Mesophyll conductance limits photosynthesis in fluctuating light under combined drought and heat stresses.

Drought and heat stresses usually occur concomitantly in nature, with increasing frequency and intensity of both stresses expected due to climate change. The synergistic agricultural impacts of these compound climate extremes are much greater than those of the individual stresses. However, the mechanisms by which drought and heat stresses separately and concomitantly affect dynamic photosynthesis have not been thoroughly assessed. To elucidate this, we used tomato (Solanum lycopersicum) seedlings to measure dynamic photosynthesis under individual and compound stresses of drought and heat. Individual drought and heat stresses limited dynamic photosynthesis at the stages of diffusional conductance to CO2 and biochemistry, respectively. However, the primary limiting factor for photosynthesis shifted to mesophyll conductance under the compound stresses. Compared with the control, photosynthetic carbon gain in fluctuating light decreased by 38%, 73%, and 114% under the individual drought, heat, and compound stresses, respectively. Therefore, compound stresses caused a greater reduction in photosynthetic carbon gain in fluctuating light conditions than individual stress. These findings highlight the importance of mitigating the effects of compound climate extremes on crop productivity by targeting mesophyll conductance and improving dynamic photosynthesis.

Within-branch photosynthetic gradients are more related to the coordinated investments of nitrogen and water than leaf mass per area.

Nitrogen (N) and water are key resources for leaf photosynthesis and the growth of whole plants. Within-branch leaves need different amounts of N and water to support their differing photosynthetic capacities according to light exposure. To test this scheme, we measured the within-branch investments of N and water and their effects on photosynthetic traits in two deciduous tree species Paulownia tomentosa and Broussonetia papyrifera. We found that leaf photosynthetic capacity gradually increased from branch bottom to top (i.e. from shade to sun leaves). Concomitantly, stomatal conductance (gs) and leaf N content gradually increased, owing to the symport of water and inorganic mineral from root to leaf. Variation of leaf N content led to large gradients of mesophyll conductance, maximum velocity of Rubisco for carboxylation, maximum electron transport rate and leaf mass per area (LMA). Correlation analysis indicated that the within-branch difference in photosynthetic capacity was mainly related to gs and leaf N content, with a relatively minor contribution of LMA. Furthermore, the simultaneous increases of gs and leaf N content enhanced photosynthetic N use efficiency (PNUE) but hardly affected water use efficiency. Therefore, within-branch adjustment of N and water investments is an important strategy used by plants to optimize the overall photosynthetic carbon gain and PNUE.

Variation in Photosynthetic Efficiency under Fluctuating Light between Rose Cultivars and its Potential for Improving Dynamic Photosynthesis.

Photosynthetic efficiency under both steady-state and fluctuating light can significantly affect plant growth under naturally fluctuating light conditions. However, the difference in photosynthetic performance between different rose genotypes is little known. This study compared the photosynthetic performance under steady-state and fluctuating light in two modern rose cultivars (Rose hybrida), "Orange Reeva" and "Gelato", and an old Chinese rose plant Rosa chinensis cultivar, "Slater's crimson China". The light and CO2 response curves indicated that they showed similar photosynthetic capacity under steady state. The light-saturated steady-state photosynthesis in these three rose genotypes was mainly limited by biochemistry (60%) rather than diffusional conductance. Under fluctuating light conditions (alternated between 100 and 1500 µmol photons m-2 m-1 every 5 min), stomatal conductance gradually decreased in these three rose genotypes, while mesophyll conductance (gm) was maintained stable in Orange Reeva and Gelato but decreased by 23% in R. chinensis, resulting in a stronger loss of CO2 assimilation under high-light phases in R. chinensis (25%) than in Orange Reeva and Gelato (13%). As a result, the variation in photosynthetic efficiency under fluctuating light among rose cultivars was tightly related to gm. These results highlight the importance of gm in dynamic photosynthesis and provide new traits for improving photosynthetic efficiency in rose cultivars.

Photoinhibition of Photosystem I Induced by Different Intensities of Fluctuating Light Is Determined by the Kinetics of ∆pH Formation Rather Than Linear Electron Flow.

Fluctuating light (FL) can cause the selective photoinhibition of photosystem I (PSI) in angiosperms. In nature, leaves usually experience FL conditions with the same low light and different high light intensities, but the effects of different FL conditions on PSI redox state and PSI photoinhibition are not well known. In this study, we found that PSI was highly reduced within the first 10 s after transition from 59 to 1809 µmol photons m-2 s-1 in tomato (Solanum lycopersicum). However, such transient PSI over-reduction was not observed by transitioning from 59 to 501 or 923 µmol photons m-2 s-1. Consequently, FL (59-1809) induced a significantly stronger PSI photoinhibition than FL (59-501) and FL (59-923). Compared with the proton gradient (∆pH) level after transition to high light for 60 s, tomato leaves almost formed a sufficient ∆pH after light transition for 10 s in FL (59-501) but did not in FL (59-923) or FL (59-1809). The difference in ∆pH between 10 s and 60 s was tightly correlated to the extent of PSI over-reduction and PSI photoinhibition induced by FL. Furthermore, the difference in PSI photoinhibition between (59-923) and FL (59-1809) was accompanied by the same level of linear electron flow. Therefore, PSI photoinhibition induced by different intensities of FL is more related to the kinetics of ∆pH formation rather than linear electron flow.

Exogenous melatonin strongly affects dynamic photosynthesis and enhances water-water cycle in tobacco.

Melatonin (MT), an important phytohormone synthesized naturally, was recently used to improve plant resistance against abiotic and biotic stresses. However, the effects of exogenous melatonin on photosynthetic performances have not yet been well clarified. We found that spraying of exogenous melatonin (100 µM) to leaves slightly affected the steady state values of CO2 assimilation rate (A N ), stomatal conductance (g s ) and mesophyll conductance (g m ) under high light in tobacco leaves. However, this exogenous melatonin strongly delayed the induction kinetics of g s and g m , leading to the slower induction speed of A N . During photosynthetic induction, A N is mainly limited by biochemistry in the absence of exogenous melatonin, but by CO2 diffusion conductance in the presence of exogenous melatonin. Therefore, exogenous melatonin can aggravate photosynthetic carbon loss during photosynthetic induction and should be used with care for crop plants grown under natural fluctuating light. Within the first 10 min after transition from low to high light, photosynthetic electron transport rates (ETR) for A N and photorespiration were suppressed in the presence of exogenous melatonin. Meanwhile, an important alternative electron sink, namely water-water cycle, was enhanced to dissipate excess light energy. These results indicate that exogenous melatonin upregulates water-water cycle to facilitate photoprotection. Taking together, this study is the first to demonstrate that exogenous melatonin inhibits dynamic photosynthesis and improves photoprotection in higher plants.

Photosynthesis under fluctuating light in the CAM plant Vanilla planifolia.

Photosynthetic induction after a sudden increase in illumination affects carbon gain. Photosynthetic dynamics under fluctuating light (FL) have been widely investigated in C3 and C4 plants but are little known in CAM plants. In our present study, the chlorophyll fluorescence, P700 redox state and electrochromic shift signals were measured to examine photosynthetic characteristics under FL in the CAM orchid Vanilla planifolia. The light use efficiency was maximized in the morning but was restricted in the afternoon, indicating that the pool of malic acid dried down in the afternoon. During photosynthetic induction in the morning, electron flow through photosystem I rapidly reached the 95% of the maximum value in 4-6 min, indicating that V. planifolia showed a fast photosynthetic induction when compared with C3 and C4 plants reported previously. Upon a sudden transition from dark to actinic light, a rapid re-oxidation of P700 was observed in V. planifolia, indicating the fast outflow of electrons from PSI to alternative electron acceptors, which was attributed to the O2 photo-reduction mediated by water-water cycle. The functioning of water-water cycle prevented photosystem I over-reduction after transitioning from low to high light and thus protected PSI under FL. In the afternoon, cyclic electron flow was stimulated under FL to fine-tune photosynthetic apparatus when photosynthetic CO2 was restricted. Therefore, water-water cycle cooperates with cyclic electron flow to regulate the photosynthesis under FL in the CAM orchid V. planifolia.

Regulation of Leaf Angle Protects Photosystem I under Fluctuating Light in Tobacco Young Leaves.

Fluctuating light is a typical light condition in nature and can cause selective photodamage to photosystem I (PSI). The sensitivity of PSI to fluctuating light is influenced by the amplitude of low/high light intensity. Tobacco mature leaves are tended to be horizontal to maximize the light absorption and photosynthesis, but young leaves are usually vertical to diminish the light absorption. Therefore, we tested the hypothesis that such regulation of the leaf angle in young leaves might protect PSI against photoinhibition under fluctuating light. We found that, upon a sudden increase in illumination, PSI was over-reduced in extreme young leaves but was oxidized in mature leaves. After fluctuating light treatment, such PSI over-reduction aggravated PSI photoinhibition in young leaves. Furthermore, the leaf angle was tightly correlated to the extent of PSI photoinhibition induced by fluctuating light. Therefore, vertical young leaves are more susceptible to PSI photoinhibition than horizontal mature leaves when exposed to the same fluctuating light. In young leaves, the vertical leaf angle decreased the light absorption and thus lowered the amplitude of low/high light intensity. Therefore, the regulation of the leaf angle was found for the first time as an important strategy used by young leaves to protect PSI against photoinhibition under fluctuating light. To our knowledge, we show here new insight into the photoprotection for PSI under fluctuating light in nature.

An imprinted PEG1/MEST antisense expressed predominantly in human testis and in mature spermatozoa.

PEG1 (or MEST) is an imprinted gene located on human chromosome 7q32 that is expressed predominantly from the paternal allele. In the mouse, Peg1/Mest is associated with embryonic growth and maternal behavior. Human PEG1 is transcribed from two promoters; the transcript from promoter P1 is derived from both parental alleles, and the transcript from P2 is exclusively from the paternal allele. We characterized the P1 and P2 transcripts in various normal and neoplastic tissues. In the normal tissues, PEG1 was transcribed from both promoters P1 and P2, whereas in six of eight neoplastic tissues, PEG1 was transcribed exclusively from promoter P1. Bisulfite sequencing demonstrated high levels of CpG methylation in the P2 region of DNA from a lung tumor. In the region between P1 and P2, we identified a novel transcript, PEG1-AS, in an antisense orientation to PEG1. PEG1-AS is a spliced transcript and was detected as a strong 2.4-kilobase band on a Northern blot. PEG1-AS and PEG1 P2-sense transcript were expressed exclusively from the paternal allele. Fragments of DNA from within the 1.5-kilobase region between PEG1-AS and the P2 exon were ligated to a pGL3 luciferase reporter vector and transfected into NCI H23 cells. This DNA exhibited strong promoter activity in both the sense and antisense directions, indicating that PEG1-AS and P2 exon share a common promoter region. Treatment of the transfected DNA fragments with CpG methylase abolished the promoter activity. Of interest, PEG1-AS was expressed predominantly in testis and in mature motile spermatozoa, indicating a possible role for this transcript in human sperm physiology and fertilization.