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It has long been held that tree species/leaves from shaded environments show faster rate of photosynthetic induction than species/leaves from sunlit environments, but the evidence so far is conflicting and the underlying mechanisms are still under debate. To address the debate, we compiled a dataset for 87 tree species and compared the initial increasing slope during the first 2-min induction (SA) and stomatal and biochemical characteristics between sun and shade species from the same study, and those between sun and shade leaves within the same species. In 77% of between-species comparisons, the species with high steady-state photosynthetic rate in the high light (Af) exhibited a larger SA than the species with low Af. In 67% within-species comparisons, the sun leaves exhibited a larger SA than the shade leaves. However, in only a few instances did the sun species/leaves more rapidly achieve 50% of full induction, with an even smaller SA, than the shade species/leaves. At both the species and leaf level, SA increased with increasing initial stomatal conductance before induction (gsi). Despite exhibiting reduced intrinsic water-use efficiency in low light, a large SA proportionally enhances photosynthetic carbon gain during the first 2-min induction in the sun species and leaves. Thus, in terms of the increase in absolute rate of photosynthesis, tree species/leaves from sunlit environments display faster photosynthetic induction responses than those from shaded environments. Our results call for re-consideration of contrasting photosynthetic strategies in photosynthetic adaption/acclimation to dynamic light environments across species., (© The Author(s) 2024. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

Plant growth is a complex process affected by a multitude of genetic and environmental factors and their interactions. To identify genetic factors influencing plant performance under different environmental conditions, vegetative growth was assessed in Arabidopsis thaliana cultivated under constant or fluctuating light intensities, using high-throughput phenotyping and genome-wide association studies. Daily automated non-invasive phenotyping of a collection of 382 Arabidopsis accessions provided growth data during developmental progression under different light regimes at high temporal resolution. Quantitative trait loci (QTL) for projected leaf area, relative growth rate, and PSII operating efficiency detected under the two light regimes were predominantly condition-specific and displayed distinct temporal activity patterns, with active phases ranging from 2 d to 9 d. Eighteen protein-coding genes and one miRNA gene were identified as potential candidate genes at 10 QTL regions consistently found under both light regimes. Expression patterns of three candidate genes affecting projected leaf area were analysed in time-series experiments in accessions with contrasting vegetative leaf growth. These observations highlight the importance of considering both environmental and temporal patterns of QTL/allele actions and emphasize the need for detailed time-resolved analyses under diverse well-defined environmental conditions to effectively unravel the complex and stage-specific contributions of genes affecting plant growth processes., (© The Author(s) 2023. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Photorespiration is a dynamic process that is intimately linked to photosynthetic carbon assimilation. There is a growing interest in understanding carbon assimilation during dynamic conditions, but the role of photorespiration under such conditions is unclear. In this review, we discuss recent work relevant to the function of photorespiration under dynamic conditions, with a special focus on light transients. This work reveals that photorespiration is a fundamental component of the light induction of assimilation where variable diffusive processes limit CO2 exchange with the atmosphere. Additionally, metabolic interactions between photorespiration and the C3 cycle may help balance fluxes under dynamic light conditions. We further discuss how the energy demands of photorespiration present special challenges to energy balancing during dynamic conditions. We finish the review with an overview of why regulation of photorespiration may be important under dynamic conditions to maintain appropriate fluxes through metabolic pathways related to photorespiration such as nitrogen and one-carbon metabolism., (© The Author(s) 2022. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Plants growing in nature often experience fluctuating irradiance. However, in the laboratory, the dynamics of photosynthesis are usually explored by instantaneously exposing dark-adapted plants to constant light and examining the dark-to-light transition, which is a poor approximation of natural phenomena. With the aim creating a better approximation, we exposed leaves of pea (Pisum sativum) to oscillating light and measured changes in the functioning of PSI and PSII, and of the proton motive force at the thylakoid membrane. We found that the dynamics depended on the oscillation period, revealing information about the underlying regulatory networks. As demonstrated for a selected oscillation period of 60 s, the regulation tries to keep the reaction centers of PSI and PSII open. We present an evaluation of the data obtained, and discuss the involvement of particular processes in the regulation of photosynthesis. The forced oscillations provided an information-rich fingerprint of complex regulatory networks. We expect future progress in understanding these networks from experiments involving chemical interventions and plant mutants, and by using mathematical modeling and systems identification and control tools., (© The Author(s) 2022. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

Increasing tropospheric ozone (O3) is well-known to decrease leaf photosynthesis under steady-state light through reductions in biochemical capacity. However, the effects of O3 on photosynthetic induction and its biochemical limitations in response to fluctuating light remain unclear, despite the rapid fluctuations of light intensity occurring under field conditions. In this study, two hybrid poplar clones with different O3 sensitivities were exposed to elevated O3. Dynamic photosynthetic CO2 response measurements were conducted to quantify the impact of elevated O3 and exposure duration on biochemical limitations during photosynthetic induction. We found that elevated O3 significantly reduced the steady-state light-saturated photosynthetic rate, the maximum rate of carboxylation (Vcmax) and Rubisco content. In addition, elevated O3 significantly decreased the time constants for slow phases and weighting of the fast phase of the Vcmax induction in poplar clone '546' but not in clone '107'. However, elevated O3 did not affect the time, it took to reach a given percentage of full Vcmax activation or photosynthetic induction in either clone. Overall, photosynthetic induction was primarily limited by the activity of Rubisco rather than the regeneration of ribulose-1,5-biphosphate regardless of O3 concentration and exposure duration. The lack of O3-induced effects on the activation of Rubisco observed here would simplify the simulation of impacts of O3 on nonsteady-state photosynthesis in dynamic photosynthetic models., (© The Author(s) 2022. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

NaCl stress affects stomatal behavior and photosynthesis by a combination of osmotic and ionic components, but it is unknown how these components affect stomatal and photosynthetic dynamics. Tomato (Solanum lycopersicum) plants were grown in a reference nutrient solution [control; electrical conductivity (EC)=2.3 dS m-1], a solution containing additional macronutrients (osmotic effect; EC=12.6 dS m-1), or a solution with additional 100 mM NaCl (osmotic and ionic effects; EC=12.8 dS m-1). Steady-state and dynamic photosynthesis, and leaf biochemistry, were characterized throughout leaf development. The osmotic effect decreased steady-state stomatal conductance while speeding up stomatal responses to light intensity shifts. After 19 d of treatment, photosynthetic induction was reduced by the osmotic effect, which was attributable to lower initial stomatal conductance due to faster stomatal closing under low light. Ionic effects of NaCl were barely observed in dynamic stomatal and photosynthetic behavior, but led to a reduction in leaf photosynthetic capacity, CO2 carboxylation rate, and stomatal conductance in old leaves after 26 d of treatment. With increasing leaf age, rates of light-triggered stomatal movement and photosynthetic induction decreased across treatments. We conclude that NaCl impacts dynamic stomatal and photosynthetic kinetics by osmotic effects and reduces photosynthetic capacity by ionic effects., (© The Author(s) 2022. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Under field environments, fluctuating light conditions induce dynamic photosynthesis, which affects carbon gain by crop plants. Elucidating the natural genetic variations among untapped germplasm resources and their underlying mechanisms can provide an effective strategy to improve dynamic photosynthesis and, ultimately, improve crop yields through molecular breeding approaches. In this review, we first overview two processes affecting dynamic photosynthesis, namely (i) biochemical processes associated with CO2 fixation and photoprotection and (ii) gas diffusion processes from the atmosphere to the chloroplast stroma. Next, we review the intra- and interspecific variations in dynamic photosynthesis in relation to each of these two processes. It is suggested that plant adaptations to different hydrological environments underlie natural genetic variation explained by gas diffusion through stomata. This emphasizes the importance of the coordination of photosynthetic and stomatal dynamics to optimize the balance between carbon gain and water use efficiency under field environments. Finally, we discuss future challenges in improving dynamic photosynthesis by utilizing natural genetic variation. The forward genetic approach supported by high-throughput phenotyping should be introduced to evaluate the effects of genetic and environmental factors and their interactions on the natural variation in dynamic photosynthesis., (© The Author(s) 2022. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

The induction and relaxation of photochemistry and non-photochemical quenching (NPQ) are not instantaneous and require time to respond to fluctuating environments. There is a lack of integrated understanding on how photochemistry and NPQ influence photosynthesis in fluctuating environments. We measured the induction and relaxation of chlorophyll a fluorescence and gas exchange in poplar and cotton at varying temperatures under saturating and fluctuating lights. When the light shifted from dark to high, the fraction of open reaction centers in photosystem II (qL) gradually increased while NPQ increased suddenly and then remained stable. Temperature significantly changed the response of qL but not that of NPQ during the dark to high light transition. Increased qL led to higher photosynthesis but their precise relationship was affected by NPQ and temperature. qL was significantly related to biochemical capacity. Thus, qL appears to be a strong indicator of the activation of carboxylase, leading to the similar dynamics between qL and photosynthesis. When the light shifted from high to low intensity, NPQ is still engaged at a high level, causing a stronger decline in photosynthesis. Our finding suggests that the dynamic effects of photochemistry and NPQ on photosynthesis depend on the phases of environmental fluctuations and interactive effects of light and temperature. Across the full spectra of light fluctuation, the slow induction of qL is a more important limiting factor than the slow relaxation of NPQ for photosynthesis in typical ranges of temperature for photosynthesis. The findings provided a new perspective to improve photosynthetic productivity with molecular biology under natural fluctuating environments., (© The Author(s) 2021. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

It has been reported that stomatal conductance often limits the steady-state photosynthetic rate. On the other hand, the stomatal limitation of photosynthesis in fluctuating light remains largely unknown, although in nature light fluctuates due to changes in sun position, cloud cover, and the overshadowing canopy. In this study, we analysed three mutant lines of Arabidopsis with increased stomatal conductance to examine to what extent stomatal opening limits photosynthesis in fluctuating light. The slac1 (slow anion channel-associated 1) and ost1 (open stomata 1) mutants with stay-open stomata, and the PATROL1 (proton ATPase translocation control 1) overexpression line with faster stomatal opening responses exhibited higher photosynthetic rates and plant growth in fluctuating light than the wild-type, whereas these four lines showed similar photosynthetic rates and plant growth in constant light. The slac1 and ost1 mutants tended to keep their stomata open in fluctuating light, resulting in lower water-use efficiency (WUE) than the wild-type. However, the PATROL1 overexpression line closed stomata when needed and opened stomata immediately upon irradiation, resulting in similar WUE to the wild-type. The present study clearly shows that there is room to optimize stomatal responses, leading to greater photosynthesis and biomass accumulation in fluctuating light in nature., (© The Author(s) 2020. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

It is well known that far-red light (FR; >700 nm) drives PSI photochemistry, but its effect on photosynthetic performance has received little attention. In this study, the effects of the addition of FR to red fluctuating light (FL) have on photosynthesis were examined in the leaves of Arabidopsis thaliana. Light-activated leaves were illuminated with FL [alternating high light/low light (HL/LL) at 800/30 μmol m-2 s-1] for 10-15 min without or with FR at intensities that reflected natural conditions. The CO2 assimilation rates upon the transition from HL to LL were significantly greater with FR than without FR. The enhancement of photosynthesis by FR was small under the steady-state conditions and in the HL phases of FL. Proton conductivity through the thylakoid membrane (gH+) in the LL phases of FL, estimated from the dark relaxation kinetics of the electrochromic absorbance shift, was greater with FR than without FR. The relaxation of non-photochemical quenching (NPQ) in the PSII antenna system and the increase in PSII photochemistry in the LL phases accelerated in the presence of FR. Similar FR-effects in FL were confirmed in typical sun and shade plants. On the basis of these results, we concluded that FR exerted beneficial effects on photosynthesis in FL by exciting PSI and accelerating NPQ relaxation and PSII-yield increase. This was probably because of the increased gH+, which would reflect faster ΔpH dissipation and ATP synthesis., (� The Author(s) 2019. Published by Oxford University Press on behalf of Japanese Society of Plant Physiologists. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Long-term high CO2 exposure accelerates photosynthetic induction response due to rapid light increase. However, it is unclear whether the acceleration is caused by acclimation of photosynthetic components (long-term CO2 effect) and/or by the sufficient substrate under high CO2 at the measurement (short-term CO2 effect). Populus koreana × trichocarpa cv. Peace has wide-open stomata almost not responding to changes of photon flux density. Using this species, we examined the long- and short-term CO2 effects on photosynthetic induction by focusing on biochemical components. We grew the plants under [CO2] of 380, 700 and 1020 μmol CO2 mol-1 air and measured the photosynthetic induction response under [CO2] of 380 and 1020 μmol CO2 mol-1 air. Despite significant reduction in Rubisco content and light-saturated photosynthetic rate in the leaves from the high growth CO2, the photosynthetic induction time was similar in leaves from different growth CO2 plants when measurement [CO2] was the same. The induction, however, was significantly fast at the higher than at the lower measurement [CO2], regardless of growth CO2 of the plants. These results demonstrate that the acceleration of apparent photosynthetic induction under high CO2 environment was mainly contributed by a short-term CO2 effect rather than by a long-term acclimation effect when stomatal limitation is not the major factor., (© The Author(s) 2018. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Leaves are often exposed to fluctuating irradiance, which limits assimilation. Elevated CO2 enhances dynamic photosynthesis (i.e. photosynthesis in fluctuating irradiance) beyond its effects on steady-state photosynthesis rates. Studying the role of CO2 in dynamic photosynthesis is important for understanding plant responses to changing atmospheric CO2 partial pressures. The rise of photosynthesis after a step-wise increase to 1000 μmol m-2 s-1, the loss of photosynthetic induction after irradiance decreases, and rates of photosynthesis during sinusoidal changes in irradiance were studied in tomato (Solanum lycopersicum L.) leaves, using three CO2 partial pressures (200, 400, and 800 µbar). Initial irradiance was set to 0, 50, 100, and 200 μmol m-2 s-1 to vary the initial induction state. Most responses at 200 µbar were not different from those at 400 µbar. In contrast, CO2 at 800 µbar increased the relative carbon gain by 12% after an increase in irradiance, decreased the loss of photosynthetic induction by 14%, and increased dynamic photosynthesis during sine waves by 17%, compared with 400 µbar. These effects were additional to steady-state effects of elevated CO2 on photosynthesis. The enhancement of dynamic photosynthesis rates by elevated CO2 may therefore additionally increase photosynthesis in a future, CO2-enriched climate., (© The Author 2017. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

Plants have mechanisms allowing them to acclimate to intense light conditions, which involves the dissipation of excess light energy. These mechanisms allow plants to perform photosynthesis efficiently and, therefore, must be accurately and precisely controlled. However, how plants dissipate excess light energy has yet to be fully elucidated. Herein we report the identification of a gene, which we named Fluctuating-Light-Acclimation Protein1 (FLAP1), that is conserved in oxygenic phototrophs. We show that Arabidopsis FLAP1 is associated with chloroplast thylakoid and envelope membranes and that the flap1 mutant shows delayed non-photochemical quenching (NPQ) relaxation during induction of photosynthesis at moderate light intensity. Under fluctuating light conditions, NPQ levels in the flap1 mutant were higher than those in the wild type during the high light period, and the mutant exhibited a pale-green phenotype. These findings suggest that FLAP1 is involved in NPQ control, which is important for an acclimation response to fluctuating light., (© The Author 2017. Published by Oxford University Press on behalf of Japanese Society of Plant Physiologists. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

It has been reported that PSI photoinhibition is induced even in wild-type plants of Arabidopsis thaliana, rice and other species by exposure of leaves to fluctuating light (FL) for a few hours. Because plants are exposed to FL in nature, they must possess protective mechanisms against the FL-induced photodamage. Here, using A. thaliana grown at various irradiances, we examined PSI photoprotection by far-red (FR) light at intensities comparable with those observed in nature. Dark-treated leaves were illuminated by red FL alternating high/low light at 1,200/30 µmol m-2 s-1 for 800 ms/10 s. By this FL treatment without FR light for 120 min, the level of photo-oxidizable P700 was decreased by 30% even in the plants grown at high irradiances. The addition of continuous FR light during the FL suppressed this damage almost completely. With FR light, P700 was kept in a more oxidized state in both low- and high-light phases. The protective effect of FR light was diminished more in mutants of the NADH dehydrogenase-like complex (NDH)-mediated cyclic electron flow around PSI (CEF-PSI) than in the PGR5 (proton gradient regulation 5)-mediated CEF-PSI, indicating that the NDH-mediated CEF-PSI would be a major contributor to PSI photoprotection in the presence of FR light. We also confirmed that PSI photoinhibition decreased with the increase in growth irradiance in A. thaliana and field-grown plants, and that this PSI photodamage was largely suppressed by addition of FR light. These results clearly indicate that the most effective PSI protection is realized in the presence of FR light., (© The Author 2017. Published by Oxford University Press on behalf of Japanese Society of Plant Physiologists. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

It has been claimed that the cyclic electron flow around PSI (CEF-PSI) plays an important role in protection of PSI against fluctuating light photoinhibition. However, the photoprotective mechanism of PSI is not fully elucidated. Here, we examined the mechanism, using two CEF-PSI mutants of Arabidopsis thaliana, and antimycin A, an inhibitor of the PGR5 (proton gradient regulation 5)-mediated CEF-PSI. Dark-adapted leaves in these plants were illuminated in fluctuating light alternating between high light at 1,200 µmol m -2 s -1 and low light at 30 µmol m -2 s -1 every 2 min, and PSI and PSII parameters were simultaneously measured for 160 min with 830 nm absorption and Chl fluorescence, respectively. When CEF-PSI, especially PGR5-mediated CEF-PSI, did not operate, the acceptor-side limitation of PSI, Y(NA), increased stepwise, leading to marked PSI photoinhibition. The deficiency of CFE-PSI decreased not only the electron transport rate through PSI but also the donor-side limitation of PSI, Y(ND), in high light phases. These results showed that the large Y(ND), observed only when CEF-PSI operated, contributed to suppression of PSI photoinhibition. Taken together with our previous report that high Y(NA) was alleviated by the enhancement of CEF-PSI, a model for the protective mechanisms of PSI is proposed. In this model, both alleviation of Y(NA) and acceleration of Y(ND) are indispensable, and for realization of such a situation, regulation of the electron flows, especially the PGR5-mediated CEF-PSI, plays a key role. It is important for effective protection to regulate the balance of Y(ND) and Y(NA) through CEF-PSI., (© The Author 2016. Published by Oxford University Press on behalf of Japanese Society of Plant Physiologists. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

The flavodiiron proteins (FDPs) Flv1 and Flv3 in cyanobacteria function in photoreduction of O 2 to H 2 O, without concomitant formation of reactive oxygen species, known as the Mehler-like reaction. Both Flv1 and Flv3 are essential for growth under fluctuating light (FL) intensities, providing protection for PSI. Here we compared the global transcript profiles of the wild type (WT), Δflv1 and Δflv1/Δflv3 grown under constant light (GL) and FL. In the WT, FL induced the largest down-regulation in transcripts involved in carbon-concentrating mechanisms (CCMs), while those of the nitrogen assimilation pathways increased as compared with GL. Already under GL the Δflv1/Δflv3 double mutant demonstrated a partial down-regulation of transcripts for CCM and nitrogen metabolism, while in FL conditions the transcripts for nitrogen assimilation were strongly down-regulated. Many alterations were specific only for Δflv1/Δflv3, and not detected in Δflv1, suggesting that certain transcripts are affected primarily because of the lack of flv3 By constructing the strains overproducing solely either Flv1 or Flv3, we demonstrate that the homo-oligomers of these proteins also function in acclimation of cells to FL, by catalyzing reactions with as yet unidentified components, while the presence of both Flv1 and Flv3 is a prerequisite for the Mehler-like reaction and thus the electron transfer to O 2 Considering the low expression of flv1, it is unlikely that the Flv1 homo-oligomer is present in the WT., (© The Author 2016. Published by Oxford University Press on behalf of Japanese Society of Plant Physiologists.)

To assess the roles of the cyclic electron flow around PSI (CEF-PSI) and O₂-dependent alternative pathways including the water-water cycle in fluctuating light, we grew the wild type and pgr5 mutant of Arabidopsis thaliana in constant light, and measured Chl fluorescence and P700 parameters in their leaves in the fluctuating light alternating between 240 (HL) and 30 µmol photons m⁻² s⁻² (LL) every 2 min. At 20% O₂, the photochemical quantum yield of PSII decreased, in particular in the pgr5 plants, soon after the start of the fluctuating light treatment. PSI of the pgr5 plants was markedly photoinhibited by this treatment for 42 min. Slight PSI photoinhibition was also observed in the wild type. We measured energy sharing between PSII and PSI and estimated the PSI and PSII electron transport rates (ETRs). pgr5 showed larger energy allocation to PSI. In contrast to the wild type, the ratio of the PSI to the PSII ETR in pgr5 was higher in LL but lower in HL at 20% O₂ due to PSI acceptor-side limitation. At 2.7% or 0% O₂, the CEF-PSI of the pgr5 plants was enhanced, the acceptor-side limitation of PSI electron flow was released and PSI photoinhibition was not observed. The results suggest that the light fluctuation is a potent stress to PSI and that the CEF-PSI is essential to protect PSI from this stress.