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Non-photochemical quenching (NPQ) has been regarded as a safety valve to dissipate excess absorbed light energy not used for photochemistry. However, there exists no general consensus on the photoprotective role of NPQ. In the present study, we quantified the Photosystem II (PSII) photo-susceptibilities (mpi) in the presence of lincomycin, under red light given to five shade-acclimated tree species grown in the field. Photosynthetic energy partitioning theory was applied to investigate the relationships between mpi and each of the regulatory light-induced NPQ [Y(NPQ)], the quantum yield of the constitutive nonregulatory NPQ [Y(NO)] and the PSII photochemical yield in the light-adapted state [Y(PSII)] under different red irradiances. It was found that in the low to moderate irradiance range (50–800 μmol m–2 s–1) when the fraction of open reaction centers (qP) exceeded 0.4, mpi exhibited no association with Y(NPQ), Y(NO) and Y(PSII) across species. However, when qP

The quantum yield of photosynthesis (QY, CO

Alternative electron fluxes such as the cyclic electron flux (CEF) around photosystem I (PSI) and Mehler reaction (Me) are essential for efficient photosynthesis because they generate additional ATP and protect both photosystems against photoinhibition. The capacity for Me can be estimated by measuring O2 exchange rate under varying irradiance and CO2 concentration. In this study, mass spectrometric measurements of O2 exchange were made using leaves of representative species of C3 and C4 grasses grown under natural light (control; PAR ~ 800 µmol quanta m−2 s−1) and shade (~ 300 µmol quanta m−2 s−1), and in representative species of gymnosperm, liverwort and fern grown under natural light. For all control grown plants measured at high CO2, O2 uptake rates were similar between the light and dark, and the ratio of Rubisco oxygenation to carboxylation (Vo/Vc) was low, which suggests little potential for Me, and that O2 uptake was mainly due to photorespiration or mitochondrial respiration under these conditions. Low CO2 stimulated O2 uptake in the light, Vo/Vc and Me in all species. The C3 species had similar Vo/Vc, but Me was highest in the grass and lowest in the fern. Among the C4 grasses, shade increased O2 uptake in the light, Vo/Vc and the assimilation quotient (AQ), particularly at low CO2, whilst Me was only substantial at low CO2 where it may contribute 20–50% of maximum electron flow under high light.

Light energy causes damage to Photosystem I (PSI) and Photosystem II (PSII). The majority of the previous photoinhibition studies have been conducted with PSII, which shows much larger photoinhibition than PSI; therefore, relatively little is known about the mechanism of PSI photoinhibition so far. A previous report showed that the photoinhibition action spectrum measured with PSI activity of isolated thylakoid is similar to the absorption spectrum of chlorophyll. However, it is known that the extent of PSI photoinhibition is much smaller in vivo compared to in vitro. It is also possible that the different extent of PSII photoinhibition, caused by different spectral light qualities, can affect the photoinhibition of PSI in vivo because PSI receives electrons from PSII. In the present research, to study the effect of light quality and the effect of the extent of PSII photoinhibition on the PSI photoinhibition in vivo, intact leaves were photoinhibited under four different light qualities. The rate coefficient of PSI photoinhibition was significantly higher in blue and red light compared to white light. The rate of PSI photoinhibition at the same photon-exposure was the largest in blue and red light and followed by white and green light. These results support the notion that light absorption by chlorophyll is responsible for the PSI photoinhibition, even in intact leaves. The variation among light colors in the relationships between the extent of photoinhibition of PSII and that of PSI indicate that PSI and PSII are independently photoinhibited with different mechanisms in the early stage of in vivo photoinhibition.

Robert John Porra (7.8.1931–16.5.2019) is probably best known for his substantial practical contributions to plant physiology and photosynthesis by addressing the problems of both the accurate spectroscopic estimation and the extractability of chlorophylls in many organisms. Physiological data and global productivity estimates, in particular of marine primary productivity, are often quoted on a chlorophyll basis. He also made his impact by work on all stages of tetrapyrrole biosynthesis: he proved the C5 pathway to chlorophylls, detected an alternative route to protoporphyrin in anaerobes and the different origin of the oxygen atoms in anaerobes and aerobes. A brief review of his work is supplemented by personal memories of the authors.

It is hypothesized that plant submergence tolerance could be assessed from the decline of plant biomass due to submergence, as biomass integrates all eco-physiological processes leading to fitness. An alternative hypothesis stated that the consumption rate of carbohydrate is essential in differing tolerance to submergence. In the present study, the responses of biomass, biomass allocation, and carbohydrate content to simulated long-term winter submergence were assessed in four tolerant and four sensitive perennials. The four tolerant perennials occur in a newly established riparian ecosystem created by The Three Gorges Dam, China. They had 100% survival after 120 days’ simulated submergence, and had full photosynthesis recovery after 30 days’ re-aeration, and the photosynthetic rate was positively related to the growth during the recovery period. Tolerant perennials were characterized by higher carbohydrate levels, compared with the four sensitive perennials (0% survival) at the end of submergence. Additionally, by using a method which simulates posterior estimates, and bootstraps the confidence interval for the difference between strata means, it was found that the biomass response to post-hypoxia, rather than that to submergence, could be a reliable indicator to assess submergence tolerance. Interestingly, the differences of changes in carbohydrate content between tolerant and sensitive perennials during submergence were significant, which were distinct from the biomass response, supporting the hypothesis that tolerant perennials could sacrifice non-vital components of biomass to prioritize the saving of carbohydrates for later recovery. Our study provides some insight into the underlying mechanism(s) of perennials’ tolerance to submergence in ecosystems such as temperate wetland and reservoir riparian.

The hybrid rice (Oryza sativa L.) cultivar Peiai64S/E32, which is a hybrid of E32 and Peiai64S, has a super high yield compared with that of its parents. Changes in photosynthetic pigments and chlorophyll (Chl) fluorescence parameters were investigated in the flag leaves during senescence. The content of Chl and carotenoids in the flag leaves of Peiai64S/E32 was more stable than those of their parents during senescence. Furthermore, Peiai64S/E32 had a more stable PSII activity and a lower nonphotochemical quenching and deepoxidation state than that of its parents in the later stage of senescence. These results demonstrated that Peiai64S/E32 functioned longer than its parents did during the leaf senescence and grain-filling period. This occurred because there was more excitation energy to drive photosynthetic CO2 fixation in Peiai64S/E32, as the pigments supplying the energy survived longer in the high-yielding rice compared to the parents.

Anthocyanins are water-soluble pigments in plants known for their photoprotective role against photoinhibitory and photooxidative damage under high light (HL). However, it remains unclear whether light-shielding or antioxidant activity plays a major role in the photoprotection exerted by anthocyanins under HL stress. To shed light on this question, we analyzed the physiological and biochemical responses to HL of three Arabidopsis thaliana lines (Col, chi, ans) with different light absorption and antioxidant characteristics. Under HL, ans had the highest antioxidant capacity, followed by Col, and finally chi; Col had the strongest light attenuation capacity, followed by chi, and finally ans. The line ans had weaker physiological activity of chloroplasts and more severe oxidative damage than chi after HL treatment. Col with highest photoprotection of light absorption capacity had highest resistance to HL among the three lines. The line ans with high antioxidant capacity could not compensate for its disadvantages in HL caused by the absence of the light-shielding function of anthocyanins. In addition, the expression level of the Anthocyanin Synthase (ANS) gene was most upregulated after HL treatment, suggesting that the conversion of colorless into colored anthocyanin precursors was necessary under HL. The contribution of anthocyanins to flavonoids, phenols, and antioxidant capacity increased in the late period of HL, suggesting that plants prefer to synthesize red anthocyanins (a group of colored antioxidants) over other colorless antioxidants to cope with HL. These experimental observations indicate that the light attenuation role of anthocyanins is more important than their antioxidant role in photoprotection.

Biological invasion is a hot topic in ecological research. Most studies on the physiological mechanisms of plants focus on leaves, but few studies focus on stems. To study the tolerance of invasive plant (Sphagneticola trilobata L.) to low temperature, relevant physiological indicators (including anthocyanin and chlorophyll) in different organs (leaves and stems) were analyzed, using a native species (Sphagneticola calendulacea L.) as the control. The results showed that, upon exposure to low temperature for 15 days, the stems of two Sphagneticola species were markedly reddened, their anthocyanin content increased, chlorophyll and chlorophyll fluorescence parameters decreased, and the accumulation of reactive oxygen species in the stem increased. The percentage increases of antioxidants and total antioxidant capacities in stems were significantly higher in S. trilobata than in S. calendulacea. This showed that S. trilobata had higher cold tolerance in stems while leaves were opposite. To further verify the higher cold tolerance of the stem of S. trilobata, a defoliation experiment was designed. We found that the defoliated stem of S. trilobata reduced anthocyanin accumulation and increased chlorophyll content, while alleviating membrane lipid damage and electrical conductivity, and the defoliated stem still showed an increase in stem diameter and biomass under low temperature. The discovery of the physiological and adaptive mechanisms of the stem of S. trilobata to low temperature will provide a theoretical basis for explaining how S. trilobata maintains its annual growth in South China. This is of great significance for predicting the future spread of cloned and propagated invasive plants.

Cyclic electron flow (CEF) around photosystem I (PSI) is essential for generating additional ATP and enhancing efficient photosynthesis. Accurate estimation of CEF requires knowledge of the fractions of absorbed light by PSI (fI) and PSII (fII), which are only known for a few model species such as spinach. No measures of fI are available for C4 grasses under different irradiances. We developed a new method to estimate (1) fII in vivo by concurrently measuring linear electron flux through both photosystems $$\left( {{\text{LEF}}_{{{\text{O}}_{ 2} }} } \right)$$LEFO2 in leaf using membrane inlet mass spectrometry (MIMS) and total electron flux through PSII (ETR2) using chlorophyll fluorescence by a Dual-PAM at low light and (2) CEF as ETR1—$${\text{LEF}}_{{{\text{O}}_{ 2} }}$$LEFO2. For a C3 grass, fI was 0.5 and 0.4 under control (high light) and shade conditions, respectively. C4 species belonging to NADP-ME and NAD-ME subtypes had fI of 0.6 and PCK subtype had 0.5 under control. All shade-grown C4 species had fI of 0.6 except for NADP-ME grass which had 0.7. It was also observed that fI ranged between 0.3 and 0.5 for gymnosperm, liverwort and fern species. CEF increased with irradiance and was induced at lower irradiances in C4 grasses and fern relative to other species. CEF was greater in shade-grown plants relative to control plants except for C4 NADP-ME species. Our study reveals a range of CEF and fI values in different plant functional groups. This variation must be taken into account for improved photosynthetic calculations and modelling.

Photosynthetic induction, a gradual increase in photosynthetic rate on a transition from darkness or low light to high light, has ecological significance, impact on biomass accumulation in fluctuating light and relevance to photoprotection in strong light. However, the experimental quantification of the component electron fluxes in and around both photosystems during induction has been rare. Combining optimized chlorophyll fluorescence, the redox kinetics of P700 [primary electron donor in Photosystem I (PSI)] and membrane inlet mass spectrometry in the absence/presence of inhibitors/mediator, we partially estimated the components of electron fluxes in spinach leaf disks on transition from darkness to 1,000 �mol photons�m−2�s−1 for up to 10 min, obtaining the following findings: (i) the partitioning of energy between both photosystems did not change noticeably; (ii) in Photosystem II (PSII), the combined cyclic electron flow (CEF2) and charge recombination (CR2) to the ground state decreased gradually toward 0 in steady state; (iii) oxygen reduction by electrons from PSII, partly bypassing PSI, was small but measurable; (iv) cyclic electron flow around PSI (CEF1) peaked before becoming somewhat steady; (v) peak magnitudes of some of the electron fluxes, all probably photoprotective, were in the descending order: CEF1 > CEF2 + CR2 > chloroplast O2 uptake; and (vi) the chloroplast NADH dehydrogenase-like complex appeared to aid the antimycin A-sensitive CEF1. The results are important for fine-tuning in silico simulation of in vivo photosynthetic electron transport processes; such simulation is, in turn, necessary to probe partial processes in a complex network of interactions in response to environmental changes.

To finish this special issue, some friends, colleagues and students of Prof. Chow (Emeritus Professor, the Research School of Biology, the Australian National University) have written small tributes to acknowledge not only his eminent career but to describe his wonderful personality.

Robert John Porra (7.8.1931-16.5.2019) is probably best known for his substantial practical contributions to plant physiology and photosynthesis by addressing the problems of both the accurate spectroscopic estimation and the extractability of chlorophylls in many organisms. Physiological data and global productivity estimates, in particular of marine primary productivity, are often quoted on a chlorophyll basis. He also made his impact by work on all stages of tetrapyrrole biosynthesis: he proved the C

Induction of non-photochemical quenching (NPQ) of chlorophyll fluorescence in leaves affords photoprotection to the photosynthetic apparatus when, for whatever reason, photon capture in the antennae of photosystems exceeds their capacity to utilise this excitation in photochemistry and ultimately in CO2 assimilation. Here we augment traditional monitoring of NPQ using the fast time resolution, remote and relatively non-intrusive light induced fluorescence transient (LIFT) technique (Kolber et al . 2005 ; Osmond et al . 2017 ) that allows direct measurement of functional (σ'PSII ) and optical cross-sections (a 'PSII ) of PSII in situ , and calculates the half saturation light intensity for ETR (E k ). These parameters are obtained from the saturation and relaxation phases of fluorescence transients elicited by a sequence of 270, high intensity 1 μs flashlets at controlled time intervals over a period of 30 ms in the QA flash at intervals of a few seconds. We report that although σ'PSII undergoes large transient increases after transfer from dark to strong white light (WL) it declines little in steady-state as NPQ is induced in shade- and sun-grown spinach and Arabidopsis genotypes Col , OEpsbs , pgr 5bkg , stn 7 and stn 7/8. In contrast, σ'PSII increases by ~30% when induction of NPQ in spinach is inhibited by dithiothreitol and by inhibition of NPQ in Arabidopsis npq 1, npq 4 and pgr 5. We propose this increase in σ'PSII arises as some excitation from closed PSII reaction centres is transferred to open centres when excitation partitioning to photochemistry (Y II ) and NPQ (Y NP ) declines, and is indicated by an increased excitation dissipation from closed PSII centres (Y NO , including fluorescence emission). Although E k increases following dissipation of excitation as heat when NPQ is engaged, it declines when NPQ is inhibited. Evidently photochemistry becomes more easily light saturated when excitation is transferred from closed RCIIs to open centres with larger σ'PSII . The NPQ mutant pgr 5 is an exception; E k increases markedly in strong light as electron transport QA → PQ and PQ → PSI accelerate and the PQ pool becomes strongly reduced. These novel in situ observations are discussed in the context of contemporary evidence for functional and structural changes in the photosynthetic apparatus during induction of NPQ.

Despite my humble beginnings in rural China, I had the good fortune of advancing my career and joining an international community of photosynthesis researchers to work on the ‘light reactions’ that are a fundamental process in Nature. Along with supervisors, mentors, colleagues, students and lab assistants, I worked on ionic redistributions across the photosynthetic membrane in response to illumination, photophosphorylation, forces that regulate the stacking of photosynthetic membranes, the composition of components of the photosynthetic apparatus during acclimation to the light environment, and the failure of the photosynthetic machinery to acclimate to too much light or even to cope with moderate light due to inevitable photodamage. These fascinating underlying mechanisms were investigated in vitro and in vivo. My career path, with its ups and downs, was never secure, but the reward of knowing a little more of the secret of Nature offset the job uncertainty.

Alternative electron fluxes such as the cyclic electron flux (CEF) around photosystem I (PSI) and Mehler reaction (Me) are essential for efficient photosynthesis because they generate additional ATP and protect both photosystems against photoinhibition. The capacity for Me can be estimated by measuring O

Increasing amounts of experimental evidence show that anthocyanins provide physiological protection to plants under stress. However, the difference in photoprotection mediated by anthocyanins and other photoprotective substances in different seasons is still uncertain. To determine the relationship between anthocyanin accumulation and the photoprotective effects in different seasons, Castanopsis chinensis and Acmena acuminatissima, whose anthocyanin accumulation patterns differ in different seasons, were used as materials to explain how plants adapt to different seasons; as such, their physiological and biochemical responses were analyzed. Young leaves of C. chinensis and A. acuminatissima presented different colors in the different seasons. In summer, the young leaves of C. chinensis were purplish red, while those of A. acuminatissima were light green. In winter, the young leaves of C. chinensis were light green, while those of A. acuminatissima were red. Compared with the young red leaves, the young light green leaves that did not accumulate anthocyanins had higher flavonoid and phenolics contents, total antioxidant capacity, non-photochemical quenching (NPQ), and relative membrane leakage, and a slower recovery rate in the maximum photochemical efficiency (Fv/Fm) after high-light treatment. In addition, the net photosynthesis rate (Pn), transpiration rate (Tr), stomatal conductance (gs), and the effective quantum yield of PSII (ΦPSII) of the young leaves in winter were significantly lower than those in summer, while the activities of catalase (CAT, EC 1.11.1.6), peroxidase (POD, EC 1.11.1.7), and superoxide dismutase (SOD, EC 1.15.1.1) were significantly higher than those in summer. These data indicate that to adapt to seasonal changes anthocyanins, other antioxidative substances and antioxidative enzymes, as well as components involved in the safe dissipation of excitation energy as heat need to cooperate with one another.

The success of growing vegetables indoors requires the most appropriate selection of lighting spectrum. This mini review discusses the impacts of LED spectral quality on different leafy vegetables with a focus on the studies of Chinese broccoli (Brassica alboglabra), ice plants (Mesembryanthem crystallinum) and lettuce (Lactuca sativa L. cv. Canasta). For each species, plants exposed to different spectral LED lights were all under the same light intensity and same photoperiod. Chinese broccoli grown under red(R):blue(B)-LED ratio of 84:16 (16B) had the highest light-saturated photosynthetic CO2 assimilation rate (Asat) and stomatal conductance (gs sat) compared to plants grown under other R:B-LED ratios. It was also shown that 16B is the most appropriate selection for Chinese broccoli to achieve the highest shoot productivity with a rapid leaf number and leaf area development. The highest concentrations of photosynthetic pigments, soluble and Rubisco protein on a leaf area basis were also observed in 16B plants. The results conclusively affirmed that the highest productivity of Chinese broccoli grown under 16B is closely linked to the highest photosynthetic performance on a leaf area basis. For ice plants grown under R:B-LED ratios of 90:10 (10B), they had the highest shoot biomass with a faster leaf development compared to plants grown under other RB-LED combinations. However, there were no differences in Asat, gs sat, photosynthetic pigments, soluble and Rubisco proteins on a leaf area basis. In the case of lettuce plants, it was a surprise to observe that plants grown under 0B and 20G (20% green (G)-LED and 80% R-LED) had the highest shoot biomass, and largest total leaf area and light interception area but the lowest net maximal photosynthetic rate on a leaf area basis, compared to other plants. The combined RB-LED enhanced other photosynthetic parameters while 0B and 20G conditions had inhibitory effects on maximum quantum efficiency of PS II with lower photosynthetic pigments, total soluble protein and Rubisco protein. These results suggest that impacts of LED light quality on productivity of lettuce (L. sativa L. cv. Canasta) is closely linked to leaf traits not associated with photosynthetic performance on a leaf area basis. Keywords: leafy vegetable, leaf traits, LED spectral quality, photosynthetic performance, productivity DOI: 10.25165/j.ijabe.20191206.5178 Citation: He J, Qin L, Chow W S. Impacts of LED spectral quality on leafy vegetables: Productivity closely linked to photosynthetic performance or associated with leaf traits? Int J Agric & Biol Eng, 2019; 12(6): 16–25.

Plants in their natural environment are often exposed to fluctuating light because of self-shading and cloud movements. As changing frequency is a key characteristic of fluctuating light, we speculated that rapid light fluctuation may induce rapid photosynthetic responses, which may protect leaves against photoinhibition. To test this hypothesis, maize seedlings were grown under fluctuating light with various frequencies (1, 10, and 100 cycles of fluctuations/10 h), and changes in growth, chlorophyll content, gas exchange, chlorophyll a fluorescence, and P700 were analyzed carefully. Our data show that though the growth and light-saturated photosynthetic rate were depressed by rapidly fluctuating light, photosynthesis induction was clearly speeded up. Furthermore, more rapid fluctuation of light strikingly reduced the chlorophyll content, while thermal dissipation was triggered and enhanced. The chlorophyll a fluorescence induction kinetics and P700 absorption results showed that the activities of both photosystem II and photosystem I decreased as the frequency of the fluctuating light increased. In all treatments, the light intensities of the fluctuating light were kept constant. Therefore, rapid light fluctuation frequency itself induced the acceleration of photosynthetic induction and the enhancement of photoprotection in maize seedlings, which play important roles in protecting photosynthetic apparatus against fluctuating high light to a certain extent.

Previous studies have suggested that high light (HL) stress causes photoinhibition in plants, while anthocyanins could protect the photosynthetic apparatus against photoinhibition. However, the photoprotection mechanism of anthocyanins is still ambiguous. We studied physiological responses and molecular changes for CHS-overexpression lines (CHS1, CHS2, CHS3), Arabidopsis thaliana ecotype Columbia (Col), and T-DNA insertion lines of CHS (tt4) under HL (200 μmol m−2 s−1) to explore the photoprotection mechanism of anthocyanins. The results showed that HL induced anthocyanin synthesis and accumulation. The leaves of CHS-overexpression lines turned reddest and the genes, including CHS, DFR, ANS, were expressed at highest levels. Thus, the CHS-overexpression lines maintained the highest photosynthetic capacity and suffered the least damage from HL of the three phenotypes. However, the CHS enzyme and anthocyanins were undetectable in tt4 during the experiment. Correspondingly, chlorophyll fluorescence parameters of tt4 declined greatly. The photosynthetic apparatus and cell membranes were also impaired dramatically. The physiological characteristics of Col were compared between CHS-overexpression lines and tt4. Together, the results suggest that over-expression of CHS gene enhances HL resistance by synthesizing more anthocyanins, that anthocyanins enhance the adaptability of plants to HL and that they maintain photosynthetic capacity via both antioxidation and attenuation of light.

The successful development of photosynthetic organs is the basis of plant growth and community development. To reveal photo-acclimation to high irradiance in tree species during the course of leaf development and plant succession of subtropical forests, photosynthetic efficiency and photoprotective compounds were analyzed in young and mature leaves of three mid-successional tree species (Castanopsis fissa, Castanopsis chinensis and Schima superba) and three late-successional tree species (Machilus chinensis, Cryptocarya chinensis and Cryptocarya concinna), grown in 100% full sunlight (FL) or 30% of FL (low light, LL). Young leaves of the two species groups exhibited lower chlorophyll (Chl) content, Rubisco content, net photosynthetic rate (Pn), carboxylation efficiency (CE), effective photochemical yield (ΦPSII), photorespiratory electron flow (JO), but higher dark respiration (Rd), and ratios of carotenoids/chlorophylls (Car/Chl), anthocyanins/chlorophylls (Anth/Chl), flavonoids/chlorophylls (Flav/Chl), phenols/chlorophylls (Phen/Chl) and total antioxidant capacity/chlorophylls (TAC/Chl) than those of mature leaves, regardless of growth irradiance. Young leaves of both species groups demonstrated a higher flexibility of Anth/Chl, Flav/Chl, Phen/Chl and TAC/Chl in response to different light conditions than mature leaves. Flav/Chl in young leaves of late-successional group was remarkably higher than that of mid-successional group under the same light conditions. There was a negative correlation between antioxidant-dependent photoprotective potential and photosynthetic efficiency in young and mature leaves of the six tree species grown under either FL or LL. Our results explain partial mechanisms that lie behind the replacement of communities in subtropical forests: highly integrated photoprotective potential allows young leaves of shade-tolerant late-successional species to develop smoothly into mature organs under high irradiance.

Joan Mary (Jan) Anderson pioneered the investigation of the molecular organization of the plant thylakoid membrane, making seminal discoveries that laid the foundations for the current understanding of photosynthesis. She grew up in Queenstown, New Zealand, obtaining a BSc and MSc at the University of Otago in Dunedin. After completing her PhD at the University of California, she embarked on a glittering career at CSIRO and then the Australian National University in Canberra. Not only a gifted experimentalist, Jan was a creative thinker, not afraid to put her insightful and prophetic hypotheses into the public domain. Her many notable achievements include establishing the details and the physiological significance of lateral heterogeneity in the distribution of the two photosystems between stacked and unstacked thylakoid membranes and the dynamic changes in the extent of stacking that occur in response to changes in the light environment. Her investigations brought her into collaboration with leading researchers throughout the world. Recognized with many honours as a leading scientist in Australia, international recognition included the Lifetime Achievement Award from the International Society of Photosynthesis Research and honorary fellowships at universities in the UK and USA.

Anthocyanins and nonphotochemical quenching (NPQ) are two important tools that provide photoprotection in plant leaves. In order to understand how plants use these tools for acclimation to changing seasonal conditions, we investigated pigments, antioxidative capacity, and photosynthesis in leaves of an evergreen tree (Acmena acuminatissima) in two contrasting seasons. Young leaves of A. acuminatissima appeared in distinct colors, being light green in summer and red in winter due to the presence of anthocyanins. In the winter young leaves, anthocyanins contributed less than 2% to the antioxidant pool. In the summer, young leaves had higher NPQ than that of mature leaves, but in the winter, they did not derive any NPQ-related advantage over mature leaves. These results suggest that the accumulation of anthocyanins in young leaves in the winter may compensate for the insufficient photoprotection afforded by NPQ and that anthocyanins function as a light attenuator to protect the photochemical apparatus against excess light.

Ascorbate is an important antioxidant involved in both enzymatic and nonenzymatic reactions in plant cells. To reveal the function of ascorbate associated with defense against photo-oxidative damage, responses of the ascorbate-deficient mutant vtc2-1 of Arabidopsis thaliana to high-light stress were investigated. After high-light treatment at 1,600 μmol(photon) m–2 s–1 for 8 h, the vtc2-1 mutant exhibited visible photo-oxidative damage. The total ascorbate content was lower, whereas accumulation of H2O2 was higher in the vtc2-1 mutant than that in the wild type. The chlorophyll (Chl) content and PSII Chl fluorescence parameters, such as maximal quantum yield of PSII photochemistry, yield, and electron transport rate, in vtc2-1 mutant decreased more than that in the wild type, whereas the nonphotochemical quenching coefficient increased more in the wild type than that in vtc2-1 mutant. Therefore, the vtc2-1 mutant was more sensitive to high-light stress than the wild type. Accumulation of reactive oxygen species was mainly responsible for the damage of PSII in the vtc2-1 mutant under high light. The results indicate that ascorbate plays a critical role in maintaining normal photosynthetic function in plants under high-light stress.

Addition of K+ improves the tolerance of the photosynthetic apparatus to salt stress, but the primary, underlying mechanism is unclear, prompting our study. The main aim of this study was to elucidate the primary mechanism by which supplemental K+ alleviates the effects of salinity stress on photosynthetic parameters in mulberry seedlings. K+ contents in leaves and chloroplasts, plasma-membrane ATPase activity, photosynthetic properties and expression of genes encoding photosynthetic proteins and ion channels/transporters were measured. K+ addition during salt stress improved H+-ATPase activity in plasma-membranes, restored photosynthetic linear and cyclic electron flow, restored the activity of key Photosystem I (PSI) and PSII proteins, and increased dark-adapted PSII photochemical yield Fv/Fm, while reducing non-photochemical quenching. Among the genes encoding ion channels/transporters, that of the Two-Pore-K+ channel in thylakoid membranes (TPK3) was notably responsive to NaCl/K+ treatments: relative to the control (CK) treatment, it decreased slightly in the CK + K treatment, but increased in the NaCl treatment and increased further in the NaCl + K treatment. The above observations are consistent with our hypothesis that K+ supplementation in salt stress restores a high stromal [K+] which, by utilizing TPK3, minimizes a transient, proton diffusion potential that otherwise develops as protons exit the thylakoid lumen through the ATP synthase; thus, rapid charge compensation by K+ movement accelerates both the proton efflux from the lumen and the associated ATP synthesis to enhance carbon assimilation, thereby improving linear as well as cyclic electron flow. The hypothesis pinpoints the primary physicochemical mechanism by which potassium mitigates salt-stress impacts on photosynthesis. It rationalizes both the existence of a K+ channel in thylakoid membranes and the maintenance of a high stromal [K+].

Phototropic leaf movement of plants is an effective mechanism for adapting to light conditions. Light is the major driver of plant photosynthesis. Leaf N is also an important limiting factor on leaf photosynthetic potential. Cotton (Gossypium hirsutum L.) exhibits diaheliotropic leaf movement. Here, we compared the long-term photosynthetic acclimation of fixed leaves (restrained) and free leaves (allowed free movement) in cotton. The fixed leaves and free leaves were used for determination of PAR, leaf chlorophyll concentration, leaf N content and leaf gas exchange. The measurements were conducted under clear sky conditions at 0, 7, 15 and 30 days after treatment (DAT). The results showed that leaf N allocation and partitioning among different components of the photosynthetic apparatus were significantly affected by diaheliotropic leaf movement. Diaheliotropic leaf movement significantly increased light interception per unit leaf area, which in turn affected leaf mass per area (LMA), leaf N content (N

To clarify the influence of water deficit on the functionality of the photosynthetic apparatus of cotton plants, leaf gas exchange, chlorophyll a fluorescence, and P700 redox state were examined in field-grown cotton Gossypium hirsutum L. cv. Xinluzao 45. In addition, we measured changes in the P515 signal and analyzed the activity of ATP synthase and the trans-thylakoid proton gradient (ΔpH). With increasing water deficit, the net CO2 assimilation rate (AN) and stomatal conductance (gs) significantly decreased, but the maximum quantum efficiency of PSII photochemistry (Fv/Fm) did not change. The photochemical activity of photosystem II (PSII) was reflected by the photochemical quenching coefficient (qP), quantum efficiency of photosystem II [Y(II)], and electron transport rate through PSII [ETR(II)], while the activity of photosystem I (PSI) was reflected by the quantum efficiency of photosystem I [Y(I)] and the electron transport rate through PSI [ETR(I)]. Both activities were maintained under mild water deficit, but were slightly decreased under moderate water deficit. Under moderate water deficit, cyclic electron flow (CEF), the fraction of absorbed light dissipated thermally via the ΔpH- and xanthophyll-regulated process [Y(NPQ)], and the fraction of P700 oxidized under a given set of conditions [Y(ND)] increased. Our results suggest that the activities of both photosystems are stable under mild water deficit and decrease only slightly under moderate water deficit. Moderate water deficit stimulates CEF, and the stimulation of CEF is essential for protecting PSI and PSII against photoinhibition.

Photodamage to Photosystem II (PSII) has been attributed either to excessive excitation of photosynthetic pigments or by direct of light absorption by Mn4CaO5 cluster. Here we investigated the time course of PSII photodamage and release of Mn in PSII-enriched membranes under high light illumination at 460 nm and 660 nm. We found that the loss of PSII activity, assayed by chlorophyll fluorescence, is faster than release of Mn from the Mn4CaO5 cluster, assayed by EPR. Loss of PSII activity and Mn release was slower during illumination in the presence of exogenous electron acceptors. Recovery of PSII activity was observed, after 30 min of addition of electron donor post illumination. The same behavior was observed under 460 and 660 nm illumination, suggesting stronger correlation between excessive excitation and photodamage compared to direct light absorption by the cluster. A unified model of PSII photodamage that takes into account present and previous literature reports is presented.

The presence of anthocyanins in young leaves plays an important role in mitigation against photodamage and allows leaves to grow and develop normally. Many studies have reported that foliar anthocyanins are distributed within the vacuoles of mesophyll cells, so we explored the novel defence style of anthocyanin-coated young leaves of Castanopsis fissa, a dominant subtropical forest tree species, via removable trichomes. Anthocyanins were distributed in C. fissa leaf trichomes, which produced a red coating for the young leaves. As young leaves developed and then matured, the thickness and density of the anthocyanin trichomes progressively decreased, the coating finally disappearing, allowing greater utilization of light by mature leaves. In addition to anthocyanins, the trichomes contained a remarkably high amount of phenolics, which enable the red coating to be more efficient in screening ultraviolet light. Compared with mature leaves, the young leaves exhibited lower photosynthetic ability, which was attributable to the reduced chlorophyll and Rubisco contents. Removal of the red coating had little effect on the photosynthetic capacity of young leaves. However, the young leaves without the coating suffered greater light-induced photoinhibition due to greater excess light entering the chloroplast and the production of H2O2 Our results suggest that the anthocyanin coating is photoprotective and this anthocyanin defence style may be a metabolically cost-effective way of adjusting the anthocyanin content in response to demand.

The responses of gas exchange, chlorophyll fluorescence and the anti-oxidative system of cotton leaves were studied during water deficit and recovery. The results show that water deficit led to a reversible reduction in the photosynthetic rate. This reduction was mainly accompanied by stomatal limitation. The activity of photosystem II (PSII) and photosystem I (PSI) was relatively stable during water deficit and recovery. Water deficit caused an enhanced production of reactive oxygen species (ROS) and increased lipid peroxidation. Proline accumulation and the anti-oxidative enzymes such as superoxide dismutase (SOD), ascorbate peroxidase (APX) and peroxidase (POD), along with the antioxidant ascorbate (AsA), increased during water deficit. On re-watering, the ROS generation rate, anti-oxidative enzymes activities and the extent of the lipid peroxidation returned to near control values. Overall, rapid recovery of the photosynthetic rate is related to the stability of the photosystems which appears to be a critical mechanism allowing cotton plants to withstand and survive drought environments.

The quenching of chlorophyll fluorescence caused by photodamage of Photosystem II (qI) is a well recognized phenomenon, where the nature and physiological role of which are still debatable. Paradoxically, photodamage to the reaction centre of Photosystem II is supposed to be alleviated by excitation quenching mechanisms which manifest as fluorescence quenchers. Here we investigated the time course of PSII photodamage in vivo and in vitro and that of picosecond time-resolved chlorophyll fluorescence (quencher formation). Two long-lived fluorescence quenching processes during photodamage were observed and were formed at different speeds. The slow-developing quenching process exhibited a time course similar to that of the accumulation of photodamaged PSII, while the fast-developing process took place faster than the light-induced PSII damage. We attribute the slow process to the accumulation of photodamaged PSII and the fast process to an independent quenching mechanism that precedes PSII photodamage and that alleviates the inactivation of the PSII reaction centre.

Oxygen effects have long been ambiguous: exacerbating, being indifferent to, or ameliorating the net photoinactivation of Photosystem II (PS II). We scrutinized the time course of PS II photoinactivation (characterized by rate coefficient k i) in the absence of repair, or when recovery (characterized by k r) occurred simultaneously in CO2 ± O2. Oxygen exacerbated photoinactivation per se, but alleviated it by mediating the utilization of electrons. With repair permitted, the gradual net loss of functional PS II during illumination of leaves was better described phenomenologically by introducing τ, the time for an initial k r to decrease by half. At 1500 μmol photons m(-2) s(-1), oxygen decreased the initial k r but increased τ. Similarly, at even higher irradiance in air, there was a further decrease in the initial k r and increase in τ. These observations are consistent with an empirical model that (1) oxygen increased k i via oxidative stress but decreased it by mediating the utilization of electrons; and (2) reactive oxygen species stimulated the degradation of photodamaged D1 protein in PS II (characterized by k d), but inhibited the de novo synthesis of D1 (characterized by k s), and that the balance between these effects determines the net effect of O2 on PS II functionality.

Action spectra of photoinactivation of Photosystem II (PS II) in wild-type and chlorophyll b-less barley leaf segments were obtained. Photoinactivation of PS II was monitored by the delivery of electrons from PS II to PS I following single-turnover flashes superimposed on continuous far-red light, the time course of photoinactivation yielding a rate coefficient k i. Susceptibility of PS II to photoinactivation was quantified as the ratio of k i to the moderate irradiance (I) of light at each selected wavelength. k i/I was very much higher in blue light than in red light. The experimental conditions permitted little excess light energy absorbed by chlorophyll (not utilized in photochemical conversion or dissipated in controlled photoprotection) that could lead to photoinactivation of PS II. Therefore, direct absorption of light by Mn in PS II, rather than by chlorophyll, was more likely to have initiated the much more severe photoinactivation in blue light than in red light. Mutant leaves were ca. 1.5-fold more susceptible to photoinactivation than the wild type. Neither the excess-energy mechanism nor the Mn mechanism can explain this difference. Instead, the much lower chlorophyll content of mutant leaves could have exerted an exacerbating effect, possibly partly due to less mutual shading of chloroplasts in the mutant leaves. In general, which mechanism dominates depends on the experimental conditions.

The cyclic electron flux (CEF) around photosystem I (PSI) was discovered in isolated chloroplasts more than six decades ago, but its quantification has been hampered by the absence of net formation of a product or net consumption of a substrate. We estimated in vivo CEF in leaves as the difference (ΔFlux) between the total electron flux through PSI (ETR1) measured by a near infrared signal, and the linear electron flux through both photosystems by optimised measurement of chlorophyll a fluorescence (LEFfl). Chlorophyll fluorescence was excited by modulated green light from a light-emitting diode at an optimal average irradiance, and the fluorescence was detected at wavelengths >710 nm. In this way, LEFfl matched the gross rate of oxygen evolution multiplied by 4 (LEFO2) in broad-spectrum white actinic irradiance up to half (spinach, poplar and rice) or one third (cotton) of full sunlight irradiance. This technique of estimating CEF can be applied to leaves attached to a plant.

Joan Mary (Jan) Anderson pioneered the investigation of the molecular organization of the plant thylakoid membrane, making seminal discoveries that laid the foundations for the current understanding of photosynthesis. She grew up in Queenstown, New Zealand, obtaining a BSc and MSc at the University of Otago in Dunedin. After completing her PhD at the University of California, she embarked on a glittering career at CSIRO and then the Australian National University in Canberra. Not only a gifted experimentalist, Jan was a creative thinker, not afraid to put her insightful and prophetic hypotheses into the public domain. Her many notable achievements include establishing the details and the physiological significance of lateral heterogeneity in the distribution of the two photosystems between stacked and unstacked thylakoid membranes and the dynamic changes in the extent of stacking that occur in response to changes in the light environment. Her investigations brought her into collaboration with leading researchers throughout the world. Recognized with many honours as a leading scientist in Australia, international recognition included the Lifetime Achievement Award from the International Society of Photosynthesis Research and honorary fellowships at universities in the UK and USA.

© 2017 Szabó, Larkum, Suggett, Vass, Sass, Osmond, Zavafer, Ralph and Chow. Reef building corals (phylum Cnidaria) harbor endosymbiotic dinoflagellate algae (genus Symbiodinium) that generate photosynthetic products to fuel their host's metabolism. Non-invasive techniques such as chlorophyll (Chl) fluorescence analyses of Photosystem II (PSII) have been widely used to estimate the photosynthetic performance of Symbiodinium in hospite. However, since the spatial origin of PSII chlorophyll fluorescence in coral tissues is uncertain, such signals give limited information on depth-integrated photosynthetic performance of the whole tissue. In contrast, detection of absorbance changes in the near infrared (NIR) region integrates signals from deeper tissue layers due to weak absorption and multiple scattering of NIR light. While extensively utilized in higher plants, NIR bio-optical techniques are seldom applied to corals. We have developed a non-intrusive measurement method to examine photochemistry of intact corals, based on redox kinetics of the primary electron donor in Photosystem I (P700) and chlorophyll fluorescence kinetics (Fast-Repetition Rate fluorometry, FRRf). Since the redox state of P700 depends on the operation of both PSI and PSII, important information can be obtained on the PSII-PSI intersystem electron transfer kinetics. Under moderate, sub-lethal heat stress treatments (33◦ C for~20 min), the coral Pavona decussata exhibited down-regulation of PSII electron transfer kinetics, indicated by slower rates of electron transport from QA to plastoquinone (PQ) pool, and smaller relative size of oxidized PQ with concomitant decrease of a specifically-defined P700 kinetics area, which represents the active pool of PSII. The maximum quantum efficiency of PSII (Fv /Fm ) and functional absorption cross-section of PSII (σPSII ) remained unchanged. Based on the coordinated response of P700 parameters and PSII-PSI electron transport properties, we propose that simple P700 kinetics parameters as employed here serve as indicators of the integrity of PSII-PSI electron transfer dynamics in corals.

Background Polyploidy is an important phenomenon in plants because of its roles in agricultural and forestry production as well as in plant tolerance to environmental stresses. Tetraploid black locust (Robinia pseudoacacia L.) is a polyploid plant and a pioneer tree species due to its wide ranging adaptability to adverse environments. To evaluate the ploidy-dependent differences in leaf mitochondria between diploid and tetraploid black locust under salinity stress, we conducted comparative proteomic, physiological, biochemical and ultrastructural profiling of mitochondria from leaves. Results Mitochondrial proteomic analysis was performed with 2-DE and MALDI-TOF-MS, and the ultrastructure of leaf mitochondria was observed by transmission electron microscopy. According to 2-DE analysis, 66 proteins that responded to salinity stress significantly were identified from diploid and/or tetraploid plants and classified into 9 functional categories. Assays of physiological characters indicated that tetraploids were more tolerant to salinity stress than diploids. The mitochondrial ultrastructure of diploids was damaged more severely under salinity stress than that of tetraploids. Conclusions Tetraploid black locust possessed more tolerance of, and ability to acclimate to, salinity stress than diploids, which may be attributable to the ability to maintain mitochondrial structure and to trigger different expression patterns of mitochondrial proteins during salinity stress. Electronic supplementary material The online version of this article (doi:10.1186/s12864-017-4038-2) contains supplementary material, which is available to authorized users.

Many flooding-tolerant species are clonal plants; however, the effects of physiological integration on plant responses to flooding have received limited attention. We hypothesise that flooding can trigger changes in metabolism of carbohydrates and ROS (reactive oxygen species) in clonal plants, and that physiological integration can ameliorate the adverse effects of stress, subsequently restoring the growth of flooded ramets. In the present study, we conducted a factorial experiment combining flooding to apical ramets and stolon severing (preventing physiological integration) between apical and basal ramets of Cynodon dactylon, which is a stoloniferous perennial grass with considerable flooding tolerance. Flooding-induced responses including decreased root biomass, accumulation of soluble sugar and starch, as well as increased activity of superoxide dismutase (SOD) and ascorbate peroxidase (APX) in apical ramets. Physiological integration relieved growth inhibition, carbohydrate accumulation and induction of antioxidant enzyme activity in stressed ramets, as expected, without any observable cost in unstressed ramets. We speculate that relief of flooding stress in clonal plants may rely on oxidising power and electron acceptors transferred between ramets through physiological integration.

During leaf development, the increase in stomatal conductance cannot meet photosynthetic demand for CO2, thus leading to stomatal limitation of photosynthesis (Ls). Considering the crucial influences of stomatal development on stomatal conductance, we speculated whether stomatal development limits photosynthesis to some extent. To test this hypothesis, stomatal development, stomatal conductance and photosynthesis were carefully studied in both Syringa oblata (normal greening species) and Euonymus japonicus Thunb (delayed greening species). Our results show that the size of stomata increased gradually with leaf expansion, resulting in increased stomatal conductance up to the time of full leaf expansion. During this process, photosynthesis also increased steadily. Compared to that in S. oblata, the development of chloroplasts in E. japonicus Thunb was obviously delayed, leading to a delay in the improvement of photosynthetic capacity. Further analysis revealed that before full leaf expansion, stomatal limitation increased rapidly in both S. oblata and E. japonicus Thunb; after full leaf expansion, stomatal limitation continually increased in E. japonicus Thunb. Accordingly, we suggested that the enhancement of photosynthetic capacity is the main factor leading to stomatal limitation during leaf development but that stomatal development can alleviate stomatal limitation with the increase of photosynthesis by controlling gas exchange.

Elucidation of the mechanisms by which plants adapt to elevated CO2 is needed; however, most studies of the mechanisms investigated the response of plants adapted to current atmospheric CO2. The rapid respiration rate of cotton (Gossypium hirsutum) fruits (bolls) produces a concentrated CO2 microenvironment around the bolls and bracts. It has been observed that the intercellular CO2 concentration of a whole fruit (bract and boll) ranges from 500 to 1300 µmol mol(-1) depending on the irradiance, even in ambient air. Arguably, this CO2 microenvironment has existed for at least 1·1 million years since the appearance of tetraploid cotton. Therefore, it was hypothesized that the mechanisms by which cotton bracts have adapted to elevated CO2 will indicate how plants will adapt to future increased atmospheric CO2 concentration. Specifically, it is hypothesized that with elevated CO2 the capacity to regenerate ribulose-1,5-bisphosphate (RuBP) will increase relative to RuBP carboxylation.To test this hypothesis, the morphological and physiological traits of bracts and leaves of cotton were measured, including stomatal density, gas exchange and protein contents.Compared with leaves, bracts showed significantly lower stomatal conductance which resulted in a significantly higher water use efficiency. Both gas exchange and protein content showed a significantly greater RuBP regeneration/RuBP carboxylation capacity ratio (Jmax/Vcmax) in bracts than in leaves.These results agree with the theoretical prediction that adaptation of photosynthesis to elevated CO2 requires increased RuBP regeneration. Cotton bracts are readily available material for studying adaption to elevated CO2.

Using radioactively labelled amino acids to investigate repair of photoinactivated photosystem II (PS II) gives only a relative rate of repair, while using chlorophyll fluorescence parameters yields a repair rate coefficient for an undefined, variable location within the leaf tissue. Here, we report on a whole-tissue determination of the rate coefficient of photoinactivation k i , and that of repair k r in cotton leaf discs. The method assays functional PS II via a P700 kinetics area associated with PS I, as induced by a single-turnover, saturating flash superimposed on continuous background far-red light. The P700 kinetics area, directly proportional to the oxygen yield per single-turnover, saturating flash, was used to obtain both k i and k r . The value of k i , directly proportional to irradiance, was slightly higher when CO2 diffusion into the abaxial surface (richer in stomata) was blocked by contact with water. The value of k r , sizable in darkness, changed in the light depending on which surface was blocked by contact with water. When the abaxial surface was blocked, k r first peaked at moderate irradiance and then decreased at high irradiance. When the adaxial surface was blocked, k r first increased at low irradiance, then plateaued, before increasing markedly at high irradiance. At the highest irradiance, k r differed by an order of magnitude between the two orientations, attributable to different extents of oxidative stress affecting repair (Nishiyama et al., EMBO J 20: 5587-5594, 2001). The method is a whole-tissue, convenient determination of the rate coefficient of photoinactivation k i and that of repair k r .

The prototype light-induced fluorescence transient (LIFT) instrument provides continuous, minimally intrusive, high time resolution (~2 s) assessment of photosynthetic performance in terrestrial plants from up to 2 m. It induces a chlorophyll fluorescence transient by a series of short flashes in a saturation sequence (180 ~1μs flashlets in

Joan Mary Anderson, known to most people as Jan, was born on May 12, 1932 in Dunedin, New Zealand. She died on August 28, 2015 in Canberra, Australia. To celebrate her life, we present here a brief biography, some comments on her discoveries in photosynthesis during a career spanning more than half a century, and reminiscences from family and friends. We remember this wonderful person who had an unflagging curiosity, creative ability to think laterally, enthusiasm, passion, generosity and love of color and culture.