Photosynthesis is a highly integrated and regulated process which is highly sensitive to any change in environmental conditions, because it needs to balance the light energy absorbed by the photosystems with the energy consumed by metabolic sinks of the plant. Low temperatures exacerbate an imbalance between the source of energy and the metabolic sink, thus requiring adjustments of photosynthesis to maintain the balance of energy flow. Photosynthesis itself functions as a sensor of this imbalance through the redox state of photosynthetic electron-transport components and regulates photophysical, photochemical and metabolic processes in the chloroplast. Recent progress has been made in understanding how plants sense the low temperature signal. It is clear that photosynthesis interacts with other processes during cold acclimation involving crosstalk between photosynthetic redox, cold acclimation and sugar-signalling pathways to regulate plant acclimation to low temperatures.

Photosynthetic carbon gain in plants using the C3 photosynthetic pathway is substantially inhibited by photorespiration in warm environments, particularly in atmospheres with low CO2 concentrations. Unlike C4 plants, C3 plants are thought to lack any mechanism to compensate for the loss of photosynthetic productivity caused by photorespiration. Here, for the first time, we demonstrate that the C3 plants rice and wheat employ a specific mechanism to trap and reassimilate photorespired CO2. A continuous layer of chloroplasts covering the portion of the mesophyll cell periphery that is exposed to the intercellular air space creates a diffusion barrier for CO2 exiting the cell. This facilitates the capture and reassimilation of photorespired CO2 in the chloroplast stroma. In both species, 24-38% of photorespired and respired CO2 were reassimilated within the cell, thereby boosting photosynthesis by 8-11% at ambient atmospheric CO2 concentration and 17-33% at a CO2 concentration of 200 mmol mol -1 . Widespread use of this mechanism in tropical and subtropical C3 plants could explain why the diversity of the world's C3 flora, and dominance of terrestrial net primary productivity, was maintained during the Pleistocene, when atmospheric CO2 concentrations fell below 200 mmol mol -1 .

Contents 986I.987II.987III.988IV.991V.992VI.995VII.997VIII.998References998 Summary It has been 75 yr since leaf respiratory metabolism in the light (day respiration) was identified as a low‐flux metabolic pathway that accompanies photosynthesis. In principle, it provides carbon backbones for nitrogen assimilation and evolves CO2 and thus impacts on plant carbon and nitrogen balances. However, for a long time, uncertainties have remained as to whether techniques used to measure day respiratory efflux were valid and whether day respiration responded to environmental gaseous conditions. In the past few years, significant advances have been made using carbon isotopes, ‘omics’ analyses and surveys of respiration rates in mesocosms or ecosystems. There is substantial evidence that day respiration should be viewed as a highly dynamic metabolic pathway that interacts with photosynthesis and photorespiration and responds to atmospheric CO2 mole fraction. The view of leaf day respiration as a constant and/or negligible parameter of net carbon exchange is now outdated and it should now be regarded as a central actor of plant carbon‐use efficiency.

Photorespiration is a major bioengineering target for increasing crop yields as it is often considered a wasteful process. Photorespiratory metabolism is integrated into leaf metabolism and thus may have certain benefits. Here, we show that plants can increase their rate of photosynthetic CO uptake when assimilating nitrogen de novo via the photorespiratory pathway by fixing carbon as amino acids in addition to carbohydrates. Plants fed NO had higher rates of CO assimilation under photorespiratory than low-photorespiratory conditions, while plants lacking NO nutrition exhibited lower stimulation of CO uptake. We modified the widely used Farquhar, von Caemmerer and Berry photosynthesis model to include the carbon and electron requirements for nitrogen assimilation via the photorespiratory pathway. Our modified model improves predictions of photosynthetic CO uptake and of rates of photosynthetic electron transport. The results highlight how photorespiration can improve photosynthetic performance despite reducing the efficiency of Rubisco carboxylation.

Rubisco catalyzes a rate-limiting step in photosynthesis and has long been a target for improvement due to its slow turnover rate. An alternative to modifying catalytic properties of Rubisco is to increase its abundance within C 4 plant chloroplasts, which might increase activity and confer a higher carbon assimilation rate. Here, we overexpress the Rubisco large (LS) and small (SS) subunits with the Rubisco assembly chaperone RAF1. While overexpression of LS and/or SS had no discernable impact on Rubisco content, addition of RAF1 overexpression resulted in a >30% increase in Rubisco content. Gas exchange showed a 15% increase in CO 2 assimilation (A SAT ) in UBI-LSSS-RAF1 transgenic plants, which correlated with increased fresh weight and in vitro V cmax calculations. The divergence of Rubisco content and assimilation could be accounted for by the Rubisco activation state, which decreased up to 23%, suggesting that Rubisco activase may be limiting V cmax , and impinging on the realization of photosynthetic potential from increased Rubisco content.

Summary Photorespiratory metabolism is essential for plants to maintain functional photosynthesis in an oxygen‐containing environment. Because the oxygenation reaction of Rubisco is followed by the loss of previously fixed carbon, photorespiration is often considered a wasteful process and considerable efforts are aimed at minimizing the negative impact of photorespiration on the plant’s carbon uptake. However, the photorespiratory pathway has also many positive aspects, as it is well integrated within other metabolic processes, such as nitrogen assimilation and C1 metabolism, and it is important for maintaining the redox balance of the plant. The overall effect of photorespiratory carbon loss on the net CO2 fixation of the plant is also strongly influenced by the physiology of the leaf related to CO2 diffusion. This review outlines the distinction between Rubisco oxygenation and photorespiratory CO2 release as a basis to evaluate the costs and benefits of photorespiration.

Temperature and daylength act as environmental signals that determine the length of the growing season in boreal evergreen conifers. Climate change might affect the seasonal development of these trees, as they will experience naturally decreasing daylength during autumn, while at the same time warmer air temperature will maintain photosynthesis and respiration. We characterized the down-regulation of photosynthetic gas exchange and the mechanisms involved in the dissipation of energy in Jack pine (Pinus banksiana) in controlled environments during a simulated summer-autumn transition under natural conditions and conditions with altered air temperature and photoperiod. Using a factorial design, we dissected the effects of daylength and temperature. Control plants were grown at either warm summer conditions with 16-h photoperiod and 22°C or conditions representing a cool autumn with 8 h/7°C. To assess the impact of photoperiod and temperature on photosynthesis and energy dissipation, plants were also grown under either cold summer (16-h photoperiod/7°C) or warm autumn conditions (8-h photoperiod/22°C). Photosynthetic gas exchange was affected by both daylength and temperature. Assimilation and respiration rates under warm autumn conditions were only about one-half of the summer values but were similar to values obtained for cold summer and natural autumn treatments. In contrast, photosynthetic efficiency was largely determined by temperature but not by daylength. Plants of different treatments followed different strategies for dissipating excess energy. Whereas in the warm summer treatment safe dissipation of excess energy was facilitated via zeaxanthin, in all other treatments dissipation of excess energy was facilitated predominantly via increased aggregation of the light-harvesting complex of photosystem II. These differences were accompanied by a lower deepoxidation state and larger amounts of b-carotene in the warm autumn treatment as well as by changes in the abundance of thylakoid membrane proteins compared to the summer condition. We conclude that photoperiod control of dormancy in Jack pine appears to negate any potential for an increased carbon gain associated with higher temperatures during the autumn season.

Mitochondrial respiration often appears to be inhibited in the light when compared with measurements in the dark. This inhibition is inferred from the response of the net CO assimilation rate (A) to absorbed irradiance (I), changing slope around the light compensation point (I ). We suggest a model that provides a plausible mechanistic explanation of this 'Kok effect'. The model uses the mathematical description of photosynthesis developed by Farquhar, von Caemmerer and Berry; it involves no inhibition of respiration rate in the light. We also describe a fitting technique for quantifying the Kok effect at low I. Changes in the chloroplastic CO partial pressure (C ) can explain curvature of A vs I, its diminution in C plants and at low oxygen concentrations or high carbon dioxide concentrations in C plants, and effects of dark respiration rate and of temperature. It also explains the apparent inhibition of respiration in the light as inferred by the Laisk approach. While there are probably other sources of curvature in A vs I, variation in C can largely explain the curvature at low irradiance, and suggests that interpretation of day respiration compared with dark respiration of leaves on the basis of the Kok effect needs reassessment.