Photosynthetic Acclimation of Erythrophleum guineense and Dalbergia odorifera to Winter Low Temperature in a Marginal Tropical Area

In marginal tropical areas, air temperature in winter usually decreases by 10℃ compared with summer at night/day. Although tropical plants are sensitive to low temperature, the mechanism underlying photosynthetic acclimation of tropical trees to winter low temperature is unclear. To address this question, the photosystem I (PSI) and photosystem II (PSII) activities, and energy distribution in PSI and PSII were examined in summer and winter in two tropical high quality timber tree species Erythrophleum guineense and Dalbergia odorifera grown in a marginal tropical area (21°54′N, 101°46′E). Our results indicated that the photosynthetic apparatus of Eguineense and Dodorifera was maintained stable in winter. The effective quantum yield of PSII decreased significantly in winter, but non photochemical quenching (NPQ) significantly increased. In winter, cyclic electron flow (CEF) was significantly stimulated in both species, which was significantly and positively correlated with NPQ. Meanwhile, the stimulation of CEF led to an increase in P700 oxidation ratio and the over reduction of PSI acceptor side was prevented. Antimycin A (a specific inhibitor of PGR5 dependent CEF) significantly aggravated PSII photoinhibition under high light in both species. These results suggested that stimulation of CEF is an important mechanism for photosynthetic acclimation to winter low temperature in a marginal tropical area in the two tropical tree species.     

几内亚格木和降香黄檀对热带北缘地区冬季低温的光合适应

在热带北缘地区,冬季气温较夏季下降10℃左右,虽然热带植物对零上低温敏感,但是大部分热带树木能够适应热带北缘地区的冬季气温,其光合生理机制并不清楚。我们通过测定种植在热带北缘地区（21°54′N,101°46′E）的两种热带树木（几内亚格木和降香黄檀）的光系统Ⅰ和Ⅱ活性以及光系统Ⅰ和Ⅱ的能量分配的季节变化,发现这两个树种的光系统Ⅰ和Ⅱ活性在冬季并没有下降。两个树种的光系统Ⅱ的有效量子产额在冬季明显下降,同时伴随着热耗散激发。在冬季,环式电子传递的激发与热耗散的激发呈现显著的正相关。环式电子传递的激发使得氧化态P700比例的上升,从而避免了光系统Ⅰ受体端的过度还原。化学试剂抗霉素A（PGR5途径环式电子传递的一种特异性抑制剂）处理过的叶片较对照组表现出更强光损伤程度。这些结果表明环式电子传递的激发是热带树木适应热带北缘地区冬季低温的一个重要的光合生理机制。     

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.     

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.     

The different effects of chilling stress under moderate light intensity on photosystem II compared with photosystem I and subsequent recovery in tropical tree species

Tropical plants are sensitive to chilling temperatures above zero but it is still unclear whether photosystem I (PSI) or photosystem II (PSII) of tropical plants is mainly affected by chilling temperatures. In this study, the effect of 4°C associated with various light densities on PSII and PSI was studied in the potted seedlings of four tropical evergreen tree species grown in an open field, Khaya ivorensis, Pometia tomentosa, Dalbergia odorifera, and Erythrophleum guineense. After 8 h chilling exposure at the different photosynthetic flux densities of 20, 50, 100, 150 μmol m⁻² s⁻¹, the maximum quantum yield of PSII (F _v /F _m) in all of the four species decreased little, while the quantity of efficient PSI complex (P _m) remained stable in all species except E. guineense. However, after chilling exposure under 250 μmol m⁻² s⁻¹ for 24 h, F _v /F _m was severely photoinhibited in all species whereas P _m was relative stable in all plants except E. guineense. At the chilling temperature of 4°C, electron transport from PSII to PSI was blocked because of excessive reduction of primary electron acceptor of PSII. F _v /F _m in these species except E. guineense recovered to ~90% after 8 h recovery in low light, suggesting the dependence of the recovery of PSII on moderate PSI and/or PSII activity. These results suggest that PSII is more sensitive to chilling temperature under the moderate light than PSI in tropical trees, and the photoinhibition of PSII and closure of PSII reaction centers can serve to protect PSI.     

Different responses of photosystem I and photosystem II in three tropical oilseed crops exposed to chilling stress and subsequent recovery

Key message

Different responses of photosystem I and II to chilling.

Abstract

Tropical crops are sensitive to chilling stress, but the underlying physiological mechanisms are unclear. We investigated the maximum quantum yield of PSII (F _v/F _m), the maximum photo-oxidizable P700 (P _m), the energy distribution in PSII, and the redox state of P700 in leaves of seedlings of three promising oilseed crops originating from tropical regions, Plukenetia volubilis, Jatropha curcas and Ricinus communis, during chilling treatment and subsequent recovery under a photon flux density of 450 μmol m^?2 s^?1. Our results showed that F _v/F _m decreased progressively and significantly to about 44.7, 62.2 and 77.0 % of the control after chilling treatment for 3 days in P. volubilis, J. curcas and R. communis, respectively, mainly due to the decrease in F _m (maximum fluorescence of PSII). After recovery under 18 °C for 6 days, F _v/F _m recovered to 81.4 and 94.9 % of the control in J. curcas and R. communis, but only to 26.3 % in P. volubilis. Under chilling stress and subsequent recovery, P _m remained stable in J. curcas and R. communis, whereas it decreased slightly in P. volubilis. These results indicated that PSII was more sensitive to chilling stress than PSI under moderate light for all three species, and that P. volubilis was the most susceptible. Cyclic electron flow around PSI and effective quantum yield of photosystem II [Y _(CEF)/Y _(II)] ratio were stimulated much more in J. curcas and R. communis compared with that in P. volubilis under chilling conditions, resulting in more severe injury as indicated by higher accumulation of hydrogen peroxide and malondialdehyde. There was a significantly negative relationship between F _v/F _m and Y _(CEF)/Y _(II), suggesting that stimulation of Y _(CEF)/Y _(II) plays a pivotal role in protecting PSI and PSII from photoinhibition caused by chilling stress.     

Exogenous Calcium Alleviates Low Night Temperature Stress on the Photosynthetic Apparatus of Tomato Leaves

The effect of exogenous CaCl₂ on photosystem I and II (PSI and PSII) activities, cyclic electron flow (CEF), and proton motive force of tomato leaves under low night temperature (LNT) was investigated. LNT stress decreased the net photosynthetic rate (Pn), effective quantum yield of PSII [Y(II)], and photochemical quenching (qP), whereas CaCl₂ pretreatment improved Pn, Y(II), and qP under LNT stress. LNT stress significantly increased the non-regulatory quantum yield of energy dissipation [Y(NO)], whereas CaCl₂ alleviated this increase. Exogenous Ca²⁺ enhanced stimulation of CEF by LNT stress. Inhibition of oxidized PQ pools caused by LNT stress was alleviated by CaCl₂ pretreatment. LNT stress reduced zeaxanthin formation and ATPase activity, but CaCl₂ pretreatment reversed both of these effects. LNT stress caused excess formation of a proton gradient across the thylakoid membrane, whereas CaCl₂ pretreatment decreased the said factor under LNT. Thus, our results showed that photoinhibition of LNT-stressed plants could be alleviated by CaCl₂ pretreatment. Our findings further revealed that this alleviation was mediated in part by improvements in carbon fixation capacity, PQ pools, linear and cyclic electron transports, xanthophyll cycles, and ATPase activity.     

Differences in the stimulation of cyclic electron flow in two tropical ferns under water stress are related to leaf anatomy

Cyclic electron flow (CEF) plays an important role in photoprotection for angiosperms under environmental stresses. However, ferns are more sensitive to drought and their water transport systems are not as efficient as those of angiosperms, it is unclear whether CEF also contributes to photoprotection in these plants. Using Microsorum punctatum and Paraleptochillus decurrens, we studied the electron fluxes through both photosystem I (PSI) and photosystem II (PSII) under water stress and their leaf anatomies. Our goal was to determine if CEF functions in the photoprotection of these ferns and, if so, whether CEF stimulation is related to leaf anatomy. Compared with P. decurrens, M. punctatum had thicker leaves and cuticles and higher water storage capacity, but lower stomatal density and slower rate of water loss. During induced drought, the decrease in leaf water potential (Ψ_leaf) was more pronounced in P. decurrens than in M. punctatum. For both species, the decline in Ψ_leaf was associated with a lower effective PSII quantum yield, photochemical quantum yield of PSI and electron transport rate (ETR), whereas increases were found in the quantum yield of regulated energy dissipation, CEF and CEF/ETR(II) ratio. Values for CEF and the CEF/ETR(II) ratio peaked in M. punctatum at a light intensity of 500–600 µmol m^?2 s^?1 vs only 150–200 µmol m^?2 s^?1 in P. decurrens. Therefore, our results indicate that the stimulation of CEF in tropical ferns contributes to their photoprotection under water stress, and is related to their respective drought tolerance and leaf anatomy.     

低温弱光胁迫对野生大豆和大豆栽培种光系统功能的影响

以野生大豆和栽培大豆为材料,通过同时测定大豆叶片的叶绿素荧光快速诱导动力学曲线和对820nm光的吸收曲线,以及测定超氧化物歧化酶（SOD）和抗坏血酸过氧化物酶（APX）的活性,分析了低温弱光胁迫及常温弱光恢复下这2种大豆光系统Ⅱ（PSⅡ）和光系统Ⅰ（PSI）功能的变化。结果表明,低温弱光胁迫对这2种大豆的PSI和PSⅡ的功能都造成伤害;在低温弱光胁迫下,维持较高的SOD和APX活性和维持PSI和PSⅡ的协调性是野生大豆比栽培大豆耐低温的一个重要原因。     

Cyclic electron flow plays an important role in protection of spinach leaves under high temperature stress

Heat stress is one of the major abiotic stresses and affects plant productivity in a negative manner. Photosynthetic processes are largely influenced by heat stress. In spinach (Spinacia oleracea L.) leaves at 40°C the decrease in PSII activity was mainly due to the decreased efficiency to capture excitation energy, increased yield of regulatory energy dissipation mechanism Y(NPQ), and decreased quantum yield Y(II). According to the results below 45°C PSI is stable and protected while at a higher temperature stability of PSI was reduced and protection was not sufficient. Therefore, we conclude that cyclic electron flow plays an important role in protecting PSI from heat stress.     

The Arabidopsis szl1 Mutant Reveals a Critical Role of β-Carotene in Photosystem I Photoprotection

Carotenes and their oxygenated derivatives, the xanthophylls, are structural determinants in both photosystems (PS) I and II. They bind and stabilize photosynthetic complexes, increase the light-harvesting capacity of chlorophyll-binding proteins, and have a major role in chloroplast photoprotection. Localization of carotenoid species within each PS is highly conserved: Core complexes bind carotenes, whereas peripheral light-harvesting systems bind xanthophylls. The specific functional role of each xanthophyll species has been recently described by genetic dissection, however the in vivo role of carotenes has not been similarly defined. Here, we have analyzed the function of carotenes in photosynthesis and photoprotection, distinct from that of xanthophylls, by characterizing the suppressor of zeaxanthin-less (szl) mutant of Arabidopsis (Arabidopsis thaliana) which, due to the decreased activity of the lycopene-β-cyclase, shows a lower carotene content than wild-type plants. When grown at room temperature, mutant plants showed a lower content in PSI light-harvesting complex I complex than the wild type, and a reduced capacity for chlorophyll fluorescence quenching, the rapidly reversible component of nonphotochemical quenching. When exposed to high light at chilling temperature, szl1 plants showed stronger photoxidation than wild-type plants. Both PSI and PSII from szl1 were similarly depleted in carotenes and yet PSI activity was more sensitive to light stress than PSII as shown by the stronger photoinhibition of PSI and increased rate of singlet oxygen release from isolated PSI light-harvesting complex I complexes of szl1 compared with the wild type. We conclude that carotene depletion in the core complexes impairs photoprotection of both PS under high light at chilling temperature, with PSI being far more affected than PSII.     

The difficulty of estimating the electron transport rate at photosystem I

It is still a controversial issue how the electron transport reaction is carried out around photosystem I (PSI) in the photosynthetic electron transport chain. The measurable component in PSI is the oxidized P700, the reaction center chlorophyll in PSI, as the absorbance changes at 820–830 nm. Previously, the quantum yield at PSI [Y(I)] has been estimated as the existence probability of the photo-oxidizable P700 by applying the saturated-pulse illumination (SP; 10,000–20,000 µmol photons m^?2 s^?1). The electron transport rate (ETR) at PSI has been estimated from the Y(I) value, which was larger than the reaction rate at PSII, evaluated as the quantum yield of PSII, especially under stress-conditions such as CO₂-limited and high light intensity conditions. Therefore, it has been considered that the extra electron flow at PSI was enhanced at the stress condition and played an important role in dealing with the excessive light energy. However, some pieces of evidence were reported that the excessive electron flow at PSI would be ignorable from other aspects. In the present research, we confirmed that the Y(I) value estimated by the SP method could be easily misestimated by the limitation of the electron donation to PSI. Moreover, we estimated the quantitative turnover rate of P700⁺ by the light-to-dark transition. However, the turnover rate of P700 was much slower than the ETR at PSII. It is still hard to quantitatively estimate the ETR at PSI by the current techniques.

     

Low temperature and high light dependent dynamic photoprotective strategies in Arabidopsis thaliana

Arabidopsis thaliana has been recognized as a chilling tolerant species based on analysis of resistance to low temperature stress, however, the mechanisms involved in this tolerance are not yet clarified. The low temperature-induced effects are exacerbated when plants are exposed to low temperatures in the presence of high light irradiance but the experimental data on the impact of light intensity during cold stress and its influence during recovery from stress are rather limited. The main objective of this study was to re-examine the photosynthetic responses of A. thaliana plants to short term (6 days) low temperature stress (12/10°C) under optimal (150 μmol m⁻² s⁻¹) and high light (500 μmol m⁻² s⁻¹) intensity and the subsequent recovery from the stress. Simultaneous measurements of the in vivo and in vitro functional performance of both photosystem II (PSII) and photosystem I (PSI), as well as, net photosynthesis, low temperature (77 K) chlorophyll fluorescence and immunoblot analysis of the relative abundance of PSII and PSI reaction center proteins were used to evaluate the role of light in the development of possible protective mechanisms during low temperature stress and the consequent recovery from exposure to low temperature and different light intensities. The results presented clearly suggest that Arabidopsis plants can employ a number of highly dynamic photoprotective strategies depending on the light intensity. These strategies include one based on LHCII quenching and two other quenching mechanisms localized within the PSII and PSI reaction centers, which are all expressed to different extent depending on the severity of the photoinhibitory treatments under low temperature stress conditions.     

Inhibition of photosystems I and II and enhanced back flow of photosystem I electrons in cucumber leaf discs chilled in the light

Pre-illumination of cucumber leaf discs at a chilling temperature in low-irradiance white light resulted in accelerated re-reduction of P700(+) [the special Chl pair in the photosystem I (PSI) reaction centre] when the far-red measuring light was turned off. Measurements (in +/- methyl viologen or +/- DCMU conditions) of the re-reduction half time suggest that accelerated re-reduction of P700(+) appeared to be predominantly due to charge recombination and only partly due to reductants sustained by previous cyclic electron flow around PSI. Apparently, charge recombination in PSI was greatly enhanced by inhibition of forward, linear electron flow. Inhibition of PSII electron transport was observed to occur to a lesser extent than that of PSI, but only if the measurement of PSII functionality was free from complications due to downstream accumulation of electrons in pools. We suggest that promotion of controlled charge recombination and cyclic electron flow round PSI during chilling of leaves in the light may partly prevent further damage to both photosystems.     

The chilling injury induced by high root temperature in the leaves of rice seedlings

Root temperature is found to be a very important factor forleaves to alter the response and susceptibility to chillingstress. Severe visible damage was observed in the most activeleaves of seedlings of a japonica rice (Oryza sativa cv. Akitakomachi),e.g. the third leaf at the third-leaf stage, after the treatmentwhere only leaves but not roots were chilled (L/H). On the otherhand, no visible damage was observed after the treatment whereboth leaves and roots were chilled simultaneously (L/L). Thechilling injury induced by L/H, a novel type of chilling injury,required the light either during or after the chilling in orderto develop the visible symptoms such as leaf bleaching and tissuenecrosis. Chlorophyll fluorescence parameters measured aftervarious lengths of chilling treatments showed that significantchanges were induced before the visible injury. The effectivequantum yield and photochemical quenching of PSII dropped dramaticallywithin 24 h in both the presence and absence of a 12 h lightperiod. The maximal quantum yield and non-photochemical quenchingof PSII decreased significantly only in the presence of light.On the other hand, L/H chilling did not affect the functionof PSI, but caused a significant decrease in the electron availabilityfor PSI. These results suggest that the leaf chilling with highroot temperature destroys some component between PSII and PSIwithout the aid of light, which causes the over-reduction ofPSII in the light, and thereby the visible injury is inducedonly in the light.     

Low and High Temperature Limits to PSII : A Survey Using trans-Parinaric Acid, Delayed Light Emission, and F(o) Chlorophyll Fluorescence

Many studies have shown that membrane lipids of chilling-sensitive plants begin lateral phase separation (i.e. a minor component begins freezing) at chilling temperatures and that chilling-sensitive plants are often of tropical origin. We tested the hypothesis that membranes of tropical plants begin lateral phase separation at chilling temperatures, and that plants lower the temperature of lateral phase separation as they invade cooler habitats. To do so we studied plant species in one family confined to the tropics (Piperaceae) and in three families with both tropical and temperate representatives (Fabaceae [Leguminosae], Malvaceae, and Solanaceae). We determined lateral phase separation temperatures by measuring the temperature dependence of fluorescence from trans-parinaric acid inserted into liposomes prepared from isolated membrane phospholipids. In all families we detected lateral phase separations at significantly higher temperatures, on average, in species of tropical origin. To test for associated physiological effects we measured the temperature dependence of delayed light emission (DLE) by discs cut from the same leaves used for lipid analysis. We found that the temperature of maximum DLE upon chilling was strongly correlated with lateral phase separation temperatures, but was on average approximately 4°C lower. We also tested the hypothesis that photosystem II (PSII) (the most thermolabile component of photosynthesis) of tropical plants tolerates higher temperatures than PSII of temperate plants, using DLE and F_o chlorophyll fluorescence upon heating to measure the temperature at which PSII thermally denatured. We found little difference between the two groups in PSII denaturation temperature. We also found that the temperature of maximum DLA upon heating was not significantly different from the critical temperature for F_o fluorescence. Our results indicate that plants lowered their membrane freezing temperatures as they radiated from their tropical origins. One interpretation is that the tendency for membranes to begin freezing at chilling temperatures is the primitive condition, which plants corrected as they invaded colder habitats. An alternative is that membranes which freeze at temperatures only slightly lower than the minimum growth temperature confer an advantage.     

A complex containing PGRL1 and PGR5 is involved in the switch between linear and cyclic electron flow in Arabidopsis

During photosynthesis, two photoreaction centers located in the thylakoid membranes of the chloroplast, photosystems I and II (PSI and PSII), use light energy to mobilize electrons to generate ATP and NADPH. Different modes of electron flow exist, of which the linear electron flow is driven by PSI and PSII, generating ATP and NADPH, whereas the cyclic electron flow (CEF) only generates ATP and is driven by the PSI alone. Different environmental and metabolic conditions require the adjustment of ATP/NADPH ratios and a switch of electron distribution between the two photosystems. With the exception of PGR5, other components facilitating CEF are unknown. Here, we report the identification of PGRL1, a transmembrane protein present in thylakoids of Arabidopsis thaliana. Plants lacking PGRL1 show perturbation of CEF, similar to PGR5-deficient plants. We find that PGRL1 and PGR5 interact physically and associate with PSI. We therefore propose that the PGRL1-PGR5 complex facilitates CEF in eukaryotes.     

Impairment of the photosynthetic apparatus by oxidative stress induced by photosensitization reaction of protoporphyrin IX

Treatment with the herbicide acifluorfen-sodium (AF-Na), an inhibitor of protoporphyrinogen oxidase, caused an accumulation of protoporphyrin IX (Proto IX) , light-induced necrotic spots on the cucumber cotyledon within 12-24 h, and photobleaching after 48-72 h of light exposure. Proto IX-sensitized and singlet oxygen ((1)O(2))-mediated oxidative stress caused by AF-Na treatment impaired photosystem I (PSI), photosystem II (PSII) and whole chain electron transport reactions. As compared to controls, the F(v)/F(m) (variable to maximal chlorophyll a fluorescence) ratio of treated samples was reduced. The PSII electron donor NH(2)OH failed to restore the F(v)/F(m) ratio suggesting that the reduction of F(v)/F(m) reflects the loss of reaction center functions. This explanation is further supported by the practically near-similar loss of PSI and PSII activities. As revealed from the light saturation curve (rate of oxygen evolution as a function of light intensity), the reduction of PSII activity was both due to the reduction in the quantum yield at limiting light intensities and impairment of light-saturated electron transport. In treated cotyledons both the Q (due to recombination of Q(A)(-) with S(2)) and B (due to recombination of Q(B)(-) with S(2)/S(3)) band of thermoluminescence decreased by 50% suggesting a loss of active PSII reaction centers. In both the control and treated samples, the thermoluminescence yield of B band exhibited a periodicity of 4 suggesting normal functioning of the S states in centers that were still active. The low temperature (77 K) fluorescence emission spectra revealed that the F(695) band (that originates in CP-47) increased probably due to reduced energy transfer from the CP47 to the reaction center. These demonstrated an overall damage to the PSI and PSII reaction centers by (1)O(2) produced in response to photosensitization reaction of protoporphyrin IX in AF-Na-treated cucumber seedlings.     

CO2 response of cyclic electron flow around PSI (CEF-PSI) in tobacco leaves--relative electron fluxes through PSI and PSII determine the magnitude of non-photochemical quenching (NPQ) of Chl fluorescence

We hypothesized that cyclic electron flow around photosystem I (CEF-PSI) participates in the induction of non-photochemical quenching (NPQ) of chlorophyll (Chl) fluorescence when the rate of photosynthetic linear electron flow (LEF) is electron-acceptor limited. To test this hypothesis, the relationships among photosynthesis rate, electron fluxes through both PSI and PSII [Je(PSI) and Je(PSII)] and Chl fluorescence parameters were analyzed simultaneously in intact leaves of tobacco plants at several light intensities and partial pressures of ambient CO2 (Ca). At low light intensities, decreasing Ca lowered the photosynthesis rate, but Je(PSI) and Je(PSII) remained constant. Je(PSI) was larger than Je(PSII), indicating the existence of CEF-PSI. Increasing the light intensity enhanced photosynthesis and both Je(PSI) and Je (PSII). Je(PSI)/Je(PSII) also increased at high light and at high light and low Ca combined, showing a strong, positive relationship with NPQ of Chl fluorescence. These results indicated that CEF-PSI contributed to the dissipation of photon energy in excess of that consumed by photosynthesis by driving NPQ of Chl fluorescence. The main physiological function of CEF-PSI in photosynthesis of higher plants is discussed.     

The function of chloroplastic NAD(P)H dehydrogenase in tobacco during chilling stress under low irradiance

The function of chloroplastic NAD(P)H dehydrogenase (NDH) was examined by comparing a tobacco transformant (DeltandhB) in which the ndhB gene had been disrupted with its wild type, upon exposure to chilling temperature (4 degrees C) under low irradiance (100 micro mol m(-2) s(-1) PFD). During the chilling stress, the maximum photochemical efficiency of PSII (F(v)/F(m)) decreased markedly in both the wild type and DeltandhB. However, both F(v)/F(m) and P700(+), as well as the PSII-driven electron transport rate (ETR), in DeltandhB were lower than that in the wild type, implying that NDH-dependent cyclic electron flow around PSI functioned to protect the photosynthetic apparatus from chilling stress under low irradiance. Under the stress, non-photochemical quenching (NPQ), particularly the fast relaxing NPQ component (qf) and the de-epoxidized ratio of the xanthophyll cycle pigments, (A+Z)/(V+A+Z), were distinguishable in DeltandhB from those in the wild type. The lower NPQ in DeltandhB might be related to an inefficient proton gradient across thylakoid membranes (DeltapH) because of lacking an NDH-dependent cyclic electron flow around PSI at chilling temperature under low irradiance.