大豆叶片衰老过程中PSⅡ功能和光破坏防御机制的变化
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摘要
本文利用大豆叶片为实验材料，通过测定叶绿素荧光参数、气体交换、蛋白及色素等，着重研究了其衰老过程中PSII功能及光破坏防御机制的动态变化规律。主要结果如下：
1．随着衰老的进行，光呼吸与光合都呈下降趋势，米尔反应却呈波动式变化，用乙烯利诱导衰老的前3天内，依赖米尔反应的电子传递逐渐上升，3天后又逐渐下降，但在整个衰老过程中依赖米尔反应的电子传递与PSII中总电子传递的比值不断增加。
2．NPQ，qf和(Z+A)/(Z+A+V)在整个衰老过程中并不呈现单一变化趋势。用乙烯利诱导衰老的前3天内，虽然PSII实际光化学效率（ФPSII）下降，但NPQ，qf和(Z+A)/(Z+A+V)逐渐增加；诱导衰老3天后，这3个参数又呈下降趋势。在整个衰老过程中，qI一直呈现上升趋势。
3．在衰老初期，虽然ФPSII和qP下降了，但叶片中过剩光能（通过光化学途径和依赖NPQ耗散后所剩的光能，用 (1-qP)/NPQ来衡量）与对照（功能叶）相比变化不大；然而在极度衰老叶片中，过剩光能急剧增加。
4．在光系统II中，随着衰老，PSII原初光化学反应能力逐渐下降，而QA的氧化还原程度却逐渐增加。另外通过瞬时荧光曲线和Vt曲线可知衰老叶片中PSII反应中心QB位点发生了变化，并且我们认为该位点的变化导致了QA到QB的电子传递受到抑制（受体端抑制），并最终引起PSII原初光化学反应的降低。
5．单位激发截面所吸收的光量子（ABS/CSo）以及用于电子传递的光量子(ETO/CSO)随着衰老也逐渐下降。用乙烯利诱导衰老过程中，单位激发截面活性反应中心数目（RC/CSo）逐渐下降，相反单位激发截面所
耗散的光量子（DIO/CSO）逐渐增加。
6. 随着衰老，虽然PSII平均原初光化学反应降低，但单位反应中心捕获的光量子（TRo/RC）和用于电子传递的光量子(ETo/RC)增加了，这可能是由于失活反应中心将其捕获的激发能传给具有活性的反应中心并导致电荷分离所致。
7．叶绿素，可溶性蛋白，类囊体膜脂脂肪酸不饱和度和紫黄质脱环氧化酶（VDE）蛋白量在衰老过程中逐渐下降。
8．自然光照条件下，田间大豆叶片光抑制主要是由PSII反应中心可逆失活引起的，但可逆失活并没有随衰老增加。
总之，用乙烯利诱导大豆叶片衰老过程中，其PSII反应中心的状态和功能发生了变化，并导致了受体端抑制；在此过程中，光破坏防御机制也随之发生变化，与叶黄素循环有关的耗散并不能自始至终地起到重要的耗散作用，叶片热耗散位点也由天线逐渐转移到反应中心。
By measurement of chlorophyll fluorescence, gas exchange, protein and pigment, etc., changes of PSII function and photoprotection mechanism in soybean leaves during senescence were investigated in this paper. The main results are as follows:
1. With the senescence in soybean leaves, photorespiration and photosynthesis decreased markedly. In contrast, Mehler reaction dependent electron transport increased during the first three days of ethylene treatment then decreased to a low level after 3 days of ethylene treatment, and the ratio of Mehler reaction dependent electron transport to the total electron transport increased throughout the senescence.
2. NPQ, qf and (Z+A)/(Z+A+V) exhibited two distinct phases, with an initial increase (during the first 3 days) followed by a dramatically decrease after 3 days of ethylene treatment during senescence. However, increase of qI was observed from beginning to the end.
At the beginning of senescence, (1-qP)/NPQ (used to estimate the fraction of excess photons) showed a little difference compared to the control; however, it increased to a high level when leaf became
3. severely senescent.
4. In PSII, the decline of primary photochemical activity was observed during senescence. The rate and extent of QA reduction increased markedly. The QB site was changed reflected by the fluorescence transient and Vt curve, which was the key factor leading to inhibition of electron transporting to QB from QA resulting in the decline of primary photochemistry.
5. Photons absorbed by per excited cross-section (ABS/CSo) and used to move electron transport beyond QA- per excited cross-section (ETO/ CSo) decreased throughout the senescence. Accompanied with the decline in the density of active reaction center per excited cross-section (RC/CSO), the dissipated flux per excited cross-section (DIO/CSO) was enhanced.
6. It was clear to observe that the fraction of photons used to move electron transport beyond QA- per reaction center (ETO/RC) and trapped photons per reaction center(TRo/RC) increased markedly during the senescence, which was attribute to that the absorbed photons by inactive reaction center was transported to active reaction centers and resulted in charge separation.
7. Besides, chlorophyll content, soluble protein, thylakoid membrane lipids fatty acid instauration and activity of Violaxanthin de-epoxidase（VDE）were all decreased with the progress of senescence.
Photoinhibition in soybean leaves grown in the field was mainly attributed to reversible inactivation of PSII reaction center. However,
8. the inactivated degree did not increase with the senescence.
In summary, structure and function of PSII changed, and this resulted in inhibition of PSII acceptor side in soybean leaves during senescence induced by ethylene. Photoprotective mechanisms exhibited dynamic change: xanthophyll cycle related thermal dissipation showed a biphasic change, with an initial increase followed a final decrease, throughout the senescence; energy quenching site shifted gradually from antenna to reaction center.
1．随着衰老的进行，光呼吸与光合都呈下降趋势，米尔反应却呈波动式变化，用乙烯利诱导衰老的前3天内，依赖米尔反应的电子传递逐渐上升，3天后又逐渐下降，但在整个衰老过程中依赖米尔反应的电子传递与PSII中总电子传递的比值不断增加。
2．NPQ，qf和(Z+A)/(Z+A+V)在整个衰老过程中并不呈现单一变化趋势。用乙烯利诱导衰老的前3天内，虽然PSII实际光化学效率（ФPSII）下降，但NPQ，qf和(Z+A)/(Z+A+V)逐渐增加；诱导衰老3天后，这3个参数又呈下降趋势。在整个衰老过程中，qI一直呈现上升趋势。
3．在衰老初期，虽然ФPSII和qP下降了，但叶片中过剩光能（通过光化学途径和依赖NPQ耗散后所剩的光能，用 (1-qP)/NPQ来衡量）与对照（功能叶）相比变化不大；然而在极度衰老叶片中，过剩光能急剧增加。
4．在光系统II中，随着衰老，PSII原初光化学反应能力逐渐下降，而QA的氧化还原程度却逐渐增加。另外通过瞬时荧光曲线和Vt曲线可知衰老叶片中PSII反应中心QB位点发生了变化，并且我们认为该位点的变化导致了QA到QB的电子传递受到抑制（受体端抑制），并最终引起PSII原初光化学反应的降低。
5．单位激发截面所吸收的光量子（ABS/CSo）以及用于电子传递的光量子(ETO/CSO)随着衰老也逐渐下降。用乙烯利诱导衰老过程中，单位激发截面活性反应中心数目（RC/CSo）逐渐下降，相反单位激发截面所
耗散的光量子（DIO/CSO）逐渐增加。
6. 随着衰老，虽然PSII平均原初光化学反应降低，但单位反应中心捕获的光量子（TRo/RC）和用于电子传递的光量子(ETo/RC)增加了，这可能是由于失活反应中心将其捕获的激发能传给具有活性的反应中心并导致电荷分离所致。
7．叶绿素，可溶性蛋白，类囊体膜脂脂肪酸不饱和度和紫黄质脱环氧化酶（VDE）蛋白量在衰老过程中逐渐下降。
8．自然光照条件下，田间大豆叶片光抑制主要是由PSII反应中心可逆失活引起的，但可逆失活并没有随衰老增加。
总之，用乙烯利诱导大豆叶片衰老过程中，其PSII反应中心的状态和功能发生了变化，并导致了受体端抑制；在此过程中，光破坏防御机制也随之发生变化，与叶黄素循环有关的耗散并不能自始至终地起到重要的耗散作用，叶片热耗散位点也由天线逐渐转移到反应中心。
By measurement of chlorophyll fluorescence, gas exchange, protein and pigment, etc., changes of PSII function and photoprotection mechanism in soybean leaves during senescence were investigated in this paper. The main results are as follows:
1. With the senescence in soybean leaves, photorespiration and photosynthesis decreased markedly. In contrast, Mehler reaction dependent electron transport increased during the first three days of ethylene treatment then decreased to a low level after 3 days of ethylene treatment, and the ratio of Mehler reaction dependent electron transport to the total electron transport increased throughout the senescence.
2. NPQ, qf and (Z+A)/(Z+A+V) exhibited two distinct phases, with an initial increase (during the first 3 days) followed by a dramatically decrease after 3 days of ethylene treatment during senescence. However, increase of qI was observed from beginning to the end.
At the beginning of senescence, (1-qP)/NPQ (used to estimate the fraction of excess photons) showed a little difference compared to the control; however, it increased to a high level when leaf became
3. severely senescent.
4. In PSII, the decline of primary photochemical activity was observed during senescence. The rate and extent of QA reduction increased markedly. The QB site was changed reflected by the fluorescence transient and Vt curve, which was the key factor leading to inhibition of electron transporting to QB from QA resulting in the decline of primary photochemistry.
5. Photons absorbed by per excited cross-section (ABS/CSo) and used to move electron transport beyond QA- per excited cross-section (ETO/ CSo) decreased throughout the senescence. Accompanied with the decline in the density of active reaction center per excited cross-section (RC/CSO), the dissipated flux per excited cross-section (DIO/CSO) was enhanced.
6. It was clear to observe that the fraction of photons used to move electron transport beyond QA- per reaction center (ETO/RC) and trapped photons per reaction center(TRo/RC) increased markedly during the senescence, which was attribute to that the absorbed photons by inactive reaction center was transported to active reaction centers and resulted in charge separation.
7. Besides, chlorophyll content, soluble protein, thylakoid membrane lipids fatty acid instauration and activity of Violaxanthin de-epoxidase（VDE）were all decreased with the progress of senescence.
Photoinhibition in soybean leaves grown in the field was mainly attributed to reversible inactivation of PSII reaction center. However,
8. the inactivated degree did not increase with the senescence.
In summary, structure and function of PSII changed, and this resulted in inhibition of PSII acceptor side in soybean leaves during senescence induced by ethylene. Photoprotective mechanisms exhibited dynamic change: xanthophyll cycle related thermal dissipation showed a biphasic change, with an initial increase followed a final decrease, throughout the senescence; energy quenching site shifted gradually from antenna to reaction center.
引文
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Melis A, Homann PH. Heterogeneity of the photochemical centers in system II of chloroplasts. Photochem Photobiol 1976, 23: 343-350
Miller KW. Structural and functional responses of wheat mitochondrial membranes to growth at low temperature. Plant physiology 1974, 53: 426-433
Misra AN, Biswal UC. Differential changes in the electron properties of thylakoid membranes during aging of attached and detached leaves, and isolated chloroplasts. Plant Cell and Environment 1982, 51:27-30
Munné-Bosch, S. and Alegre, L. The xanthophyll cycle is induced by light irrespective of water status in field-grown lavender (Lavandula stoechas). Physiologica Plantarum 2000, 108: 147-151
Nedbal L, Masojidek J, Komenda J, Prasil O, Setlik I. Photosynth Res 1990, 24: 89-97
Neubauer C, Schreiber U. The polyphsic rise of chlorophyll fluorescence upon onset of strong continuous illumination. I.Saturation characteristics and partial control by the photosystem acceptor side. Zeitschrift für Naturforschung 1987, 42c:1246-1254
Neubauer C, Yamamoto HY. Mehler-peroxidase reactive mediates zeaxanthin formation and zeaxanthin-related fluorescence quenching in intact chloroplasts. Plant Physiol 1992, 99:1354－1361
Osmond CB. Photorespiration and photoinhibition, some implications for the energetics of photosynthesis. Biochim Biophys Acta 1981, 639:77-85
Osmond CB, Grace SC. Perspectives on photoinhibition and photorespiration in the field: quintessential inefficiencies of the light and dark reactions of photorespiration? Journal of Experimental Botany 1995, 46 :1351-1362
Owens TG, Shreve AP, Albrecht AC. Dynamics and mechanism of singlet energy transfer between carotenoids and chlorophylls: light harvesting and nonphotochemical fluorescence quenching. In Murata N(ed). Rearch in photosynthesis. Dordrecht: Kluwer Academic 1992, vol 5:179-186
?quist G, Chow WS, Anderson JM. Photoinhibition of photosynthesis represents a mechanism for long-term regulation of photosystem II. Planta. 1992, 186: 450-460
Patterson T G, Moss D N, Brun W A. Enzymatic changes during the senescence of field-grown wheat. Crop Sci. 1980, 20(1):15-18
Pfündel EE, Diley RA. The pH dependent of violaxanthin deepoxidase in isolated pea chloroplasts. Plant Physiol 1993, 101:65-71
Robert C B, Chang S-H, Yamamoto HY. Developmental expression of violaxanthin de-epoxidase in leaves of tobacco growing under high and low light. Plant Physiology 1999, 121: 207-213
Raven JA, Samuelsson G. Photoinhibition at low quantum flux densities in a marine dinoflagellate (Amphidinium carterae). Maringe Biology 1986, 70: 21-26
Schreiber U, Neubauer C. The polyphasic rise of chlorophyll fluorescence upon of strong continous illumination. II. Partial control by the photosystem IIdonor side and possible ways of interpretation. Zeitschrift für Naturforschung 1987, 42c:1255-1264
Setlik I, Allakhverdiev SJ, Nedbal L, Setlikovs E , Klimov VV. Photosynth Research. 1990, 23: 39-48
Shimazaki K, Ito K, Kondo N, Sugahara K. Reversible inhibition of the photosynthetic water-splitting enzyme system by SO2-fumigation assayed by chlorophyll fluorescence and EPR signal in vivo. Plant Cell Physiolgy 1984, 25:795-803
Siegenthaler PA, Rawyler A., Smutny J. The phospholipids population which sustains the uncoupled non-cyclic electron flow activity is located in
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Srivastava A and Strasser RJ. Stress and stress management of land plants during a regular day. Journal of Plant Physiology 1996, 148: 445-455
Srivastava A, Guisse B, Greppin H, Strasser RJ. Regulation of antenna structure and electron transport in PSII of Pisum Sativum under elevated temperature probed by the fast polyphasic chlorophyll a fluorescence transient OKJIP. Biochimica Biophysica Acta. 1997, 1320:95-106
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Strasser RJ. Donor side capacity of photosystem II probed by chlorophyll a fluorescence transient. Photosynth Res. 1997, 52:147-155
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Strasser BJ and Strasser RJ. Measuring fast fluorescence transients to address environmental questions: The JIP test. In mathis, P.(ed.) Photosynthesis: from light to Biosphere Vol 5: 977-980 Kluwer Academic, The Netherlands.1995
Strasser BJ, Srivastava A, Govindjee. Polyphasic chlorophyll a fluorescence transient in plants and cyanobacteria. Photochemistry and Photobiology 1995, 61:32-42
Thomas CP，Aubrey WN. A comparison of the effects of chilling on thylakoid electron transfer in Pea（Pisum sativun L.） and Cucumber（Cucumis sativus L.）. Plant physiology 1988，86：147～151
Telfer A, De Las Rivas J, Barber J. β-carotene within the isolated photosystem II reaction center: photooxidation and irreversible bleaching of this chromophose by oxidized p680. Biochim. Biophys Acta. 1991, 1060:106-114
Thomopson LK and Brudwig GW. Cytochrome b-559 may function to protect photosystem II from photoinhibition. Biochemistry 1988, 27: 6653-6658
Verhoven AS, Adams, III WW, and Demmig-adams B. Enhanced employment of the xanthophyll cycle and thermal energy dissipation in spinach exposed to high light and N stress. Plant Physiology 1997, 113, 817-824
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Yamamoto HY. Biochemistry of the violaxanthin cycle in higher plants. Pure Appl Chen. 1979, 51:639-648
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Woolhouse H W, Batt T In: Perspective in Experimental Biology. Ed: Sunderland N, Pergamon Press. Oxford 1976: 163-175
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Misra AN, Biswal UC. Differential changes in the electron properties of thylakoid membranes during aging of attached and detached leaves, and isolated chloroplasts. Plant Cell and Environment 1982, 51:27-30
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Neubauer C, Yamamoto HY. Mehler-peroxidase reactive mediates zeaxanthin formation and zeaxanthin-related fluorescence quenching in intact chloroplasts. Plant Physiol 1992, 99:1354－1361
Osmond CB. Photorespiration and photoinhibition, some implications for the energetics of photosynthesis. Biochim Biophys Acta 1981, 639:77-85
Osmond CB, Grace SC. Perspectives on photoinhibition and photorespiration in the field: quintessential inefficiencies of the light and dark reactions of photorespiration? Journal of Experimental Botany 1995, 46 :1351-1362
Owens TG, Shreve AP, Albrecht AC. Dynamics and mechanism of singlet energy transfer between carotenoids and chlorophylls: light harvesting and nonphotochemical fluorescence quenching. In Murata N(ed). Rearch in photosynthesis. Dordrecht: Kluwer Academic 1992, vol 5:179-186
?quist G, Chow WS, Anderson JM. Photoinhibition of photosynthesis represents a mechanism for long-term regulation of photosystem II. Planta. 1992, 186: 450-460
Patterson T G, Moss D N, Brun W A. Enzymatic changes during the senescence of field-grown wheat. Crop Sci. 1980, 20(1):15-18
Pfündel EE, Diley RA. The pH dependent of violaxanthin deepoxidase in isolated pea chloroplasts. Plant Physiol 1993, 101:65-71
Robert C B, Chang S-H, Yamamoto HY. Developmental expression of violaxanthin de-epoxidase in leaves of tobacco growing under high and low light. Plant Physiology 1999, 121: 207-213
Raven JA, Samuelsson G. Photoinhibition at low quantum flux densities in a marine dinoflagellate (Amphidinium carterae). Maringe Biology 1986, 70: 21-26
Schreiber U, Neubauer C. The polyphasic rise of chlorophyll fluorescence upon of strong continous illumination. II. Partial control by the photosystem IIdonor side and possible ways of interpretation. Zeitschrift für Naturforschung 1987, 42c:1255-1264
Setlik I, Allakhverdiev SJ, Nedbal L, Setlikovs E , Klimov VV. Photosynth Research. 1990, 23: 39-48
Shimazaki K, Ito K, Kondo N, Sugahara K. Reversible inhibition of the photosynthetic water-splitting enzyme system by SO2-fumigation assayed by chlorophyll fluorescence and EPR signal in vivo. Plant Cell Physiolgy 1984, 25:795-803
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Srivastava A and Strasser RJ. Stress and stress management of land plants during a regular day. Journal of Plant Physiology 1996, 148: 445-455
Srivastava A, Guisse B, Greppin H, Strasser RJ. Regulation of antenna structure and electron transport in PSII of Pisum Sativum under elevated temperature probed by the fast polyphasic chlorophyll a fluorescence transient OKJIP. Biochimica Biophysica Acta. 1997, 1320:95-106
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Strasser BJ and Strasser RJ. Measuring fast fluorescence transients to address environmental questions: The JIP test. In mathis, P.(ed.) Photosynthesis: from light to Biosphere Vol 5: 977-980 Kluwer Academic, The Netherlands.1995
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Thomas CP，Aubrey WN. A comparison of the effects of chilling on thylakoid electron transfer in Pea（Pisum sativun L.） and Cucumber（Cucumis sativus L.）. Plant physiology 1988，86：147～151
Telfer A, De Las Rivas J, Barber J. β-carotene within the isolated photosystem II reaction center: photooxidation and irreversible bleaching of this chromophose by oxidized p680. Biochim. Biophys Acta. 1991, 1060:106-114
Thomopson LK and Brudwig GW. Cytochrome b-559 may function to protect photosystem II from photoinhibition. Biochemistry 1988, 27: 6653-6658
Verhoven AS, Adams, III WW, and Demmig-adams B. Enhanced employment of the xanthophyll cycle and thermal energy dissipation in spinach exposed to high light and N stress. Plant Physiology 1997, 113, 817-824
Verhoven AS, Adams III WW, Demmig-adams B, Groce R, Bassi R. Xanthophyll cycle pigment localization and dynamics during exposure to low temperature and light stress in Vinca major. Plant Physiology 1999, 120, 727-737
Weise C, Shi LB, Heber U. Oxygen reduction in the Mehler reaction is insufficient to protect photosystems I and II of leaves against photoinactivation. Physiologia Plantarum 1998, 102:437-446
Woolhouse H W, Batt T In: Perspective in Experimental Biology. Ed: Sunderland N, Pergamon Press. Oxford 1976: 163-175
Xu YN，Siegenthaler PA. Low temperature treatments induce an increase in the relative content of both linolenic and -trans-hexadecenoic acids in thylakoid membrane. Plant Cell Physiol，1997，38（5）：611~618
Yamamoto HY. Biochemistry of the violaxanthin cycle in higher plants. Pure Appl Chen. 1979, 51:639-648
Yamamoto HY. Xanthophyll cycles. Methyleneods Enzymol 1985, 110: 303-312.
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