Incident irradiance on plant leaves often fluctuates, causing dynamic photosynthesis. Whereas steady-state photosynthetic responses to environmental factors have been extensively studied, knowledge of dynamic modulation of photosynthesis remains scarce and scattered. This review addresses this discrepancy by summarizing available data and identifying the research questions necessary to advance our understanding of interactions between environmental factors and dynamic behaviour of photosynthesis using a mechanistic framework. Firstly, dynamic photosynthesis is separated into sub-processes related to proton and electron transport, non-photochemical quenching, control of metabolite flux through the Calvin cycle (activation states of Rubisco and RuBP regeneration, and post-illumination metabolite turnover), and control of CO₂ supply to Rubisco (stomatal and mesophyll conductance changes). Secondly, the modulation of dynamic photosynthesis and its sub-processes by environmental factors is described. Increases in ambient CO₂ concentration and temperature (up to ~35°C) enhance rates of photosynthetic induction and decrease its loss, facilitating more efficient dynamic photosynthesis. Depending on the sensitivity of stomatal conductance, dynamic photosynthesis may additionally be modulated by air humidity. Major knowledge gaps exist regarding environmental modulation of loss of photosynthetic induction, dynamic changes in mesophyll conductance, and the extent of limitations imposed by stomatal conductance for different species and environmental conditions. The study of mutants or genetic transformants for specific processes under various environmental conditions could provide significant progress in understanding the control of dynamic photosynthesis.
Diffuse light enhanced crop photosynthesis. A more uniform horizontal PPFD distribution played the most important role in this enhancement, and a more uniform vertical PPFD distribution and higher leaf photosynthetic capacity contributed more to the enhancement of crop photosynthesis than did higher values of LAI.
Greenhouse tomato (Solanum lycopersicum) yield in The Netherlands has increased tremendously over the past 50 years. The effects of breeding during this period were investigated. Eight Dutch cultivars and one typical current Japanese cultivar that were released over the past 50 years were compared in a short-term experiment conducted from summer to fall in The Netherlands. Fresh fruit yield of the Dutch cultivars significantly increased ≈0.9% per year with the year of release from 1950 to 2000. Dry weight fruit yield of the Dutch cultivars also increased with the year of release, whereas the fruit dry matter content was not correlated with the year of release. Total dry matter production of plants increased with the year of release, and the dry matter partitioning to fruit was not correlated with the year of release. An increase in dry matter production was caused not by an increase in fraction of intercepted light, but by light use efficiency based on correlations between each of them and the year of release. The light extinction coefficient in the plant canopy decreased, whereas leaf photosynthetic rate increased significantly with the year of release. Although fresh fruit yield of the Japanese cultivar was lower than that of the modern Dutch cultivars, fruit dry matter content of the Japanese cultivar was higher than that of the Dutch cultivars. An increase in yield over the past 50 years in Dutch tomato was caused by an increase in light use efficiency resulting from a decrease in light extinction coefficient (a morphological change) and an increase in leaf photosynthetic rate (a physiological change).
An important constraint for plant biomass production is the natural day length. Artificial light allows for longer photoperiods, but tomato plants develop a detrimental leaf injury when grown under continuous light-a still poorly understood phenomenon discovered in the 1920s. Here, we report a dominant locus on chromosome 7 of wild tomato species that confers continuous light tolerance. Genetic evidence, RNAseq data, silencing experiments and sequence analysis all point to the type III light harvesting chlorophyll a/b binding protein 13 (CAB-13) gene as a major factor responsible for the tolerance. In Arabidopsis thaliana, this protein is thought to have a regulatory role balancing light harvesting by photosystems I and II. Introgressing the tolerance into modern tomato hybrid lines, results in up to 20% yield increase, showing that limitations for crop productivity, caused by the adaptation of plants to the terrestrial 24-h day/night cycle, can be overcome
The environmental factors CO partial pressure, temperature and VPD had significant impacts on rates of photosynthetic induction, as well as on underlying diffusional, carboxylation and electron transport processes. Furthermore, maximizing Rubisco activation rates would increase photosynthesis by at most 6-8 % in ambient CO partial pressure (across temperatures and humidities), while maximizing rates of stomatal opening would increase photosynthesis by at most 1-3 %.
A better understanding of the metabolic and diffusional limitations of photosynthesis in fluctuating irradiance can help identify targets for improving crop yields. We used different genotypes of Arabidopsis thaliana to characterise the importance of Rubisco activase (Rca), stomatal conductance (gs), non-photochemical quenching of chlorophyll fluorescence (NPQ) and sucrose phosphate synthase (SPS) on photosynthesis in fluctuating irradiance. Leaf gas exchange and chlorophyll fluorescence were measured in leaves exposed to stepwise increases and decreases in irradiance. rwt43, which has a constitutively active Rubisco enzyme in different irradiance intensities (except in darkness), showed faster increases than the wildtype, Colombia-0, in photosynthesis rates after step increases in irradiance. rca-2, having decreased Rca concentration, showed lower rates of increase. In aba2-1, high gs increased the rate of change after stepwise irradiance increases, while in C24, low gs tended to decrease it. Differences in rates of change between Colombia-0 and plants with low levels of NPQ (npq1-2, npq4-1) or SPS (spsa1) were negligible. In Colombia-0, the regulation of Rubisco activation and of gs were therefore limiting for photosynthesis in fluctuating irradiance, while levels of NPQ or SPS were not. This suggests Rca and gs as targets for improvement of photosynthesis of plants in fluctuating irradiance.
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