The Relationship Among Photoinhibition, Photooxidation and Early Aging at a Late Development Stage in Different Varieties with High Yield

Using six high yield varieties from different ecological districts in China the parameters of Chl fluorescence, the performance of membrane lipid peroxidation and the reduction of Chl content in flag leaf under natural condition at the later developmental stages (from heading stage to mature stage). The results showed that Fv/Fm , decreased gradually, the excessive light energy led to the accumulation of active oxygen O2-, H2O2 and the product of membrane-lipid peroxidation, MDA, which resulted in the reduction of Chl content and early aging due to photooxidation during the course of senescence of flag leaf. This phenomenon varied obviously in rice varieties. When comparina japonica tolerant to photooxidation, Fv/Fm in indica shanyou 63 susceptible to photoxidation decreased significantly. An increase of active oxygen and a sharp drop of Chl content, resulted in 'yellowish' early aging and influenced filling and setting of rice grain. The mechanism on early aging in indica was related to light and temperature conditions in filling stages. On a fine day (above 251), PS Ⅱ reaction center exhibited a dynamic change on revisable inactivation. Under strong midday light, PS Ⅱ function in indica exhibited obvious down-regulation and photoinhibition; Under strong light with low temperature, PS Ⅱ resulted in photodamage, showing early aging, which were related to the degradation of PSⅡ - D1 Protein and the inhibition of the endogenous protecting system such as Xanthophyll cycle and the enzymes of scavenging active oxygen. The results suggested that for a view of high-yield breeding, on the basis of a good type-plant, giving consideration to the utilization of heterosis and resistance to early aging, selecting japonica or a sterile line with japonice genotype as maternal was a breeding strategy worthy of being paid more attention.

Wheat PROTON GRADIENT REGULATION 5 is Involved in Tolerance to Photoinhibition

Wheat （Triticum aestivum L.） often experiences photoinhibition due to strong light during the grain filling stage. As such, increasing the tolerance of wheat to photoinhibition is very desirable in breeding efforts focused on increasing grain yields. Previous reports have suggested that PROTON GRADIENT REGULATION 5 （PGR5） plays a central role in the generation of a proton gradient across the thylakoid membrane （ApH） and in acclimation to high light intensity conditions. Three PGR5 homoeologues were isolated from wheat, and mapped onto chromosomes 7A, 7B and 7D, respectively. The TaPGR5s shared highly similar genomic sequences and gene structures. The transcripts of TaPGR5s were found to be abundantly expressed in the flag leaves, and were transiently up-regulated by treatment with high light. High light treatment inhibited the net photosynthetic rate （Pn） and the maximal quantum yield ofphotosystem II （Fv/Fm）. Further, these inhibitions were more evident in the leaves with reduced expression of TaPGR5s achieved using virus-induced gene silencing methods. Moreover, reducing TaPGR5 expression impaired the induction of non-photochemical quenching （NPQ）, which caused more severe cell membrane damage and lipid peroxidation in high light. Additionally, we observed that TaPGR5s transcripts were more abundantly expressed in the wheat genotypes with higher ms-delayed light emission （ms-DLE）, a value reflecting transthylakoid ApH. These results suggested that TaPGR5s play important roles in the tolerance of wheat to photoinhibition.

Preliminary study on the molecular mechanism of photoinhibition of rice photosynthesis

Photoinhibition occurs when crops grow under strong light and are simultaneously subjected to stresses such as high or low temperature, drought,etc. It causes a decrease of crop photosynthetic efficiency leading to an obvious decrease of the yield.

Characterization of photosynthesis,photoinhibition and the activities of C_4 pathway enzymes in a superhigh-yield rice,Liangyoupeijiu

Characteristics of photosynthetic gas exchange, photoinhibition and C4 pathway enzyme activities in both flag leaves and lemma were compared between a superhigh-yield rice (Oryza sativa L.) hybrid, Liangyoupeijiu and a traditional rice hybrid, Shanyou63. Liangyoupeijiu had a similar light saturated assimilation rate (Asat) to Shanyou63, but a much higher apparent quantum yield (AQY), carboxylation efficiency (CE) and quantum yield of CO2 fixation (φCO2). Liangyoupeijiu also showed a higher resistance to photoinhibition and higher non-radiative energy dissipation associated with the xanthophyll cycle than Shanyou63 when subjected to strong light. In addition, Liangyoupeijiu had higher activities of the C4 pathway enzymes in both flag leaves and lemmas than Shanyou63. These results indicate that higher light and CO2 use efficiency, higher resistance to photoinhibition and C4 pathway in both flag leaf and lemma may contribute to the higher yield of the superhigh-yield rice hybrid, Liangyoupeijiu.

Relationships between Phosphatidylglycerol Molecular Species of Thylakoid Membrane Lipids and Sensitivities to Chilling-induced Photoinhibition in Rice

In an attempt to explore the relationships between phosphatidylglycerol （PG） molecular species of thylakoid membrane lipids and sensitivities to chilling-induced photoinhibition, PG molecular species, D1 protein, electron transport activities of thylakoid membrane and the potential quantum yield （FvlFm） in rice treated under middle and low photon flux density （PFD） at 11℃ were analyzed by high performance liquid chromatography, enzyme hydrolysis, gas phase chromatography （GC） and so on. Results showed that the major molecular species of PGs in rice thylakoid membrane were 18：3/16：0, 18：3/16：1（3t）, 18：2/16：0, 18：2/16：1（3t）, 18：1/16：0, 18：1/16：1（3t）, 16：0/16：0, 16：0/16：1（3t）. There were large differences in the contents of unsaturated PG molecular species such as 18：1-3/16：0-16：1（3t） and saturated PG molecular species like 16：0/16：0-16：1（3t） among japonica cv 9516 0-9516）, japonica-indica hybrid F1 j-9516/i-SY63 （ji-95SY） and indica cv Shanyou 63 （i-SY63）. J-9516 containing higher contents of unsaturated PG molecular species was manifest in stable D1 protein contents under chill and tolerant to chill-induced photoinhibition. In contrast to j-9516, i-SY63 with lower contents of unsaturated PG molecular species, exhibited unstable D1 protein contents under chill and was sensitive to chill-induced photoinhibition, ji-95SY containing middle contents of unsaturated PG molecular species between those of j-9516 and i-SY63, exhibited mid extent of sensitivity to chill-induced photoinhibition. The losses in D1 protein also account for the inhibition in electron transport activity of thylakoid membrane and the observed decline in FvlFm. The PG molecular species that is efficient in raising chilling-resistant capacity were those containing unsaturated fatty acids, namely, unsaturated PG molecular species. These results implied that the substrate selectivity of the glycerol-3-phosphate acyltransferase in chloroplasts towards 16：0 or 18：1 displayed greatly the difference between japonica and indica rice. Itwas possible to enhance the capacity of resistance to chilling-induced photoinhibition by improving or modifying the GPAT gene.

Relation between photorespiration and photoinhibitionin cotton leaves

Photorespiration is a well-known CO2-evolution process accompanying photosyntheticCO2 uptake, whereas its physiological significance is still not well understood. As alreadyreported, severe photoinhibition occurred in the leaves or chloroplasts exposed to stronglight in the absence of CO2 upon suppression of photorespiration, while photoinhibitioncould be prevented by the supply of CO2 at a concentration near the compensation point.

Relationship between state transition and photoinhibition of photosynthesis in wheat leaves

IT is well known that state transitions can keep even distribution of excitation energy betweenthe two photosystems (PS Ⅱ and PS Ⅰ), favoring efficient operation of the photosyntheticapparatus. However, state transitions are usually thought to be associated with low light con-ditions. Its physiological significance under high light conditions is still a controversial is-suc. Many people think that state transitions occur and take significant effect only under

Effect of calcium ions on the secondary structures of photosystemⅡ and its relations with photoinhibition

Calcium ions play an important role in the oxygen_evolving process of photosystem Ⅱ as demonstrated in many experiments. The changes of the secondary structures of PS Ⅱ induced by the depletion of Ca 2+ were reported. The results indicated that the removal of Ca 2+ led to the transition of α helix to turns and sheet structures. While Ca 2+ was re_added to the media, only the structures changed to turns could be recovered. The protein conformational changes of PS Ⅱ during the donor side photoinhibition induced by the depletion of Ca 2+ were also studied. This showed that the protein conformational changes differed between the control and Ca 2+ _depleted samples in a short period of illumination (within 10 min). However, the changes became similar when the illumination time was increased.

Effect of Mn cluster on the formation of superoxide radicals in photoinhibition of photosystem Ⅱ

To further realize the action of superoxide radicals (O2 ) in photoinhibition of photosystem II (PS II), we employed 5,5-dimethyl-l-pyrroline-N-oxide (DMPO) as a spin trap, associated with EPK spectroscopy, to study theeffect of illumination time on O2- formation during highlight photoinhibition in PS II membranes and Mn-depleted PS II membranes. Results indicated that the removal of Mn cluster from PS II membranes has a strong influence on the dynamics of superoxide formation. The relative mechanismwas also discussed. These novel findings may further promote the studies of the structure and function of PS II and the mechanism of photoinhibition.

Alleviatory effect of rare earth micro-fertilizer on photosystemⅡ(PSⅡ)photoinhibition in Pseudostellaria heterophylla leaves at photosynthetic midday depression

In order to explo re the mitigatory mechanism of rare earths on the midday depression of photosynthesis,this study used fast-phase chlorophyll fluorescence technology combined with an antioxidant enzyme system to analyze the effects of lanthanum and cerium on the photosystemⅡ(PSⅡ)function of Pseudostellaria heterophylla leaves at noon.The results show that the maximum photochemical efficiency(F_(v)/F_(m))and the photosynthetic performance index(PI_(ABS))of P.heterophylla leaves during the photo synthetic noon-break are only 0.73 and 0.86,respectively.The leaves of P.heterophylla show obvious photoinhibition.The application of two rare earth fertilizers promotes the light energy absorption,capture,conversion,and electron transfer efficiencies of PSⅡin the P.heterophylla leaves and alleviates the photoinhibition caused by excess excitation energy.Additionally,the effect of lanthanum is better than that of cerium.In the fertilization groups,P.heterophylla leaves show higher PSⅡphotochemical activity.Fertilization protects the oxygen-evolving complex(OEC)activity of P.heterophylla,thus reducing oxidative damage to the chloroplasts and preventing the dissociation of the thylakoids to maintain the normal physiological functioning of PSⅡ,thereby reducing the degree of photoinhibition.Additionally,fertilization increases the electron transfer ability from Q_(A) to Q_(B) on the PSⅡelectron receptor side of P.heterophylla.Correlation analysis shows that the fluorescence parameters V_(L),V_(K),and V_(J) are significantly negatively correlated with antioxidant enzyme activity and are significantly positively correlated with malondialdehyde content,relative electric conductivity and reactive oxygen species(ROS),which further indicates that rare earths can enhance the ROS scavenging system in plants and reduce the degree of membrane lipid peroxidation and plasma membrane damage,thereby maintaining the structure and function of the OEC on the donor side of PSⅡ,increasing the electron receptor pool,and contributing to the defense against photoinhibition.

Higher Chilling-Tolerance of Grafted-Cucumber Seedling Leaves upon Exposure to Chilling Stress

The roots of figleaf gourd （Cucurbita ficifolia, as rootstock） could improve the resistance of cucumber plants （Cucumis sativus L. cv. Jinyan 4, as scion） to low temperature. In this experiment, the root activity and photosynthetic activity of photosystems in the own-rooted and grafted-cucumber plants were studied at chilling temperature （4℃） under low irradiance （100 μmol m^2 s^-1 PFD）. Compared with dark adaptation seedlings, the chlorophyll a fluorescence transient curve and the oxidizable P700 （P700＋） of both the own-rooted and grafted seedlings decreased, and PS2 and PS1 of the own-rooted seedling leaves were more inhibited than that of grafted ones at the end of chilling stress. The reduced triphenyltetrazolium chloride （TTC）, which was used to reflect the root activity, kept stable in grafted seedling roots at the end of chilling stress, while it decreased noticeably in the own-rooted seedling roots. These results implied that the root system activity of the grafted seedling roots was higher than that of the own-rooted ones. Superoxide dismutase （SOD） activity was higher in both the grafted seedling roots and leaves than that in own-rooted seedlings at both room temperature and chilling temperature. Upon exposure to chilling stress, the malondialdehyde （MDA） content, which reflects the degree of lipid peroxidation, increased markedly in the own-rooted seedling roots and leaves and kept stable in the grafted-cucumber seedlings.

Regulation of Reversible Dissociation of LHCII from PSII by Phosphorylation in Plants

LHCII is a crucial light-harvesting pigment/protein complex in photosystem II (PSII) supercomplex. It also participates in the light energy redistribution between photosystems and in the photoprotection via its reversible dissociation with PSII and PSI (photosystem I). This reversible detachment of LHCII is regulated by phosphorylation of its own and PSII core protein. Under low light conditions, LHCII is phosphorylated and dissociated with PSII core protein complex and combined with PSI, which balances the excitation energy between PSII and PSI;Under high light environment, the phosphorylation of PSII core proteins makes LHCII detach from PSII. The dissociated LHCII presents in a free state, which involves in the thermal dissipation of excess excitation energy. During photodamage, dual phosphorylations of both PSII core proteins and LHCII complexes occur. The phosphorylation of D1 is conductive to the disintegration of photodamaged PSII and the cycle of repair. In this circumstance, the phosphorylation of LHCII is induced by reactive oxygen species (ROS) and then the phosphorylated LHCII migrates to PSI, into the repair cycle of damaged PSII. The ferredoxin (Fdr) and thioredoxin (Tdr) system may play a possible central role in the phosphorylation regulation on LHCII dissociation.

Shade shelters increase survival and photosynthetic performance of oak transplants at abandoned fields in semi-arid climates

Forest restorations conducted in semiarid, seasonally dry climates must deal with the intense drought stress that affects tree seedlings during the dry season. Although this water deficit is the most commonly invoked source of mortality for seedlings, several other environmental factors may also preclude survival of transplants. For instance, it has been widely reported that excessive light reduces the efficiency of the photosynthetic apparatus, hence decreasing plant survival, but most seedling transplants in deforested areas are conducted under these light conditions. This study is focused in determining whether excessive light affects the photosynthetic performance and survival of Quercus coccolobifolia, a Mexican oak species, when their seedlings are transplanted in semiarid deforested areas. Further, this study tests the possibility of using artificial shade shelters to improve the ecophysiological performance and survival of seedlings. Oak seedlings were transplanted under full sunlight conditions and beneath artificial shade shelters of two different colors： white and black. To reduce water stress, and hence isolate the effects of light treatments, a drip irrigation system was implemented at each experimental plot. Seedling survival was monitored weekly for 128 days and photosynthetic performance was assessed by measuring chlorophyll fluorescence at three opportunities during the experiment. Sun-exposed seedlings showed lower photosynthetic performance and survival rates than those beneath shelters of both colors. These results suggest that sunlight damage can reduce seedling survival when they are transplanted in exposed sites, and that shade shelters can improve the success of forest restoration programs in semiarid climates.

Investigation of the decline of Ulva prolifera in the Subei Shoal and Qingdao based on physiological changes

It has been a scientific consensus that Ulva prolifera green tide that break out annually between May and August in the recent decade off eastern coast of Yellow Sea.To investigate the ecological mechanism of the decline of U.prolifera green tide,we compared the physiological characteristics of U.prolifera samples collected in three stages of the green tide in Subei Shoal north Jiangsu,the initiation point,on May 10(early,the initiation),July 10(middle,the outbreak),and in Taipingjiao,Qingdao on August 10(late,the decline).Compared to the early samples,the middle,and late samples showed a lower chlorophyll content and photosynthetic e fficiency at a higher lipid peroxidation level.The middle and later samples had fewer chloroplasts in the cells of U.prolifera than those in the early samples.In addition,the cells of U.prolifera sampled in the late stage showed ruptured and deformed cell walls.Differences in physiological function were reflected in the growth rate.Both the middle and late U.prolifera samples presented negative growth rates.Correlation analysis results show that the rising temperature was mainly responsible for the local decline of U.prolifera green tide.The high light-intensity that could induce the photoinhibition was also an important factor for the decline.Therefore,U.prolifera population that remained in the Subei Shoal and those that northward drifted to Qingdao experienced the same decline processes:rising temperature and continuous high light conditions in the late phase of green tide reduced the photosynthetic capacity and destroyed the antioxidant system of the thallus,which lead to decrease of biomass.The only difference is that the temperature in the Subei Shoal rose earlier than that in Qingdao;thus,U.prolifera remaining in the Subei Shoal went disappeared first.

Photoinhibitive and Recovery Properties of Hybrid Rice Ⅱ You 129 under Field Conditions

Photoinhibitive properties of super-high-yielding hybrid rice Ⅱ you 129 and its adaptation mechanism to strong light stress were investigated by measuring the light-response curve, diurnal variations of net photosynthetic rate and chlorophyll fluorescence parameters of Ⅱ you 129 leaves and compared with Shanyou 63. Photoinhibition of rice flag leaves under field conditions mainly resulted from the increase of thermal dissipation, especially for thermal dissipation depended on the xanthophyll circle, but no destruction of photosynthetic apparatus occurred. Potentially super-high-yielding hybrid rice Ⅱ you 129 was more tolerant to photoinhibition than Shanyou 63, because it had higher light saturation intensity and maximum net photosynthetic rate; more active xanthophyll cycle, and more rapid recovery ability after photoinhibition.

Photosynthetic Diurnal Variation of Soybean Cultivars with High Photosynthetic Efficiency

The photosynthetic characters were investigated among soybean cultivars with high photosynthetic efficiency and high yield. The results indicated that: 1) There were significant differences in photosynthetic rate (Pn) and dark respiration rate (DR) under saturation light intensity and appropriate temperature. 2) There were a little difference in light compensation point among them. Photo flux density (PFD) were about 360μmol/m2·s when Pn tended to zero. 3) When PFD>1 900 μmol/m2·s, photoinhibition occuried a-mong the cultivars. Diurnal variation of Pn was shown a curve with two peaks. 4) The cultivars with high photosynthetic efficiency were subjected less to photoinhibition than that with high yield. Critical temperatures of photoinhibition in high photosynthetic efficiency cultivars were higher than that of high yield.

Study of the Protective Effects in PEPC Transgenic Rice

The diurnal course of chlorophyll fluorescence parameter and active oxygen metabolism of flag leaves in PEPC transgenic and untransformed rice Kitaake were studied. The results showed that the photosynthetic rate under high light intensity has been increased by 50% and photoinhibition of photosynthesis in PEPC transgenic was alleviated after the introduction of PEPC gene from maize into rice. It was demonstrated that the increment of photosynthesis in PEPC transgenic was related to the introduction of PEPC gene using specific inhibitor of PEPC. Photoinhibition of photosynthesis in different genotypes exists at noon under natural condition. PEPC transgenic rice exhibited a less decrease in Fv/Fm, a less photoinhibition and a higher efficiency of light energy conversion to chemical energy and lower thermal energy dissipation. These results provided the physiological basis on the mechanism of tolerance to photoinhibition and rice breeding with high photosynthetic efficiency.

Relationships between D1 protein, xanthophyll cycle and photodamage-resistant capacity in rice (Orysa sativa L.)

Relationships between D1 protein, xanthophyll cycle and subspecific difference of photodamage-resistant capacity have been studied in O. japonica rice varieties 02428 and 029 (photoinhibition-tolerance) and O. indica rice varieties 3037 and Palghar (photoinhibition-sensitivity) and their reciprocal cross F, hybrids after photoinhibitory treatment. It was shown that PS II photochemical efficiency (Fv /Fm) decreased, and xanthophyll cycle from violaxanthin (V), via anaxanthin (A), to zeaxanthin (Z) was enhanced and non-photochemical quenching (g/v) increased accordingly in SM-pretreated leaves of rice when the synthesis of D1 protein was inhibited, and that there was a decrease in qN and, as a result, more loss of D1 protein and a big decrease in Fv/Fm in DTT-pretreated leaves when xanthophyll cycle was inhibited. O. japonica subspecies had a higher maintaining capacity of D1 protein and a decrease of Fv /Fm in a more narrow range, and exhibited more resistance against photodamage, as compared with O.

Carbon assimilation characteristics of plants in oasis-desert ecotone and their response to CO_2 enrichment

Six species of more than 20-year-old desert woody plants in the oasis-desert ecotone were selected for study. The results showed that: (1) in different growing seasons δ13 C values of assimilating organ varied between -14‰ and -16‰ for Haloxylon ammodendron (HA),-14‰ - -15‰ for Calligonum mongolicum (CM) and -25‰ - -28‰ for Caragana korshinskii (CK), Nitraria sphaerocarpa (NS) and Hedysarum scoparium (HS). (2) The net photosynthetic rate (Pn) of HA and CM was significantly higher than those of the other species. With the decrease in Pn for the six species, their intercellular CO2 concentration increased, but stomatal limitation value decreased under the intensive light. At the same time, the photochemical efficiency of PS Ⅱ dropped to different degrees. (3) The CO2 enrichment experiment demonstrated that, Pn of HA and CM increased to different extent under 450 μmol/mol, but their Pn reduced or approximated to the current condition under 650 μmol/mol. Under 450 μmol/mol the efficiency for solar energy utilization of CK and HS significantly reduced and under 650 μmol/mol their respiration rate exceeded photosynthesis rate. It can be concluded that HA and CM have some function of pathway for C4, but the other three species have the function for C3. The decline in their Pn is mainly caused by non-stomatal factors. HA, CM, CK and HS exhibited photoinhibition, which disappeared in a short time. This is a kind of positive readjustment to adapting to the desert environment. HA and CM can adapt to the high CO2 environment, but CKand HS cannot. With the rise in atmospheric CO2 concentration and climate warming, the latter two species in the oasis-desert ecotone may be gradually degraded or even disappear.

Xanthophyll Cycle and Inactivation of Photosystem Ⅱ Reaction Centers Alleviating Reducing Pressure to Photosystem Ⅰ in Morning Glory Leaves under Short-term High Irradiance

Under 30-min high irradiance （1500μmol m^-2 s^-1）, the roles of the xanthophyll cycle and D1 protein turnover were investigated through chlorophyll fluorescence parameters in morning glory （Ipomoea setosa） leaves, which were dipped into water, dithiothreitol （DTT） and lincomycin （LM）, respectively. During the stress, both the xanthophyll cycle and D1 protein turnover could protect PSI from photoinhibition. In DTT leaves, non-photochemical quenching （NPQ） was inhibited greatly and the oxidation level of P700 （P700^＋） was the lowest one. However, the maximal photochemical efficiency of PSII （Fv/Fm） in DTT leaves was higher than that of LM leaves and was lower than that of control leaves. These results suggested that PSI was more sensitive to the loss of the xanthophyll cycle than PSII under high irradiance. In LM leaves, NPQ was partly inhibited, Fv/Fm was the lowest one among three treatments under high irradiance and P700^＋ was at a similar level as that of control leaves. These results implied that inactivation of PSII reaction centers could protect PSI from further photoinhibition. Additionally, the lowest of the number of active reaction centers to one inactive reaction center for a PSII cross-section （RC/CSo）, maximal trapping rate in a PSll cross-section （TRo/CSo）, electron transport in a PSll cross-section （ETo/CSo） and the highest of 1-qP in LM leaves further indicated that severe photoinhibition of PSII in LM leaves was mainly induced by inactivation of PSII reaction centers, which limited electrons transporting to PSh However, relative to the LM leaves the higher level of RC/CSo, TRo/CSo, Fv/Fm and the lower level of 1-qP in DTT leaves indicated that PSI photoinhibition was mainly induced by the electron accumulation at the PSI acceptor side, which induced the decrease of P700^＋ under high irradiance.