Photoinhibition of Photosynthetic Oxygen Production and its Recovery in the Subtidal Red Alga Polyneura hilliae

In the marine red alga Polyneura hilliae photoinhibition of photosynthesis was investigated by measuring the photosynthetic oxygen production. The extent of photoinhibition in this alga depends on the fluence rate as well as on the time of exposure. Strongly photoinhibited algae recover slowly. Full recovery is reached only in weak light after 3 days. Two phases of recovery can be distinguished: a first relatively fast phase of recovery that is independent of light, and a second slow phase which begins after about 6 hours and requires weak light. Consequently, even in complete darkness partial recovery is observed. An action spectrum of photoinhibition and its comparison with the in vivo absorption spectrum of Polyneura demonstrates that photoinhibition is mainly caused by light which is absorbed by phycobiliproteins, chlorophyll a and carotenoids. This fact and the ineffectiveness of light above 686 nm clearly indicate that the main photoinhibition site in this alga is PS II. No photosynthetic oxygen production is detectable after 3 h irradiation at 402 nm, 150 μmol quanta m^?2s^?1. As photosynthetic activity recovers slowly and to a limited extent, the suppression of oxygen production is apparently only in part the result of photoinhibition sensu strictu and in part due to permanent photodamage.     

The Effect of Photoinhibition on Photosynthetic Oxygen Production in the Brown Alga Dictyota dichotoma

Photoinhibition of photosynthesis in the brown alga, Dictyota dichotoma, was studied with a PAM fluorometer (Walz, Effeltrich, Germany) and a homemade oxygen measuring device. As a measure of fluorescence, Fv/Fm, and for the photosynthetic yield, ΔF/Fm', were used. Oxygen measurements show clearly that the observed degree, as well as the time course, of photoinhibition depends on the fluence rate of the light used to measure changes of the production rate. After photoinhibition of photosynthesis the depression of oxygen production caused by non-saturating fluence rates was generally much more pronounced than that caused by saturating or nearly saturating fluence rates. At minimal photoinhibition the initial slope and the convexity of the fluence rate-response curve of oxygen evolution decrease, whereas the level of light saturation decreases only after strong photoinhibition. Nevertheless, at different degrees of photoinhibition, changes in the degree of the upper bending of the fluence rate-response curve of oxygen production are also linearly correlated to changes in the fluorescence ratios (Fv/Fm and ΔF/Fm'). The action spectrum of photoinhibition, calculated on the basis of changes of Fv/Fm, indicates that the reaction center of PS I is not involved in photoinhibition. The lower effectiveness of blue light in comparison to effects of green and red light may be due to chloroplast displacement, as in the so-called strong light position, the light absorbed by the thalli in vivo is decreased.     

Photoinhibition and recovery of photosynthesis in Coffea arabica and C. canephora

Photosynthetic parameters were determined in disks from leaves of C. arabica cv. Red Catuaí and C. canephora cv. Kouillou grown in the field. Kouillou showed a relatively higher irradiance requirement for saturating photosynthesis, lower chlorophyll (Chl) content, and higher Chl a/b ratio than Catuaí. Photoinhibition of photosynthesis under bright irradiance was manifested by decreases in maximum photochemical efficiency (evaluated by the variable to maximum fluorescence ratio, Fv/Fm), as a consequence of an increased initial and a quenched maximum fluorescence. Restoration of Fv/Fm following photoinhibition in low irradiance was faster in Kouillou than in Catuaí. Chloramphenicol both accelerated photoinhibition (mainly in Kouillou) and blocked its recovery for at least 190 min in either cultivar. Photosynthetic oxygen evolution under photoinhibitory conditions was decreased by chloramphenicol; in control leaf disks this decrease was only observed in C. arabica, but with a rapid recovery within 90 min of low irradiance exposure. In both coffee cultivars, the depressed photochemical efficiency of photosystem 2 was not accompanied by a concomitant lowering in oxygen evolution during reversal from photoinhibition.     

Photoinhibition and recovery of photosynthesis in Coffea arabica and C. canephora

Photosynthetic parameters were determined in disks from leaves of C. arabica cv. Red Catuaí and C. canephora cv. Kouillou grown in the field. Kouillou showed a relatively higher irradiance requirement for saturating photosynthesis, lower chlorophyll (Chl) content, and higher Chl a/b ratio than Catuaí. Photoinhibition of photosynthesis under bright irradiance was manifested by decreases in maximum photochemical efficiency (evaluated by the variable to maximum fluorescence ratio, Fv/Fm), as a consequence of an increased initial and a quenched maximum fluorescence. Restoration of Fv/Fm following photoinhibition in low irradiance was faster in Kouillou than in Catuaí. Chloramphenicol both accelerated photoinhibition (mainly in Kouillou) and blocked its recovery for at least 190 min in either cultivar. Photosynthetic oxygen evolution under photoinhibitory conditions was decreased by chloramphenicol; in control leaf disks this decrease was only observed in C. arabica, but with a rapid recovery within 90 min of low irradiance exposure. In both coffee cultivars, the depressed photochemical efficiency of photosystem 2 was not accompanied by a concomitant lowering in oxygen evolution during reversal from photoinhibition.     

Photoinhibition of Photosynthesis Without Net Loss of D1 Protein in Wheat Leaves Under Field Conditions

To determine whether the net loss of D1 protein is the main cause of photoinhibition of photosynthesis in wheat leaves under field conditions in the absence of any environmental stress other than strong sunlight, the D1 protein content, photosynthetic evolution of oxygen and chlorophyll a fluorescence parameters were measured in field grown wheat leaves. After exposure to midday strong light for about 3 h, apparent photosynthetic quantum efficiency (Φ), Fv/Fm and Fo in wheat leaves declined, and these parameters recovered almost completely 1 h after transfer to the weak light of 30~40 ttmol photons · m-2 · s-1. No evident change in the D1 protein content was observed in the leaves after exposure to midday strong light for 3 h. After 3 hours exposure to strong light, the slow-relaxed fluorescence quenching in the leaves treated with streptomycin (SM) increased much more than that in the control leaves, but there was no effect SM on the recovery of Fv/Fm and F0; dithiothretol (DTT) treatment enhanced photoinhibition of photosynthesis and reduced the D1 protein content in the leaves after exposure to midday strong light. These results indicated that under field conditions with no environmental stress other than strong sunlight, photoinhibition of photosynthesis in wheat leaves was not due to the net loss of D1 protein, and it could be attributed mainly by the increased nonradiative energy dissipation.     

Effects of Strong Light and Active Oxygen on Photosynthesis in Soybean

With the use of chlorophyll fluorescence technique, it was found that the net photosynthetic oxygen evolution rate decreased after strong light (2 000 μmol· m-2·2-1 ) treatment for two hours in soybean ( Glycine max L. ) leaves. The chlorophyll fluorescence parameters, Fm/Fo, Fv/Fm, ФPSII, qp and qN decreased along with the increase of light intensity. In strong light, exogenous active oxygen H202、·OH and 'O2 were harmful to soybean leaves. The destruction of 'O2 and·OH to leaves was most evident, as was shown that Fv/Fm and PS H decreased significantly. The antioxidants DABCO, mannitol, ascorbate and histidine protected the leaves, but weakly, from strong light. In darkness, the SOD inhibitor sodium diethyldithiocar- bamate (DDC) had no significant effect on Fm/Fo and Fv/Fm, but NAN,, the ascorbate peroxidase (APX)inhibitor, significantly decreased Fm/Fo, Fv/Fm and ФPS II. In strong light, however, beth DDC and NaN3 reduced the above-mentioned fluorescence parameters, but NaN3 was more effective than DDC. The results suggested that photoinhibition did take place in soybean leaves under strong light, and it was related to active oxygen in vivo.     

EFFECTS OF UV-B-INDUCED DNA DAMAGE AND PHOTOINHIBITION ON GROWTH OF TEMPERATE MARINE RED MACROPHYTES: HABITAT-RELATED DIFFERENCES IN UV-B TOLERANCE

The sensitivity to UV-B radiation (UVBR: 280–315 nm) was tested for littoral (Palmaria palmata[L.] O. Kuntze, Chondrus crispus Stackhouse) and sublittoral (Phyllophora pseudoceranoides S. G. Gmelin, Rhodymenia pseudopalmata[Lamouroux] Silva, Phycodrys rubens[L.] Batt, Polyneura hilliae[Greville] Kylin) red macrophytes from Brittany, France. Algal fragments were subjected to daily repeated exposures of artificial UVBR that were realistic for springtime solar UVBR at the water surface in Brittany. Growth, DNA damage, photoinhibition, and UV-absorbing compounds were monitored during 2 weeks of PAR + UV-A radiation (UVAR) + UVBR, whereas PAR + UVAR and PAR treatments were used as controls. The littoral species showed a higher UV tolerance than the sublittoral species. After 2 weeks, growth of P. palmata and C. crispus was not significantly affected by UVBR, and DNA damage, measured as the number of cyclobutane-pyrimidine dimers per 10⁶ nucleotides, was negligible. Photoinhibition, determined as the decline in optimal quantum yield, was low and decreased during the course of the experiment, coinciding with the production of UV-absorbing compounds in these species. In contrast, no UV-absorbing compounds were induced in the sublittoral species. Growth rates of P. pseudoceranoides and R. pseudopalmata were reduced by 40% compared with the PAR treatment. Additionally, constant levels of DNA damage and pronounced photoinhibition were observed after the UVBR treatments. Growth was completely halted for Phycodrys rubens and Polyneura hilliae, whereas DNA damage accumulated in the course of the experiment. Because Phycodrys rubens and Polyneura hilliae showed the same degree of photoinhibition as the other sublittoral species, it appears that the accumulation of DNA damage may have been responsible for the complete inhibition of growth. The results suggest an important role of DNA repair pathways in determining the UV sensitivity in red macrophytes.     

草莓叶片光合作用对强光的响应及其机理研究

用便携式调制叶绿素荧光仪和光合仪研究了强光下草莓叶片荧光参数及表观量子效率的变化.结果表明,Fm、Fv/Fm、PSⅡ无活性反应中心数量和Q_A的还原速率在强光下降低,在暗恢复时升高;而PSⅡ反应中心非还原性Q_B的比例在强光下增加,在暗恢复时降低.上述荧光参数的变化幅度均以强光胁迫或暗恢复的前10 min最大.强光下Φ_PSII、ETR和qP先升高后降低,但qN先大幅度降低,然后小幅回升.强光处理4 h后,丰香和宝交早生的表观量子效率(AQY)分别降低了20.9%和37.5%;qE(能量依赖的非光化学猝灭)为NPQ(非光化学猝灭)的最主要成分.强光胁迫下丰香的Fo、Fm、Fv/Fm、Φ_PSII、ETR和AQY的变化幅度均明显比宝交早生小.DTT处理后,草莓叶片的Fm和Fv/Fm明显降低,Fo显著升高.可以认为,依赖叶黄素循环和类囊体膜质子梯度两种非辐射能量耗散在草莓叶片防御光损伤方面起着重要作用,丰香的光合机构比宝交早生更耐强光.     

Photoinhibition and its wavelength dependence in the cyanobacterium Anabaena variabilis

In the cyanobacterium Anabaena variabilis the dependence of photoinhibition on fluence rate, duration and wavelength of irradiation were studied by measurements of oxygen production and fluorescence emission spectra. The analysis of the photosynthetic activity revealed that photoinhibition affects exclusively photosystem II (PS II), whereas photosystem I (PS I) remained largely unimpaired. Furthermore, PS II fluorescence emission decreased much faster in bleached than in unbleached controls.Studying the wavelength dependence of photoinhibition it was found that only radiation between 520 and 680 nm causes photoinhibition. This is about the same range of wavelengths which causes photobleaching. Fluorescence emission spectra of samples exposed to high fluence rates of 582 and 662 nm, respectively, essentially agree with those samples exposed to high fluence rates of white light, whereas the fluorescence emission spectra of samples exposed to blue light resemble those exposed to dim white light.NaN₃, a substance which prevents photobleaching, inhibits the photosynthetic O₂ production of Anabaena and, hence, enhances the photoinhibitory effect.     

Tidal Dependence of Photoinhibition of Photosynthesis in Marine Macrophytes of the South China Sea*

During an expedition in spring 1992 to Hainandao, an island in the tropical zone of the South China Sea, the daily courses of photoinhibition of different brown algal species and of the seagrass Thalassia hemprichii were investigated. Experiments were carried out with the new portable chlorophyll fluorometer PAM 2000 (Walz, Germany). As a measure of photoinhibition Fv/Fm was used and as a measure of the photosynthetic yield ΔF/Fm'. Photoinhibition occurred in all algae floating near the water surface and reached its maximum between noon and the early afternoon. In the evening photosynthesis was always fully recovered. The extent of photoinhibition depended on both the depth of the algae and the course of the irradiance during the day. Algae of the sublittoral zone showed only a low degree of photoinhibition at high fluence rates when they were covered by a water column of more than 1 m, even if the water was clear. The seagrass Thalassia hemprichii grew in the middle and upper intertidal zone and showed a significant photoinhibition at low tide only when it was not water-covered. Apparently, it is able to cope with extreme high light conditions without downregulation of photosynthesis caused by photoinhibition.     

Recovery of photosynthesis in winter-stressed Scots pine

Abstract. . Winter-induced inhibition of photosynthesis in Scots pine (Pinns sylvestris L.) is caused by the combined effects of light and freezing temperatures; light causes photoinhibition of photosystem II (Strand & Oquist, 1985b, Physiologic Plantarum, 65 , 117–123), whereas frost causes inhibition of enzymatic steps of photosynthesis (Strand & Öquist, 1988, Plant, Cell & Environment, 11 , 231–238). To reveal limiting steps during recovery from winter stress, the potential of photosynthesis to recover and the actual recovery outdoors during spring, were studied in Scots pine. Studies of light dependent O₂-evolution under saturating CO₂ and recordings of room temperature fluorescence induction kinetics were used. When branches of pine, in February and March, were brought into the laboratory and kept at 18°Cand 100μmol m^?2 s^?1, light saturated rates and apparent quantum yields of photo-synthetic O₂-evolution recovered fully within approximately 48h. The photochemical efficiency of photosystem II, as measured by Fv/Fm ratios, recovered fully within 24h after an initial lag-phase of 2-3 h. Under natural winter conditions, the Fv/Fm ratio decreased more in exposed than in shaded pine, whereas the efficiency of photosynthesis was similarly inhibited in exposed and shadedpine. However, when recovery from winter stress occurred during spring, the Fv/Fm ratios of both shaded and exposed pine recovered well before photosynthesis. It is concluded that the light-induced photoinhibition component of winter stress in photosynthesis of pine recovers well before the frost induced component(s) of winter stress. In this context, reversible photoinhibition of photosynthesis in evergreen conifers is considered as a dynamic down-regulation of photosystem II to prevent more severe photodynamic damage of the thylakoid membrane when photosynthesis is inhibited by frost.     

The role of chloroplastic NAD(P)H dehydrogenase in photoprotection.

After a brief exposure to supra-saturating light, leaves of a tobacco transformant, in which chloroplastic NAD(P)H dehydrogenase (NDH) was defective, showed more severe photoinhibition than the wild-type, when judged by the parameter of chlorophyll fluorescence Fv/Fm. Repeated application of supra-saturating light eventually resulted in chlorosis in the NDH-defective mutant, while the wild-type sustained less photodamage and was able to recover from it. The mechanism of the phenomena is discussed with respect to the potential role of NDH in photosynthesis.     

Relationship between xanthophyll cycle and non-photochemical quenching in rice (Oryza sativa L.) plants in response to light stress

Thirty days old rice plants grown under low and moderate light conditions were transferred to full sunlight to observe the extent of photoinhibitory damage and protective mechanism, and the relationship between xanthophyll cycle and nonphotochemical quenching (qN) under changing light environment. Control plants (low, moderate and sun grown) exhibited similar Fv/Fm ratio, indicating similar photosynthetic efficiency prior to light stress. On exposure to the high light treatment, low light grown plants exhibited faster and higher degree of photoinhibition compared to moderate and high light grown plants. Moderate and high light grown plants showed relatively less photoinhibition and also showed higher qN, indicating better capacity of energy dissipation. Increase in qN in moderate light and sun grown plants was accompanied by conversion of violaxanthin (V) to antheraxanthin (A) and zeaxanthin (Z) indicating operation of Z-dependent thermal dissipation. Rice plants fed with ascorbate (AsA), a stimulator of the de-epoxidation state of V to Z, showed higher Fv/Fm ratio and qN than the plants fed with dithiothreitol (DTT) an inhibitor of xanthophyll cycle. This indicated that an increased amount of energy reached PS II reaction centre, due to absence of A and Z formation, thereby causing greater damage to photosynthesis in DTT fed rice plants. The present data confirmed the relationship between qN and Z in dissipating the excess light energy, thereby protecting plants against photodamage.     

不同光质对烟草叶片生长及光合作用的影响

通过对烟草植株覆盖白、红、黄、蓝、紫色滤膜获得不同光质,研究了光质对烟叶生长及光合作用的影响。结果表明,与白膜处理相比,红膜与蓝膜处理下的烟草叶片较厚,比叶面积较小,叶绿素a/b比值、净光合速率、可变荧光强度（Fv）和最大荧光强度（Fm）的比值Fv/Fm（PSⅡ最大光化学量子效率）、PSⅡ实际光化学量子效率（ΦPSⅡ）、光饱和点和CO2饱和点均较高。黄膜处理下的叶片较白膜处理的更薄,净光合速率、Fv/Fm、ΦPSⅡ、光饱和点、CO2饱和点均较低。紫膜处理的叶片比叶面积比白膜处理的小,净光合速率和Fv/Fm比白膜的大。实验结果表明红光、蓝光和紫光促进了烟叶的生长,这种促进作用是与其高光合效率紧密相连的;而黄光对烟叶的生长有一定程度的抑制作用。     

The Susceptibility of Photosynthesis to Photoinhibition and the Capacity of Recovery in High and Low Light Grown Cyanobacteria, Anacystis nidulans

The susceptibility of photosynthesis to photoinhibition and the rate of its recovery were studied in the cyanobacterium Anacystis nidulans grown at a low (10 micromoles per square meter per second) and a high (120 micromoles per square meter per second) photosynthetically active radiation. The rate of light limited photosynthetic O₂ evolution was measured to determine levels of photoinhibition and rates of recovery. Studies of photoinhibition and recovery with and without the translation inhibitor streptomycin demonstrated the importance of a recovery process for the susceptibility of photosynthesis to photoinhibition. We concluded that the approximately 3 times lower susceptibility to photoinhibition of high light than of low light grown cells, significantly depended on high light grown cells having an approximately 3 times higher recovery capacity than low light grown cells. It is suggested that these differences in susceptibility to photoinhibition and recovery depends on high light grown cells having a higher turnover rate of photosystem II protein(s) that is(are) the primary site(s) of photodamage, than have low light grown cells. Furthermore, we demonstrated that photoinhibition of A. nidulans may occur under physiological light conditions without visible harm to the growth of the cell culture. The results give support for the hypotheses that the net photoinhibitory damage of photosystem II results from the balance between the photoinhibitory process and the operation of a recovery process; the capacity of the latter determining significant differences in the susceptibility of photosynthesis to photoinhibition of high and low light grown A. nidulans.     

Photoprotective Effects of D1 Protein Turnover and the Lutein Cycle on Three Ephemeral Plants under Heat Stress

To clarify the characteristics of photoinhibition and the primary defense mechanisms of ephemeral plant leaves against photodestruction under high temperature stress, inhibitors and the technology to determine chlorophyll fluorescence were used to explore the protective effects of D1 protein turnover and the lutein cycle in the high temperature stress of the leaves of three ephemeral plants. The results showed that the maximum light conversion efficiency (Fv/Fm) of the ephemeral plant leaves decreased, and the initial fluorescence (Fo) increased under 35°C ± 1°C heat stress for 1–4 h or on sunny days in the summer. Both Fv/Fm and Fo could be recovered after 8 h of darkness or afternoon weakening of the external temperature. Streptomycin sulfate (SM) or dithiothreitol (DTT) accelerated the decrease of Fv/Fm and the photochemical quenching coefficient (qP) in the leaves of three ephemeral plants at high temperature, and the decrease was greater in the SM than in the DTT treatment. When the high temperature stress was prolonged, the Y(II) values of light energy distribution parameters of PSII decreased, and the Y(NPQ) and Y(NO) values increased gradually in all the treatment groups of the three ephemeral plants. The results showed that the leaves of the three ephemeral plants had their own highly advanced mechanisms to protect against photodamage, which inhibited the turnover of D1 protein and xanthophyll cycle. This can damage the PSII reaction center in the leaves of the three ephemeral plants under high temperature. The protective effect of D1 protein turnover on heat stress in Erodium oxyrrhynchum and Senecio subdentatus was greater than that of the lutein cycle, while the protective effect of lutein cycle was greater than that of D1 protein turnover in Heliotropium acutiflorum subjected to heat damage.     

Use of chlorophyll fluorescence to estimate the effect of photoinhibition in outdoor cultures ofSpirulina platensis

Chlorophyll fluorescence measurements were used to evaluate the effect of temperature on photoinhibition inSpirulina platensis cultures grown in tubular reactors outdoors. Cultures grown at 35 °C during the day time showed a lower reduction in the Fv/Fm ratio as compared to cultures grown at 25 °C. It is demonstrated that the lower temperature photoinhibited cells can undergo a complete recovery once transferred to low light and higher temperature. This recovery does not take place when 100 µg ml^-1 chloramphenicol is added to cells. The recovery is light dependent and cells incubated in the dark at low temperature do not show a recovery in the Fv/Fm ratio. The data presented strongly support the hypothesis that photoinhibition takes place in outdoorSpirulina cultures. At the same time it is demonstrated that fluorescence measurements can be used as a fast reliable indication for photoinhibition in outdoor algal cultures.Author for correspondencePublication No. 69 of the Microalgal Biotechnology Laboratory.     

Reversible Photoinhibition in Antarctic Moss during Freezing and Thawing

Tolerance of antarctic moss to freezing and thawing stress was investigated using chlorophyll a fluorescence. Freezing in darkness caused reductions in Fv/Fm (ratio of variable to maximum fluorescence) and Fo (initial fluorescence) that were reversible upon thawing. Reductions in Fv/Fm and Fo during freezing in darkness indicate a reduction in the potential efficiency of photosystem II that may be due to conformational changes in pigment-protein complexes due to desiccation associated with freezing. The absorption of light during freezing further reduced Fv/Fm and Fo but was also reversible. Using dithiothreitol (DTT), which inhibits the formation of the carotenoid zeaxanthin, we found reduced flurorescence quenching during freezing and reduced concentrations of zeaxanthin and antheraxanthin after freezing in DTT-treated moss. Reduced concentrations of zeaxanthin and antheraxanthin in DTT-treated moss were partially associated with reductions in nonphotochemical fluorescence quenching. The reversible photoinhibition observed in antarctic moss during freezing indicates the existence of processes that protect from photoinhibitory damage in environments where freezing temperatures occur in conjunction with high solar radiation levels. These processes may limit the need for repair cycles that require temperatures favorable for enzyme activity.     

外源甜菜碱对盐胁迫下枸杞光合功能的改善

研究了外源甜菜碱对盐胁迫下枸杞扦插苗叶片光合功能的影响。结果表明,外源甜菜碱能使盐胁迫下的枸杞叶片叶绿素荧光动力学参数Fo、Fm、Fv、Fv／Fm、Fm／Fo和Fv／Fo增高,使光合色素叶绿素a(Chla)、叶绿素b(Chlb)和类胡萝卜素(Car)含量明显增加,叶绿素a与b的比值(Chla／Chlb)升高,类胡萝卜素与叶绿素的比值(Car／Chl)降低,说明外源甜菜碱有利于植物对光能的捕获、吸收、传递和转换,提高叶片的光合活性,降低盐胁迫对植物的抑制作用。     

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 CO2 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 QB-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 QB-non-reductive PSⅡ RC could only be partly effective at the early time of photoinhibition, thus led to the accumulation of QB-non-reductive PSⅡ RC in the course of restoration under dim light.