Lower Non-photochemical Quenching Research Articles

The role of the pigment-protein complex LHCBM1 in nonphotochemical quenching in Chlamydomonas reinhardtii.

Nonphotochemical quenching (NPQ) is the process that protects photosynthetic organisms from photodamage by dissipating the energy absorbed in excess as heat. In the model green alga Chlamydomonas reinhardtii, NPQ is abolished in the knock-out mutants of the pigment-protein complexes LHCSR3 and LHCBM1. However, while LHCSR3 is a pH sensor and switches to a quenched conformation at low pH, the role of LHCBM1 in NPQ has not been elucidated yet. In this work, we combined biochemical and physiological measurements to study short-term high-light acclimation of npq5, the mutant lacking LHCBM1. In low light in the absence of this complex, the antenna size of PSII was smaller than in its presence; this effect was marginal in high light (HL), implying that a reduction of the antenna was not responsible for the low NPQ. The mutant expressed LHCSR3 at the wild-type level in HL, indicating that the absence of this complex is also not the reason. Finally, NPQ remained low in the mutant even when the pH was artificially lowered to values that can switch LHCSR3 to the quenched conformation. We concluded that both LHCSR3 and LHCBM1 are required for the induction of NPQ and that LHCBM1 is the interacting partner of LHCSR3. This interaction can either enhance the quenching capacity of LHCSR3 or connect this complex with the PSII supercomplex.

Microscale imaging sheds light on species-specific strategies for photo-regulation and photo-acclimation of microphytobenthic diatoms.

Intertidal microphytobenthic (MPB) biofilms are key sites for coastal primary production, predominantly by pennate diatoms exhibiting photo-regulation via non-photochemical quenching (NPQ) and vertical migration. Movement is the main photo-regulation mechanism of motile (epipelic) diatoms and because they can move from light, they show low-light acclimation features such as low NPQ levels, as compared to non-motile (epipsammic) forms. However, most comparisons of MPB species-specific photo-regulation have used low light acclimated monocultures, not mimicking environmental conditions. Here we used variable chlorophyll fluorescence imaging, fluorescent labelling in sediment cores and scanning electron microscopy to compare the movement and NPQ responses to light of four epipelic diatom species from a natural MPB biofilm. The diatoms exhibited different species-specific photo-regulation features and a large NPQ range, exceeding that reported for epipsammic diatoms. This could allow epipelic species to coexist in compacted light niches of MPB communities. We show that diatom cell orientation within MPB can be modulated by light, where diatoms oriented themselves more perpendicular to the sediment surface under high light vs. more parallel under low light, demonstrating behavioural, photo-regulatory response by varying their light absorption cross-section. This highlights the importance of considering species-specific responses and understanding cell orientation and photo-behaviour in MPB research.

Ripening Process of Tomato Fruits Postharvest: Impact of Environmental Conditions on Quality and Chlorophyll a Fluorescence Characteristics

This study aimed to investigate the combined effects of temperature and light conditions on tomato maturation. Tomato fruits that had completed volumetric growth at the mature green stage were harvested and matured in growth chambers composed of two temperature conditions (daytime–nighttime: 30–20 °C or 20–15 °C) and two light conditions (0 µmol·m−2·s−1 or 400 µmol·m−2·s−1), which were set for 12 h each day and night. Our findings indicate that tomato ripening was significantly influenced by both light and temperature. Tomatoes that matured under low-temperature conditions in the absence of light took more than three times longer to transition from the green stage to the breaker stage compared to those matured under high-temperature conditions with light exposure. Notably, tomato fruit maturation occurred at a faster rate under low-temperature and light conditions than under high-temperature and dark conditions. Changes in chlorophyll a fluorescence parameters were observed throughout the ripening process of tomato fruits. Tomato fruits ripened under low-temperature and dark conditions exhibited significantly lower NPQ (non-photochemical quenching) and RFD (relative fluorescence decrease) values compared to other treatments, while their FO (initial fluorescence) and FM (maximum fluorescence) values were higher. The accumulation of sugar in tomato fruits was observed to be more influenced by light than temperature. On the other hand, the highest levels of phenolic content and lycopene were observed in tomato fruits matured under high-temperature and light conditions. Antioxidant activities, as measured by ABTS and DPPH assays, were highest in mature tomato fruits under high-temperature and light conditions, while they were lowest in fruits under low-temperature and dark conditions. In conclusion, this study highlights the critical role of temperature and light as crucial environmental factors influencing tomato maturation. Understanding these factors can contribute to optimizing postharvest conditions and enhancing fruit quality in the tomato industry.

Chlorophyll fluorescence variation in two Mediterranean forest species over a 21-year drought treatment period.

The Mediterranean is among the regions predicted to be most affected by climate change due to rising temperatures and increasing frequency as well as intensity of extreme weather events, such as drought. These shifts in climatic conditions might lead to changes in species community composition by enabling the increase of drought-tolerant species at the expense of less tolerant ones. This hypothesis was tested in the current study using chlorophyll fluorescence data from a 21-year precipitation exclusion experiment in a Mediterranean forest for two co-dominant Quercus ilex and Phillyrea latifolia species with contrasting low and high levels of drought tolerance, respectively. The maximum potential quantum efficiency of photosystem II (PSII) (Fv/Fm), photochemical efficiency of PSII (yield), and non-photochemical quenching (NPQ) varied seasonally. Fv/Fm and NPQ levels were positively correlated with air temperature and Standardized Precipitation-Evapotranspiration Index (SPEI), whereas yield, which was greater under drought treatment was negatively associated with vapor pressure deficit and SPEI. Fv/Fm values showed similar increase in the two species over the 21-year study period regardless of treatment and in parallel to progressive warming. In contrast, yield values were higher in Q. ilex than in P. latifolia, while NPQ values were higher in P. latifolia than in Q. ilex. Notably, high yield values were also observed in the drought treated plots. Throughout the study, plants in the drought treated plots exhibited decreased basal area, leaf biomass, and aerial cover due to high stem mortality. In addition, a continuous increase in temperature was detected in summer and autumn, which might explain the observed increase in Fv/Fm values over the study period. Higher yield and lower NPQ detected in Q. ilex could be attributed to less competition for resources in the drought treated plots and acclimation of Q. ilex plants over the study period. Our results indicate that reduction in stem density could improve forest resilience to climate change-induced drought conditions.

The sensitivity of photosynthesis to magnesium deficiency differs between rice (Oryza sativa L.) and cucumber (Cucumis sativus L.).

Magnesium is an essential macronutrient for plant photosynthesis, and in response to Mg deficiency, dicots appear more sensitive than monocots. Under Mg deficiency, we investigated the causes of differing photosynthetic sensitivities in a dicot and a monocot species. Rice (Oryza sativa L.) and cucumber (Cucumis sativus L.) were grown in hydroponic culture to explore their physiological responses to Mg deficiency stress. Both Mg-deficient rice and cucumber plants exhibited lower biomass, leaf area, Mg concentration, and chlorophyll content (Chl) compared with Mg-sufficient plants. However, a more marked decline in Chl and carotenoid content (Car) occurred in cucumber. A lower CO2 concentration in chloroplasts (C c) was accompanied by a decrease in the maximum rate of electron transport (J max) and the maximum rate of ribulose 1,5-bisphosphate carboxylation (V cmax), restricting CO2 utilization in Mg-deficient plants. Rice and cucumber photorespiration rate (P r) increased under Mg deficiency. Additionally, for cucumber, Car and non-photochemical quenching (NPQ) were reduced under lower Mg supply. Meanwhile, cucumber Mg deficiency significantly increased the fraction of absorbed light energy dissipated by an additional quenching mechanism (Φf,D). Under Mg deficiency, suppressed photosynthesis was attributed to comprehensive restrictions of mesophyll conductance (g m), J max, and V cmax. Cucumber was more sensitive to Mg deficiency than rice due to lower NPQ, higher rates of electron transport to alternative pathways, and subsequently, photooxidation damage.

Response of photosynthesis, the xanthophyll cycle, and wax in Japanese yew (Taxus cuspidata L.) seedlings and saplings under high light conditions.

In order to understand the adaptative changes of the Japanese yew (Taxus cuspidate L.) to high light conditions, this study investigated gas-exchange, chlorophyll fluorescence, chlorophyll, and the impact of epicuticular wax on the gas-exchange and photoinhibition of Japanese yew seedlings and saplings. The chlorophyll content per unit area and photosynthetic rate in seedling leaves were significantly lower than in sapling leaves. When leaves from seedlings and saplings were exposed to 1,200 μmol·m-2·s-1 photon flux density (PFD) for 2 h, seedling leaves exhibited a greater down-regulation of maximum quantum yield (Fv/Fm) and actual photosystem II efficiency ( PSII). Non-photochemical quenching (NPQ) and high energy quenching (qE) in sapling leaves were much higher than in seedling leaves when both were exposed to 1,200 μmol·m-2·s-1 PFD for 2 h. At a low level of O2, the photorespiration rate (Pr) and the ratio of photorespiration/gross photosynthetic rate (Pr/Pg) in seedling leaves were lower than in sapling leaves when both were exposed to 1,200 μmol·m-2·s-1 PFD, but this difference did not reach statistical significance (P < 0.05). Compared with sapling leaves, seedling leaves exhibited lower levels of xanthophyll pool. Epicuticular wax content on seedling leaves was significantly lower than on sapling leaves. The results of this study showed that wax coverage on the leaf surface decreased the photosynthetic rate in sapling leaves as a consequence of decreased stomatal conductance. Epicuticular wax is related to tree age and photoinhibition prevention in the Japanese yew. It is possible that lower photosynthetic rate, lower NPQ depending on the xanthophyll cycle, and lower deposition of epicuticular wax results in seedling plants that are not adapted to high light conditions.

Melanisation in Boreal Lichens Is Accompanied by Variable Changes in Non-Photochemical Quenching.

Lichens often grow in microhabitats where they absorb more light than they can use for fixing carbon, and this excess energy can cause the formation of harmful reactive oxygen species (ROS). Lichen mycobionts can reduce ROS formation by synthesizing light-screening pigments such as melanins in the upper cortex, while the photobionts can dissipate excess energy radiationlessly using non-photochemical quenching (NPQ). An inherent problem with using fluorimetry techniques to compare NPQ in pale and melanised thalli is that NPQ is normally measured through a variously pigmented upper cortex. Here we used a dissection technique to remove the lower cortices and medullas of Lobaria pulmonaria and Crocodia aurata and then measure NPQ from the underside of the thallus. Results confirmed that NPQ can be satisfactorily assessed with a standard fluorimeter by taking measurement from above using intact thalli. However, photobionts from the bottom of the photobiont layer tend to have slightly lower rates of PSII activity and lower NPQ than those at the top, i.e., display mild “shade” characteristics. Analysis of pale and melanised thalli of other species indicates that NPQ in melanised thalli can be higher, similar or lower than pale thalli, probably depending on the light history of the microhabitat and presence of other tolerance mechanisms.

PeVDE, a violaxanthin de-epoxidase gene from moso bamboo, confers photoprotection ability in transgenic Arabidopsis under high light.

Plants employ an array of photoprotection mechanisms to alleviate the harmful effects of high light intensity. The violaxanthin cycle, which is associated with non-photochemical quenching (NPQ), involves violaxanthin de-epoxidase (VDE), and zeaxanthin epoxidase (ZEP) and is one of the most rapid and efficient mechanisms protecting plants under high light intensity. Woody bamboo is a class of economically and ecologically important evergreen grass species widely distributed in tropical and subtropical areas. However, the function of VDE in bamboo has not yet been elucidated. In this study, we found that high light intensity increased NPQ and stimulated the de-epoxidation of violaxanthin cycle components in moso bamboo (Phyllostachys edulis), whereas, samples treated with the VDE inhibitor (dithiothreitol) exhibited lower NPQ capacity, suggesting that violaxanthin cycle plays an important role in the photoprotection of bamboo. Further analysis showed that not only high light intensity but also extreme temperatures (4 and 42°C) and drought stress upregulated the expression of PeVDE in bamboo leaves, indicating that PeVDE is induced by multiple abiotic stresses. Overexpression of PeVDE under the control of the CaMV 35S promoter in Arabidopsis mutant npq1 mutant could rescue its NPQ, indicating that PeVDE functions in dissipating the excess absorbed light energy as thermal energy in bamboo. Moreover, compared with wild-type (Col-0) plants, the transgenic plants overexpressing PeVDE displayed enhanced photoprotection ability, higher NPQ capacity, slower decline in the maximum quantum yield of photosystem II (Fv/Fm) under high light intensity, and faster recovery under optimal conditions. These results suggest that PeVDE positively regulates the response to high light intensity in bamboo plants growing in the natural environment, which could improve their photoprotection ability through the violaxanthin cycle and NPQ.

The Responses of Light Reaction of Photosynthesis to Dynamic Sunflecks in a Typically Shade-Tolerant Species Panax notoginseng.

Light is highly heterogeneous in natural conditions, and plants need to evolve a series of strategies to acclimate the dynamic light since it is immobile. The present study aimed to elucidate the response of light reaction of photosynthesis to dynamic sunflecks in a shade-tolerant species Panax notoginseng and to examine the regulatory mechanisms involved in an adaptation to the simulated sunflecks. When P. notoginseng was exposed to the simulated sunflecks, non-photochemical quenching (NPQ) increased rapidly to the maximum value. Moreover, in response to the simulated sunflecks, there was a rapid increase in light-dependent heat dissipation quantum efficiency of photosystem II (PSII) (ΦNPQ), while the maximum quantum yield of PSII under light (Fv′/Fm′) declined. The relatively high fluorescence and constitutive heat dissipation quantum efficiency of PSII (Φf,d) in the plants exposed to transient high light (400, 800, and 1,600 μmol m–2 s–1) was accompanied by the low effective photochemical quantum yield of PSII (ΦPSII) after the dark recovery for 15 min, whereas the plants exposed to transient low light (50 μmol m–2 s–1) has been shown to lead to significant elevation in ΦPSII after darkness recovery. Furthermore, PSII fluorescence and constitutive heat dissipation electron transfer rate (Jf,d) was increased with the intensity of the simulated sunflecks, the residual absorbed energy used for the non-net carboxylative processes (JNC) was decreased when the response of electron transfer rate of NPQ pathway of PSII (JNPQ) to transient low light is restricted. In addition, the acceptor-side limitation of PSI [Y(NA)] was increased, while the donor-side limitation of photosystems I (PSI) [Y(ND)] was decreased at transient high light conditions accompanied with active cyclic electron flow (CEF). Meanwhile, when the leaves were exposed to transient high light, the xanthophyll cycle (V cycle) was activated and subsequently, the JNPQ began to increase. The de-epoxidation state [(Z + A)/(V + A + Z)] was strongly correlated with NPQ in response to the sunflecks. In the present study, a rapid engagement of lutein epoxide (Lx) after the low intensity of sunfleck together with the lower NPQ contributed to an elevation in the maximum photochemical quantum efficiency of PSII under the light. The analysis based on the correlation between the CEF and electron flow devoted to Ribulose-1, 5-bisphosphate (RuBP) oxygenation (JO) indicated that at a high light intensity of sunflecks, the electron flow largely devoted to RuBP oxygenation would contribute to the operation of the CEF. Overall, photorespiration plays an important role in regulating the CEF of the shade-tolerant species, such as P. notoginseng in response to transient high light, whereas active Lx cycle together with the decelerated NPQ may be an effective mechanism of elevating the maximum photochemical quantum efficiency of PSII under light exposure to transient low light.

Role of cyclic electron transport mutations pgrl1 and pgr5 in acclimation process to high light in Chlamydomonas reinhardtii.

Light is crucial for photosynthesis, but the amount of light that exceeds an organism's assimilation efficacy can lead to photo-oxidative damage and even cell death. In Chlamydomonas (C). reinhardtii cyclic electron flow (CEF) is very important for the elicitation of non-photochemical quenching (NPQ) by controlling the acidification of thylakoid lumen. This process requires the cooperation of proton gradient regulation (PGR) proteins, PGRL1 and PGR5. Here, we compared the growth pattern and photosynthetic activity between wild type (137c, t222+) and mutants impaired in CEF (pgrl1 and pgr5) under photoautotrophic and photoheterotrophic conditions. We have observed the discriminative expression of NPQ in the mutants impaired in CEF of pgrl1 and pgr5. The results obtained from the mutants showed reduced cell growth and density, Chl a/b ratio, fluorescence, electron transport rate, and yield of photosystem (PS)II. These mutants have reduced capability to develop a strong NPQ indicating that the role of CEF is very crucial for photoprotection. Moreover, the CEF mutant exhibits increased photosensitivity compared with the wild type. Therefore, we suggest that besides NPQ, the fraction of non-regulated non-photochemical energy loss (NO) also plays a crucial role during high light acclimation despite a low growth rate. This low NPQ rate may be due to less influx of protons coming from the CEF in cases of pgrl1 and pgr5 mutants. These results are discussed in terms of the relative photoprotective benefit, related to the thermal dissipation of excess light in photoautotrophic and photoheterotrophic conditions.

Exogenous putrescine alleviates photoinhibition caused by salt stress through cooperation with cyclic electron flow in cucumber.

When plants suffer from abiotic stresses, cyclic electron flow (CEF) is induced for photo-protection. Putrescine (Put), a primary polyamine in chloroplasts, plays a critical role in stress tolerance. However, the relationship between CEF and Put in chloroplasts for photo-protection is unknown. In this study, we investigated the role of Put-induced CEF for salt tolerance in cucumber plants (Cucumis sativus L). Treatment with 90mM NaCl and/or Put did not influence the maximum photochemical efficiency of PSII (Fv/Fm), but the photoactivity of PSI was severely inhibited by NaCl. Salt stress induced a high level of CEF; moreover, plants treated with both NaCl and Put exhibited much higher CEF activity and ATP accumulation than those exhibited by single-salt-treated plants to provide an adequate ATP/NADPH ratio for plant growth. Furthermore, Put decreased the trans-membrane proton gradient (ΔpH), which was accompanied by reduced pH-dependent non-photochemical quenching (NPQ) and an increased the effective quantum yield of PSII (Y(II)). The ratio of NADP+/NADPH increased significantly with Put in salt-stressed leaves compared with the ratio in leaves treated with NaCl, indicating that Put relieved over-reduction pressure at the PSI acceptor side caused by salt stress. Collectively, our results suggest that exogenous Put creates an excellent condition for CEF promotion: a large amount of pmf is predominantly stored as Δψ, resulting in moderate lumen pH and low NPQ, while maintaining high rates of ATP synthesis (high pmf).

Importance of the mitochondrial alternative oxidase (AOX) pathway in alleviating photoinhibition in cucumber leaves under chilling injury and subsequent recovery when leaves are subjected to high light intensity

ABSTRACTThe aim of this study was to assess the impact of the mitochondrial alternative oxidase (AOX) pathway on energy metabolism in chloroplast, and evaluate the importance of the AOX pathway in alleviating photoinhibition in cucumber (Cucumis sativus L.) leaves during chilling and the subsequent recovery period. Chilling (12°C) induced up-regultaion of the AOX pathway, which is important for chilled leaves to avoid overreduction of PSII and maintain photosynthetic electron transport. High light (200–300 µmol m−2 s−1) induced the decreases of maximal quantum yield of PSII photochemistry (Fv/Fm) and nonphotochemical quenching (NPQ), which was aggravated by 1 mM salicylhydroxamic acid (SHAM; AOX inhibitor), in cucumber leaves during chilling. However, Fv/Fm did not change and NPQ increased significantly in SHAM-treated leaves by chilling under low light. During recovery periods after chilling under low light, Fv/Fm, qP, ΦPSII, NPQ, and Fv’/Fm’ could return to normal degree in chilled leaves with or without SHAM. In contrast, high light induced more decrease of chlorophyll fluorescence parameters in chilled leaves during the recovery period, and SHAM treatment further aggravated the decrease. These results implied that up-regulation of AOX pathway in chilled leaves plays a critical role in protection against photoinhibition in leaves under excess light energy condition during chilling and the subsequent recovery period. However, thermal energy dissipation was more important to prevent photoinhibition in leaves exposed to chilling under low light.

Towards efficient photosynthesis: overexpression of Zea mays phosphoenolpyruvate carboxylase in Arabidopsis thaliana.

Plants with C4 photosynthesis are efficient in carbon assimilation and have an advantage over C3 photosynthesis. In C4 photosynthesis, the primary CO2 fixation is catalyzed by phosphoenolpyruvate carboxylase (PEPC). Here, we show that overexpression of Zea mays PEPC cDNA, under the control of 35S promoter, in Arabidopsis thaliana resulted in ~7-10 fold higher protein abundance and ~7-10 fold increase in PEPC activity in the transgenic lines than that in the vector control. We suggest that overexpression of PEPC played an anaplerotic role to increase the supply of 4-carbon carboxylic acids, which provided carbon skeletons for increased amino acid and protein synthesis. Higher protein content must have been responsible for increased metabolic processes including chlorophyll biosynthesis, photosynthesis, and respiration. Consequently, the PEPC-overexpressed transgenic plants had higher chlorophyll content, enhanced electron transport rate (ETR), lower non-photochemical quenching (NPQ) of chlorophyll a fluorescence, and a higher performance index (PI) than the vector control. Consistent with these observations, the rate of CO2 assimilation, the starch content, and the dry weight of PEPC-overexpressed plants increased by 14-18%, 10-18%, and 6.5-16%, respectively. Significantly, transgenics were tolerant to salt stress as they had increased ability to synthesize amino acids, including the osmolyte proline. NaCl (150mM)-treated transgenic plants had higher variable to maximum Chl a fluorescence (F v/F m) ratio, higher PI, higher ETR, and lower NPQ than the salt-treated vector controls. These results suggest that expression of C4 photosynthesis enzyme(s) in a C3 plant can improve its photosynthetic capacity with enhanced tolerance to salinity stress.

Photosynthesis and photoprotection under drought in the annual desert plant Anastatica hierochuntica

Anastatica hierochuntica is an annual desert plant, which was recently shown to have unusually low nonphotochemical quenching (NPQ) and a high PSII electron transport rate (ETR). In the current study, we examined how these unusual characteristics are related to a lack of CO2 and inhibition of net photosynthetic rate (P N). We compared the photosynthetic and photoprotective response of A. hierochuntica and sunflower (Helianthus annuus), under conditions of photosynthetic inhibition, with either low CO2 or drought. We found that under nonsteady state conditions of low CO2 availability, A. hierochuntica exhibited about half of the NPQ values and almost twice of the ETR values of H. annuus. However, the long-term inhibition of P N under drought caused a similar increase in NPQ and a decrease in ETR in both A. hierochuntica and H. annuus. These results suggest that the unusually low NPQ and high ETR in A. hierochuntica are not directly related to a response to drought conditions.

Enhanced chloroplastic generation of H2O2 in stress‐resistant Thellungiella salsuginea in comparison to Arabidopsis thaliana

In order to find some basis of salinity resistance in the chloroplastic metabolism, a halophytic Thellungiella salsuginea was compared with glycophytic Arabidopsis thaliana. In control T.s. plants the increased ratios of chlorophyll a/b and of fluorescence emission at 77 K (F730/F685) were documented, in comparison to A.t.. This was accompanied by a higher YII and lower NPQ (non-photochemical quenching) values, and by a more active PSI (photosystem I). Another prominent feature of the photosynthetic electron transport (PET) in T.s. was the intensive production of H2O2 from PQ (plastoquinone) pool. Salinity treatment (0.15 and 0.30 M NaCl for A.t. and T.s., respectively) led to a decrease in ratios of chl a/b and F730/F685. In A.t., a salinity-driven enhancement of YII and NPQ was found, in association with the stimulation of H2O2 production from PQ pool. In contrast, in salinity-treated T.s., these variables were similar as in controls. The intensive H2O2 generation was accompanied by a high activity of PTOX (plastid terminal oxidase), whilst inhibition of this enzyme led to an increased H2O2 formation. It is hypothesized, that the intensive H2O2 generation from PQ pool might be an important element of stress preparedness in Thellungiella plants. In control T.s. plants, a higher activation state of carboxylase ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco, EC 4.1.1.39) was also documented in concert with the attachment of Rubisco activase (RCA) to the thylakoid membranes. It is supposed, that a closer contact of RCA with PSI in T.s. enables a more efficient Rubisco activation than in A.t.

Growth form defines physiological photoprotective capacity in intertidal benthic diatoms.

In intertidal marine sediments, characterized by rapidly fluctuating and often extreme light conditions, primary production is frequently dominated by diatoms. We performed a comparative analysis of photophysiological traits in 15 marine benthic diatom species belonging to the four major morphological growth forms (epipelon (EPL), motile epipsammon (EPM-M) and non-motile epipsammon (EPM-NM) and tychoplankton (TYCHO)) found in these sediments. Our analyses revealed a clear relationship between growth form and photoprotective capacity, and identified fast regulatory physiological photoprotective traits (that is, non-photochemical quenching (NPQ) and the xanthophyll cycle (XC)) as key traits defining the functional light response of these diatoms. EPM-NM and motile EPL showed the highest and lowest NPQ, respectively, with EPM-M showing intermediate values. Like EPL, TYCHO had low NPQ, irrespective of whether they were grown in benthic or planktonic conditions, reflecting an adaptation to a low light environment. Our results thus provide the first experimental evidence for the existence of a trade-off between behavioural (motility) and physiological photoprotective mechanisms (NPQ and the XC) in the four major intertidal benthic diatoms growth forms using unialgal cultures. Remarkably, although motility is restricted to the raphid pennate diatom clade, raphid pennate species, which have adopted a non-motile epipsammic or a tychoplanktonic life style, display the physiological photoprotective response typical of these growth forms. This observation underscores the importance of growth form and not phylogenetic relatedness as the prime determinant shaping the physiological photoprotective capacity of benthic diatoms.

Satellite‐detected fluorescence: Decoupling nonphotochemical quenching from iron stress signals in the South Atlantic and Southern Ocean

AbstractSatellite‐detected sunlight‐induced chlorophyll fluorescence could offer valuable information about the physiological status of phytoplankton on a global scale. Realization of this potential is confounded by the considerable uncertainty that exists in deconvolving the multiple ecophysiological processes that can influence the satellite signal. A dominant source of current uncertainty arises from the extent of reductions in chlorophyll fluorescence caused by the high light intensities phytoplankton are typically exposed to when satellite images are captured. In this study, results from over 200 nonphotochemical quenching (NPQ) experiments conducted on cruises spanning from subtropical gyre to Southern Ocean waters have confirmed that satellite fluorescence quantum yields have the potential to reveal broad regions of iron (Fe) stress. However, our results suggest significant variability in phytoplankton NPQ behavior between oceanic regimes. Dynamic NPQ must therefore be considered to achieve a reliable interpretation of satellite fluorescence in terms of Fe stress. Specifically, significantly lower NPQ was found in stratified subtropical gyre‐type waters than in well‐mixed Southern Ocean waters. Such variability is suggested to result from differences in incident irradiance fluctuation experienced by phytoplankton, with highly variable irradiance conditions likely driving phytoplankton to acclimate or adapt toward a higher dynamic NPQ capacity. Sea surface temperature empirically demonstrated the strongest correlation with NPQ parameters and is presented as a means of correcting the chlorophyll fluorescence signature for the region studied. With these corrections, a decadal composite of satellite austral summer observations is presented for the Southern Ocean, potentially reflecting spatial variability in the distribution and extent of Fe stress.

Prediction of respective contribution of linear electron flow and PGR5-dependent cyclic electron flow to non-photochemical quenching induction

In chloroplasts, regulated formation of the proton gradient across the thylakoid membrane (ΔpH) is important for controlling non-photochemical quenching (NPQ), which is crucial for plants to perform photosynthesis under fluctuating light conditions. The ΔpH is generated by two electron flows: the linear electron flow (LEF) and the cyclic electron flow (CEF). The Arabidopsis CEF mutant, pgr5, showed significantly lower NPQ values than those observed in WT, indicating that ΔpH, generated by the PGR5-dependent CEF, has a crucial role in controlling NPQ. However, the respective significance of LEF and CEF for ΔpH formation is largely unknown. Here we applied computer simulation to reproduce NPQ induction kinetics and estimate the respective contribution of LEF and PGR5-dependent CEF to the dynamics of ΔpH formation. The results indicate that the contribution of CEF to total ΔpH formation for induction of NPQ varies from 60–80%. The simulation also suggested a role of the PGR5-dependent CEF in accelerating electron transfer in the cytochrome b6f complex.

Low induction of non‐photochemical quenching and high photochemical efficiency in the annual desert plant Anastatica hierochuntica

Non-photochemical quenching (NPQ) plays a major role in photoprotection. Anastatica hierochuntica is an annual desert plant found in hot deserts. We compared A. hierochuntica to three other different species: Arabidopsis thaliana, Eutrema salsugineum and Helianthus annuus, which have different NPQ and photosynthetic capacities. Anastatica hierochuntica plants had very different induction kinetics of NPQ and, to a lesser extent, of photosystem II electron transport rate (PSII ETR), in comparison to all other plants species in the experiments. The major components of the unusual photosynthetic and photoprotective response in A. hierochuntica were: (1) Low NPQ at the beginning of the light period, at various light intensities and CO2 concentrations. The described low NPQ cannot be explained by low leaf absorbance or by low energy distribution to PSII, but was related to the de-epoxidation state of xanthophylls. (2) Relatively high PSII ETR at various CO2 concentrations in correlation with low NPQ. PSII ETR responded positively to the increase of CO2 concentrations. At low CO2 concentrations PSII ETR was mostly O2 dependent. At moderate and high CO2 concentrations the high PSII ETR in A. hierochuntica was accompanied by relatively high CO2 assimilation rates. We suggest that A. hierochuntica have an uncommon NPQ and PSII ETR response. These responses in A. hierochuntica might represent an adaptation to the short growing season of an annual desert plant.

Integrating Transient Heterogeneity of Non-Photochemical Quenching in Shade-Grown Heterobaric Leaves of Avocado (Persea americana L.): Responses to CO2 Concentration, Stomatal Occlusion, Dehydration and Relative Humidity

Long-lived shade leaves of avocado had extremely low rates of photosynthesis. Gas exchange measurements of photosynthesis were of limited use, so we resorted to Chl fluorescence imaging (CFI) and spot measurements to evaluate photosynthetic electron transport rates (ETRs) and non-photochemical quenching (NPQ). Imaging revealed a remarkable transient heterogeneity of NPQ during photosynthetic induction in these hypostomatous, heterobaric leaves, but was adequately integrated by spot measurements, despite long-lasting artifacts from repeated saturating flashes during assays. Major veins (mid-vein, first- and second-order veins) defined areas of more static large-scale heterogeneous NPQ, with more dynamic small-scale heterogeneity most strongly expressed in mesophyll cells between third- and fourth-order veins. Both responded to external CO2 concentration ([CO2]), occlusion of stomata with Vaseline™, leaf dehydration and relative humidity (RH). We interpreted these responses in terms of independent behavior of stomata in adjacent areoles that was largely expressed through CO2-limited photosynthesis. Heterogeneity was most pronounced and prolonged in the absence of net CO2 fixation in 100 p.p.m. [CO2] when respiratory and photorespiratory CO2 cycling constrained the inferred ETR to ~75% of values in 400 or 700 p.p.m. [CO2]. Likewise, sustained higher NPQ under Vaseline™, after dehydration or at low RH, also restricted ETR to ~75% of control values. Low NPQ in chloroplast-containing cells adjacent to major veins but remote from stomata suggested internal sources of high [CO2] in these tissues.