Sudden Increase In Light Intensity Research Articles

The ribosome assembly GTPase EngA is involved in redox signaling in cyanobacteria.

Photosynthetic organisms must cope with environmental challenges, like those imposed by the succession of days and nights or by sudden changes in light intensities, that trigger global changes in gene expression and metabolism. The photosynthesis machinery is particularly susceptible to environmental changes and adaptation to them often involves redox-sensing proteins that are the targets of reactive oxygen species generated by photosynthesis activity. Here we show that EngA, an essential GTPase and ribosome-assembly protein involved in ribosome biogenesis in bacteria and chloroplasts, also plays a role in acclimatization to environmentally relevant stress in Synechococcus elongatus PCC7942 and that PipX, a promiscuous regulatory protein that binds to EngA, appears to fine-tune EngA activity. During growth in cold or high light conditions, the EngA levels rise, with a concomitant increase of the EngA/PipX ratio. However, a sudden increase in light intensity turns EngA into a growth inhibitor, a response involving residue Cys122 of EngA, which is part of the GD1-G4 motif NKCES of EngA proteins, with the cysteine conserved just in the cyanobacteria-chloroplast lineage. This work expands the repertoire of ribosome-related factors transmitting redox signals in photosynthetic organisms and provides additional insights into the complexity of the regulatory interactions mediated by EngA and PipX.

Flv3A facilitates O2 photoreduction and affects H2 photoproduction independently of Flv1A in diazotrophic Anabaena filaments.

The model heterocyst-forming filamentous cyanobacterium Anabaena sp. PCC 7120 (Anabaena) is a typical example of a multicellular organism capable of simultaneously performing oxygenic photosynthesis in vegetative cells and O2 -sensitive N2 -fixation inside heterocysts. The flavodiiron proteins have been shown to participate in photoprotection of photosynthesis by driving excess electrons to O2 (a Mehler-like reaction). Here, we performed a phenotypic and biophysical characterization of Anabaena mutants impaired in vegetative-specific Flv1A and Flv3A in order to address their physiological relevance in the bioenergetic processes occurring in diazotrophic Anabaena under variable CO2 conditions. We demonstrate that both Flv1A and Flv3A are required for proper induction of the Mehler-like reaction upon a sudden increase in light intensity, which is likely important for the activation of carbon-concentrating mechanisms and CO2 fixation. Under ambient CO2 diazotrophic conditions, Flv3A is responsible for moderate O2 photoreduction, independently of Flv1A, but only in the presence of Flv2 and Flv4. Strikingly, the lack of Flv3A resulted in strong downregulation of the heterocyst-specific uptake hydrogenase, which led to enhanced H2 photoproduction under both oxic and micro-oxic conditions. These results reveal a novel regulatory network between the Mehler-like reaction and the diazotrophic metabolism, which is of great interest for future biotechnological applications.

The success of the bloom-forming cyanobacteria Planktothrix: Genotypes variability supports variable responses to light and temperature stress

Cyanobacterial blooms can modify the dynamic of aquatic ecosystems and have harmful consequences for human activities. Moreover, cyanobacteria can produce a variety of cyanotoxins, including microcystins, but little is known about the role of environmental factors on the prevalence of microcystin producers in the cyanobacterial bloom dynamics. This study aimed to better understand the success of Planktothrix in various environments by unveiling the variety of strategies governing cell responses to sudden changes in light intensity and temperature. The cellular responses (photosynthesis, photoprotection, heat shock response and metabolites synthesis) of four Planktothrix strains to high-light or high-temperature were studied, focusing on how distinct ecotypes (red- or green-pigmented) and microcystin production capability affect cyanobacteria's ability to cope with such abiotic stimuli. Our results showed that high-light and high-temperature impact different cellular processes and that Planktothrix responses are heterogeneous, specific to each strain and thus, to genotype. The ability of cyanobacteria to cope with sudden increase in light intensity and temperature was not related to red- or green-pigmented ecotype or microcystin production capability. According to our results, microcystin producers do not cope better to high-light or high-temperature and microcystin content does not increase in response to such stresses.

Photosynthetic regulation in response to fluctuating light conditions under temperature stress in three mosses with different light requirements

Under natural field conditions, mosses experience fluctuating light intensities combined with temperature stress. Alternative electron flow mediated by flavodiiron proteins (FLVs) and cyclic electron flow (CEF) around photosystem I (PSI) allow mosses to growth under fluctuating light conditions. However, little is known about the roles of FLVs and CEF in the regulation of photosynthesis under temperature stress combined with fluctuating light. Here, we measured chlorophyll fluorescence and P700 redox state under fluctuating light conditions at 4 °C, 20 °C, and 35 °C in three mosses with different light requirements. Upon a sudden increase in light intensity, electron flow from photosystem II initially increased and then gradually decreased at 20 °C and 35 °C, indicating that the operation of FLV-dependent flow lasted much longer than previously thought. Furthermore, the absolute rates of FLV-dependent flow and CEF were enhanced under fluctuating light at 35 °C, pointing to their important roles in photoprotection when exposed to fluctuating light at moderate high temperature. Furthermore, the downregulation of FLV activity at 4 °C was partially compensated for by enhanced CEF activity. These results suggested the subtle coordination between FLV activity and CEF under fluctuating light and temperature stress. Racomitrium japonicum and Hypnum plumaeforme, which usually grow under relatively high light levels, exhibited higher FLV activity and CEF than the shade-grown moss Plagiomnium ellipticum. Based on our results, we conclude that photosynthetic acclimation to fluctuating light and temperature stress in different mosses is largely linked to the adjustment of FLV activity and CEF.

The water-water cycle is not a major alternative sink in fluctuating light at chilling temperature

The water-water cycle (WWC) has the potential to alleviate photoinhibition of photosystem I (PSI) in fluctuating light (FL) at room temperature and moderate heat stress. However, it is unclear whether WWC can function as a safety valve for PSI in FL at chilling temperature. In this study, we measured P700 redox state and chlorophyll fluorescence in FL at 25 °C and 4 °C in the high WWC activity plant Dendrobium officinale. At 25 °C, the operation of WWC contributed to the rapid re-oxidation of P700 upon dark-to-light transition. However, such rapid re-oxidation of P700 was not observed at 4 °C. Upon a sudden increase in light intensity, WWC rapidly consumed excess electrons in PSI and thus avoided an over-reduction of PSI at 25 °C. On the contrary, PSI was highly reduced within the first seconds after transition from low to high light at 4 °C. Therefore, in opposite to 25 °C, the WWC is not a major alternative sink in FL at chilling temperature. Upon transition from low to high light, cyclic electron transport was highly stimulated at 4 °C when compared with 25 °C. These results indicate that D. officinale enhances cyclic electron transport to partially compensate for the inactivation of WWC in FL at 4 °C.

Photodamage to photosystem in a typically shade-tolerant species <i>Panax notoginseng</i> exposed to a sudden increase in light intensity

Photodamage to photosystem in a typically shade-tolerant species <i>Panax notoginseng</i> exposed to a sudden increase in light intensity

Rice Cultivar Takanari Has Higher Photosynthetic Performance Under Fluctuating Light Than Koshihikari, Especially Under Limited Nitrogen Supply and Elevated CO2.

Plants in the field experience dynamic changes of sunlight rather than steady-state irradiation. Therefore, increasing the photosynthetic rate of an individual leaf under fluctuating light is essential for improving crop productivity. The high-yielding indica rice (Oryza sativa L.) cultivar Takanari is considered a potential donor of photosynthesis genes because of its higher steady-state photosynthesis at both atmospheric and elevated CO2 concentrations than those of several Japanese commercial cultivars, including Koshihikari. Photosynthetic induction after a sudden increase in light intensity is faster in Takanari than in Koshihikari, but whether the daily carbon gain of Takanari outperforms that of Koshihikari under fluctuating light in the field is unclear. Here we report that Takanari has higher non-steady-state photosynthesis, especially under low nitrogen (N) supply, than Koshihikari. In a pot experiment, Takanari had greater leaf carbon gain during the initial 10 min after a sudden increase in irradiation and higher daily CO2 assimilation under simulated natural fluctuating light, at both atmospheric (400 ppm) and elevated (800 ppm) CO2 concentrations. The electron transport rate during a day under field conditions with low N supply was also higher in Takanari than in Koshihikari. Although the advantages of Takanari were diminished under high N supply, photosynthetic N use efficiency was consistently higher in Takanari than in Koshihikari, under both low and high N supply. This study demonstrates that Takanari is a promising donor parent to use in breeding programs aimed at increasing CO2 assimilation in a wide range of environments, including future higher CO2 concentrations.

Genetic variation in the photosynthetic induction response in rice (Oryza sativa L.)

ABSTRACT Crop leaves growing under field conditions experience fluctuating light intensities. The photosynthetic response to a sudden increase in light intensity, which is termed the photosynthetic induction response, can potentially affect crop productivity. In the present study, the genetic variation in the photosynthetic induction response and the related gas exchange parameters in the rice diversity research set of germplasm (RDRS) were evaluated using two reference genotypes: Koshihikari and Takanari. Takanari is known to show superior induction response than Koshihikari. The photosynthetic induction response was found to be highly diverse in 59 rice genotypes. The cumulative CO2 fixation during the first 10 minutes after a transition from low to high light intensity (CCF10) differed by a factor of four between genotypes. The variation of CCF10 showed no relationship with light saturated photosynthetic rate (R = 0.21), but was significantly correlated with photosynthetic rate (A) under low light conditions (R = 0.88). Several genotypes showed even more rapid induction responses than that of Takanari. In particular, A of ARC 11094 and Rexmont both increased rapidly during the first two minutes of the induction response. The CCF10 of these two genotypes was approximately four times greater than that of Koshihikari. The rapid induction responses of Rexmont and ARC 11094 most likely resulted from the activation of CO2 fixation in mesophyll cells and CO2 diffusion from the air to the leaves, respectively. This study demonstrates that there is a large genetic variation in and, therefore, much potential for genetic improvement of, the photosynthetic induction response in rice.

A high-throughput assay for screening environmental pollutants and drugs impairing predator avoidance in Daphnia magna

This study addresses short-term habituation of the escape response in the aquatic crustacean Daphnia magna evoked by sudden changes in light intensity, using a high-throughput system. Daphnia magna exhibits a marked phototactic behaviour and swim away from light to avoid predation by fish. Currently, there is no information available on the habituation of this phototactic response. The Daphnia photomotor response assay (DPRA) measures the distance moved after a sudden increase in light intensity. Using DPRA, it is possible to determine not only the magnitude of the phototactic response, but also its habituation after repetitive cycles of light and darkness. The progressive reduction observed in response to a series of light stimuli in the proposed assay meet the criteria for habituation. Most cholinergic and serotonergic modulators enhanced photomotor responses and reduced habituation. Dopaminergic and histaminergic modulators also reduced habituation, whereas diazepam was the only compound that increased habituation. Imidacloprid, apomorphine, diphenhydramine, diazepam, and memantine decreased photomotor responses. Thus, the DPRA was also predictive in assessing the effects of neuroactive and neurotoxic environmental contaminants such as selective serotonin reuptake inhibitors, diazepam, organophosphorous, and neonicotinoid pesticides. We conclude that the proposed DPRA may be an effective screening tool for compounds that can impair predation avoidance behaviour in aquatic organisms.

Charge transfer states in phycobilisomes

Phycobilisomes (PBs) absorb light and supply downstream photosynthetic processes with excitation energy in many cyanobacteria and algae. In response to a sudden increase in light intensity, excess excitation energy is photoprotectively dissipated in PBs by means of the orange carotenoid protein (OCP)-related mechanism or via a light-activated intrinsic decay channel. Recently, we have identified that both mechanisms are associated with far-red emission states. Here, we investigate the far-red states involved with the light-induced intrinsic mechanism by exploring the energy landscape and electro-optical properties of the pigments in PBs. While Stark spectroscopy showed that the far-red states in PBs exhibit a strong charge-transfer (CT) character at cryogenic temperatures, single molecule spectroscopy revealed that CT states should also be present at room temperature. Owing to the strong environmental sensitivity of CT states, the knowledge gained from this study may contribute to the design of a new generation of fluorescence markers.

PGR5-Dependent Cyclic Electron Flow Protects Photosystem I under Fluctuating Light at Donor and Acceptor Sides.

In response to a sudden increase in light intensity, plants must cope with absorbed excess photon energy to protect photosystems from photodamage. Under fluctuating light, PSI is severely photodamaged in the Arabidopsis (Arabidopsis thaliana) proton gradient regulation5 (pgr5) mutant defective in the main pathway of PSI cyclic electron transport (CET). Here, we aimed to determine how PSI is protected by two proposed regulatory roles of CET via transthylakoid ΔpH formation: (1) reservation of electron sink capacity by adjusting the ATP/NADPH production ratio (acceptor-side regulation) and (2) down-regulation of the cytochrome b 6 f complex activity called photosynthetic control for slowing down the electron flow toward PSI (donor-side regulation). We artificially enhanced donor- and acceptor-side regulation in the wild-type and pgr5 backgrounds by introducing the pgr1 mutation conferring the hypersensitivity of the cytochrome b 6 f complex to luminal acidification and moss Physcomitrella patens flavodiiron protein genes, respectively. Enhanced photosynthetic control partially alleviated PSI photodamage in the pgr5 mutant background but restricted linear electron transport under constant high light, suggesting that the strength of photosynthetic control should be optimized. Flavodiiron protein-dependent oxygen photoreduction formed a large electron sink and alleviated PSI photoinhibition, accompanied by the induction of photosynthetic control. Thus, donor-side regulation is essential for PSI photoprotection but acceptor-side regulation also is important to rapidly induce donor-side regulation. In angiosperms, PGR5-dependent CET is required for both functions.

Polygonal motion and adaptable phototaxis via flagellar beat switching in the microswimmer Euglena gracilis

Biological microswimmers exhibit versatile strategies for sensing and navigating their environment, such as run-and-tumble and curvature modulation. Here, we report a striking phototactic behaviour of the microswimmer Euglena gracilis, where these eukaryotic cells swim in polygonal trajectories due to a sudden increase in light intensity. While smoothly curved trajectories are common for microswimmers, such quantized ones have not been reported previously. We find that this polygonal behaviour emerges from periodic switching between the flagellar beating patterns of helical swimming and spinning behaviours. We develop and experimentally validate a biophysical model that describes the phase relationship between the eyespot, cell orientation, light detection and cellular reorientation, accounting for all three behavioural states. Coordinated switching between these behaviours selects for ballistic, superdiffusive, diffusive or subdiffusive motion (including tuning the effective diffusion constant over several orders of magnitude), thereby enabling navigation in spatially structured light fields, such as edge avoidance and gradient descent. This feedback control links multiple system scales (flagellar beats, cellular behaviours and phototaxis strategies), with implications for other natural and synthetic microswimmers. A single-celled organism exhibits complex swimming behaviours in response to changes in light intensity. Modelling and experiments suggest that the swimmer exploits phase relations between its photoreceptor and orientation to enable navigation.

Switching an Individual Phycobilisome Off and On.

Photosynthetic organisms have found various smart ways to cope with unexpected changes in light conditions. In many cyanobacteria, the lethal effects of a sudden increase in light intensity are mitigated mainly by the interaction between phycobilisomes (PBs) and the orange carotenoid protein (OCP). The latter senses high light intensities by means of photoactivation and triggers thermal energy dissipation from the PBs. Due to the brightness of their emission, PBs can be characterized at the level of individual complexes. Here, energy dissipation from individual PBs was reversibly switched on and off using only light and OCP. We reveal the presence of quasistable intermediate states during the binding and unbinding of OCP to PB, with a spectroscopic signature indicative of transient decoupling of some of the PB rods during docking of OCP. Real-time control of emission from individual PBs has the potential to contribute to the development of new super-resolution imaging techniques.

Effect of a Sudden Increase in Light Intensity on Normalized Difference Vegetation Index (NDVI) Reflected from Leaves of Tobacco

Effect of a Sudden Increase in Light Intensity on Normalized Difference Vegetation Index (NDVI) Reflected from Leaves of Tobacco

LHCSR3 affects de-coupling and re-coupling of LHCII to PSII during state transitions in Chlamydomonas reinhardtii

Photosynthetic organisms have to tolerate rapid changes in light intensity, which is facilitated by non-photochemical quenching (NPQ) and involves modification of energy transfer from light-harvesting complexes (LHC) to the photosystem reaction centres. NPQ includes dissipating excess light energy to heat (qE) and the reversible coupling of LHCII to photosystems (state transitions/qT), which are considered separate NPQ mechanisms. In the model alga Chlamydomonas reinhardtii the LHCSR3 protein has a well characterised role in qE. Here, it is shown in the npq4 mutant, deficient in LHCSR3, that energy coupling to photosystem II (PSII) more akin to qT is also disrupted, but no major differences in LHC phosphorylation or LHC compositions were found in comparison to wild-type cells. The qT of wild-type cells possessed two kinetically distinguishable phases, with LHCSR3 participating in the more rapid (<2 min) phase. This LHCSR3-mediated qT was sensitive to physiological levels of H2O2, which accelerated qE induction, revealing a way that may help C. reinhardtii tolerate a sudden increase in light intensity. Overall, a clear mechanistic overlap between qE and qT is shown.

Photosystem II photoinhibition-repair cycle protects Photosystem I from irreversible damage

Photodamage of Photosystem II (PSII) has been considered as an unavoidable and harmful reaction that decreases plant productivity. PSII, however, has an efficient and dynamically regulated repair machinery, and the PSII activity becomes inhibited only when the rate of damage exceeds the rate of repair. The speed of repair is strictly regulated according to the energetic state in the chloroplast. In contrast to PSII, Photosystem I (PSI) is very rarely damaged, but when occurring, the damage is practically irreversible. While PSII damage is linearly dependent on light intensity, PSI gets damaged only when electron flow from PSII exceeds the capacity of PSI electron acceptors to cope with the electrons. When electron flow to PSI is limited, for example in the presence of DCMU, PSI is extremely tolerant against light stress. Proton gradient (ΔpH)-dependent slow-down of electron transfer from PSII to PSI, involving the PGR5 protein and the Cyt b6f complex, protects PSI from excess electrons upon sudden increase in light intensity. Here we provide evidence that in addition to the ΔpH-dependent control of electron transfer, the controlled photoinhibition of PSII is also able to protect PSI from permanent photodamage. We propose that regulation of PSII photoinhibition is the ultimate regulator of the photosynthetic electron transfer chain and provides a photoprotection mechanism against formation of reactive oxygen species and photodamage in PSI.

Light History Influences the Response of Fluvial Biofilms to Zn Exposure.

Fluvial biofilms are subject to multistress situations in natural ecosystems, such as the co-occurrence of light intensity changes and metal toxicity. However, studies simultaneously addressing both factors are rare. This study evaluated in microcosm conditions the relationship between short-term light intensity changes and Zn toxicity on fluvial biofilms with long-term photoacclimation to different light conditions. Biofilms that had long-term photoacclimation to 25μmol photons·m(-2) · s(-1) (low light [LL] biofilms), 100μmol photons· m(-2) ·s(-1) (medium light [ML] biofilms), and 500μmol photons·m(-2) ·s(-1) (high light [HL] biofilms) were characterized by different structural (Chlorophyll-a [Chl-a], total biomass-AFDW, EPS, algal groups, and diatom taxonomy) and physiological attributes (ETR-I curves and photosynthetic pigments). HL biofilms showed higher light saturation intensity and a higher production of xanthophylls than LL biofilms. In contrast, LL biofilms had many structural differences; a higher proportion of diatoms and lower AFDW and EPS contents than ML and HL biofilms. A clear effect of light intensity changes on Zn toxicity was also demonstrated. Zn toxicity was enhanced when a sudden increase in light intensity also occurred, mainly with LL biofilms, causing higher inhibition of both the Φ'PSII and the ΦPSII . A decoupling of NPQ from de-epoxidation reaction (DR) processes was also observed, indicating substantial damage to photoprotective mechanisms functioning in biofilms (i.e., xanthophyll cycle of diatoms) due to Zn toxicity. This study highlights the need to take into account environmental stress (e.g., light intensity changes) to better assess the environmental risks of chemicals (e.g., metals).

Growth and physiological changes in saplings of Minquartia guianensis and Swietenia macrophylla during acclimation to full sunlight

Low light availability under a forest canopy often limits plant growth; however, sudden increase in light intensity may induce photoinhibition of photosynthesis. The aim of this study was to evaluate the ecophysiological changes that occur in potted plants of Minquartia guianensis and Swietenia macrophylla during the acclimation process to full sunlight. We used six full-sun independent acclimation periods (30, 60, 90, 120, 150, and 180 days) and a control kept in the shade. Shading was obtained by placing plants under the canopy of a small forest. The Fv/Fm ratio, net photosynthetic rate (PN), the maximum carboxylation velocity of Rubisco (Vcmax), maximum electron transport rate (Jmax), specific leaf area (SLA), and growth were assessed at the end of each of the six acclimation periods. Plant exposure to full sunlight caused a sudden decrease in the Fv/Fm ratio (photoinhibition) particularly in Minquartia. Photooxidation (necrotic patches) of the leaf tissue was observed in upper leaves of Minquartia. The higher PN values were observed in Swietenia under full sun, about 12 μmol(CO2) m−2 s−1. Vcmax25 values were higher after 90 days of acclimation, about 14 μmol(CO2) m−2 s−1 for Minquartia, and 35 μmol(CO2) m−2 s−1 for Swietenia. At the end of a 180-d acclimation period Jmax25 was 35 μmol(electron) m−2 s−1 for Minquartia and 60 μmol(electron) m−2 s−1 for Swietenia. SLA was higher in Swietenia than in Minquartia. In Minquartia, monthly rate of leaf production per plant (MRLP) was positive (0.22 leaf month−1) after four months in the open. Whereas, in Swietenia MRLP was positive (0.56 leaf month−1) after an acclimation period of two months. After six months in the open, height growth rates were 3.5 and 28 mm month−1 for Minquartia and Swietenia, respectively. The greater acclimation capacity of Swietenia was associated to an enhanced photosynthetic plasticity under full sun. In Minquartia, transition to full-sun conditions and lack of physiological adjustment resulted in severe photoinhibition and loss of leaves.

Removing aboveground vegetation facilitates survival but slows height growth of spruce saplings in a fenced, degraded sub-alpine forest in central Japan

The effects of removing aboveground vegetation on survival and growth of Hondo spruce (Picea jezoensis var. hondoensis) saplings were examined in a dense dwarf bamboo (Sasa nipponica) field inside a deer-proof fence built in a degraded sub-alpine forest on Mt Ohdaigahara, central Japan. All bamboo culms were removed within a radius of 1 m from each sapling in a plot of 0.6 ha (removal plot), whereas no culms were removed in two control plots of 0.1 ha. We measured the height, crown depth, and main shoot elongation over 1 year for all spruce saplings, culm height and cover ratios of dwarf bamboo growing around each spruce sapling, and the light levels above each sapling. Removal of dwarf bamboo regulated the culm height and bamboo coverage until the following summer such that light conditions improved. The mortality of saplings smaller than the average culm height was lower in the removal plot (0.89%) than in a control plot (5.9%). After bamboo removal, sapling height growth was reduced with increasing bamboo cover in the previous year. Complete removal of the aboveground portions of covering bamboo improved sapling survival while regulating elongation growth, possibly because of the sudden increase in light intensity. Thus, controlling the extent of dwarf bamboo removal might be necessary to facilitate the growth of spruce saplings to restore the forest.

Photochemistry, remotely sensed physiological reflectance index and de-epoxidation state of the xanthophyll cycle in Quercus coccifera under intense drought

Quercus coccifera L. is a Mediterranean sclerophyllous shrub with a high capacity to resist intense drought stress. Therefore, it could be used in the study of physiological changes suffered by plants at very low water potentials. A remote sensing sensor was used to measure continuously the physiological reflectance index (PRI; defined as the changes in reflectance at 531 nm with respect to those at 570 nm; PRI = [(R531 - R570)/(R531 + R570)] at canopy level and under field conditions in an artificial carpet of seedlings of Q. coccifera during a drought cycle. Correlations between leaf level-measured chlorophyll fluorescence parameters as well as the de-epoxidation state of the xanthophyll cycle [(A + Z)/(V + A + Z)] and canopy level-measured PRI were reasonably good (R (2) = 0.57-0.63, P < 0.01), and quite interesting for water stress remote sensing purposes. The instrument's temporal resolution allowed us to follow the rapid response of PRI to changing photosynthetic active radiation, and to resolve, in response to cloud-induced changes in light intensity, a fast and a slow PRI component. We report the disappearance of the rapid one under conditions of intense drought in response to a sudden increase in light intensity. The underlying photoprotection mechanisms that Q. coccifera shows in response to intense drought stress periods seem to be related to the existence of a low intrathylakoid lumenal pH at the end of the drought cycle. Under intense drought, these mechanisms allow this species to avoid oxidative damage, which was evidenced by the maintenance of an unaltered photosynthetic pigment composition and constant photosystem II efficiency in the mornings. It is concluded that, contrary to early reports, PRI is a sensible, indirect, non-destructive water stress indicator, even in plants experiencing intense drought.