Effects of Drought Stress on the Photosynthesis in Maize

To clarify how the components of the entire photosynthetic electron transport chain in response to drought stress in maize. The activities of photosystem II (PSII), photosystem I (PSI), and the electron transport chain between PSII and PSI of maize were investigated by prompt fluorescence (PF), delayed fluorescence (DF) and 820 nm modulated reflection (MR). Maize (Zea mays L.) plants were subjected to different levels of soil water availability including control, moderate and severe drought stress. A significant decrease in ?E₀, Ψ₀ and PI_ABS was found in maize treated with moderate drought stress. A significant increase in ABS/RC was observed, but there were no significant change in the fast MR phase and the amplitude of DF under moderate drought stress compared to the control. Under severe drought stress, the exchange capacity between Q_A to Q_B, reoxidation capacity of plastoquinol, and the oxidation and re-reduction rates of PC and P₇₀₀ all decreased. These results demonstrated that moderate drought stress reduced the photochemical activity of PSII from Q_A to PQH₂, while the photochemical activity of PSI was unscathed. However, severe drought stress inhibited the entire electron transport chain from the donor side of PSII to PSI-end electron acceptors. In addition, the photochemical activity of PSII is more sensitive to drought stress than PSI.     

Photochemical efficiency of photosystem II,photon yield of O2 evolution,photosynthetic capacity,and carotenoid composition during the midday depression of net CO2 uptake in Arbutus unedo growing in Portugal

During the midday depression of net CO₂ exchange in the mediterranean sclerophyllous shrub Arbutus unedo, examined in the field in Portugal during August of 1987, several parameters indicative of photosynthetic competence were strongly and reversibly affected. These were the photochemical efficiency of photosystem (PS) II, measured as the ratio of variable to maximum chlorophyll fluorescence, as well as the photon yield and the capacity of photosynthetic O₂ evolution at 10% CO₂, of which the apparent photon yield of O₂ evolution was most depressed. Furthermore, there was a strong and reversible increase in the content of the carotenoid zeaxanthin in the leaves that occurred at the expense of both violaxanthin and -carotene. Diurnal changes in fluorescence characteristics were interpreted to indicate three concurrent effects on the photochemical system. First, an increase in the rate of radiationless energy dissipation in the antenna chlorophyll, reflected by changes in 77K fluorescence of PSII and PSI as well as in chlorophyll a fluorescence at ambient temperature. Second, a state shift characterized by an increase in the proportion of energy distributed to PSI as reflected by changes in PSI fluorescence. Third, an effect lowering the photon yield of O₂ evolution and PSII fluorescence at ambient temperature without affecting PSII fluorescence at 77K which would be expected from a decrease in the activity of the water splitting enzyme system, i.e. a donor side limitation.Abbreviations and symbols c_i concentration of CO₂ within the leaf - F_o instantaneous fluorescence emission - F_M maximum fluorescence emission - F_v variable fluorescence emission - PFD photon flux density (400–700 nm) - PSI, II photosystem I, II - T_L leaf temperature     

Temporal Pattern of Photosynthesis under Continuous Illumination of Radish Plants

Changes in photosynthetic activity, redox state of photosystem I (PSI) and photosystem II (PSII), as well as starch and sucrose content were studied on the source leaves of 18- to 20-day-old radish (Raphanus sativus L.) plants that were dark-adapted for 12 h and then exposed to continuous white light (170 mol quanta/(m² s)). The kinetic pattern of photosynthetic activity comprised three phases. Within the first 6 h of light adaptation (first phase), the maximum photosynthetic rate and the quantum yield of photosynthesis increased 1.6 times in the illuminated leaves compared to the leaves of plants placed in darkness. Further illumination led to the decrease of both photosynthetic indices by about 20% (12 h after the onset of light exposure, second phase) and finally increased them to the level observed after 6-h light exposure (72 h, third phase). The stationary photooxidation level of PSI primary donor was relatively low within the first 6 h of light adaptation, and then it steeply increased. The linear relationship between the amounts of photoreduced PSII primary acceptor and photooxidized PSI primary donor did not change during prolonged light adaptation, showing a highly coordinated functioning of both photosystems. The amount of sucrose in leaves attained its peak after 12 h of light adaptation and did not change further on. The starch content increased to its peak within 24 h of illumination and decreased gradually upon longer exposures. It is concluded that, despite active export of assimilates to the developing storage organ, the source leaves exhibit a nonmonotonic temporal course of endogenously regulated photosynthetic activity, which was related to changes in the effectiveness and, possibly, the number of the components of photosynthetic apparatus.     

Cyclic electron flow plays an important role in photoprotection for the resurrection plant <Emphasis Type="Italic">Paraboea</Emphasis><Emphasis Type="Italic">rufescens</Emphasis> under drought stress

Resurrection plants could survive severe drought stress, but the underlying mechanism for protecting their photosynthetic apparatus against drought stress is unclear. Cyclic electron flow (CEF) has been documented as a crucial mechanism for photoprotection in Arabidopsis and tobacco. We hypothesized that CEF plays an important role in protecting photosystem I (PSI) and photosystem II (PSII) against drought stress for resurrection plants. To address this hypothesis, the effects of mild drought stress on light energy distribution in PSII and P700 redox state were examined in a resurrection plant Paraboea rufescens. Cyclic electron flow was not activated below the photosynthetic photon flux density (PPFD) of 400 μmol m⁻² s⁻¹ in leaves without drought stress. However, CEF was activated under low light in leaves with mild drought stress, and the effective quantum yield of PSII significantly decreased. Meanwhile, non-photochemical quenching (NPQ) was significantly stimulated not only under high light but also under low light. Compared with the control, the fraction of overall P700 that cannot be oxidized in a given state (PSI acceptor side limitation) under high light was maintained at low level of 0.1 in leaves with water deficit, indicating that the over-reduction of the PSI acceptor side was prevented by the significant stimulation of CEF. Furthermore, methyl viologen could significantly increase the PSII photo-inhibition induced by high light compared with chloramphenicol. These results suggested that CEF is an important mechanism for protecting PSI and PSII from drought stress in resurrection plants.     

Reversible changes in structure and function of photosynthetic apparatus of pea (Pisum sativum) leaves under drought stress

The effects of drought on photosynthesis have been extensively studied, whereas those on thylakoid organization are limited. We observed a significant decline in gas exchange parameters of pea (Pisum sativum) leaves under progressive drought stress. Chl a fluorescence kinetics revealed the reduction of photochemical efficiency of photosystem (PS)II and PSI. The non-photochemical quenching (NPQ) and the levels of PSII subunit PSBS increased. Furthermore, the light-harvesting complexes (LHCs) and some of the PSI and PSII core proteins were disassembled in drought conditions, whereas these complexes were reassociated during recovery. By contrast, the abundance of supercomplexes of PSII-LHCII and PSII dimer were reduced, whereas LHCII monomers increased following the change in the macro-organization of thylakoids. The stacks of thylakoids were loosely arranged in drought-affected plants, which could be attributed to changes in the supercomplexes of thylakoids. Severe drought stress caused a reduction of both LHCI and LHCII and a few reaction center proteins of PSI and PSII, indicating significant disorganization of the photosynthetic machinery. After 7 days of rewatering, plants recovered well, with restored chloroplast thylakoid structure and photosynthetic efficiency. The correlation of structural changes with leaf reactive oxygen species levels indicated that these changes were associated with the production of reactive oxygen species.     

Adaptation potential of photosynthesis in wheat cultivars with a capability of leaf rolling under high temperature conditions

For the first time, the adaptive role of the rolling leaf trait for tolerance of wheat plants (Triticum aestivum L.) to the main factor of drought, high temperature, was demonstrated. Cv. Otan with high degree of this trait expression was more tolerant to temperature stress (40°C, 4 h during 2 days (2h/day)). Changes in parameters of chlorophyll fluorescence, F _v/F _m and R _Fd690, suggest that cv. Otan was tolerant to inhibition of photochemical activities of photosystem II (PSII) and photosystem I (PSI). Furthermore, high temperature had no effect on the rate of net photosynthesis (P _N) in cv. Otan, although it decreased this parameter in the other wheat cultivars. The main factors, which provid for this tolerance, were adaptation of the photosynthetic pigment system by active accumulation of carotenoids, more stable structural organization of PSII and PSI, and their high photosynthetic activities, as well as efficient stomatal regulation of transpiration and supplying of mesophyll cells with CO₂. It is hypothesized that the physiological role of the rolling leaf trait is the maintenance of adaptation potential by increasing the efficiency of water metabolism in the flag leaves of wheat under high temperature.     

Moderate water stress causes different stomatal and non‐stomatal changes in the photosynthetic functioning of Phaseolus vulgaris L. genotypes

The impact of moderate water deficit on the photosynthetic apparatus of three Phaseolus vulgaris L. cultivars, Plovdiv 10 (P10), Dobrudjanski Ran (DR) and Prelom (Prel), was investigated. Water shortage had less impact on leaf hydration, RWC (predawn and midday) and predawn water potential in Prel. RWC and Ψ_p were more reduced in P10, while there was no osmotic adjustment in any cultivar. Although drought drastically reduced stomatal opening in P10 and DR, reduced A_max indicated non‐stomatal limitations that contributed to the negligible P_n. These limitations were on potential thylakoid electron transport rates of PSI and II, pointing to photosystem functioning as a major limiting step in photosynthesis. This agrees with decreases in actual photochemical efficiency of PSII (F_v′/F_m′), quantum yield of photosynthetic non‐cyclic electron transport (?_e) and energy‐driven photochemical events (q_P), although the impact on these parameters would also include down‐regulation processes. When compared to DR, Prel retained a higher functional state of the photosynthetic machinery, justifying reduced need for photoprotective mechanisms (non‐photochemical quenching, zeaxanthin, lutein, β‐carotene) and maintenance of the balance between energy capture and dissipative pigments. The highest increases in fructose, glucose, arabinose and sorbitol in Prel might be related to tolerance to a lower oxidative state. All cultivars had reduced A_max due to daytime stomatal closure in well‐watered conditions. Under moderate drought, Prel had highest tolerance, higher leaf hydration and maintenance of important photochemical use of energy. However, water shortage caused appreciable non‐stomatal limitations to photosynthesis linked to regulation/imbalance at the metabolic level (and growth) in all cultivars. This included damage, as reflected in decreased potential photosystem functioning, pointing to higher sensitivity of photosynthesis to drought than is commonly assumed.     

Photosynthetic electron transport and specific photoprotective responses in wheat leaves under drought stress

The photosynthetic responses of wheat (Triticum aestivum L.) leaves to different levels of drought stress were analyzed in potted plants cultivated in growth chamber under moderate light. Low-to-medium drought stress was induced by limiting irrigation, maintaining 20 % of soil water holding capacity for 14 days followed by 3 days without water supply to induce severe stress. Measurements of CO₂ exchange and photosystem II (PSII) yield (by chlorophyll fluorescence) were followed by simultaneous measurements of yield of PSI (by P700 absorbance changes) and that of PSII. Drought stress gradually decreased PSII electron transport, but the capacity for nonphotochemical quenching increased more slowly until there was a large decrease in leaf relative water content (where the photosynthetic rate had decreased by half or more). We identified a substantial part of PSII electron transport, which was not used by carbon assimilation or by photorespiration, which clearly indicates activities of alternative electron sinks. Decreasing the fraction of light absorbed by PSII and increasing the fraction absorbed by PSI with increasing drought stress (rather than assuming equal absorption by the two photosystems) support a proposed function of PSI cyclic electron flow to generate a proton-motive force to activate nonphotochemical dissipation of energy, and it is consistent with the observed accumulation of oxidized P700 which causes a decrease in PSI electron acceptors. Our results support the roles of alternative electron sinks (either from PSII or PSI) and cyclic electron flow in photoprotection of PSII and PSI in drought stress conditions. In future studies on plant stress, analyses of the partitioning of absorbed energy between photosystems are needed for interpreting flux through linear electron flow, PSI cyclic electron flow, along with alternative electron sinks.     

Protective mechanism of desiccation tolerance in <Emphasis Type="Italic">Reaumuria soongorica</Emphasis>: Leaf abscission and sucrose accumulation in the stem

Reaumuria soongorica (Pall.) Maxim., a perennial semi-shrub, is widely found in semi-arid areas in northwestern China and can survive severe desiccation of its vegetative organs. In order to study the protective mechanism of desiccation tolerance in R. soongorica, diurnal patterns of net photosynthetic rate (Pn), water use efficiency (WUE) and chlorophyll fluorescence parameters of Photosystem II (PSII), and sugar content in the source leaf and stem were investigated in 6-year-old plants during progressive soil drought imposed by the cessation of watering. The results showed that R. soongorica was characterized by very low leaf water potential, high WUE, photosynthesis and high accumulation of sucrose in the stem and leaf abscission under desiccation. The maximum Pn increased at first and then declined during drought, but intrinsic WUE increased remarkably in the morning with increasing drought stress. The maximal photochemical efficiency of PSII (Fv/Fm) and the quantum efficiency of noncyclic electric transport of PSII(Φ_PSII) decreased significantly under water stress and exhibited an obvious phenomenon of photoinhibition at noon. Drought stressed plants maintained a higher capacity of dissipation of the excitation energy (measured as NPQ) with the increasing intensity of stress. Conditions of progressive drought promoted sucrose and starch accumulation in the stems but not in the leaves. However, when leaf water potential was less than −21.3 MPa, the plant leaves died and then abscised. But the stem photosynthesis remained and, afterward the plants entered the dormant state. Upon rewatering, the shoots reactivated and the plants developed new leaves. Therefore, R. soongorica has the ability to reduce water loss through leaf abscission and maintain the vigor of the stem cells to survive desiccation.     

Differential responses of photosystems I and II to seasonal drought in two Ficus species

Hemiepiphytic Ficus species exhibit more conservative water use strategy and are more drought-tolerant compared with their non-hemiepiphytic congeners, but a difference in the response of photosystem I (PSI) and photosystem II (PSII) to drought stress has not been documented to date. The enhancement of non-photochemical quenching (NPQ) and cyclic electron flow (CEF) have been identified as important mechanisms that protect the photosystems under drought conditions. Using the hemiepiphytic Ficus tinctoria and the non-hemiepiphytic Ficus racemosa, we studied the water status and the electron fluxes through PSI and PSII under seasonal water stress. Our results clearly indicated that the decline in the leaf predawn water potential (ψ_pd), the maximum photosynthetic rate (A_max) and the predawn maximum quantum yield of PSII (F_v/F_m) were more pronounced in F. racemosa than in F. tinctoria at peak drought. The F_v/F_m of F. racemosa was reduced to 0.69, indicating net photoinhibition of PSII. Concomitantly, the maximal photo-oxidizable P700 (P_m) decreased significantly in F. racemosa but remained stable in F. tinctoria. The fraction of non-photochemical quenching [Y(NPQ)] and the ratio of effective quantum yield of PSI to PSII [Y(I)/Y(II)] increased for both Ficus species at peak drought, with a stronger increase in F. racemosa. These results indicated that the enhancement of NPQ and the activation of CEF contributed to the photoprotection of PSI and PSII for both Ficus species under seasonal drought, particularly for F. racemosa.     

Effects of 5-aminolevulinic acid treatment on photosynthesis of strawberry

Effects of root treatment with 5-aminolevulinic acid (ALA) on leaf photosynthesis in strawberry (Fragaria ananassa Duch.) plants were investigated by rapid chlorophyll fluorescence and modulated 820 nm reflection using 3-(3,4-dichlorophenyl)-1,1-dimethyl urea (DCMU) and methyl viologen (MV). Our results showed that ALA treatments increased the net photosynthetic rate and decreased the intercelluar CO₂ concentration in strawberry leaves. Under DCMU treatment, trapping energy for Q_A reduction per PSII reaction center increased greatly, indicating DCMU inhibited electron transfer from Q_A ^?. The maximum photochemical efficiency of PSII (F_v/F_m) decreased under the DCMU treatment, while a higher F_v/F_m remained in the ALA-pretreated plants. Not only the parameters related to a photochemical phase, but also that one related to a heat phase remained lower after the ALA pretreatment, compared to the sole DCMU treatment. The MV treatment decreased PSI photochemical capacity. The results of modulated 820 nm reflection analysis showed that DCMU and MV treatments had low re-reduction of P700 and plastocyanin (PSI). However, the strawberry leaf discs pretreated with ALA exhibited high re-reduction of PSI under DCMU and MV treatments. The results of this study suggest that the improvement of photosynthesis by ALA in strawberry was not only related to PSII, but also to PSI and electron transfer chain.     

Physiological and anatomical changes induced by drought in two olive cultivars (cv Zalmati and Chemlali)

Photosynthetic gas exchange, vegetative growth, water relations and fluorescence parameters as well as leaf anatomical characteristics were investigated on young plants of two Olea europaea L. cultivars (Chemlali and Zalmati), submitted to contrasting water availability regimes. Two-year-old olive trees, grown in pots in greenhouse, were not watered for 2 months. Relative growth rate (RGR), leaf water potential (Ψ_LW) and the leaf relative water content (LWC) of the two cultivars decreased with increasing water stress. Zalmati showed higher values of RGR and LWC and lower decreased values of Ψ_LW than Chemlali, in response to water deficit, particularly during severe drought stress. Water stress also caused a marked decline on photosynthetic capacity and chlorophyll fluorescence. The net photosynthetic rate, stomatal conductance, transpiration rate, the maximal photochemical efficiency of PSII (F _v/F _m) and the intrinsic efficiency of open PSII reaction centres (F′ _v/F′ _m) decreased as drought stress developed. In addition, drought conditions, reduced leaf chlorophyll and carotenoids contents especially at severe water stress. However, Zalmati plants were the less affected when compared with Chemlali. In both cultivars, stomatal control was the major factor affecting photosynthesis under moderate drought stress. At severe drought-stress levels, the non-stomatal component of photosynthesis is inhibited and inactivation of the photosystem II occurs. Leaf anatomical parameters show that drought stress resulted in an increase of the upper epidermis and palisade mesophyll thickness as well as an increase of the stomata and trichomes density. These changes were more characteristic in cv. ‘Zalmati’. Zalmati leaves also revealed lower specific leaf area and had higher density of foliar tissue. From the behaviour of Zalmati plants, with a smaller reduction in relative growth rate, net assimilation rate and chlorophyll fluorescence parameters, and with a thicker palisade parenchyma, and a higher stomatal and trichome density, we consider this cultivar more drought-tolerant than cv. Chemlali and therefore, very promising for cultivation in arid areas.     

Interactive effects of photon fluence rates and drought on CAM‐cycling in Delosperma tradescantioides (Mesembryanthemaceae)

The ability of photosynthesis and CAM to acclimate to low (220 µmol m^?2 s^?1; LL) and relatively high (550 µmol m^?2 s^?1; HL) photosynthetic photon flux densities (PPFD) was investigated in the CAM-cycling species Delosperma tradescantioides by means of CO₂ gas exchange and chlorophyll fluorescence analysis. Furthermore, the influence of short-term drought on malic acid accumulation and the activity of photosystem II (PSII) was studied to assess the possible interactions between drought and the prevailing PPFD in this species. HL plants showed features of sun versus shade acclimation relative to LL plants. Nocturnal malic acid accumulation (Δ-malate) and leaf water content also tended to be higher in HL plants. Irrespective of the PPFD during growth, the weak Δ-malate doubled within 3 days of drought. Despite largely restricted CO₂ uptake, photosynthetic activity as estimated from fluorescence analysis declined only ca 5%. After 7 days of drought, when plants showed CAM-idling and Δ-malate had decreased again, potential carbon assimilation was still ca 84% of that in well-watered plants and remained relatively constant throughout the day. Decarboxylation of malic acid accounted for ca 23% of potential assimilation assuming total oxidation of a maximum portion of this organic acid. Drought did not affect predawn maximum photochemical efficiency (F_v/F_m). Nonphotochemical quenching (qN) increased (24%) in response to desiccation and resulted in a more or less constant reduction state of PSII. This increase in qN resulted mainly from the change in its fast-relaxing component (qNF), while the slow component (qNS) was significant only at or above saturating PPFD in both HL and LL plants. The photon response characteristics of PSII, which differed between LL and HL plants, were unaffected by short-term drought. Photon harvesting and photon use were always adjusted to guarantee a low reduction state of PSII. Results suggest that in both LL and HL plants CAM-cycling may help to stabilize photosynthesis but to a large extent by other means than simply providing internally derived CO₂.     

Evaluation of photosynthetic activities in thylakoid membranes by means of Fourier transform infrared spectroscopy

Light-induced Fourier transformed infrared (FTIR) difference spectroscopy is a powerful method to study the structures and reactions of redox cofactors involved in the photosynthetic electron transport chain. So far, most of the FTIR studies of the reactions of oxygenic photosynthesis have been performed using isolated photosystem I (PSI) and photosystem II (PSII) preparations, which, however, could be modified during isolation procedures. In this study, we developed a methodology to evaluate the photosynthetic activities of thylakoids using FTIR spectroscopy. FTIR difference spectra upon successive flashes using thylakoids from spinach exhibited signals typical of the S-state cycle at the Mn₄CaO₅ cluster and Q_B reactions in PSII with period-four and -two oscillations, respectively. Similar measurement in the presence of an artificial quinone as an exogenous electron acceptor showed features specific to the S-state cycle. Simulations of the oscillation patterns provided the quantum efficiencies of the S-state cycle and electron transfer in PSII. Moreover, FTIR measurement under continuous illumination on thylakoids in the presence of DCMU showed signals due to Q_A reduction and P700 oxidation simultaneously. From the relative amplitudes of marker bands of Q_A^? and P700⁺, the molar ratio of photoactive PSII and PSI centers in thylakoids was estimated. FTIR analyses of the photo-reactions in thylakoids, which are more intact than isolated photosystems, will be useful in investigations of the photosynthetic mechanism especially by genetic modification of photosystem proteins.     

Protective mechanism of desiccation tolerance in Reaumuria soongorica: Leaf abscission and sucrose accumulation in the stem

Reaumuria soongorica (Pall.) Maxim., a perennial semi-shrub, is widely found in semi-arid areas in northwestern China and can survive severe desiccation of its vegetative organs. In order to study the protective mechanism of desiccation tolerance in R. soongorica, diurnal patterns of net photosynthetic rate (Pn), water use efficiency (WUE) and chlorophyll fluorescence parameters of Photosystem II (PSII), and sugar content in the source leaf and stem were investigated in 6-year-old plants during progressive soil drought imposed by the cessation of watering. The results showed that R. soongorica was characterized by very low leaf water potential, high WUE, photosynthesis and high accumulation of sucrose in the stem and leaf abscission under desiccation. The maximum Pn increased at first and then declined during drought, but intrinsic WUE increased remarkably in the morning with increasing drought stress. The maximal photochemical efficiency of PSII (Fv/Fm) and the quantum efficiency of noncyclic electric transport of PSII(Φ_PSII) decreased significantly under water stress and exhibited an obvious phenomenon of photoinhibition at noon. Drought stressed plants maintained a higher capacity of dissipation of the excitation energy (measured as NPQ) with the increasing intensity of stress. Conditions of progressive drought promoted sucrose and starch accumulation in the stems but not in the leaves. However, when leaf water potential was less than −21.3 MPa, the plant leaves died and then abscised. But the stem photosynthesis remained and, afterward the plants entered the dormant state. Upon rewatering, the shoots reactivated and the plants developed new leaves. Therefore, R. soongorica has the ability to reduce water loss through leaf abscission and maintain the vigor of the stem cells to survive desiccation. Supported by the Program of the Research of Vegetation Restoration in Arid Areas of Lanzhou (Grant No. 03-2-27) and the National Natural Science Foundation of China (Grant No. 30270243)     

Inhibition of photosynthetic reactions under water stress: interaction with light level

When the shrub Nerium oleander L., growing under full natural daylight outdoors, was subjected to water stress, stomatal conductance declined, and so did non-stomatal components of photosynthesis, including the CO₂-saturated rate of CO₂ uptake by intact leaves and the activity of electron transport by chloroplasts isolated from stressed plants. This inactivation of photosynthetic activity was accompanied by changes in the fluorescence characteristics determined at 77 K (-196°C) for the upper leaf surface and from isolated chloroplasts. The maximum (F _M) and the variable (F _V) fluorescence yield at 692 nm were strongly quenched but there was little effect on the instantaneous (F _O) fluorescence. There was a concomitant quenching of the maximum and variable fluorescence at 734 nm. These results indicate an inactivation of the primary photochemistry associated with photosystem II. The lower, naturally shaded surfaces of the same leaves were much less affected than the upper surfaces and water-stress treatment of plants kept in deep shade had little or no effect on the fluorescence characteristics of either surface, or of chloroplasts isolated from the water-stressed leaves. The effects of subjecting N. oleander plants, growing in full daylight, to water stress are indistinguishable from those resulting when plants, grown under a lower light regime, are exposed to full daylight (photoinhibition). Both kinds of stress evidently cause an inactivation of the primary photochemistry associated with photosystem II. The results indicate that water stress predisposes the leaves to photoinhibition. Recovery from this inhibition, following restoration of favorable water relations, is very slow, indicating that photoinhibition is an important component of the damage to the photosynthetic system that takes place when plants are exposed to water stress in the field. The underlying causes of this water-stress-induced susceptibility to photoinhibition are unknown; stomatal closure or elevated leaf temperature cannot explain the increased susceptibility.Abbreviations and symbols Chl chlorophyll - PFD photon flux area density - PSI, PSII photosystem I, II - F _M, F _O, F _V maximum, instantaneous, variable fluorescence emission - leaf water potential C.I.W.-D.P.B. Publication No. 775     

Influence of variation potential on resistance of the photosynthetic machinery to heating in pea

Electrical signals [action potentials (APs) and variation potentials (VPs)] induced by local stimuli are a mechanism that underlies rapid plant response to environmental factors. Such signals induce a number of functional responses, including changes in photosynthesis. Ultimately, these responses are considered to increase plant resistance to stress factors, but this question has been poorly investigated. We studied the influence of VP on photosynthesis and resistance of the photosynthetic machinery to heating in leaves of pea (Pisum sativum). Localized burning induced a VP that decreased photosynthesis parameters [CO₂ assimilation rate and quantum yields of photosystem I (PSI) and photosystem II (PSII)]. The photosynthetic response was initiated by a decrease in photosynthesis dark‐stage activity, which in turn increased resistance of PSI to heating. Three results supported this hypothesized mechanism: (1) the magnitude of VP‐induced decrease in CO₂ assimilation and enhanced PSI resistance to heating were highly correlated; (2) the VP influence on PSI resistance to heating was suppressed under a low external CO₂ concentration and (3) decreasing external CO₂ concentration imitated the VP‐induced photosynthetic response and increased PSI resistance to heating.     

Photosynthetic CO<Subscript>2</Subscript>/H<Subscript>2</Subscript>O gas exchange and dynamics of carbohydrates content in maize leaves under drought

We studied the temporal sequence of changes in the photosynthetic CO₂/H₂O gas exchange intensity, as well as leaf water status, contents of soluble carbohydrates, starch, proline, pigments, and MDA, in maize seedlings (Zea mays L., cv. Luchistaya) under adaptation to increasing water deficit. The duration of drought was 2, 3, 5, and 6 days. Withholding water from maize plants caused gradual increase in the intensity of water deficit: from mild (2 or 3 days) to moderate (5 days) and nearly severe (6 days) water stress. After 6 days, relative leaf water content decreased by 19.8% as compared to the control. On the second day after the onset of drought, slight reduction in the photosynthetic CO₂/H₂O gas exchange intensity of the treated plants was observed. After 6 days, photosynthesis and transpiration of leaves synchronously reduced almost threefold due to stomatal closure. The progressive soil drought had substantial impact on the carbohydrate metabolism. After 2 days of water deficit, the content of reducing sugars and sucrose increased slightly, whereas after 6 days, it increased ten and four times, respectively. After 2, 3, and 5 days of drought, the starch content declined slightly; however, under severe drought (6 days), it increased by 30% as compared to the control. Simultaneously with the increase in the content of soluble sugars, proline content increased significantly and it was the highest on the sixth day of drought. At all stages of water deficit, the proline content increased more significantly than the content of reducing carbohydrates and sucrose. Under increasing water deficit (5 and 6 days), the content of MDA was found to rise. At the initial drought stage (2 or 3 days) and under severe water deficit (6 days), no significant changes in the pigment content were observed. Thus, at the initial stages of progressive drought, in the leaves of this maize cultivar, a decline in photosynthetic activity proceeded simultaneously with accumulation of reducing sugars, sucrose, and proline. The results obtained showed that, at the first stages of adaptation of maize seedlings to drought, the changes in carbohydrate and proline metabolism have been observed, which have increased upon further plant dehydration.     

After more than a decade of soil moisture deficit,tropical rainforest trees maintain photosynthetic capacity,despite increased leaf respiration

Determining climate change feedbacks from tropical rainforests requires an understanding of how carbon gain through photosynthesis and loss through respiration will be altered. One of the key changes that tropical rainforests may experience under future climate change scenarios is reduced soil moisture availability. In this study we examine if and how both leaf photosynthesis and leaf dark respiration acclimate following more than 12 years of experimental soil moisture deficit, via a through‐fall exclusion experiment (TFE) in an eastern Amazonian rainforest. We find that experimentally drought‐stressed trees and taxa maintain the same maximum leaf photosynthetic capacity as trees in corresponding control forest, independent of their susceptibility to drought‐induced mortality. We hypothesize that photosynthetic capacity is maintained across all treatments and taxa to take advantage of short‐lived periods of high moisture availability, when stomatal conductance (g_s) and photosynthesis can increase rapidly, potentially compensating for reduced assimilate supply at other times. Average leaf dark respiration (R_d) was elevated in the TFE‐treated forest trees relative to the control by 28.2 ± 2.8% (mean ± one standard error). This mean R_d value was dominated by a 48.5 ± 3.6% increase in the R_d of drought‐sensitive taxa, and likely reflects the need for additional metabolic support required for stress‐related repair, and hydraulic or osmotic maintenance processes. Following soil moisture deficit that is maintained for several years, our data suggest that changes in respiration drive greater shifts in the canopy carbon balance, than changes in photosynthetic capacity.     

Leaf <Emphasis Type="Italic">vs</Emphasis>. inflorescence: differences in photosynthetic activity of grapevine

Using measures of gas exchange and photosynthetic chain activity, we found some differences between grapevine inflorescence and leaf in terms of photosynthetic activity and photosynthesis regulations. Generally, the leaf showed the higher net photosynthesis (P _N) and lower dark respiration than that of the inflorescence until the beginning of the flowering process. The lower (and negative) P _N indicated prevailing respiration over photosynthesis and could result from a higher metabolic activity rather than from a lower activity of the photosynthetic apparatus. Considerable differences were observed between both organs in the functioning and regulation of PSI and PSII. Indeed, in our conditions, the quantum yield efficiency and electron transport rate of PSI and PSII were higher in the inflorescence compared to that of the leaf; nevertheless, protective regulatory mechanisms of the photosynthetic chain were clearly more efficient in the leaf. This was in accordance with the major function of this organ in grapevine, but it highlighted also that inflorescence seems to be implied in the whole carbon balance of plant. During inflorescence development, the global PSII activity decreased and PSI regulation tended to be similar to the leaf, where photosynthetic activity and regulations remained more stable. Finally, during flowering, cyclic electron flow (CEF) around PSI was activated in parallel to the decline in the thylakoid linear electron flow. Inflorescence CEF was double compared to the leaf; it might contribute to photoprotection, could promote ATP synthesis and the recovery of PSII.