Photoinhibition in Shaded Cotton Leaves After Exposing to High Light and the Time Course of Its Restoration

Chlorophyll fluorescence emission, pigment composition and photosynthetic rate of shade-grown cotton ( Gossypium hirsutum L.) plants were measured immediately after suddenly exposing to full sunlight and at regular intervals there after within 15 d. Photoinhibition occurred in shade-grown cotton leaves immediately after exposed to full sunlight. The chlorophyll fluorescence parameter F-v/F-m and PhiPS II, which reflect the efficiency of PS II,obviously decreased in shade-grown leaves, much lower than that of the full sunlight-grown leaves. On the contrary, F-o value was sharply increased. Neither of these parameters could completely recover till next morning. The photoinhibition was chronic and continued for about 4 d, while the F-v/F-m and the net photosynthetic rate ( P-n) continued to decline, then began to increase gradually 6 d later and turned stable after 10 - 12 d, appearing as an acclimation phenomenon. However, the final value of F-v/F-m and P-n did not reach the level as in those leaves grown in the full sunlight ever before. The final P-n was higher by 60% than that before exposure, but lower for more than 40% than that of the full sunlight-grown leaves. The most notable response of chloroplast pigment composition was a pronounced increase in the pool size of carotenoids in xanthophyll cycle over a period of 3 d. The results indicated that when shade-grown cotton seedlings were suddenly transferred to the full sunlight, the decline of F-v/F-m and P-n might associate with the damage of the PS II reaction center. During the light acclimation, photoprotective mechanisms such as the xanthophyll cycle-dependent energy dissipation were increased, so that photodamage in leaves transferred from low to high light might be reduced.

Photoinhibition and Photoprotection in Ginkgo biloba leaves: Influence of Temperature, CO 2 and O 2

In midday ginkgo ( Ginkgo biloba L.) leaves have to bear photon flux density over 1400 μmol·m -2 ·s -1 in combination with high temperatures around 35 ℃ at natural habitat. They show typical midday depression of stomatal conductance and of CO 2 assimilation rate. The zeaxanthin changes with light intensity during the day. The influence of the combination of strong light and temperature on photoinhibition was also examined in the laboratory. A low CO 2 internal conductance (31 mmol·m -2 ·s -1 ) was found in ginkgo leaves, which had been exposed to excessive light at temperature between 15 ℃ and 35 ℃ with reduced CO 2 (80 μL·L -1 ) or oxygen (2%) for 2 h, causing a low CO 2 concentration at the carboxylation site and a high proportion of photorespiration. The ratio of electron transport to CO 2 fixation was rather high in ginkgo (16 e -/CO 2 at 25 ℃) as compared with other plants. It increased with temperature also in 2% O 2 which could not be explained solely as due to change of photorespiration. The reduction of oxygen in 340 or 80 μL·L -1 CO 2 had no effect on the extent of photoinhibition at all temperatures, which indicated that electron flow caused by photorespiration in excess light was negligible in protective effect in ginkgo leaves. However, a decreased CO 2 concentration increased photoinhibition, especially at high temperature. It is concluded that the dissipation of excessive excitation energy in the PSⅡ antennae through the xanthophyll cycle may be the major protective mechanism to preventing from the deteriorated effects of strong light in ginkgo leaves.

Effects of Photoinhibition and Its Recovery on Photosynthetic Functions of Winter Wheat Under Salt Stress

Effects of photoinhibition and its recovery on photosynthetic functions of winter wheat (Triticum aestivum L.) under salt stress were studied. The results showed that several parameters associated with PSⅡ functions, e.g. Fv/Fo?Fv/Fm and qP were not influenced by lower salt concentration (200 mmol/L NaCl) while CO 2 assimilation rate decreased significantly. When exposed to higher salt concentration (400 mmol/L NaCl), PSⅡ functions were significantly inhibited which led to the decrease of carbon assimilation. These results suggest that different concentrations of salt stress affected photosynthesis by different modes. Salt stress made photosynthesis more sensitive to strong light and led to more serious photoinhibition. Under lower concentration of salt stress, the Q B-non-reductive PSⅡ reaction centers formed at the beginning of photoinhibition could be effectively used to compose active PSⅡ reaction center (RC) and repair the reversible inactivated PSⅡ RC. Under higher concentration of salt stress, PSⅡ reaction centers were seriously damaged during photoinhibition, the Q B-non-reductive PSⅡ RC could only be partly effective at the early time of photoinhibition, thus led to the accumulation of Q B-non-reductive PSⅡ RC in the course of restoration under dim light.

PSⅡ Photoinhibition and O_2~ Production

To know more about the site of O -· 2 production in thylakoids, the F v/F m ratio was measured in parallel with the oxygen free_radical production detected by electron spin resonance (ESR) using Tiron as a probe in the isolated thylakoids of Scots pine ( Pinus sylvestris L.). As the light intensity was strong enough to induce photoinhibition leading to reduction in the F v/F m ratio, the intensity of the TH · signal representing the superoxide production was enhanced. The TH · signal was also stimulated by 3_(3,4_dichloropheneyl)_1,1_dimethylurea but was suppressed by exogenous scavengers or by 2,6_dichlorophenol indophenol. Therefore, the superoxide production is closely related to photoinhibition in PSⅡ. The novel results may further promote research on the mechanism of photoinhibition and on the structure and function of PSⅡ as well.

The Possible Mechanism of Photoinhibition of Purified PSⅠ Complexes

The protecting effect of histidine on the photodamage of pigments and proteins of the isolated PSⅠ particles from the chloroplast of Spinacia oleracea L. during the strong illumination (2 300 μmol·m -2 ·s -1 ) was studied by spectroscopy and SDS_PAGE. The absorbance of PSⅠ particles decreased during the strong illumination treatment, but the decrease would be slowed down in the presence of externally added histidine after 30 min illumination. The decrease of CD (circular dichroism) signal intensities of PSⅠ particles also was slowed down by the added histidine after about 10 min illumination. The retarded protecting effect of the added histidine on the photobleaching of pigments of PSⅠ complexes implied that the mechanisms of photoinhibition of isolated PSⅠ complexes are different from early stage to later stage during the strong illumination treatment. In addition, the added histidine suppressed the decrease of 77 K fluorescence yield of PSⅠ particles during the illumination. SDS_PAGE showed that the added histidine not only protected the reaction center proteins of PSⅠ particles, but also protected other subunits of PSⅠ particles from degradation.

Comparative Study on Photoinhibition Between Two Wheat Genotypes

Changes in the efficiency of the primary light energy conversion, fluorescence quenching parameters and contents of photosynthetic pigments were compared between two wheat ( Triticum aestivum L.) genotypes in response to high light stress. The contents of chlorophyll and carotenoid in “Jing_411' were slightly higher than those in “Xiaoyan_54'. Under high light stress, photoinhibition as indicated by a sustained decrease in PSⅡ photochemical efficiency was more pronounced in “Jing_411' than in “Xiaoyan_54'. The content of ascorbate and the activity of the deepoxidase were higher in “Xiaoyan_54' than in “Jing_411'. The genotypic difference in resistance to photoinhibition is related to the capacity to dissipate the excess energy nonradiatively.

PSⅡ Photochemistry and Xanthophyll Cycle in Two Superhigh-yield Rice Hybrids, Liangyoupeijiu and Hua-an 3 During Photoinhibition and Subsequent Restoration

PSⅡ photochemistry and xanthophyll cycle during photoinhibition (exposed to strong light of 2 000 μmol photons·m -2 ·s -1 ) and the subsequent restoration were compared between two superhigh_yield rice hybrids (Liangyoupeijiu and Hua_an 3, the newly developed rice hybrids from two parental lines) and the traditional rice hybrid Shanyou 63 developed from three parental lines. The results showed that the maximal efficiency of PSⅡ photochemistry ( Fv/Fm ), the efficiency of excitation energy capture by open PSⅡ centers ( Fv′/Fm′ ), and the yield of PSⅡ electron transport ( Φ PSⅡ ) of the three rice hybrids decreased during photoinhibition. However, a greater decrease in Fv/Fm , Fv′/Fm′ , and Φ PSⅡ was observed in Shanyou 63 than in Liangyoupeijiu and Hua_an 3. At the same time, the components of xanthophyll cycle, antherxanthin (A) and zeathanxin (Z) increased rapidly while violaxanthin (V) decreased considerably. Both the rate of accumulation and the amount of A and Z in the two superhigh_yield rice hybrids were higher than that in Shanyou 63. The de_epoxidation state (DES) of xanthophyll cycle increased rapidly with the fast accumulation of A and Z, and reached the maximal level after first 30 min during photoinhibition. Of the three hybrids, the increasing rate of DES in Liangyoupeijiu and Hua_an 3 was higher than that in Shanyou 63. After photoinhibition treatment, the plant materials were transferred to a dim light (70 μmol photons·m -2 ·s -1 ) for restoration. During restoration, both chlorophyll fluorescence parameters and xanthophyll cycle relaxed gradually, but the rate and level of restoration in the two superhigh_yield rice hybrids were higher than those in Shanyou 63. Our results suggest that Liangyoupeijiu and Hua_an 3 had higher resistance to photoinhibition and higher capacity of non_radiative energy dissipation associated with xanthophyll cycle, as well as higher rate of restoration after photoinhibition, than Shanyou 63 when subjected to strong light.

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.

Photosynthetic Characteristics of Two Superhigh-yield Hybrid Rice

The photosynthetic functions and the sensitivity to photoinhibition were compared between two superhigh_yield hybrid rice (Oryza sativa L.) Liangyoupeijiu and X07S/Zihui 100, the newly developed from two parental lines and traditional hybrid rice Shanyou 63 developed from three parental lines. The results showed that, as compared to Shanyou 63, the net photosynthetic rate of Liangyoupeijiu and X07S/Zihui 100 was 9.1% and 11.9% higher, the transpiration rate was 37.4% and 31.4% lower, and their water use efficiency was 74.2% and 63.5% higher respectively. After strong light (2 000 μmol photons·m -2 ·s -1 ) treatment for 2 h, the photochemical quantum yield and the photochemical quenching increased by 37.0% and 18.0% respectively in Liangyoupeijiu, 28.3% and 46.2% in X07S/Zihui 100, but decreased a little in Shanyou 63. The non_photochemical quenching decreased in Liangyoupeijiu and X07S/Zihui 100 (about 50%) but increased greatly in Shanyou 63 (about 50%). Better photosynthetic functions, higher water use efficiency and stronger resistance to photoinhibition, may be the physiological basis for the super high_yield of the two hybrid rice under study.

Photosynthetic Characteristics of Transgenic Rice Plants Overexpressing Maize Phosphoenopyruvate Carboxylase

用转PEPC、PPDK、NADP_ME、PEPC +PPDK 双基因水稻 (OryzasativaL .)及原种为材料 ,以光合酶活性、饱和光合速率及PSⅡ光化学效率 (Fv/Fm)为指标 ,研究了转PEPC基因水稻的光合生理特征。结果如下 :转PEPC基因水稻PEPC活性比原种提高 2 0倍 ;饱和光合速率比原种高 5 5 % ;用高光强或人工光氧化剂甲基紫精 (MV)处理后 ,与原种相比 ,转PEPC基因水稻光化学效率下降较少 ,证明其耐光抑制、耐光氧化能力增强 ,测定其光合日变化看出 :在 1d中不同时间 ,转PEPC基因水稻的光合速率均高于原种 ,且与PEPC活性的日变化有相似的趋势。上述结果为转PEPC基因水稻的生理机制和育种研究提供了依据和途径。

Responses of Chlorophyll Fluorescence and Carotenoids Biosynthesis to High Light Stress in Rice Seedling Leaves at Different Leaf Position

In the present study, we investigated the changes of photosynthesis, chlorophyll fluorescence and the content of carotenoid pigments in rice (Oryza sativa L.) seedling leaves and their responses to high light. The results showed that the rate of photosynthesis, the contents of individual and total carotenoids and the pool size of xanthophyll cycle decreased with age increasing of the leaf. When the leaves were exposed to high light for 2 h, the qN of mature leaf (5th leaf) increased more significantly than that of younger (6th leaf) and older leaves (3rd and 4th leaf). Comparing with the leaves before exposure to high light, the excitation pressure on PSⅡ (1- qP ) increased by 44%, 57%, 19% and 45% in the 3rd, 4th, 5th and 6th leaf under high light, respectively. The highest content of carotenoids and the greatest conversion of violaxanthin to zeaxanthin were found in the 5th leaf, and it was consistent with the 5th leaf exhibiting the strongest resistance to high light. Our results suggested that the ability of rice leaf to resist photoinhibition is related to the level of carotenoids and the ability of carotenoids biosynthesis.