Understanding the Low Photosynthetic Rates of Sun and Shade Coffee Leaves: Bridging the Gap on the Relative Roles of Hydraulic,Diffusive and Biochemical Constraints to Photosynthesis

It has long been held that the low photosynthetic rates (A) of coffee leaves are largely associated with diffusive constraints to photosynthesis. However, the relative limitations of the stomata and mesophyll to the overall diffusional constraints to photosynthesis, as well as the coordination of leaf hydraulics with photosynthetic limitations, remain to be fully elucidated in coffee. Whether the low actual A under ambient CO₂ concentrations is associated with the kinetic properties of Rubisco and high (photo)respiration rates also remains elusive. Here, we provide a holistic analysis to understand the causes associated with low A by measuring a variety of key anatomical/hydraulic and photosynthetic traits in sun- and shade-grown coffee plants. We demonstrate that leaf hydraulic architecture imposes a major constraint on the maximisation of the photosynthetic gas exchange of coffee leaves. Regardless of the light treatments, A was mainly limited by stomatal factors followed by similar limitations associated with the mesophyll and biochemical constraints. No evidence of an inefficient Rubisco was found; rather, we propose that coffee Rubisco is well tuned for operating at low chloroplastic CO₂ concentrations. Finally, we contend that large diffusive resistance should lead to large CO₂ drawdown from the intercellular airspaces to the sites of carboxylation, thus favouring the occurrence of relatively high photorespiration rates, which ultimately leads to further limitations to A.     

Carbon dioxide diffusion across stomata and mesophyll and photo-biochemical processes as affected by growth CO2 and phosphorus nutrition in cotton

Nutrients such as phosphorus may exert a major control over plant response to rising atmospheric carbon dioxide concentration (CO₂), which is projected to double by the end of the 21st century. Elevated CO₂ may overcome the diffusional limitations to photosynthesis posed by stomata and mesophyll and alter the photo-biochemical limitations resulting from phosphorus deficiency. To evaluate these ideas, cotton (Gossypium hirsutum) was grown in controlled environment growth chambers with three levels of phosphate (Pi) supply (0.2, 0.05 and 0.01 mM) and two levels of CO₂ concentration (ambient 400 and elevated 800 μmol mol⁻¹) under optimum temperature and irrigation. Phosphate deficiency drastically inhibited photosynthetic characteristics and decreased cotton growth for both CO₂ treatments. Under Pi stress, an apparent limitation to the photosynthetic potential was evident by CO₂ diffusion through stomata and mesophyll, impairment of photosystem functioning and inhibition of biochemical process including the carboxylation efficiency of ribulose-1,5-bisphosphate carboxylase/oxyganase and the rate of ribulose-1,5-bisphosphate regeneration. The diffusional limitation posed by mesophyll was up to 58% greater than the limitation due to stomatal conductance (g_s) under Pi stress. As expected, elevated CO₂ reduced these diffusional limitations to photosynthesis across Pi levels; however, it failed to reduce the photo-biochemical limitations to photosynthesis in phosphorus deficient plants. Acclimation/down regulation of photosynthetic capacity was evident under elevated CO₂ across Pi treatments. Despite a decrease in phosphorus, nitrogen and chlorophyll concentrations in leaf tissue and reduced stomatal conductance at elevated CO₂, the rate of photosynthesis per unit leaf area when measured at the growth CO₂ concentration tended to be higher for all except the lowest Pi treatment. Nevertheless, plant biomass increased at elevated CO₂ across Pi nutrition with taller plants, increased leaf number and larger leaf area.     

Photosynthetic response of Tempranillo grapevine to climate change scenarios

Photosynthesis in C₃ plants is CO₂ limited and therefore any increase in Rubisco carboxylation substrate may increase net CO₂ fixation, unless plants experience acclimation or other limitations. These aspects are largely unexplored in grapevine. Photosynthesis analysis was used to assess the stomatal, mesophyll, photochemical and biochemical contributions to the decreasing photosynthesis observed in Tempranillo grapevines (Vitis vinifera) from veraison to ripeness, modulated by CO₂, temperature and water availability. Photosynthesis and photosystem II photochemistry decreased from veraison to ripeness. The elevated CO₂ and temperature increased photosynthesis, but transiently, in both well irrigated (WI) and water‐stressed plants. Photosynthetic rates were maxima 1 week after the start of elevated CO₂ and temperature treatments, but differences with treatments of ambient conditions disappeared with time. There were not marked changes in leaf water status, leaf chlorophyll or leaf protein that could limit photosynthesis at ripeness. Leaf total soluble sugars remained at ripeness as high as 2 weeks after the start of treatments. On the other hand, and as expected, CO₂ diffusional limitations impaired photosynthesis in grapevine plants grown under water scarcity, stomatal and mesophyll conductances to CO₂ decreased and in turn low chloroplastic CO₂ concentrations limited photosynthetic CO₂ fixation. In summary, photochemistry and photosynthesis from veraison to ripeness in Tempranillo grapevine were dominated by a developmental‐related decreasing trend that was only transiently influenced by elevated CO₂ concentrations.     

Disentangling the contributions of ontogeny and water stress to photosynthetic limitations in almond trees

Very few studies have attempted to disentangle the respective role of ontogeny and water stress on leaf photosynthetic attributes. The relative significance of both effects on photosynthetic attributes has been investigated in leaves of field‐grown almond trees [Prunus dulcis (Mill.) D. A. Webb] during four growth cycles. Leaf ontogeny resulted in enhanced leaf dry weight per unit area (W_a), greater leaf dry‐to‐fresh weight ratio and lower N content per unit of leaf dry weight (N_w). Concomitantly, area‐based maximum carboxylation rate (V_cmax), maximum electron transport rate (J_max), mesophyll conductance to CO₂ diffusion (gm)′ and light‐saturated net photosynthesis (A_max) declined in both well‐watered and water‐stressed almond leaves. Although g_m and stomatal conductance (g_s) seemed to be co‐ordinated, a much stronger coordination in response to ontogeny and prolonged water stress was observed between g_m and the leaf photosynthetic capacity. Under unrestricted water supply, the leaf age‐related decline of A_max was equally driven by diffusional and biochemical limitations. Under restricted soil water availability, A_max was mainly limited by g_s and, to a lesser extent, by photosynthetic capacity and g_m. When both ontogeny and water stress effects were combined, diffusional limitations was the main determinant of photosynthesis limitation, while stomatal and biochemical limitations contributed similarly.     

Leaf anatomy does not explain apparent short‐term responses of mesophyll conductance to light and CO2 in tobacco

Mesophyll conductance to CO₂ (g_m), a key photosynthetic trait, is strongly constrained by leaf anatomy. Leaf anatomical parameters such as cell wall thickness and chloroplast area exposed to the mesophyll intercellular airspace have been demonstrated to determine g_m in species with diverging phylogeny, leaf structure and ontogeny. However, the potential implication of leaf anatomy, especially chloroplast movement, on the short‐term response of g_m to rapid changes (i.e. seconds to minutes) under different environmental conditions (CO₂, light or temperature) has not been examined. The aim of this study was to determine whether the observed rapid variations of g_m in response to variations of light and CO₂ could be explained by changes in any leaf anatomical arrangements. When compared to high light and ambient CO₂, the values of g_m estimated by chlorophyll fluorescence decreased under high CO₂ and increased at low CO₂, while it decreased with decreasing light. Nevertheless, no changes in anatomical parameters, including chloroplast distribution, were found. Hence, the g_m estimated by analytical models based on anatomical parameters was constant under varying light and CO₂. Considering this discrepancy between anatomy and chlorophyll fluorescence estimates, it is concluded that apparent fast g_m variations should be due to artefacts in its estimation and/or to changes in the biochemical components acting on diffusional properties of the leaf (e.g. aquaporins and carbonic anhydrase).     

Photosynthetic responses to slowly decreasing leaf water potentials in Encelia frutescens

The importance of reduced leaf conductance (stomatal and boundary layer) in limiting photosynthetic rates during water stress was studied in Encelia frutescens, a drought-deciduous leaved subshrub of the Mohave and Sonoran Deserts. Light-saturated CO₂ assimilation rates of greenhouse grown plants decreased from 42.6±1.6mol CO₂ m^-2 s^-1 (x±s.e.) to 1.7±1.7 mol CO₂ m^-2s^-1 as leaf water potential decreased from-1.5 MPa to-4.0 MPa. The dependence of light saturated, CO₂ assimilation rate on leaf intercellular CO₂ concentrations between 60 and 335 l l^-1 was also determined as leaf water potential decline. This enabled us to compare the effects of leaf water potentials on limitations to carbon assimilation imposed by leaf conductance and by intrinsic photosynthetic capacity. Both leaf conductance and intrinsic photosynthetic capacity decreased with decreasing leaf water potential, but the decrease in leaf conductance was proportionately greater. The relative stomatal limitation, defined as the percent limitation in photosynthetic rate due to the presence of gas-phase diffusional barriers, increased from (x±s.e.) to 41±3% as water potentials became more negative. Since both leaf conductance and intrinsic photosynthetic capacity were severely reduced in an absolute sense, however, high photosynthetic rates could not have been restored at low leaf water potentials without simultaneous increases in both components.     

Photosynthesis in intact leaves of C3 plants: Physics,physiology and rate limitations

The instantaneous rate of photosynthetic CO₂ assimilation in C₃ plants has generally been studied in model systems such as isolated chloroplasts and algae. From these studies and from theoretical analyses of gas exchange behavior it is now possible to study the biochemistry of photosynthesis in intact leaves using a combination of methods, most of which are nondestructive. The limitations to the rate of photosynthesis can be divided among three general classes: (1) the supply or utilization of CO₂, (2) the supply or utilization of light, and (3) the supply or utilization of phosphate. The first limitation is most readily studied by determining how the CO₂ assimilation rate varies with the partial pressure of CO₂ inside the leaf. The second limitation can be studied by determining the quantum requirement of photosynthesis. The third limitation is most easily detected as a loss of O₂ sensitivity of photosynthesis. Measurement of fluorescence from intact leaves can give additional information about the various limitations. These methods are all non-destructive and so can be observed repeatedly as the environment of a leaf is changed. In addition, leaves can be quick-frozen and metabolite concentrations then measured to give more information about the limitations to intact leaf photosynthesis rates. In this review the physics and biochemistry of photosynthesis in intact C₃ leaves, and the interface between physiology and photosynthesis—triose phosphate utilization—are discussed.     

Growth Kinetics, Carbohydrate, and Leaf Phosphate Content of Clover (Trifolium subterraneum L.) after Transfer to a High CO(2) Atmosphere or to High Light and Ambient Air

Intact air-grown (photosynthetic photon flux density, 400 microeinsteins per square meter per second) clover plants (Trifolium subterraneum L.) were transfered to high CO₂ (4000 microliters CO₂ per liter; photosynthetic photon flux density, 400 microeinsteins per square meter per second) or to high light (340 microliters CO₂ per liter; photosynthetic photon flux density, 800 microeinsteins per square meter per second) to similarly stimulate photosynthetic net CO₂ uptake. The daily increment of net CO₂ uptake declined transiently in high CO₂, but not in high light, below the values in air/standard light. After about 3 days in high CO₂, the daily increment of net CO₂ uptake increased but did not reach the high light values. Nightly CO₂ release increased immediately in high light, whereas there was a 3-day lag phase in high CO₂. During this time, starch accumulated to a high level, and leaf deterioration was observed only in high CO₂. After 12 days, starch was two- to threefold higher in high CO₂ than in high light, whereas sucrose was similar. Leaf carbohydrates were determined during the first and fourth day in high CO₂. Starch increased rapidly throughout the day. Early in the day, sucrose was low and similar in high CO₂ and ambient air (same light). Later, sucrose increased considerably in high CO₂. The findings that (a) much more photosynthetic carbon was partitioned into the leaf starch pool in high CO₂ than in high light, although net CO₂ uptake was similar, and that (b) rapid starch formation occurred in high CO₂ even when leaf sucrose was only slightly elevated suggest that low sink capacity was not the main constraint in high CO₂. It is proposed that carbon partitioning between starch (chloroplast) and sucrose (cytosol) was perturbed by high CO₂ because of the lack of photorespiration. Total phosphate pools were determined in leaves. Concentrations based on fresh weight of orthophosphate, soluble esterified phosphate, and total phosphate markedly declined during 13 days of exposure of the plants to high CO₂ but changed little in high light/ambient air. During this time, the ratio of orthophosphate to soluble esterified phosphate decreased considerably in high CO₂ and increased slightly in high light/ambient air. It appears that phosphate uptake and growth were similarly stimulated by high light, whereas the coordination was weak in high CO₂.     

Low CO2 does not remove diffusional limitation to photosynthesis in salt stressed tomato during osmotic phase

Salinity is one of the main environmental stress for crops such as tomato which is widely spread in the Mediterranean region. It is now widely considered that in plant adaptation to salts, the time scale of the response is essential. During the initial phase of response to salinity, when the osmotic effect predominates, plant response to salts is regulated by hormones, and between them abscisic acid plays a crucial role. We showed that on tomato crops during the first phase (osmotic one) low CO₂ conditioning treatment before photosynthetic response to carbon dioxide concentration (A–Ci curve) is not applicable because diffusional limitations to photosynthesis is not removed by low CO₂ air concentration. Conditioning at low CO₂ is not sufficient to remove diffusional limitation of photosynthesis in salt stressed tomatoes during osmotic phase.     

Leaf photosynthesis of Haberlea rhodopensis before and during drought

Haberlea rhodopensis is a homoiochlorophyllous resurrection plant that shows a low rate of leaf net CO₂ uptake (4–6 μmol m^?2 s^?1) under saturating photosynthetic photon flux densities in air (21% O₂ and about 390 ppm CO₂). However, leaf net CO₂ uptake reaches values of 17–18 μmol m^?2 s^?1 under saturating CO₂ and light. H. rhodopensis leaves have a very low mesophyll CO₂ conductance that can partly explain the low rate of leaf net CO₂ uptake in normal air. Experimental evidences suggest that mesophyll conductance is not sensitive to temperature in the 20–35 °C range. In addition, it is shown that the (1) transpiration rate of H. rhodopensis is nearly linearly related to the vapour pressure difference between the leaf and the ambient air within the interval from 0.5 kPa to 2.5 kPa at a leaf temperature of 25 °C and (2) leaf net CO₂ uptake in normal air under saturating light does not change much with leaf temperature (between 20 °C and 30 °C). At a leaf relative water content of between 90% and 30%, the decrease of leaf net CO₂ assimilation during drought can be explained by a decrease of leaf CO₂ diffusional conductance. Accordingly the non-photochemical chlorophyll fluorescence quenching decreases only at relative water contents lower than 20%, indicating that photosynthetic activity maintains a trans-thylakoidal proton gradient over a wide range of leaf water contents. Moreover, PSII photochemistry (as estimated by the Fv/Fm ratio and the thermoluminescence B band intensity) is only affected at leaf relative water contents lower than about 20%, thus confirming that primary photosynthetic reactions are resistant to drought. Interestingly, the effect of leaf desiccation on photosynthetic capacity, measured at very high ambient CO₂ molar ratios under saturating PPFD, is identical to that observed for three non-resurrection C₃ mesophytes. This demonstrates that the photosynthetic apparatus of H. rhodopensis is not more resistant to desiccation when compared to other C₃ plants. Since the leaf area decreases by more than 50% when the leaf relative water content is reduced to about 40% during drought it is supposed, following Farrant et al. [Farrant, J.M., Vander, W.C., Lofell, D.A., Bartsch, S., Whittaker, A., 2003. An investigation into the role of light during desiccation of three angiosperms resurrection plants. Plant Cell Environ. 26, 1275–1286], that H. rhodopensis leaf cells avoid mechanical stress.     

Three‐dimensional microscale modelling of CO2 transport and light propagation in tomato leaves enlightens photosynthesis

We present a combined three‐dimensional (3‐D) model of light propagation, CO₂ diffusion and photosynthesis in tomato (Solanum lycopersicum L.) leaves. The model incorporates a geometrical representation of the actual leaf microstructure that we obtained with synchrotron radiation X‐ray laminography, and was evaluated using measurements of gas exchange and leaf optical properties. The combination of the 3‐D microstructure of leaf tissue and chloroplast movement induced by changes in light intensity affects the simulated CO₂ transport within the leaf. The model predicts extensive reassimilation of CO₂ produced by respiration and photorespiration. Simulations also suggest that carbonic anhydrase could enhance photosynthesis at low CO₂ levels but had little impact on photosynthesis at high CO₂ levels. The model confirms that scaling of photosynthetic capacity with absorbed light would improve efficiency of CO₂ fixation in the leaf, especially at low light intensity.     

Photosynthetic and anatomical responses of Eucalyptus grandis leaves to potassium and sodium supply in a field experiment

Although vast areas in tropical regions have weathered soils with low potassium (K) levels, little is known about the effects of K supply on the photosynthetic physiology of trees. This study assessed the effects of K and sodium (Na) supply on the diffusional and biochemical limitations to photosynthesis in Eucalyptus grandis leaves. A field experiment comparing treatments receiving K (+K) or Na (+Na) with a control treatment (C) was set up in a K‐deficient soil. The net CO₂ assimilation rates were twice as high in +K and 1.6 times higher in +Na than in the C as a result of lower stomatal and mesophyll resistance to CO₂ diffusion and higher photosynthetic capacity. The starch content was higher and soluble sugar was lower in +K than in C and +Na, suggesting that K starvation disturbed carbon storage and transport. The specific leaf area, leaf thickness, parenchyma thickness, stomatal size and intercellular air spaces increased in +K and +Na compared to C. Nitrogen and chlorophyll concentrations were also higher in +K and +Na than in C. These results suggest a strong relationship between the K and Na supply to E. grandis trees and the functional and structural limitations to CO₂ assimilation rates.     

Phosphoglycerate dehydrogenase from soybean nodules : partial purification and some kinetic properties

The relationship between single leaf photosynthesis and conductance was examined in cotton (Gossypium hirsutum L.) across a range of environmental conditions. The purpose of this research was to separate and define the degree of stomatal and nonstomatal limitations in the photosynthetic process of field-grown cotton.

Photosynthetic rates were related to leaf conductance of upper canopy leaves in a curvilinear manner. Increases in leaf conductance of CO₂ in excess of 0.3 to 0.4 mole per square meter per second did not result in significant increases in gross or net photosynthetic rates. No tight coupling between environmental influences on photosynthetic rates and those affecting conductance levels was evident, since photosynthesis per unit leaf conductance did not remain constant. Slowly developing water stress caused greater reductions in photosynthesis than in leaf conductance, indicating nonstomatal limitations of photosynthesis.

Increases in external CO₂ concentration to levels above ambient did not produce proportional increases in photosynthesis even though substomatal or intercellular CO₂ concentration increased. The lack of a linear increase in photosynthetic rate in response to increases in leaf conductance and in response to increases in external CO₂ concentration demonstrated that nonstomatal factors are major photosynthetic rate determinants of cotton under field conditions.

     

The winter-red-leaf syndrome in Pistacia lentiscus: evidence that the anthocyanic phenotype suffers from nitrogen deficiency, low carboxylation efficiency and high risk of photoinhibition

Recent evidence indicates that winter-red leaf phenotypes in the mastic tree (Pistacia lentiscus) are more vulnerable to chronic photoinhibition during the cold season relative to winter-green phenotypes occurring in the same high light environment. This was judged by limitations in the maximum quantum yield of photosystem II (PSII), found in previous studies. In this investigation, we asked whether corresponding limitations in leaf gas exchange and carboxylation reactions could also be manifested. During the cold (“red”) season, net CO₂ assimilation rates (A) and stomatal conductances (g_s) in the red phenotype were considerably lower than in the green phenotype, while leaf internal CO₂ concentration (Ci) was higher. The differences were abolished in the “green” period of the year, the dry summer included. Analysis of A versus Ci curves indicated that CO₂ assimilation during winter in the red phenotype was limited by Rubisco content and/or activity rather than stomatal conductance. Leaf nitrogen levels in the red phenotype were considerably lower during the red-leaf period. Consequently, we suggest that the inherently low leaf nitrogen levels are linked to the low net photosynthetic rates of the red plants through a decrease in Rubisco content. Accordingly, the reduced capacity of the carboxylation reactions to act as photosynthetic electron sinks may explain the corresponding loss of PSII photon trapping efficiency, which cannot be fully alleviated by the screening effect of the accumulated anthocyanins.     

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.     

Changes in photosynthesis caused by adaptation of maize seedlings to short-term drought

Detached leaf is in the state of increasing water deficit; it is a good experimental model for looking into the hardening effect of adaptation of eight-day-old maize (Zea mays L.) seedlings to short-term drought (five days without watering). The light stage of photosynthesis and photosynthetic CO₂/H₂O exchange in detached leaves were studied. Specific surface density of leaf tissue (SSDL), the content of chlorophylls a and b, proline, MDA as well as photosynthetic parameters: quantum yield of photosystem II fluorescence, assimilation of CO₂, and transpiration at room temperature and light saturation (density of PAR quantum flux of 2000 μmol/(m² s)) at normal and half atmospheric CO₂ concentration were determined. The leaves of seedlings exposed to short-term drought differed from control material by a greater SSDL and higher content of proline. The hardening effect of the stress agent on the dark stage of photosynthesis was detected; it was expressed in the maintenance of the higher photosynthetic CO₂ assimilation against control material due to the elevation of stomatal conductance for CO₂ diffusing into the leaf. Judging from the lack of differences in the MDA content, short-term drought did not injure photosynthetic membranes. In detached leaves of experimental maize seedlings, photosynthesis was maintained on a higher level than in control material.     

The phenology of leaf quality and its within‐canopy variation is essential for accurate modeling of photosynthesis in tropical evergreen forests

Leaf quantity (i.e., canopy leaf area index, LAI), quality (i.e., per‐area photosynthetic capacity), and longevity all influence the photosynthetic seasonality of tropical evergreen forests. However, these components of tropical leaf phenology are poorly represented in most terrestrial biosphere models (TBMs). Here, we explored alternative options for the representation of leaf phenology effects in TBMs that employ the Farquahar, von Caemmerer & Berry (FvCB) representation of CO₂ assimilation. We developed a two‐fraction leaf (sun and shade), two‐layer canopy (upper and lower) photosynthesis model to evaluate different modeling approaches and assessed three components of phenological variations (i.e., leaf quantity, quality, and within‐canopy variation in leaf longevity). Our model was driven by the prescribed seasonality of leaf quantity and quality derived from ground‐based measurements within an Amazonian evergreen forest. Modeled photosynthetic seasonality was not sensitive to leaf quantity, but was highly sensitive to leaf quality and its vertical distribution within the canopy, with markedly more sensitivity to upper canopy leaf quality. This is because light absorption in tropical canopies is near maximal for the entire year, implying that seasonal changes in LAI have little impact on total canopy light absorption; and because leaf quality has a greater effect on photosynthesis of sunlit leaves than light limited, shade leaves and sunlit foliage are more abundant in the upper canopy. Our two‐fraction leaf, two‐layer canopy model, which accounted for all three phenological components, was able to simulate photosynthetic seasonality, explaining ~90% of the average seasonal variation in eddy covariance‐derived CO₂ assimilation. This work identifies a parsimonious approach for representing tropical evergreen forest photosynthetic seasonality in TBMs that utilize the FvCB model of CO₂ assimilation and highlights the importance of incorporating more realistic phenological mechanisms in models that seek to improve the projection of future carbon dynamics in tropical evergreen forests.     

Light-dependent damage to photosynthesis in olive leaves during chilling and high temperature stress

Abstract The leaves of olive are long lived and likely to experience both chilling and high temperature stress during their life. Changes in photosynthetic CO₂ assimilation resulting from chilling and high temperature stress, in both dim and high light, are investigated. The quantum yield (φ) of photosynthesis at limiting light levels was reduced following chilling (at 5°C for 12 h), in dim light by approximately 10%, and in high light by 75%; the difference being attributed to photoinhibition. Similar reductions were observed in the light-saturated rate of CO₂ uptake (A_max). Decrease in A_max correlated with a halving of the leaf internal CO₂ concentration (c_i), suggesting an increased limitation by stomata following photoinhibition. Leaves were apparently more susceptible to photoinhibitory damage if the whole plant, rather than the leaf alone, was chilled. On return to 26 °C, I he photosynthetic capacity recovered to pre-stress levels within a few hours if leaves had been chilled in high light for 8 h or less, but did not fully recover from longer periods of chilling when loss of chlorophyll occurred. Leaves which were recovering from chilling in high light showed far more damage on being chilled a second time in high light. Three hours in high light at 38 °C reduced φ by 80%, but φ recovered within 4h of return to 26 °C. Although leaves of Olive are apparently less susceptible to photoinhibitory damage during chilling stress than the short-lived leaves of chilling-sensitive annual? crops, the results nevertheless show that photoinhibition during temperature stress is potentially a major factor influencing the photosynthetic productivity of Olive in the field.     

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].