Photodamage to photosystem in a typically shade-tolerant species Panax notoginseng exposed to a sudden increase in light intensity
Hong-Min WU,1,2, Sheng-Pu SHUANG1,2, Jin-Yan ZHANG1,2, Zhu CUN1,2, Zhen-Gui MENG1,2, Long-Gen LI1,2, Ben-Cai SHA1, Jun-Wen CHEN,1,2,*1National & Local Joint Engineering Research Center on Germplasm Innovation & Utilization of Chinese Medicinal Materials in Southwestern China, Key Laboratory of Medicinal Plant Biology of Yunnan Proince, Yunnan Agricultural University, Kunming 650201, China 2College of Agronomy and Biotechnology, Yunnan Agricultural University, Kunming 650201, China
National Natural Science Foundation of China(81860676) National Natural Science Foundation of China(81360609) Key Science and Technology Project of Yunnan Province(2016ZF001) Key Science and Technology Project of Yunnan Province(2017ZF001)
Abstract Aims Photodamage to a shade-tolerant species is common due to a sudden increase in growth light intensity. However, it is unknown about the underlying mechanism of the sensitivity of the shade-tolerant species to high light. The objective of the present study was to elucidate the mechanisms involved in the inability of the typically shade-tolerant species Panax notoginseng to survive under natural full-light condition. Methods The relative chlorophyll content (SPAD), photosynthetic parameters and chlorophyll fluorescence parameters were continuously examined in P. notoginseng when transferred from shade (10% of full sunlight) to full sunlight for three days. Important findings The net photosynthetic rate (Pn) of P. notoginseng exposed to full sunlight condition showed a “double-peaked” diurnal curve, and Pn decreased with the prolonged days of full light treatment. The SPAD value, water utilization efficiency and light use efficiency were significantly decreased under full sunlight condition. Furthermore, the maximum fluorescence signal of the P700 reaction center, electron transfer rate of photosystem II (PSII), the maximum quantum efficiency of PSII under dark adaptation and maximum quantum efficiency of PSII under light adaptation were significantly lower in full sunlight than those under shading condition, while the fraction of energy passively dissipated in the forms of heat and fluorescence, energy dissipation due to acceptor side limitation of PSI, and cyclic electron flow were significantly higher under the full light condition. Moreover, the sudden increase in growth light intensity caused a significant change in the fluorescence induction kinetic curve and significantly increased the fluorescence yield on the donor and acceptor side of PSII. The oxygen-evolving complex activity in the donor side of PSII was impaired under full sunlight. Furthermore, the electron transfer in the acceptor side of PSII was inhibited and the over-reduction of the acceptor side of PSI was caused by PSI photoinhibition. The study reveals that the full sunlight might induce the irreversible damage to PSII and the moderate photoinhibition to PSI in shade-tolerant species, and it may be an important underlying mechanisms why the shade-tolerant speciesP. notoginseng cannot survive under full sunlight. Keywords:shade-tolerant species;full sunlight;photosynthesis;photosystem;photoprotection;Panax notoginseng
PDF (1348KB)元数据多维度评价相关文章导出EndNote|Ris|Bibtex收藏本文 引用本文 武洪敏, 双升普, 张金燕, 寸竹, 孟珍贵, 李龙根, 沙本才, 陈军文. 短期生长环境光强骤增导致典型阴生植物三七光系统受损的机制. 植物生态学报, 2021, 45(4): 404-419. DOI: 10.17521/cjpe.2021.0013 WU Hong-Min, SHUANG Sheng-Pu, ZHANG Jin-Yan, CUN Zhu, MENG Zhen-Gui, LI Long-Gen, SHA Ben-Cai, CHEN Jun-Wen. Photodamage to photosystem in a typically shade-tolerant species Panax notoginseng exposed to a sudden increase in light intensity . Chinese Journal of Plant Ecology, 2021, 45(4): 404-419. DOI: 10.17521/cjpe.2021.0013
Fig. 3Diurnal pattern of relative chlorophyll content (SPAD value) in shade treatment and full sunlight treatment (mean ± SE, n = 7). *, significant difference between the two treatments at same measurement time ( p< 0.05).
Fig. 4Diurnal pattern of net photosynthetic rate (A), intercellular CO2 concentration (B) and stomatal conductance (C) in shade treatment and full sunlight treatment (mean ± SE, n = 7). *, significant difference in gas exchange parameters between the two treatments at same measurement time ( p< 0.05).
Fig. 5Water use efficiency (A) and light use efficiency (B) of Panax notoginseng exposed to shade and full sunlight conditions on the first, second and third day of the experiment, respectively (mean ± SE, n = 7). Different lowercase letters between shade and full sunlight conditions indicate significant difference (p< 0.05).
Fig. 6Diurnal pattern of maximum quantum efficiency under light adaptation (A), maximum quantum efficiency under dark adaptation (B) and non-photochemical quenching of PSII (C) in shade treatment and full sunlight treatment (mean ± SE, n = 7). *, significant difference between the two treatments at same measurement time ( p< 0.05).
Fig. 7The maximum quantum efficiency under dark adaptation (A), potential activity of PSII reaction center (B) and the maximum fluorescence signal of the P700 reaction center (C) of Panax notoginseng exposed to shade and full sunlight conditions on the first, second and third day of the experiment, respectively (mean ± SE, n = 7). *, significant difference between the two treatments at same measurement day ( p< 0.05).
Fig. 8Changes in rapid fluorescence induction kinetic curves (WO-J and Δ WO-J) in leaves of Panax notoginseng transferred from a shade environment with 10% transmittance to full sunlight. The abscissa is plotted on a linear time scale. W,the characteristic point at 100 μs (K point). Day 0, Sunlight-day 1, Sunlight-day 2, Sunlight-day 3 represent the exposure to low light, and the transfer to full sunlight for one, two and three days. Different lowercase letters indicate significant differences ( p< 0.05).
Fig. 9Daily pattern of parameters related to the rapid fluorescence induction kinetic curve in leaves of Panax notoginsengtransferred from a shade environment with 10% transmittance to full sunlight (mean ± SE, n = 7). Different letters indicate significant difference between shade and full sunlight conditions (p< 0.05).
Fig. 10Effect of shade and full sunlight treatment on electron transfer and photosystem energy partitioning (mean ± SE, n = 7). LL, 10% transmittance-grown Panax notoginseng (230 μmol·m -2·s-1, 13:00); HL, full sunlight-grown Panax notoginseng (2 300 μmol·m -2·s-1, 13:00). Y(I), effective quantum yield of PSI; Y(ND), heat dissipation efficiency at the donors quantum yield of PSI; Y(NA), quantum yield of PSI non-photochemical energy dissipation due to acceptor; Y(II), effective quantum yield of PSII; Y(NPQ), quantum yield of energy dissipation in PSII; Y(NO), fraction of energy passively dissipated in forms of heat and fluorescence. ETR(I), electron transport rate of PSI; ETR(II), electron transport rate of PSII;CEF, cyclic electron flow around PSI. *, significant difference between the two treatments at same measurement day ( p< 0.05).
Fig. 11Changes in electron transfer ratio in leaves of Panax notoginseng exposed to shade and full sunlight conditions on the first, second and third day of the experiment, respectively (mean ± SE, n = 7). ETR(I), electron transport rate of PSI; ETR(II), electron transport rate of PSII;CEF, cyclic electron flow around PSI. *, significant difference between the two treatments ( p< 0.05).
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ZhengF, LiZJ, QiuZJ, ZhaoHB, ZhouGY ( 2020). Effects of understory light on functional traits of evergreen broad- leaved forest saplings in Nanling Mountains, Guangdong Province Acta Ecologica Sinica, 40, 4516-4527. [本文引用: 1]
ZivcakM, BresticM, KunderlikovaK, SytarO, AllakhverdievSI (2015). Repetitive light pulse-induced photoinhibition of photosystem I severely affects CO2 assimilation and photoprotection in wheat leaves Photosynthesis Research, 126, 449-463. DOI:10.1007/s11120-015-0121-1PMID:25829027 [本文引用: 1] It was previously found that photosystem I (PSI) photoinhibition represents mostly irreversible damage with a slow recovery; however, its physiological significance has not been sufficiently characterized. The aim of the study was to assess the effect of PSI photoinhibition on photosynthesis in vivo. The inactivation of PSI was done by a series of short light saturation pulses applied by fluorimeter in darkness (every 10 s for 15 min), which led to decrease of both PSI (~60 %) and photosystem II (PSII) (~15 %) photochemical activity. No PSI recovery was observed within 2 days, whereas the PSII was fully recovered. Strongly limited PSI electron transport led to an imbalance between PSII and PSI photochemistry, with a high excitation pressure on PSII acceptor side and low oxidation of the PSI donor side. Low and delayed light-induced NPQ and P700(+) rise in inactivated samples indicated a decrease in formation of transthylakoid proton gradient (ΔpH), which was confirmed also by analysis of electrochromic bandshift (ECSt) records. In parallel with photochemical parameters, the CO2 assimilation was also strongly inhibited, more in low light (~70 %) than in high light (~45 %); the decrease was not caused by stomatal closure. PSI electron transport limited the CO2 assimilation at low to moderate light intensities, but it seems not to be directly responsible for a low CO2 assimilation at high light. In this regard, the possible effects of PSI photoinhibition on the redox signaling in chloroplast and its role in downregulation of Calvin cycle activity are discussed. Systematic analysis of the relation of electron transport and ATP synthesis to the photodamage and repair of photosystem II in Synechocystis 1 2005
... 自然环境中生长的植物, 其PSI通常对高光环境不敏感(Barth et al., 2001 ).一般只有在低温弱光、常温频闪光或是饱和脉冲处理植物时, PSI才会发生光抑制(Zhang & Scheller, 2004; 梁英等, 2018; Yamamoto & Shikanai, 2019).而滇南九节(Psychotria henryi)和镰叶肾蕨(Nephrolepis falciformis)等喜阴植物叶片的PSI却在恒定高光处理下发生光抑制(Huang et al., 2017 , 2018a).PSI发生光抑制的显著特征是PSI活性下降(Huang et al., 2018b ; 梁英等, 2018).Huang等(2018b)研究发现, 三七在高光下处理30和60 min时, Pm不受影响, 说明三七的PSI对高光胁迫不敏感.但本研究中, 短时环境强光导致三七的Pm显著下降, 该结果与上述的结果不一致, 可能是因为本研究采用的是全日照下的全天高光连续处理3天, 而Huang等(2018b)研究的高光处理时间较短(处理时间仅为30和60 min)从而尚未影响到PSI的活性.另有研究表明, Y(NA)增加也可作为PSI发生光抑制的指标(Kim et al., 2005 ; Yamori et al., 2016 ; Yang et al., 2019a ).本研究中, 正午强光下三七具有较高的Y(NA)(图10A), 该结果与拟南芥(Arabidopsis thaliana)和白及(Bletilla striata)从低到高光转变后Y(NA)升高的结果(Huang et al., 2019 ; Yang et al., 2019a )一致, 由此推测, 环境光强增加后, 因无法形成足够的pH, 类囊体管腔酸化水平低, 从PSII到PSI的电子流引起受体侧羟基自由基的积累, 最终导致PSI发生光抑制(黄伟, 2012; Yang et al., 2019a ).这说明, 短时环境光强的骤增导致典型阴生植物三七的PSI发生光抑制.与PSII不同, PSI发生光抑制通常需要一周或更长的时间才能得以恢复, 若是造成光损伤则会直接导致植株的死亡(Zivcaket al., 2015 ; Yang et al., 2019a ).因此, 植物必须具有可行的机制来保护PSI在波动光下不受损.围绕PSI的循环电子流对PSI起到重要的光保护作用, 这在从弱光转到强光环境下的九节、三七和高盆樱桃(Cerasus cerasoides)中均得到了证实(Huang et al., 2015b , 2018b; Yang et al., 2019b ).本研究中, 短期环境光照骤增导致围绕PSI的循环电子流被大量激活(图10, 图11), 不仅接收了来自PSI的电子, 还能使P700保持高氧化态以保护PSI不受损(黄伟等, 2012; Huang et al., 2018c ).但围绕PSI的循环电子流的激发也不能完全保护强光下的PSI免受光抑制. ...
Steady-state and dynamic photosynthetic performance and nitrogen partitioning in the shade-demanding plant Panax notoginseng under different levels of growth irradiance 2 2014
Photosynthesis, light energy partitioning, and photoprotection in the shade-demanding species Panax notoginseng under high and low level of growth irradiance 1 2016
Photosynthetic acclimation to light in woody and herbaceous species: a comparison of leaf structure, pigment content and chlorophyll fluorescence characteristics measured in the field 2 2012
... 自然环境中生长的植物, 其PSI通常对高光环境不敏感(Barth et al., 2001 ).一般只有在低温弱光、常温频闪光或是饱和脉冲处理植物时, PSI才会发生光抑制(Zhang & Scheller, 2004; 梁英等, 2018; Yamamoto & Shikanai, 2019).而滇南九节(Psychotria henryi)和镰叶肾蕨(Nephrolepis falciformis)等喜阴植物叶片的PSI却在恒定高光处理下发生光抑制(Huang et al., 2017 , 2018a).PSI发生光抑制的显著特征是PSI活性下降(Huang et al., 2018b ; 梁英等, 2018).Huang等(2018b)研究发现, 三七在高光下处理30和60 min时, Pm不受影响, 说明三七的PSI对高光胁迫不敏感.但本研究中, 短时环境强光导致三七的Pm显著下降, 该结果与上述的结果不一致, 可能是因为本研究采用的是全日照下的全天高光连续处理3天, 而Huang等(2018b)研究的高光处理时间较短(处理时间仅为30和60 min)从而尚未影响到PSI的活性.另有研究表明, Y(NA)增加也可作为PSI发生光抑制的指标(Kim et al., 2005 ; Yamori et al., 2016 ; Yang et al., 2019a ).本研究中, 正午强光下三七具有较高的Y(NA)(图10A), 该结果与拟南芥(Arabidopsis thaliana)和白及(Bletilla striata)从低到高光转变后Y(NA)升高的结果(Huang et al., 2019 ; Yang et al., 2019a )一致, 由此推测, 环境光强增加后, 因无法形成足够的pH, 类囊体管腔酸化水平低, 从PSII到PSI的电子流引起受体侧羟基自由基的积累, 最终导致PSI发生光抑制(黄伟, 2012; Yang et al., 2019a ).这说明, 短时环境光强的骤增导致典型阴生植物三七的PSI发生光抑制.与PSII不同, PSI发生光抑制通常需要一周或更长的时间才能得以恢复, 若是造成光损伤则会直接导致植株的死亡(Zivcaket al., 2015 ; Yang et al., 2019a ).因此, 植物必须具有可行的机制来保护PSI在波动光下不受损.围绕PSI的循环电子流对PSI起到重要的光保护作用, 这在从弱光转到强光环境下的九节、三七和高盆樱桃(Cerasus cerasoides)中均得到了证实(Huang et al., 2015b , 2018b; Yang et al., 2019b ).本研究中, 短期环境光照骤增导致围绕PSI的循环电子流被大量激活(图10, 图11), 不仅接收了来自PSI的电子, 还能使P700保持高氧化态以保护PSI不受损(黄伟等, 2012; Huang et al., 2018c ).但围绕PSI的循环电子流的激发也不能完全保护强光下的PSI免受光抑制. ...
Photosystem I acceptor side limitation is a prerequisite for the reversible decrease in the maximum extent of P700 oxidation after short-term chilling in the light in four plant species with different chilling sensitivities 1 2005
... 自然环境中生长的植物, 其PSI通常对高光环境不敏感(Barth et al., 2001 ).一般只有在低温弱光、常温频闪光或是饱和脉冲处理植物时, PSI才会发生光抑制(Zhang & Scheller, 2004; 梁英等, 2018; Yamamoto & Shikanai, 2019).而滇南九节(Psychotria henryi)和镰叶肾蕨(Nephrolepis falciformis)等喜阴植物叶片的PSI却在恒定高光处理下发生光抑制(Huang et al., 2017 , 2018a).PSI发生光抑制的显著特征是PSI活性下降(Huang et al., 2018b ; 梁英等, 2018).Huang等(2018b)研究发现, 三七在高光下处理30和60 min时, Pm不受影响, 说明三七的PSI对高光胁迫不敏感.但本研究中, 短时环境强光导致三七的Pm显著下降, 该结果与上述的结果不一致, 可能是因为本研究采用的是全日照下的全天高光连续处理3天, 而Huang等(2018b)研究的高光处理时间较短(处理时间仅为30和60 min)从而尚未影响到PSI的活性.另有研究表明, Y(NA)增加也可作为PSI发生光抑制的指标(Kim et al., 2005 ; Yamori et al., 2016 ; Yang et al., 2019a ).本研究中, 正午强光下三七具有较高的Y(NA)(图10A), 该结果与拟南芥(Arabidopsis thaliana)和白及(Bletilla striata)从低到高光转变后Y(NA)升高的结果(Huang et al., 2019 ; Yang et al., 2019a )一致, 由此推测, 环境光强增加后, 因无法形成足够的pH, 类囊体管腔酸化水平低, 从PSII到PSI的电子流引起受体侧羟基自由基的积累, 最终导致PSI发生光抑制(黄伟, 2012; Yang et al., 2019a ).这说明, 短时环境光强的骤增导致典型阴生植物三七的PSI发生光抑制.与PSII不同, PSI发生光抑制通常需要一周或更长的时间才能得以恢复, 若是造成光损伤则会直接导致植株的死亡(Zivcaket al., 2015 ; Yang et al., 2019a ).因此, 植物必须具有可行的机制来保护PSI在波动光下不受损.围绕PSI的循环电子流对PSI起到重要的光保护作用, 这在从弱光转到强光环境下的九节、三七和高盆樱桃(Cerasus cerasoides)中均得到了证实(Huang et al., 2015b , 2018b; Yang et al., 2019b ).本研究中, 短期环境光照骤增导致围绕PSI的循环电子流被大量激活(图10, 图11), 不仅接收了来自PSI的电子, 还能使P700保持高氧化态以保护PSI不受损(黄伟等, 2012; Huang et al., 2018c ).但围绕PSI的循环电子流的激发也不能完全保护强光下的PSI免受光抑制. ...
Relative importance of photosynthetic traits and allocation patterns as correlates of seedling shade tolerance of 13 tropical trees 1 1994
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 1 2005
Overexpressed superoxide dismutase and catalase act synergistically to protect the repair of PSII during photoinhibition in Synechococcus elongatus PCC 7942 1 2016
Regulation of antenna structure and electron transport in Photosystem II of Pisum sativum under elevated temperature probed by the fast polyphasic chlorophyll a fluorescence transient: OKJIP 1 1997
Stimulation of cyclic electron flow around photosystem I upon a sudden transition from low to high light in two angiosperms Arabidopsis thaliana and Bletilla striata 4 2019a
... 自然环境中生长的植物, 其PSI通常对高光环境不敏感(Barth et al., 2001 ).一般只有在低温弱光、常温频闪光或是饱和脉冲处理植物时, PSI才会发生光抑制(Zhang & Scheller, 2004; 梁英等, 2018; Yamamoto & Shikanai, 2019).而滇南九节(Psychotria henryi)和镰叶肾蕨(Nephrolepis falciformis)等喜阴植物叶片的PSI却在恒定高光处理下发生光抑制(Huang et al., 2017 , 2018a).PSI发生光抑制的显著特征是PSI活性下降(Huang et al., 2018b ; 梁英等, 2018).Huang等(2018b)研究发现, 三七在高光下处理30和60 min时, Pm不受影响, 说明三七的PSI对高光胁迫不敏感.但本研究中, 短时环境强光导致三七的Pm显著下降, 该结果与上述的结果不一致, 可能是因为本研究采用的是全日照下的全天高光连续处理3天, 而Huang等(2018b)研究的高光处理时间较短(处理时间仅为30和60 min)从而尚未影响到PSI的活性.另有研究表明, Y(NA)增加也可作为PSI发生光抑制的指标(Kim et al., 2005 ; Yamori et al., 2016 ; Yang et al., 2019a ).本研究中, 正午强光下三七具有较高的Y(NA)(图10A), 该结果与拟南芥(Arabidopsis thaliana)和白及(Bletilla striata)从低到高光转变后Y(NA)升高的结果(Huang et al., 2019 ; Yang et al., 2019a )一致, 由此推测, 环境光强增加后, 因无法形成足够的pH, 类囊体管腔酸化水平低, 从PSII到PSI的电子流引起受体侧羟基自由基的积累, 最终导致PSI发生光抑制(黄伟, 2012; Yang et al., 2019a ).这说明, 短时环境光强的骤增导致典型阴生植物三七的PSI发生光抑制.与PSII不同, PSI发生光抑制通常需要一周或更长的时间才能得以恢复, 若是造成光损伤则会直接导致植株的死亡(Zivcaket al., 2015 ; Yang et al., 2019a ).因此, 植物必须具有可行的机制来保护PSI在波动光下不受损.围绕PSI的循环电子流对PSI起到重要的光保护作用, 这在从弱光转到强光环境下的九节、三七和高盆樱桃(Cerasus cerasoides)中均得到了证实(Huang et al., 2015b , 2018b; Yang et al., 2019b ).本研究中, 短期环境光照骤增导致围绕PSI的循环电子流被大量激活(图10, 图11), 不仅接收了来自PSI的电子, 还能使P700保持高氧化态以保护PSI不受损(黄伟等, 2012; Huang et al., 2018c ).但围绕PSI的循环电子流的激发也不能完全保护强光下的PSI免受光抑制. ...
... ; Yang et al., 2019a )一致, 由此推测, 环境光强增加后, 因无法形成足够的pH, 类囊体管腔酸化水平低, 从PSII到PSI的电子流引起受体侧羟基自由基的积累, 最终导致PSI发生光抑制(黄伟, 2012; Yang et al., 2019a ).这说明, 短时环境光强的骤增导致典型阴生植物三七的PSI发生光抑制.与PSII不同, PSI发生光抑制通常需要一周或更长的时间才能得以恢复, 若是造成光损伤则会直接导致植株的死亡(Zivcaket al., 2015 ; Yang et al., 2019a ).因此, 植物必须具有可行的机制来保护PSI在波动光下不受损.围绕PSI的循环电子流对PSI起到重要的光保护作用, 这在从弱光转到强光环境下的九节、三七和高盆樱桃(Cerasus cerasoides)中均得到了证实(Huang et al., 2015b , 2018b; Yang et al., 2019b ).本研究中, 短期环境光照骤增导致围绕PSI的循环电子流被大量激活(图10, 图11), 不仅接收了来自PSI的电子, 还能使P700保持高氧化态以保护PSI不受损(黄伟等, 2012; Huang et al., 2018c ).但围绕PSI的循环电子流的激发也不能完全保护强光下的PSI免受光抑制. ...
... ; Yang et al., 2019a ).这说明, 短时环境光强的骤增导致典型阴生植物三七的PSI发生光抑制.与PSII不同, PSI发生光抑制通常需要一周或更长的时间才能得以恢复, 若是造成光损伤则会直接导致植株的死亡(Zivcaket al., 2015 ; Yang et al., 2019a ).因此, 植物必须具有可行的机制来保护PSI在波动光下不受损.围绕PSI的循环电子流对PSI起到重要的光保护作用, 这在从弱光转到强光环境下的九节、三七和高盆樱桃(Cerasus cerasoides)中均得到了证实(Huang et al., 2015b , 2018b; Yang et al., 2019b ).本研究中, 短期环境光照骤增导致围绕PSI的循环电子流被大量激活(图10, 图11), 不仅接收了来自PSI的电子, 还能使P700保持高氧化态以保护PSI不受损(黄伟等, 2012; Huang et al., 2018c ).但围绕PSI的循环电子流的激发也不能完全保护强光下的PSI免受光抑制. ...
... ; Yang et al., 2019a ).因此, 植物必须具有可行的机制来保护PSI在波动光下不受损.围绕PSI的循环电子流对PSI起到重要的光保护作用, 这在从弱光转到强光环境下的九节、三七和高盆樱桃(Cerasus cerasoides)中均得到了证实(Huang et al., 2015b , 2018b; Yang et al., 2019b ).本研究中, 短期环境光照骤增导致围绕PSI的循环电子流被大量激活(图10, 图11), 不仅接收了来自PSI的电子, 还能使P700保持高氧化态以保护PSI不受损(黄伟等, 2012; Huang et al., 2018c ).但围绕PSI的循环电子流的激发也不能完全保护强光下的PSI免受光抑制. ...
Photosynthetic regulation under fluctuating light in field-grownCerasus cerasoides: a comparison of young and mature leaves 1 2019b
... 自然环境中生长的植物, 其PSI通常对高光环境不敏感(Barth et al., 2001 ).一般只有在低温弱光、常温频闪光或是饱和脉冲处理植物时, PSI才会发生光抑制(Zhang & Scheller, 2004; 梁英等, 2018; Yamamoto & Shikanai, 2019).而滇南九节(Psychotria henryi)和镰叶肾蕨(Nephrolepis falciformis)等喜阴植物叶片的PSI却在恒定高光处理下发生光抑制(Huang et al., 2017 , 2018a).PSI发生光抑制的显著特征是PSI活性下降(Huang et al., 2018b ; 梁英等, 2018).Huang等(2018b)研究发现, 三七在高光下处理30和60 min时, Pm不受影响, 说明三七的PSI对高光胁迫不敏感.但本研究中, 短时环境强光导致三七的Pm显著下降, 该结果与上述的结果不一致, 可能是因为本研究采用的是全日照下的全天高光连续处理3天, 而Huang等(2018b)研究的高光处理时间较短(处理时间仅为30和60 min)从而尚未影响到PSI的活性.另有研究表明, Y(NA)增加也可作为PSI发生光抑制的指标(Kim et al., 2005 ; Yamori et al., 2016 ; Yang et al., 2019a ).本研究中, 正午强光下三七具有较高的Y(NA)(图10A), 该结果与拟南芥(Arabidopsis thaliana)和白及(Bletilla striata)从低到高光转变后Y(NA)升高的结果(Huang et al., 2019 ; Yang et al., 2019a )一致, 由此推测, 环境光强增加后, 因无法形成足够的pH, 类囊体管腔酸化水平低, 从PSII到PSI的电子流引起受体侧羟基自由基的积累, 最终导致PSI发生光抑制(黄伟, 2012; Yang et al., 2019a ).这说明, 短时环境光强的骤增导致典型阴生植物三七的PSI发生光抑制.与PSII不同, PSI发生光抑制通常需要一周或更长的时间才能得以恢复, 若是造成光损伤则会直接导致植株的死亡(Zivcaket al., 2015 ; Yang et al., 2019a ).因此, 植物必须具有可行的机制来保护PSI在波动光下不受损.围绕PSI的循环电子流对PSI起到重要的光保护作用, 这在从弱光转到强光环境下的九节、三七和高盆樱桃(Cerasus cerasoides)中均得到了证实(Huang et al., 2015b , 2018b; Yang et al., 2019b ).本研究中, 短期环境光照骤增导致围绕PSI的循环电子流被大量激活(图10, 图11), 不仅接收了来自PSI的电子, 还能使P700保持高氧化态以保护PSI不受损(黄伟等, 2012; Huang et al., 2018c ).但围绕PSI的循环电子流的激发也不能完全保护强光下的PSI免受光抑制. ...
Photosynthetic characteristics and light energy conversions under different light environments in five tree species occupying dominant status at different stages of subtropical forest succession 1 2015