salinity on photosynthesis and growth of the single-cell C<Subscript>4</Subscript> species <Emphasis Type="Italic">Bienertia sinuspersici</Emphasis> (Chenopodiaceae)

Recent research on the photosynthetic mechanisms of plant species in the Chenopodiaceae family revealed that three species, including Bienertia sinuspersici, can carry out C₄ photosynthesis within individual photosynthetic cells, through the development of two cytoplasmic domains having dimorphic chloroplasts. These unusual single-cell C₄ species grow in semi-arid saline conditions and have semi-terete succulent leaves. The effects of salinity on growth and photosynthesis of B. sinuspersici were studied. The results show that NaCl is not required for development of the single-cell C₄ system. There is a large enhancement of growth in culture with 50–200 mM NaCl, while there is severe inhibition at 400 mM NaCl. With increasing salinity, the carbon isotope values (δ¹³C) of leaves increased from −17.3^o/_oo (C₄-like) without NaCl to −14.6^o/_oo (C₄) with 200 mM NaCl, possibly due to increased capture of CO₂ from the C₄ cycle by Rubisco and reduced leakiness. Compared to growth without NaCl, leaves of plants grown under saline conditions were much larger (~2 fold) and more succulent, and the leaf solute levels increased up to ~2000 mmol kg solvent⁻¹. Photosynthesis on an incident leaf area basis (CO₂ saturated rates, and carboxylation efficiency under limiting CO₂) and stomatal conductance declined with increasing salinity. On a leaf area basis, there was some decline in Rubisco content with increasing salinity up to 200 mM NaCl, but there was a marked increase in the levels of pyruvate, Pi dikinase, and phosphoenolpyruvate carboxylase (possibly in response to sensitivity of these enzymes and C₄ cycle function to increasing salinity). The decline in photosynthesis on a leaf area basis was compensated for on a per leaf basis, up to 200 mM NaCl, by the increase in leaf size. The influence of salinity on plant development and the C₄ system in Bienertia is discussed.     

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.     

A Possible Role for Jasmonic Acid in Adaptation of Barley Seedlings to Salinity Stress

The changes caused by NaCl salinity and jasmonic acid (JA) treatment (8 days) on growth and photosynthesis of barley plants (Hordeum vulgare L., var. Alfa) have been studied. Gas exchange measurements and analysis of enzyme activities were used to study the reactions of photosynthesis to salinity and JA. Both 100 mm NaCl and 25 μm JA treatment led to a noticeable decrease in both the initial slope of the curves representing net photosynthetic rate vs intercellular CO₂ concentration and the maximal rate of photosynthesis. The calculated values of the intercellular CO₂ concentration, CO₂ compensation point, and maximal carboxylating efficiency of ribulose-1,5-bisphosphate carboxylase support the suggestion that biochemical factors are involved in the response of photosynthesis to JA and salinity stress. The activities of phosphoenolpyruvate carboxylase and carbonic anhydrase increased more than twofold. Pretreatment with JA for 4 days before salinization diminished the inhibitory effect of high salt concentration on the growth and photosynthesis. The results are discussed in terms of a possible role of JA in increasing salinity tolerance of the barley plants. Received September 8, 1997; accepted May 19, 1998     

Influence of water stress on photosynthetic characteristics in barley plants under ambient and elevated CO<Subscript>2</Subscript> concentrations

We evaluated the combined effects of elevated CO₂ and water availability on photosynthesis in barley. Soil and plant water content decreased with water stress, but less under elevated CO₂ concentration (EC) compared with ambient CO₂ concentration (AC). During water stress, stomatal conductance, carboxylation rate, RuBP regeneration, and the rate of triose phosphate utilisation (TPU) were decreased but less when plants grew under EC. Drought treatments caused only a slight effect on maximum photochemical efficiency (variable to maximum fluorescence ratio, F_v/F_m), whereas the actual quantum yield (Φ_PS2), maximum electron transport rate (J_max) and photochemical quenching (qP) were decreased and the non photochemical quenching (NPQ) was enhanced. Under water deficit, the allocation of electrons to CO₂ assimilation was diminished by 49 % at AC and by 26 % at EC while the allocation to O₂ reduction was increased by 15 % at AC and by 12 % at EC.     

Photosynthesis and fluorescence responses of C<Subscript>4</Subscript> plant <Emphasis Type="Italic">Andropogon gerardii</Emphasis> acclimated to temperature and carbon dioxide

Increase in both atmospheric CO₂ concentration [CO₂] and associated warming are likely to alter Earths’ carbon balance and photosynthetic carbon fixation of dominant plant species in a given biome. An experiment was conducted in sunlit, controlled environment chambers to determine effects of atmospheric [CO₂] and temperature on net photosynthetic rate (P _N) and fluorescence (F) in response to internal CO₂ concentration (C _i) and photosynthetically active radiation (PAR) of the C₄ species, big bluestem (Andropogon gerardii Vitman). Ten treatments were comprised of two [CO₂] of 360 (ambient, AC) and 720 (elevated, EC) μmol mol⁻¹ and five day/night temperature of 20/12, 25/17, 30/22, 35/27 and 40/32 °C. Treatments were imposed from 15 d after sowing (DAS) through 130 DAS. Both F-P _N/C _i and F-P _N/PAR response curves were measured on top most fully expanded leaves between 55 and 75 DAS. Plants grown in EC exhibited significantly higher CO₂-saturated net photosynthesis (P _sat), phosphoenolpyruvate carboxylase (PEPC) efficiency, and electron transport rate (ETR). At a given [CO₂], increase in temperature increased P _sat, PEPC efficiency, and ETR. Plants grown at EC did not differ for dark respiration rate (R _D), but had significantly higher maximum photosynthesis (P _max) than plants grown in AC. Increase in temperature increased Pmax, R _D, and ETR, irrespective of the [CO₂]. The ability of PEPC, ribulose-1,5-bisphosphate carboxylase/oxygenase, and photosystem components, derived from response curves to tolerate higher temperatures (>35 °C), particularly under EC, indicates the ability of C₄ species to sustain photosynthetic capacity in future climates.     

New insights into the cellular mechanisms of plant growth at elevated atmospheric carbon dioxide concentrations

Rising atmospheric carbon dioxide concentration ([CO₂]) significantly influences plant growth, development, and biomass. Increased photosynthesis rate, together with lower stomatal conductance, has been identified as the key factors that stimulate plant growth at elevated [CO₂] (e[CO₂]). However, variations in photosynthesis and stomatal conductance alone cannot fully explain the dynamic changes in plant growth. Stimulation of photosynthesis at e[CO₂] is always associated with post‐photosynthetic secondary metabolic processes that include carbon and nitrogen metabolism, cell cycle functions, and hormonal regulation. Most studies have focused on photosynthesis and stomatal conductance in response to e[CO₂], despite the emerging evidence of e[CO₂]'s role in moderating secondary metabolism in plants. In this review, we briefly discuss the effects of e[CO₂] on photosynthesis and stomatal conductance and then focus on the changes in other cellular mechanisms and growth processes at e[CO₂] in relation to plant growth and development. Finally, knowledge gaps in understanding plant growth responses to e[CO₂] have been identified with the aim of improving crop productivity under a CO₂ rich atmosphere.     

The growth of soybean under free air [CO<Subscript>2</Subscript>] enrichment (FACE) stimulates photosynthesis while decreasing in vivo Rubisco capacity

Down-regulation of light-saturated photosynthesis (A_sat) at elevated atmospheric CO₂ concentration, [CO₂], has been demonstrated for many C₃ species and is often associated with inability to utilize additional photosynthate and/or nitrogen limitation. In soybean, a nitrogen-fixing species, both limitations are less likely than in crops lacking an N-fixing symbiont. Prior studies have used controlled environment or field enclosures where the artificial environment can modify responses to [CO₂]. A soybean free air [CO₂] enrichment (FACE) facility has provided the first opportunity to analyze the effects of elevated [CO₂] on photosynthesis under fully open-air conditions. Potential ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) carboxylation (V_c,max) and electron transport through photosystem II (J_max) were determined from the responses of A_sat to intercellular [CO₂] (C_i) throughout two growing seasons. Mesophyll conductance to CO₂ (g_m) was determined from the responses of A_sat and whole chain electron transport (J) to light. Elevated [CO₂] increased A_sat by 15–20% even though there was a small, statistically significant, decrease in V_c,max. This differs from previous studies in that V_c,max/J_max decreased, inferring a shift in resource investment away from Rubisco. This raised the C_i at which the transition from Rubisco-limited to ribulose-1,5-bisphosphate regeneration-limited photosynthesis occurred. The decrease in V_c,max was not the result of a change in g_m, which was unchanged by elevated [CO₂]. This first analysis of limitations to soybean photosynthesis under fully open-air conditions reveals important differences to prior studies that have used enclosures to elevate [CO₂], most significantly a smaller response of A_sat and an apparent shift in resources away from Rubisco relative to capacity for electron transport.Abbreviations FACE Free air [CO₂] enrichment - Rubisco Ribulose-1,5-bisphosphate carboxylase/oxygenase - RuBP Ribulose-1,5-bisphosphate - SoyFACE Soybean free air [CO₂] enrichment - VPD Vapor pressure deficit     

Altered physiological function,not structure,drives increased radiation-use efficiency of soybean grown at elevated CO<Subscript>2</Subscript>

Previous studies of elevated carbon dioxide concentration ([CO₂]) on crop canopies have found that radiation-use efficiency is increased more than radiation-interception efficiency. It is assumed that increased radiation-use efficiency is due to changes in leaf-level physiology; however, canopy structure can affect radiation-use efficiency if leaves are displayed in a manner that optimizes their physiological capacity, even though the canopy intercepts the same amount of light. In order to determine the contributions of physiology and canopy structure to radiation-use and radiation-interception efficiency, this study relates leaf-level physiology and leaf display to photosynthetic rate of the outer canopy. We used a new imaging approach that delivers three-dimensional maps of the outer canopy during the growing season. The 3D data were used to model leaf orientation and mean photosynthetic electron transport of the outer canopy to show that leaf orientation changes did not contribute to increased radiation-use; i.e. leaves of the outer canopy showed similar diurnal leaf movements and leaf orientation in both treatments. Elevated [CO₂] resulted in an increased maximum electron transport rate (ETR_max) of light reactions of photosynthesis. Modeling of canopy light interception showed that stimulated leaf-level electron transport at elevated [CO₂], and not alterations in leaf orientation, was associated with stimulated radiation-use efficiency and biomass production in elevated [CO₂]. This study provides proof of concept of methodology to quantify structure–function relationships in combination, allowing a quantitative estimate of the contribution of both effects to canopy energy conversion under elevated [CO₂].     

Carbon source–sink limitations differ between two species with contrasting growth strategies

Understanding how carbon source and sink strengths limit plant growth is a critical knowledge gap that hinders efforts to maximize crop yield. We investigated how differences in growth rate arise from source–sink limitations, using a model system comparing a fast‐growing domesticated annual barley (Hordeum vulgare cv. NFC Tipple) with a slow‐growing wild perennial relative (Hordeum bulbosum). Source strength was manipulated by growing plants at sub‐ambient and elevated CO₂ concentrations ([CO₂]). Limitations on vegetative growth imposed by source and sink were diagnosed by measuring relative growth rate, developmental plasticity, photosynthesis and major carbon and nitrogen metabolite pools. Growth was sink limited in the annual but source limited in the perennial. RGR and carbon acquisition were higher in the annual, but photosynthesis responded weakly to elevated [CO₂] indicating that source strength was near maximal at current [CO₂]. In contrast, photosynthetic rate and sink development responded strongly to elevated [CO₂] in the perennial, indicating significant source limitation. Sink limitation was avoided in the perennial by high sink plasticity: a marked increase in tillering and root:shoot ratio at elevated [CO₂], and lower non‐structural carbohydrate accumulation. Alleviating sink limitation during vegetative development could be important for maximizing growth of elite cereals under future elevated [CO₂].