摘要
紫茎泽兰(Eupatorium adenophorum Spreng)是一种世界性的外来入侵恶性杂草,目前仍没有找到有效的方法来控制它。令人振奋的是强胜等在1999年从紫茎泽兰致病型链格孢菌(Alternaria alternata(Fr.) Keissler)中分离得到了一种毒素(命名为AAC-toxin),这种毒素能够引起包括紫茎泽兰在内的许多杂草的褐斑病。这为紫茎泽兰的防除提供了巨大的新的希望.后来证明AAC-toxin就是细交链格孢菌酮酸(Tenuazonic acid,TeA)。自从1957年首次从链格孢菌菌株上分离以来,已经先后在橄榄、向日葵、柑橘、胡椒、高梁、烟草、水稻和紫茎泽兰上发现了这种TeA毒素。长期以来,关于TeA的研究主要集中在来源和动物毒性方面,关于植物活性方面研究十分少见,更没有关于除草活性方面的报道.我们的前期研究表明TeA对广泛的寄主表现出很高的生物活性,能够迅速杀死许多单双子叶植物,这些寄主植物既有杂草也有作物。本论文主要研究TeA的作用靶标和它引起叶片坏死的机理,目的是为利用TeA开发生物源除草剂提供理论依据。
首先,我们利用TeA的粗毒素研究了TeA和光合作用之间的关系。结果表明在叶片出现明显的外观伤害之前,TeA处理使叶片的光合放氧速率和表观量子效率明显降低。进一步研究发现TeA主要作用于光合作用光反应的光系统Ⅱ(PSⅡ),能够完全抑制PSⅡ电子传递活性,对光Ⅰ(PSⅠ)活性影响不大。而且,TeA对光合作用暗反应关键酶RuBPcase、叶绿素含量和叶片总可溶性蛋白含量则没有明显的影响。叶绿素荧光试验结果表明TeA抑制了PSⅡ受体侧电子的传递从而导致PSⅡ光化学效率的降低,但对PSⅡ供体侧的天线色素分子和放氧复合体没有明显的影响。另外,TeA能够强烈的降低叶绿体Mg~(2+)-ATPase活性,对线粒体并没有影响。因此,可以推断PSⅡ受体侧和叶绿体ATPase可能是TeA的2个最重要的作用靶标。这也意味着TeA可能是一个PSⅡ抑制剂。
为了进一步研究TeA在PSⅡ上的精确作用靶标,我们应用了快速叶绿素荧光动力学和同位素标记光亲合技术。纯TeA毒素能够完全抑制紫茎泽兰类囊体PSⅡ电子传递活性,半抑制浓度是238μM。来自不同植物的快速叶绿素动力学结果表明TeA在PSⅡ上有几个作用位点.首先,TeA处理导致OJIP曲线发生的最大变化在于TeA加速了J点的快速上升,这一点和传统的PSⅡ除草剂敌草隆作用方式是相同的。这说明TeA最关键和最重要的作用是阻断PSⅡ受体侧电子向QA以后的位置传递。而且进一步证实了TeA对PSⅡ供体侧的天线色素分子和放氧复合体没有影响。与经典的PSⅡ除草剂不同的是TeA处理导致了热反应中心的上升。另外,TeA也可能对FNR活性产生抑制作用。此外,来自碳14标记阿特拉津的竞争结合试验证据表明:TeA和阿特拉津在Q_B绑定区域上作用的方式是不同的,虽然它们有着共同的作用靶标-Q_B绑定区域。这说明TeA可能是一种新的天然的PSⅡ抑制剂。除了最主要的位点Q_B绑定区域外,TeA也可能对叶绿体ATPase和FNR的活性产生一定的抑制作用。
十分有意思的是,在链格孢菌引起寄主植物发病过程中为什么它产生的是TeA而不是其它的TeA类似物呢?在植物体内有20多种氨基酸可供选择去合成TeA的吡咯环,然而链格孢菌却利用异亮氨酸合成了这个在5位是异丁基长链的物质—TeA。这个问题要得到解决,我们必须先弄清除TeA在PSⅡ上精确靶标的分子基础。因此我们选择了莱茵藻(包括野生型和5个psbA突变体)这种除草剂靶标研究和光合作用研究的模式材料来探讨TeA同Q_B绑定区域相互作用的分子基础。叶绿素荧光试验显示TeA抑制了光合电子在PSⅡ受体侧Q_A到Q_B之间的传递。来自莱茵藻D1蛋白突变体的证据表明在TeA和Q_B位点相互作用过程中D1蛋白上的256位氨基酸起着关键的作用。从碳14标记阿特拉津的竞争结合实验、JIP-test分析和D1系列突变体实验结果可以看出:在同Q_B绑定位点相互作用的过程中,TeA与其它已知的PSⅡ抑制剂有着不同的绑定行为,进一步证明了TeA可能是一种新型的PSⅡ抑制剂。TeA和它的类似物生物活性试验表明3-酰基-5-烷基吡咯烷酮甚至四氨酸或许提供了一种新型光合抑制剂的结构骨架。
在植物病害和许多非生物胁迫中细胞死亡总是伴随着活性氧(ROS)的产生而发生。为了进一步阐明TeA引起叶片坏死的机理,我们进行了一系列试验来探究在TeA致病过程中TeA、活性氧产生和细胞死亡的关系。我们的结果表明在TeA引起紫茎泽兰叶片发病的过程中的确产生了活性氧。激光共聚焦的试验结果表明TeA诱导叶肉细胞活性氧产生的最初位点在叶绿体,这个结果在细胞化学染色(CeCl_3染色)试验中得了到进一步证实。电子自旋共振(EPR)和活性氧清除剂试验结果表明TeA激发的叶绿体活性氧的最初种类是羟自由基(OH)、单线态氧(~1O_2)、超氧自由基(O_2~(.~-))和过氧化氢(H2O_2)。最初产生的活性氧首先对叶绿体结构造成伤害,但没有对细胞中其他细胞器造成氧化伤害。随着ROS弥散到细胞,高浓度的活性氧能够与细胞中许多重要的生物分子象膜脂、色素、蛋白质和氨基酸等发生反应,接着一系列氧化伤害如膜脂过氧化、叶绿素降解、染色质浓缩、细胞结构和功能丧失等相继发生,最终导致细胞坏死。象经典的光合作用除草剂一样,TeA所引起的光氧化伤害是细胞死亡和叶片坏死的主要原因。
总之,TeA是一种新型的天然的光合作用抑制剂。TeA能够通过与Q_B位点结合而诱发叶绿体活性氧的爆发,进而直接导致叶片快速坏死,这个过程主要是一个光氧化损伤的过程。TeA对叶绿体ATPase和FNR活性的影响可能也参入了这个过程。相对于敌草隆和百草枯等单一位点的传统光合作用除草剂而言,一旦TeA进入叶肉细胞后,它的这种多位点作用机制也许能够导致细胞在短时间内产生更多种类和更高水平的活性氧,从而迅速毁坏细胞而杀死杂草。
Nowdays,there is no quite efficacious methods to be used to Crofton weed (Eupatorium adenophorum Spreng) control,which is one of the most troublesome invasive weed with worldwide distribution.Excitingly,Qiang et al.(1999) found a phytotoxin (named AAC-toxin) produced by Alternaria alternata(Fr.) Keissler Crofton weed pathotype,which can cause brown leaf spot disease in many weeds,especially host Crofton weed,which supplies a big new opportunity to Crofton weed control.AAC-toxin is later proven to be tenuazonic acid(TeA) through isolation and identification.Since its first isolation in 1957 from the culture filtrates of Alternaria tenuis,TeA has been found in olives,sunflower seeds,tangerines,peppers,sorghum,tobacco,rice and Crofton weed.So far,interest in TeA is mainly focused on its source and its toxicity to animals.However, only few studies have addressed the bioactivity of TeA to plants with few purposes for weed control.From our previous studies,TeA shows high bioactivity to a wide range of plants,from weeds to crops,and then quickly kills the seedlings of mono- and dicotyledonous weeds.However,mechanism of killing weeds has not been known before. Here,action target of TeA and the mechanism of leaf necrosis induced by TeA are investigated in order to supply the theory basis for development of bio-based herbicides using TeA.
Firstly,we studied the relationship between TeA(crude toxin) and photosynthesis.The results show that TeA greatly inhibited their oxygen evolution rate and declines evidently quantum efficiency before any distinct change in leaf appearance was observed.The effect of TeA on the electron transfer reaction of chloroplasts shows that the activity of photosystemⅡ(PSⅡ),but not photosystemⅠ(PSI),was completely inhibited by TeA.This inhibition mainly focuses on the light reaction not the dark reaction of photosynthesis.No evident difference of the key enzyme RuBPcase of dark reaction and chlorophyll content was found under TeA treatment.Evidences from chlorophyll fluorescence further indicate that TeA decreased the efficiency of photochemistry of whole PSⅡattributed to inhibit electron transport in PSⅡacceptor side,but had no effect on the donor side of PSⅡsuch as antenna pigment molecule and Oxygen Evolving Complex(OEC).In the addition,TeA led to a distinct decrease of Mg~(2+)-ATPase activity of chloroplast,whereas,had no influence on mitochondria.Thus,it is concluded that the acceptor side of PSⅡand ATPase of chloroplast are possible two most important action target of TeA until now.These results mean that TeA may be a PSⅡinhibitor.
To further study the precise action site of TeA on PSⅡ,the modern techniques of fast chlorophyll fluorescence kinetics and photoaffinity labeling with radioactive technique were utilized.Results demonstrate that TeA(pure toxin) inhibited completely the activity of PSⅡelectron transport of Crofton weed;the pI_(50)-value was 240μM.Evidences from chlorophyll fluorescence transients OJIP curves of the various species plants show that TeA has several sites of action.As the classical PSⅡherbicides(eg.diuron and atrazine),the biggest change of OJIP curves of all plants is a fast rise of step-J after TeA treatment.It is concluded that the most key and important action site of TeA is that it interrupts electron transport beyond Q_A on the acceptor side of PSⅡ.Moreover,TeA has no affect on the antenna pigments,OEC at the donor side of PSⅡ.On the other hand,TeA treatment causes the increase of heat sink centers and inactivation of the FNR system and the reduction of NADP~+,which is different from a classical inhibitor.Evidence from the competitive displacement with[~(14)C]atrazine shows that TeA does not share the same binding environment with atrazine despite they possess a common action target:the Q_B site.It is showed that TeA is possible a novel natural PSⅡinhibitor.Moreover,TeA maybe also effect on FNR and ATPase of chloroplast besides the most main Q_B site.
Interestingly,why does A.alternata only synthesize TeA with pyrrole ring as phytotoxin against the resistance of host plant during its infection? The fungi easily use all 20 kinds of natural amino acid available as materials to synthesize pyrrole ring as phytotoxin.However,A.alternata only selectively use isoleucine,having butyl side chain, to synthesize into TeA with long side chain at 5-position.To answer the question,we must clarify the precise molecular target of TeA in PSⅡ.Chlamydononas reinhardtti,a typical model organism used in the field of herbicide target and photosynthesis research,is utilized to confirm the molecule basis of TeA binding to the Q_B site.Results from chlorophyll fluorescence of C.reinhardtti further prove that TeA blocks electron flow from Q_A to Q_B at PSⅡacceptor side.On the basis of studies with D1-mutants of C.reinhardtii,the No.256 amino acid plays a key role in TeA binding to the Q_B-niche.The results of competitive replacement with[~(14)C]atrazine combined with JIP-test and D1-mutant show that TeA should be considered as a new type of PSⅡinhibitor because it has a different binding behavior within Q_B site from other known PSⅡinhibitors.Bioassay of TeA and its analogues indicates 3-acyl-5-alkyltetramic and even tetramic acid compounds may present a new structural framework with pyrrole ring for photosynthetic inhibitors.
Cell death,along with the production of ROS,occurs during plant-pathogen and many abiotic stresses.In order to further elucidate the mechanism of TeA-induced leaf necrosis, we carried out a series of experiments to investigate the relationship between TeA,ROS generation and cell death.Our studies show that TeA induces ROS burst in leaves during TeA-induced brown leaf spot disease of E.adenophorum.According to the data of laser scanning confocal microscope,it is suggested the primary site of TeA-induced ROS generation in mesophyll cells is chloroplasts.This result is further proved by the evidence from the experiments of CeCl_3 staining.The results of EPR and ROS scavenger experiments suggest that the original sources of TeA-induced ROS production in chloroplasts are hydroxyl radicals(OH),singlet oxygen(~1O_2),superoxide radical(O_2~-) and hydrogen peroxide(H_2O_2).The initial generation of ROS firstly results in damage in the structure of chloroplasts.Latter production of ROS is dispersed into cells and reacts with many important biomolecules such as pigments,lipids,proteins and nucleic acids,causing lipid peroxidation,chlorophyll breakdown,chromatin condensation,the loss of structure and function of cells,and resulting in cell destruction.Like classical photosynthetic herbicides,this injury triggered by TeA is a result of the directly oxidative damage of high ROS level in cell.
In conclusion,TeA is a novel natural photosynthetic inhibitor.It induces oxidative burst in chloroplasts by acting with the Q_B site,resulting in leaf necrosis.In addition,the inhibition of the activity of ATPase and FNR of chloroplasts possible play some role in this process.Photo-oxidative damage is responsible for the process of TeA-induced leaf necrosis.Compared with the general photosynthetic inhibitor herbicides such as DCMU and paraquat,such action mode of TeA makes possibly cells generate more reactive oxygen species with a faster rate and higher level and kill quickly many weeds once mesophyll cells touch directly TeA.
引文
1. Abbas HK, Tanaka T, Duke SO, Porter JK, Wray EM, Hodges L, Sessions AE, Wang E, Merrill AH Jr, and Riley RT (1994) Fumonisin- and AAL-toxin-induced disruption of sphingolipid metabolism with accumulation of free sphingoid bases. Plant Physiology 106: 1085-1093
2. Abbas HK, Paul RN, Riley RT, Tanaka T, Shier WT (1998) Ultrastructural effects of AAL-toxin TA from the fungus Alternaria alternata on black nightshade {Solarium nigrum L.) leaf discs and correlation with biochemical measures of toxicity. Toxicon 36: 1821-1832
3. Abbas HK, Tanaka T, Shier WT (1995) Biological activities of synthetic analogues of Alternaria alternata toxin (AAL-toxin) and fumonisin in plant and mammalian cell cultures. Phytochemistry 40: 1681-1689
4. Abel S, Theologis A (1994) Transient transformation of Arabidopsis leaf protoplasts: a versatile experimental system to study gene expression. Plant Journal 5: 421-427
5. Ajlani G, Meyer I, Vernotte C, Astier C (1989) Mutation in phenol-type herbicide resistance maps within the psbA gene in Synechocystis 6714. FEBS Letters 246: 207-210
6. Allan A, Fluhr R (1997) Two distinct sources of elicited reactive oxygen species in tobacco epidermal cells. Plant Cell 9: 1559-1572
7. Alscher RG, Erturk N, Heath LS (2002) Role of superoxide dismutases (SODs) in controlling oxidative stress in plants. Journal of Experimental Botany 53: 1331-1341
8. Amesz J, Fork DC (1967) Quenching of chlorophyll fluorescence by quinones in algae and chloroplasts. Biochimica et Biophysica Acta 143: 97-107
9. An CF (2003) On potential of metabolite of Alternaria alternata as bio-source herbicide. Master Thesis, Nanjing Agricultural University, Nanjing
10. Ananyev G, Renger G, Wacker U, and Klimov V (1994) The photoproduction of superoxide radicals and the superoxide dismutase activity of PSII: the possible involvement of cytochrome b559. Photosynthesis Research 41: 327-338
11. Apel K, Hirt H (2004) Reactive oxygen species: metabolism, oxidative stress, and signal transduction. Annual Review of Plant Biology 55: 373-399
12. Appenroth KJ, Stockel J, Srivastava A, Strasser RJ (2001) Multiple effects of chromate on the photosynthetic apparatus of Spirodela polyrhiza as probed by OJIP chlorophyll α fluorescence measurements.Environmental Pollution 115:49-64
13.Armstrong D(1998) Free radicals and antioxidant protocols.Human Press,Totowa-New Jersey.
14.Arnon D(1949) Copper enzymes in isolated chloroplasts:Polyphenoloxidase in Beta vulgaris.Plant Physiology 24:1-15
15.Asada K(1984) Chloroplasts:formation of active oxygen and its scavenging.Methods in Enzymology 105:422-429
16.Asada K(2006) Production and scavenging of reactive oxygen species in chloroplasts and their functions.Plant Physiology 141:391-396
17.Astier C,Boussac A,Etienne AL(1984) Evidence for different binding sites on the 33-kDa protein for DCMU,atrazine and Q_B.FEBS Letters 167:321-326
18.Babbs CF,Pham JA,Coolbaugh RC(1989) Lethal hydroxyl radical production in paraquat-treated plants.Plant Physiology 90:1267-1270
19.Bais HP,Vepachedu R,Gilroy S,Callaway RM,Vivanco JM(2003) Allelopathy and exotic plant invasion:from molecules and genes to species interactions.Science 301:1377-1380
20.Bakeeva LE,Dzyubinskaya EV,Samuilov VD(2005) Programmed death in plants:ultrastructural changes in pea guard cells.Biochemistry(Moscow) 70:972-979
21.Baker NR,Percival MP(1991) Herbicides.Elsevier Press,Amsterdam-London-New York-Tokyo.
22.Barros LF,Hermosilla T,Castro J(2001) Necrotic volume increase and the early physiology of necrosis.Comparative Biochemistry and Physiology Part A 130:401-409
23.Barna B,Fodor J,Pogany M,Kiraly Z(2003) Role of reactive oxygen species and antioxidants in plant disease resistance.Pest Management Science 59:459-64
24.Berg JM,Tymoczko JL,Stryer L(2007) The light reactions of photosynthesis.Biochemistry(Sixth Edition),Chapter 19.W.H.Freeman and Company,New York,pp 541-564
25.Bertold D,Babcock GT,Yocum CA(1981) A highly resolved oxygen evolving photosystem Ⅱ preparation from the spinach thylakiod membranes.FEBS Letters 134:231-234
26.Bacon K(2001) Photosynthesis-Photobioehemistry and Photobiophysics,Kluwer Academic Publishers,Dordrecht-Boston-London,pp 301-303
27. Beligni MV, Lamattina L (1999) Nitric oxide protects against cellular damage produced by methylviologen herbicides in potato plants. Nitric Oxide: Biology and Chemistry 3: 199-208
28. Beligni MV, Lamattina L (2002) Nitric oxide interferes with plant photo-oxidative stress by detoxifying reactive oxygen species. Plant Cell and Environment 25: 737-748
29. Bestwick CS, Brown IR, Bennett MHR, Mansfield JW (1997) Localization of hydrogen peroxide accumulation during the hypersensitive reaction of lettuce cells to Pseudomonas syringae pv phaseolicola. Plant Cell 9: 209-221
30. Bethke P, Jones RL (2001) Cell death of barley aleurone protoplasts is mediated by reactive oxygen species. Plant Journal 25: 19-29
31. Bowes J, Crofts AR, Arntzen CJ (1980) Redox reaction on the reducing side of photosystem II in chloroplasts with altered herbicide binding properties. Arch. Biochimica et Biophysica Acta 200: 303-308
32. Bowyer J, Hilton M, Whitelegge J (1990) Molecular Modelling Studies on the Binding of Phenylurea Inhibitors to the Dl Protein of Photosystem I [J]. Zeitschrift fur Naturforschung c-a Journal of Biosciences 45: 379-387
33. Buchanan BB, Balmer Y (2005) Redox regulation: a broadening horizon. Annual Review of Plant Biology 56: 187-220
34. Bueno M, Fillat M, Strasser RJ, Maldonado-Rodriguez R, Marina N, Smienk H, G6mez-Moreno C, Barja F (2004) Effect of lindane on the photosynthetic apparatus of the Cyanobacterium Anabaena fluorescence induction studies and immunolocalization of ferredoxin-NADP~+ reductase. Environmental Science and Pollution Research 11: 98-106
35. Busheva M, Velitchkova M, Markova T, Zanev Y (1998) Effect of cholesterol and benzyl alcohol on fluorescence transients of pea thylakoids. Journal of Photochemistry and Photobiology B: Biology 42: 240-244
36. Camadro JM, Matringe M, Thome F, Brouillet N, Mornet R, Labbe P (1995) Photoaffinity labeling of protoporphyrinogen oxidase, the molecular target of diphenylether-type herbicides. European Journal of Biochemistry 229: 669-674
37. Camara-Artigas A, Allenl JP (2004) Comparative analyses of three-dimensional models of bacterial reaction centers. Photosynthesis Research 81: 227-237
38. Chelkowski J, Visconti A (1992) Alternaria Biology, Plant Diseases and Metabolites. Elsevier, Amsterdam-London-New York-Tokyo, pp 449-541
39.Chen SG,Dai XB,Qiang S,Tang YL(2005) Effect of a nonhost-selective toxin from Alternaria alternate on chloroplast-electron transfer activity in Eupatorium adenophorum.Plant Pathology 54:671-677
40.Chen C,Dickman MB(2005) Proline suppresses apoptosis in the fungal pathogen Colletotrichum trifolii.Proceedings of the National Academy of Science of the United State of America 102:3459-3464
41.Chen SG,Xu XM,Dai XB,Yang CL,Qiang S(2007 a) Identification of tenuazonie acid as a novel type of natural photosystem Ⅱ inhibitor binding in Q_B-site of Chlamydomonas reinhardtii.Biochimiea et Biophysica Acta 1767:306-318.
42.Chert SG,Yin CY,Dai XB,Qiang S,Xu XM(2007 b) Tenuazonic acid,a natural phytotoxin,on photosystem Ⅱ of spinach.Environmental and Experimental Botany,doi:10.1016/j.envexpbot.2007.10.002
43.Chivasa S,Ndimba BK,Simon WJ,Lindsey K,Slabas AR(2005) Extracellular ATP functions as an endogenous external metabolite regulating plant cell viability.Plant Cell 17:3019-3034
44.Cleland RE,Grace SC(1999) Voltammetrie detection of superoxide production by photosystem Ⅱ.FEBS Letters 457:348-352
45.Cobb AH,Kirkwood RC(2000) Herbicides and their mechanisms of action.CRC Press,UK.
46.Conno M C,Vurro M(1999) Effect of fungal toxins on germination of Striga hermonthica seeds.Weed Research 39:15-20
47.Coombs J,Hall DO,Long SP,Scurlock JMO(1986) Techniques in bioproductivity and photosynthesis,Science Press,Beijing,pp.142-144
48.Dacero A M,Combina M,Etcheverry M(1997) Evaluation of Alternaria and its myeotoxins during ensiling of sunflower seeds.Natural Toxins 5:20-23
49.Dahnhardt D,Falk J,Appel J,van der Kooij TAW,Schulz-Friedrich R,Krupinska K(2002) The hydroxyphenylpyruvate dioxygenase from Synechocystis sp.PCC 6803 is not required for plastoqttinone biosynthesis.FEBS Letters 523:177-181
50.Dai XB,Chert SG,Qiang S,An CF,Zhang RX(2004) Effect of Toxin from Alternaria alternate(Fr.) Keissler on leaf photosynthesis of Eupatorium adenophorum Spreng.Aeta Phytopathologiea Sinica 34:55-60
51.Dai XB,Yu Y,Zhang RX,Yu X J,He P,Xu CH(2001) Relationship among photosystem Ⅱ carbonic anhydrase,extrinsic pholypeptides and manganese cluster. Chinese Science Bulletin 46: 406-408
52. Dalla Chiesa M, Friso G, Deak Z, Vass I, Barber J, Nixon PJ (1997) Reduced turnover of the D1 polypeptide and photoactivation of electron transfer in novel herbicide resistant mutants of Synechocystis sp. PCC 6803. European Journal of Biochemistry 248: 731-40
53. Danon A, Coll NS, Apel K (2006) Crytochrome-1-dependent execution of programmed cell death induced by singlet oxygen in Arabidopsis thaliana. Proceedings of the National Academy of Science of the United State of America 103: 17036-17041
54. Davies ND, Diner UL, Morgan-Jones G (1977) Tenuazonic acid production by Alternaria alternate and Alternaria tenuissima isolated from cotton. Applied and Environmental Microbiology 34: 155-157
55. Demetriou G, Neonaki C, Navakoudis E, Kotzabasis (2007) Salt stress impact on the molecular structure and function of the photosynthetic apparatus-the protective role of polyamines. Biochimica et Biophysica Acta 1767: 272-280
56. Demming-Adams B, Adams WW (1996) Xanthophyll cycle and light stress in nature. Planta 198: 460-470
57. De Ronde JA, Cress WA, Kurger GHJ, Strasser RJ, Van Staden (2001) Photosynthetic reponse of transgenic soybean plants, containing an Arabidopsis P5CR gene, during heat and drought stress. Journal of Plant Physiology 161: 1211-1224
58. Dodge AD (1989) Society for Experimental Biology Seminar Series 38: Herbicides and plant metabolism. Cambridge University Press, New York - Port Chester -Melbourne - Sydney.
59. Dostatni R, Meyer HE, Oettmeier W (1988) Mapping of two tyrosine residues involved in the quinine-(QB) binding site of the D-1 reaction center polypeptide of photosystem II. FEBS Letters 239: 207-210
60. Eckardt NA (2005) Ins and outs of programmed cell death and toxin action. Plant Cell 17: 2849-2851
61. Edinger AL, Thompson CB (2004) Death by design: apoptosis and autophagy. Current Opinion in Cell Biology 16: 663-669
62. Enans N, Mcroberts N, Hill RA, Marshall GM (1996) Phytotoxin production by Alternaria linicola and phytoalexin production by the linseed host. Annals of Applied Biology 129: 415-431
63. Fath A, Bethke PC, Jones RL (1999) Barley aleurone cell death is not apoptotic: characterization of nuclease activities and DNA degradation.Plant journal 20:305-315
64.Ferreira KN,Iverson TM,Maghlaoui K,Barber J,Iwata S(2004) Architecture of the photosynthetic oxygen-evolving center.Science 303:1782-1784
65.Festjens N,Berghe TV,Vandenabeele(2006) Necrosis,a well-orchestrated from of cell demise:signaling cascades,important mediators and concomitant immune response.Biochimica et BiophysicaActa 1757:1371-1387
66.Foreman J,Bemidchik V,Bothwell JHF,Mylona P,Miedema H,Torres MA,Linstead P,Costa S,Brownlee C,Jone JDG,Davies JM,Dolan L(2003) Reactive oxygen species produced by NADPH oxidase regulate plant cell growth.Nature 422:442-446
67.Foster B,Heifetz PB,Lardans A,Boynton J,Gillham NW(1997) Herbicide resistance and growth of D1 Ala251 mutant in Chlamydomonas.Zeitschrift fur Naturforschung c-a Journal of Biosciences 52c:654-664
68.Foster B,Osmond CB,Pogsou BJ(2005) Improved survival of very high light and oxidative stress is conferred by spontaneous gain-of-function mutations in Chlamydomonas.Biochimica et Biophysica Acta 1709:45-57
69.Friedman MA,Aggarwal V,Lester GE(1975) Inhibition of epidermal DNA synthesis by cycloheximide and other inhibitors of protein synthesis.Research Communications in Molecular Pathology and Pharmacology 11:311-318
70.Fuerst P,Norman MA(1991) Interaction of Herbicides with Photosynthetic Electron Transport.Weed Science 39:458-464
71.Fufezan C,Rutherford AW,Krieger-Liszkay A(2002) Singlet oxygen production in herbicide-treated photosystem Ⅱ.FEBS letters 532:407-410
72.Funk C,Wiklund R,Sehrder WP,Jansson C(2001) D1' centers are less efficient than normal photosystem Ⅱ centers.FEBS Letters 505:113-117
73.Gallardo GL,Pea NI,Chaeaua P,Terzolo HR,Cabrera GM(2004) L &hyphen-TeA,a new inhibitor of Paenibacillus larva.World Journal of Microbiology and Biotechnology 20:609-612
74.Geehev TS,Gadjiev IZ,Hille J(2004) An extensive microarray analysis of AAL-toxin-induced cell death in Arabidopsis thaliana brings new insights into the complexity of programmed cell death in plants.CMLS Cellular and Molecular Life Science 61:1185-1197
75.Genty B,Harbinson J,Briantais JM,Baker N(1990) The relationship between non-photochemical quenching of chlorophyll fluorescence and the rate of photosystem 2 photochemistry in leaves. Photosynthesis Research 25: 249-257
76. Golstein P, Kroemer G (2006) Cell death by necrosis: towards a molecular definition. Trends in Biochemical Science 32: 37-43
77. Govindjee (1995) Sixty-three years since kautsky: chlorophyll a fluorescence. Australian Journal of Plant Physiology 22: 131 -160
78. Govindjee, Eggenberg P, Pfister K, Strasser RJ (1992) Chlorophyll a fluorescence decay in herbicide-resistant D1 mutants of Chlamydomonas reinhardtii and the formate effect. Biochimica et Biophysica Acta 1101: 353-358
79. Grene R (2002) Oxidative stress and acclimation mechanisms in plants. The Arabidopsis Book.
80. Greenberg JT (1996) Programmed cell death: a way of life for plants. Proceedings of the National Academy of Science of the United State of America 93: 12094-12097
81. Gronwald JW (1994) Resistance to photosystem II inhibiting herbicides. In S. B. Powles and J.A.M. Holtum, eds. Herbicide Resistance in Plants: Biology and Biochemistry. Lewis Publishers, Boca Raton, pp 27-60
82. Haldimann P, Strasser RJ (1999) Effect of anaerobiosis as probed by the polyphasic chlorophyll a fluorescence rise kinetic in pea (Pisum sativum L.) Photosynthesis Research 62: 67-83
83. Haldimann P, Tsimilli-Michael M (2005) Non-photochemical quenching of chlorophyll a fluorescence by oxidised plastoquinone: new evidences based on modulation of the redox state of the endogenous plastoquinone pool in broken spinach chloroplasts. Biochimica et Biophysica Acta 1706: 239-249
84. Halliwell B, Gutteridge JMC (1984) Oxygen toxicity, oxygen radicals, transition metals and disease. Biochemistry Journal 219: 1-14
85. Halloin JM, Hagedorn DJ (1975) Effects of tentoxin on enzymic activities in cucumber and cabbage cotyledons. Mycopathologia 55: 159-162
86. Hasan HA (2001) Phytotoxicity of pathogenic fungi and their mycotoxins to cereal seedling viability. Acta Microbiologica et Immunologica Hungarica 48: 27-37
87. Hatta R, Ito K, Hosaki Y, Tanaka T, Tanaka A, Yamamoto M, Akimitsu K, Tsuge T (2002) A conditionally dispensable chromosome controls host-specific pathogenicity in the fungal plant pathogen Alternaria alternata. Genetics 161: 59-70
88. Havaux M, Tardy F (1997) Thermostability and photostability of photosystem II in leaves of the Chlorina-f2 barley mutant deficient in lightharvesting chlorophyll a/b protein complexes.Physiology 113:913-923
89.Heap IM(2002) International survey of herbicide resistant weeds.Online Internet:www.weedscienee.com.
90.Hernandez JA,Ferrer MA,Jimenez A,Barcelo AR,Sevilla F(2001) Antioxidant systems and O_2~-/H_2O_2 production in the apoplast of pea leaves.Its relation with salt-induced necrotic in minor veins.Plant Physiology 127:817-831
91.Heredia P,Rival J D L(2003) Fluorescence induction of photosystem Ⅱ membrane shows the steps till reduction and protonation of the quinone pool.Plant Physiology 160:1499-1506
92.Hideg E,Spetea C,Vass I(1995) Superoxide radicals are not the main promoters of acceptor-side-induced photoinhibitory damage in spinach thylakoids.Photosynthesis Research 46:399-407
93.Hideg E,Spetea C,Vass I(1994) Singlet oxygen production in thylakoid membranes during photoinhibition as detected by EPR spectroscopy.Photosynthesis Research 39:191-199
94.Hill R,Larkum AWD,Frankart C,Kuhl M,Ralph PJ(2004) Loss of functional photosystem Ⅱ reaction centers in zooxanthellae of corals exposed to bleaching conditions:using fluorescence rise kinetics.Photosynthesis Research 82:59-72
95.Hiraki M,Vredenberg WJ,van Rensen JJS,Wakabayashi K(2004) A modified fluorometric method to quantify the concentration effect(pI_(50)) of photosystem Ⅱ-inhibiting herbicides.Pesticide Biochemistry and Physiology 80:183-191
96.Hirschberg J,Bleecker A,Kyle DJ,McIntosh L,and Arntzen CJ(1984) The molecular basis of triazine-herbicide resistance in higher-plant chloroplast.Zeitschrift fur Naturforschung c-a Journal of Biosciences 39c:412-420
97.Hoar TX,Nakayashild H,Tosa Y,Mayama S(2006) Specific cleavage of ribosomal RNA and mRNA during victorin-induced apoptotic cell death in oat.Plant Journal 46:922-933
98.Hock B,Elstner EF(2005) Plant Toxicology(fourth edition).Marcel Dekker,New York.
99.Holland N,Evron Y,Jansen MAK,Edelman M,Pick U(1997) Involvement of thylakoid overenergization in tentoxin-induced chlorosis in Nicotiana spp.Plant Physiology 114:887-892
100.Houot V,Etienne P,Petitot AS,Barbier S,Biein JP,Suty L(2001) Hydrogen peroxide induces programmed cell death features in cultured tobacco BY-2 cells,in a dose-dependent manner.Journal of Experimental Botany 52:1721-1730
101.Ikeda Y,Ohld S,Koizumi K,Tanaka A,Watanabe H,Kohno H,van Rensen JJS,Boger P,Wakabayashi K(2003) Binding site of novel 2-benzylamino-4-methyl-6-trifluoromethyl -1,3,5-triazine herbicides in the D1 protein of photosystem Ⅱ.Photosynthesis Research 77:35-43
102.Ilik P,Schansker G,Kotabova E,Vaczi P,Strasser RJ,Bartak M(2006) A dip in the chlorophyll fluorescence induction at 0.2-2 s in Trebouxia-possessing lichens reflects a fast reoxidation of photosystem I,A comparison with higher plants.Biocbimica et Biophysica Acta 1757:12-20
103.Jacobson MD(1996) Reactive oxygen species and programmed cell death.Trends in Biochemical Science 21:83-86
104.Janardhanan KK,Husain A(1983) Studies on isolation,purification and identification of tenuazonic acid,a phytotoxin produced by Alternaria alternata(Fr.)Keissler causing leaf blight of Datura innoxia Mill.Mycopathologia 83:135-140.
105.Joo JH,Wang SY,Chen JG,Jones AM,Fedoroff NV(2005) Different signaling and cell death roles of heterotrimeric G protein α and β subunits in the Arabidopsis oxidative stress responsed to ozone.Plant Cell 17:957-970
106.Jung IL,Kim IG(2003) Thiamine protects against paraquat-induced damage:scavenging activity of reactive oxygen species.Environmental Toxicology and Pharmacology 15:19-26
107.Jursinie PA(1978) Flash polarographic detection of superoxide production as a means of monitoring electron flow between photosystems Ⅰ and Ⅱ.FEBS Letters 90:15-20
108.Jursinie PA,McCarthy SA,Bricher TM,Stemler A(1991) Characteristics of two atrazine-binding sites that specifically inhibit photosystem Ⅱ function.Biochimica et Biophysica Acta 1059:312-322
109.Kamiya N,Shen JR(2003) Crystal structure of oxygen-evolving photosystem Ⅱ from Thermosynechococcus vuleanus at 3.7-(?) resolution.Proceedings of the National Academy of Science of the United State of America 100:98-103
110.Kaur G,Singh US,Garg GR(1987) Mode of action of toxin isolated from Fusarium oxysporumf,sp.Ciceri.Indian Phytopathology 40:76-84
111.Kern J,Loll B,Luneberg C,DiFiore D,Biesiadka J,Irrgang KD,Zouni A(2005) Purification,characterisation and crystallisafion of photosystem Ⅱ from Thermosynechococcus elongatus cultivated in a new type of photobioreactor.Biochimica et Biophysiea Acta 1706:147-157
112.Khurana SMP,Pandey SK,Sarkar D,Chanemougasoundharam A(2005)Apoptosis in plant disease response:a close encounter of the pathogen kind.Current Science 88:740-752
113.Kingston-Smith AH,Foyer CH(2000) Bundle sheath proteins are more sensitive to oxidative damage than those of the mesophyll in maize leaves exposed to paraquat or low temperatures.Journal of Experimental Botany 51:123-130
114.Kleier DA,Andrea TA,Hegedus TKJ,Gardner GM and Cohen B,1987.The topology of the 32 kDa herbicide binding protein of photosystem Ⅱ in the thylakoid membrane.Zeitschrift fur Naturforschung c-a Journal of Biosciences 42c:733-738.
115.Kless H,Vermaas W(1995) Tandem sequence duplications functionally complement deletions in the D1 protein ofphotosystem Ⅱ.The Journal of Biological Chemistry 270:16536-16541
116.Kobayashi M,Ohura I,Kawakita K,Yokota N,Fujiwara M,Shimamoto K,Doke N,Yoshioka H(2007) Calcium-dependent protein kinases regulate the production of reactive oxygen species by potato NADPH oxidase.Plant Cell 19:1065-1080
117.Kohmoto K,Itoh,Shimomura N(1993) Isolation and biological activities of two host-specific toxins from the tangerine pathotype of Alternaria alternata.Phytopathology 83:495-502
118.Krause GH,Weis E(1984) Chlorophyll fluorescence as a tool in plant physiology.Ⅱ.Interpretation of fluorescence signals.Photosynthesis Research 5:139-157
119.Krause GH,Weis E(1991) Chlorophyll fluorescence and photosynthesis:the basics.Annu.Rev.Plant Physiol.Plant Molecular Biology 42:313-49
120.Krieger-Liszkay A,Trebst A(2006) Tocopherol is the scavenger of singlet oxygen produced by triplet states of chlorophyll in the PSⅡ reaction centre.Journal of Experimental Botany 57:1677-1684
121.Kroemer G,EI-Deiry WS,Golstein P,Peter M,Vaux D,Vandenabeele P,Zhivotovsky B,Blagosklonny MV,Malorni W,Knight PA,Piacentini M,Nagata S,Melino G(2005) Cell Death and Differentiation 12:1463-1467
122.Krüger GHJ,Tsimilli-Michanel M,Strasser RJ(1997) Light stress provokes plastic and elastic modifications in structure and function ofphotosystem Ⅱ in camellia leaves. Physiologia Plantarum 101: 265-277
123.Laloi C, Przybyla D, Apel K (2006) A genetic approach towards elucidating the biological activity of different reactive oxygen species in Arabidopsis thaliana. Journal of Experimental Botany 57: 1719-1724
124.Lancaster CRD, Michel H (1997) The coupling of light-induced electron transfer and proton uptake as derived from crystal structures of reaction centres from Rhodopseudomonas viridis modified at the binding site of the secondary quinone, Q_B. Structure 5: 1339-1359
125. Lancaser CRD, Michel H (1999) Refined crystal structures of reaction centres from Rhodopseudomonas viridis in complexes with the herbicide atrazine and two chiral atrazine derivatives also lead to a new model of the bound carotenoid. Journal of Molecular Biology 286: 883-898
126.Lancaster CRD, Bibikovai MV, Sabatino P, Oesterhelt D, Michel H (2000) Structural basis of the drastically increased initial electron transfer rate in the reaction renter from a Rhodopseudomonas viridis mutant described at 2.00-A resolution. Journal of Biological Chemistry 275: 39364-39368
127.La Rocca N, Rascio N, Oster U, Rudiger W (2001). Amitrole treatment of etiolated barley seedlings leads to deregulation of tetrapyrrole synthesis and to reduced expression of Lhc and RbcS genes. Planta 213: 101-108
128.Lax AR, Shepherd HS, Edwards JV (1988) Tentoxin, a chlorosis-inducing toxin from Alternaria as a potential herbicide. Weed Technology 2: 540-544
129.Lazar D (1999) Chlorophyll a fluorescence induction. Biochimica et Biophysica Acta 1412: 1-28
130.Lazar D (2003) Chlorophyll a fluorescence rise induced by high light illumination of dark-adapted plant tissue studied by means of a model of photosystem II and considering photosystem II heterogeneity. Journal of Theoretical Biology 220: 469-503
131.Lazar D, Brokes M, Naus J, Dvorak L (1998) Mathematical modeling of 3-(3',4'-dichlorophenyl)-1,1-dimethylurea action in plant leaves. Journal of Theoretical Biology 191: 79-86
132.Lazar D, Naus J, Matouskova M, Flasarova M (1997) Mathematical modeling of changes in chlorophyll fluorescence induction caused by herbicides. Pesticide Biochemistry and Physiology 57: 200-210
133.Le Baron HM (1991) Distribution and seriousness of herbicide resistant weed infestations world wide.In Herbicide Resistance in Weeds and Crop,Oxford Press,Butterwarth-Henemarm.
134.Lincoln JE,Riehael C,Overduin B,Smith K,Bostock R,Gilehrist DG(2002)Expression of the antiapoptotic baculovirus p35 gene in tomato blocks programmed cell death and provides broad-spectrum resistance to disease.Proceedings of the National Academy of Science of the United State of America 99:15217-15221
135.Lin K,Sun J,Song YG,Lin B,Xu YK,Zhang SX,Tian Q,Liu Y(2004) Superoxide,hydrogen peroxide and hydroxyl radical in D1/D2/cytochrome b-559 photosystem Ⅱreaction center complex.Photosynthesis Research 81:41-47
136.Liu HY,Lou RJ,Yang HZ(1997) Research progress on structure-activity relationship of photosystem Ⅱ inhibitor.World Pesticide 19:20-24
137.Liu YD,Ren D,Pike S,Pallardy S,Gassmann W,Zhang S(2007)Chloroplast-generated reactive oxygen species are involved in hypersensitive response-like cell death mediated by a mitogen-activated protein kinase cascade.Plant Journal 51:941-954
138.Loiseau N,Cavelier F,Noel JP,Gomis JM(2002) High yield synthesis of tentoxin,a cyclic tetrapeptide.Journal of Peptide Science 8:335-346
139.Lu M,Wu MQ,Wang JY(1995) Ultrastuctural studies on leaf cells of susceptible apple inoculated with conidia of Alternaria mali roberts.Acta Phytopathologiea Sinica 25:47-50
140.Lupinkova L,Metz JG,Diner BA,Vass I,Komenda J(2002) Hisfidine residue 252of the photosystem Ⅱ D1 polypeptide is involved in a light-induced cross-linking of the polypeptide with the α subunit of cytochrome b-559:study of a site-directed mutant of Synechocystis PCC 6803.Biochimiea et Biophysiea Acta 1554:192-201
141.Mackay SP,O'Malley PJ(1993 a) Molecular modeling of the interactions between optically active triazine herbicides and photosystem Ⅱ.Zeitschrift fur Naturforschung c-a Journal of Biosciences 48c:474-481
142.Mackay SP,O'Malley PJ(1993 b) Molecular modeling of the interactions between DCMU and the Q_B-binding site of photosystem Ⅱ.Zeitschrift fur Naturforschung c-a Journal of Biosciences 48c:191-198
143.Mackay SP,O'Malley PJ(1993 c) Molecular modelling of the interaction of Cyanoaerylate inhibitors with photosystem Ⅱ,Part 2.the effect of stereochemistry of inhibitor binding,Zeitschrift fur Naturforschung c-a Journal of Biosciences 48c: 782-787
144.Maekawa N, Yamamoto M, Nishimura S, Kohmoto K, Kawada M, Watanebe Y (1984) Studies on host-specific AF-toxins produced by Alternaria alternata strawberry pathotype causing Alternaria black spot of strawberry. (1) Production of host-specific toxins and their biological activities. Annual review of Phytopathology Society of Japan 50: 600-609
145.Malkin S, Armond PA, Mooney HA, Fork DC (1981) Photosystem II photosynthetic unit sizes from fluorescence induction in leaves. Correlation to photosynthetic capacity. Plant Physiology 67: 570-579
146.Marfori EC, Kajiyama SI, Fukusaki EI, Kobayashi A (2003) Phytotoxicity of the tetramic acid metabolite trichosetin. Photochemistry 62: 715-721
147Mattoo AK, Pick U, Hoffmann-Falk H, Edelman M (1981) The rapidly metabolized 32,000-dalton polypeptide of the chloroplast is the 'proteinaceous shield' regulating photosystem II electron transport and mediating diuron herbicide sensitivity. Proceedings of the National Academy of Science of the United State of America 78: 1572-1576
148.Meazza G, Scheffler BE, Tellez MR, Rimando AM, Romagni JG, Duke SO, Nanayakkara D, Khan IA, Abourashed EA, Dayan FE (2002) The inhibitory activity of natural products on plant p-hydroxyphenylpyruvate dioxygenase. Phytochemistry 59: 281-288
149.Mehler AH (1951) Studies on reactions of illuminated chloroplasts. I. Mechanism of the reduction of oxygen and other Hill reagents. Achives of biochemistry 33: 65-77
150.Melis A (1999) Photosystem-II damage and repair cycle in chloroplasts: what modulates the rate of photodamage in vivo? Trends in Plant Science 4: 130-135
151.Mete JG, Pakrasi HB, Seibert M, Arntzen CJ (1986) Evidence for a dual function of the herbicide-binding D1 protein in photosystem II. FEBS Letters 205: 269-274
152.Millter R, Lam E (1995) Indification, characterization, and purification of a tobacco endonuclease activity induced upon hypersensitive response cell death. Plant Cell 7: 1951-1962
153.Mirocha CJ, GUchrist DG, Shier WT, Abbas HK, Wen Y, Vesonder RF (1992) AAL toxins, fumonisins (biology and chemistry) and host-specificity concepts. Mycopathologia 117: 47-56
154.Misra AN, Srivastava A, Strasser RJ (2001) Utilization of fast chlorophyll a fluorescence technique in assessing the salt/ion sensitivity of mung bean and Brassica seedlings.Journal of Plant Physiology 158:1173-1181
155.Mitsuhara I,Malik KA,Miura M,Ohashi Y(1999) Animal cell-death suppressors Bcl-x_L and Ced-9 inhibit cell death in tobacco plants.Current Biology 9:775-778
156.Mittler R(2002) Oxidative stress,antioxidants and stress tolerance.Trends in Plant Science 7:405-410
157.Mittler R,Simon L,Lam E(1997) Pathogen-induced programmed cell death in tobacco.Journal of Cell Science 110:1333-1344
158.Mittler R,Vanderauwera S,Gallery M,Van Breusegem F(2004) Reactive oxygen gene network of plants.Trends in Plnat Science 9:490-498
159.Miyashita M,Nakamori T,Murai T,Yonemoto T,Miyagawa H,Akamatsu M,and Ueno T(2001) Structure-activity relationship study of host-specific phytotoxins (AM-toxin analogs) using a new assay method with leaves from apple meristem culture Zeitschrift fur Naturforschung c-a Journal of Biosciences 56:1029-1037
160.Moller IM,Jensen PE,Hansson A(2007) Oxidative modifications to cellular components in plants.Annual Review of Plant Biology 58:459-481
161.Morgan C L,Austin RB(1986) Analysis of fluorescence transients of DCMU-treated leaves of Triticum species to provide estimates of the densities of photosystem Ⅱreaction centers.Photosynthesis Research 7:203-219
162.Motta SD,Soares LMV(2000) Simultaneous determination of tenuazonic and cyclopiazonic acids in tomato products.Food Chemistry 71:111-116
163.Mubarakshina M,Khorobrykh S,Ivanov B(2006) Oxygen reduction in chloroplast thylakoids results in production of hydrogen peroxide inside the membrane.Biochimica et Biophysiea Aeta 1757:1496-1503
164.Munne-Bosch S,Jubauy-Mari T,Alegre L(2001) Drought-induced senescence is characterized by a loss of antioxidant defenees in chloroplast.Plant Cell and Environment 24:1319-1327
165.Nakatsuta S,Namiki F,Okamoto H,Katou K,Yamamoto M,Nishimura S,Nakatsuka S,Goto T,Kohmoto K,Otomi H,Novacky A(1986) Studies on host-specific AF-toxins produced by Alternaria alternata strawberry pathotypc causing Alternaria black spot of strawberry(5).Effect of toxins on membranee potential of susceptible plants as assessed by eletrophysiologieal method.Annual Phytopathology Society of Japan 52:610-619
166.Narusaka Y, Narusaka M, Kobayashi H, Satoh K (1998) The herbicide-resistant species of the cyanobacterial D1 protein obtained by thorough and random in vitro mutagenesis. Plant Cell Physiology 39: 620-626
167.Navarre DA, Wolpert T (1999) Victorin induction of an apoptotic/senescence-like response in oats. Plant Cell 11: 237-249
168.Neill SJ, Desikan R, Hancock J (2002) Hydrogen peroxide signaling. Current Opinion in Plant Biology 5: 388-395
169.Nishimura S, Kohmoto K (1983) Host-specific toxins and chemical structures from Alternaria species. Annual Review of Phytopathology 21: 87-119
170.Oettmeier W (1999) Herbicide resistance and supersensitivity in photosystem II. CMLS Cellular and Molecular Life Sciences 55: 1255-1277
171.Oettmeier W, Masson K, Donner A (1988) Anthraquinone inhibitors of photosystem II electron transport. FEBS Letters 231: 259-262
172.Oettmeier W, Masson K, Hecht HJ (2001) Heterocyclic ortho-quinones, a novel type of photosystem II inhibitors. Biochimica et Biophysica Acta 1504: 346-351
173.Oettmeier W, Masson K (1980) Synthesis and thylakoid membrane binding of the radioactively labeled herbicide dinoseb. Pesticide Biochemistry and Physiology 14: 86-97
174.Oettmeier W, Masson K, Johanningmeier U (1982) Evidence for two different herbicide-binding proteins at the reducing side of photosystem II. Biochimica et Biophysica Acta 679: 376-383
175.Oettmeier W, Soll HJ (1983) Competition between plastoquinone and 3-(3,4-dichlorphenyl) -1,1 -dimethylurea at the acceptor side of photosystem II. Biochimica et Biophysica Acta 724: 287-297
176.Ohad N, Hirschberg J (1990) A similar structure of the herbicide binding site in photosystem II of plants and cyanobacteria is demonstrated by site specific mutagenesis of the psbA gene. Photosynthesis Research 23: 73-79
177.Ohad N, Hirschberg J (1992) Mutations in the Dl subunit of photosystem II distinguish between quinine and herbicide binding sites. Plant Cell 4: 273-282
178.Ohta Y, Yoshioka T, Mochimaru M, Hisabori T, Sakurai H (1993) Tentoxin inhibits both photophosphorylation in thylakoids and the ATPase activity of isolated coupling factor F1 from the Cyanobacterium Anacystis nidulans. Plant Cell and Physiology 34: 523-529
179.Ohtani K,Yamamoto H,Akimitsu K(2002) Sensitivity to Alternaria alternata toxin in citrus because of altered mitochondrial RNA processing.Proceedings of the National Academy of Science of the United State of America 99:2439-2444
180.Omura S,Iwai Y,Takahashi Y,Sadakane N,Nakagawa A,Oiwa H,Hasegawa Y,Ikai T(1979) Herbimycin,a new antibiotic produced by a strain of Streptomyces.Journal of Antibiotics(Tokyo) 32:255-261
181.Omura S,Murata M,Hanaki H,Hinotozawa K,Oiwa R,Tanaka H(1984)Phosalacine,a new herbicidal antibiotic containing phosphinothricin.Fermentation,isolation,biological activity and mechanism of action.Journal of Antibiotics(Tokyo)37:829-835
182.Op den Camp RGL,Przybyla D,Oehsenhein C,Laloi C,Kim C,Danon A,Wagner D,Hideg E,Gobel C,Feussner I,Nater M,Apel K(2003) rapid induction of distinct stress responses after the release of singlet oxygen in Arabidopsis.Plant Cell 15:2320-2332
183.Orozco-Cardenas M,Ryan CA(1999) Hydrogen peroxide is generated systemically in plant leaves by wounding and systemin via the octadecanoid pathway.Proceedings of the National Academy of Science of the United State of America 96:6553-6557
184.Ortega X,Perez LM(2001) Participation of the phosphoinositide metabolism in the hypersensitive response of Citrus limon against Alternaria alternata.Biological Research 34:43-50
185.Ortega X,Polanco R,Castaneda P,Perez LM(2002) Signal transduction in lemon seedlings in the hypersensitive response against Alternaria alternata:participation of calmodulin,G-protein and protein kinases.Biological Research 35:373-383
186.Ouchane S,Picaud M,Astier C(1995) A new mutation in the pufL gene responsible for the terbutryn resistance phenotype in Rubrivivax gelatinosus.FEBS Letters 374:130-134
187.Ouzounidou G,Moustakas M,Strasser RJ(1997) Sites of action of copper in the photosynthetic apparatus of maize leaves:kinetic analysis of chlorophyll fluorescence,oxygen evolution,absorption changes and thermal dissipation as monitored by photoacoustie signals.Australian Journal of Plant Physiology 24:81-90
188.Palatnik J,Tognetti VB,Poli HO,Rodrignez RE,Blaneo N,Gattuso M,Hajirezaei MR,Sonnewald U,Vaile EM,Carrillo N(2003) Transgenic tobacco plants expressing antisense ferredoxin-NADP(H) reducase transcripts display increased susceptibility to photo-oxidative damage. Plant Journal 35: 332-341
189.Panda D, Rao DN, Sharma SG, Strasser RJ, Sarkar RK (2006) Submergence effect on rice genotypes during seedling stage: probing of submergence driven changes of photosystem 2 by chlorophyll a fluorescence induction O-J-I-P transients. Photosynthetica 44: 69-75
190.Peixoto F, Vicente J, Madeira VMC (2004) A comparative study of plant and animal mitochondria exposed to paraquat reveals that hydrogen peroxide is not related to the observed toxicity. Toxicology in Vitro 18: 733-739
191.Perchorowicz JT, Raynes DA, Jensen RG (1982) Measurement and preservation of the in vivo activation of ribulose 1,5-bisphosphate carboxylase in leaf extracts. Plant Physiology 69:1165-1168
192.Pfannschmidt T (2003) Chloroplast redox signal: how photosynthesis controls its own genes. Trends in Plant Science 8: 33-41
193.Pfister K, Arntzen CJ (1979) The mode of action of photosystem II -specific inhibitors in herbicide-resistant weed biotypes. Zeitschrift fur Naturforschung c-a Journal of Biosciences. 34c: 996-1009
194.Pfister K, Steinback KE, Gardner G, Arntzen J (1981) Photoaffinity labeling ofan herbicide receptor protein in chloroplast membranes. Proceedings of the National Academy of Science of the United State of America 78: 981-985
195.Pfister K, Radosevich SR, Arntzen C (1979) Modification of herbicide binding to photosystem II in two biotype of Senecio vulgaris L. Plant Physiology 64: 995-999
196.Philip H, John B (2000) Changes in chlorophyll fluorescence during exposure of Dunaliella tertiolecta to UV radiation indicate a dynamic interaction between damage and repair processes. Photosynthesis Research 63: 123-134
197.Pinto MCD, Paradiso A, Leonetti P, Gara LD (2006) Hydrogen peroxide, nitric oxide and cytosolic ascorbate peroxidase at the crossroad between defence and cell death. Plant Journal 48: 784-795
198.Porra RJ, Thompson WA, Kriedemann PE (1989) Determination of accurate extinction coefficients and simultaneous equations for assaying chlorophylls a and b extracted with four different solvents: verification of the concentration of chlorophyll standards by atomic absorption spectroscopy. Biochimica et Biophysica Acta 975: 384-394
199.Pospisil P,Dau H(2002) Valinomycin sensitivity proves that light-induced thylakoid voltages result in millisecond phase of chlorophyll fluorescence transients.Biochimiea et Biophysica Acta 1554:94-100
200.Powles SB,Shaner DL(2001) Herbicide resistant and world gains.CRC Press,Boca raton-London-New York-Washington,D.C.
201.Prado RD,Jorrín J,García-Torres L(1997) Weed and crop resistance to herbicides.Kluwer Academic Publishers,Dordreeht-Boston-London.
202.Prakash JSS,Srivastava A,Strasser RJ,Mohanty P(2003) Senescence-induced alterations in the photosystem Ⅱ functions of Cucumis sativus cotyledons:probing of senescence driven alterations ofphotosystem Ⅱ by chlorophyll α fluorescence induction O-J-I-P transients.Indian Journal of Biochemistry 40:160-168
203.Przibilla E,Heiss S,Johanningmeier U,Trebst(1991) Site-specific mutagenesis of the D1 subunit ofphotosystem Ⅱ in wild-type Chlamydomonas.Plant Cell 3:169-174
204.Qiang S(1998 a) On the potential of Alternaria Alternata as a mycoherbicide for Crofton weed.Doctor Thesis,Nanjing Agricultural University,Nanjing
205.Qiang S(1998 b) The history,status and prospect of the study on Crofton weed (Eupatorium adenophorum Spreng.),a worst worldwide weed.Wuhan Botanical Research 16:354-360
206.Qiang S,Chang Y,Wan ZX,Li YH(2002) Comparison on pathogenicity and other characteristics of five isolates of Alternaria alternata from Eupatorium adenophorum.Journal of Nanjing Agricultural University 25:23-27
207.Qiang S,Dong YF,An CF,Zhou B,Zhu YZ,Chen SG,Dai XB,Dai BJ,Cai JG.(2005) Biological control of weeds using the metabolites of Alternaria alternata(Fr.)Keissler.China Patent Application No.CN1644046,PCTICN2005/002367
208.Qiang S,Wan ZX,Dong YF,Li YH(1999) Phytotoxicity of rude metabolites produced by Alternaria alternata to Croften weed.In the Sustainable Management of Weeds Meeting the 21st Century in China.Guangxi Nationality Press,Nanning,pp 158-165
209.Ramm K,Ramm M,Liebermann B,Reuter G(1994) Studies of the biosynthesis of tentoxin by Altemaria alternata.Microbiology 140:3257-3266
210.Richter M,Wild A,and Ruble W(1994) Studies on the mechanism of photosystem Ⅱphotoinhibition.Ⅱ.The involvement of toxic oxygen species.Photosynthesis Reaearch 24:237-243
211. Rodriguez RE, Lodeyro A, Poli HO, Zurbriggen M, Peisker M, Palatnik JF, Tognetti VB, Tschiersch H, Hajirezaei MR, Valle EM, Carrillo N (2007) Transgenic tobacco plants overexpressing chloroplastic ferredoxin-NADP(H) reductase display normal rates of photosynthesis and increased tolerance to oxidative stress. Plant Physiology 143: 639-649
212.Rosen T, Fernandes PB, Marovich MA, Shen L, Mao J, Pernet AG (1989) Aromatic dienoyl tetramic acids. Novel antibacterial agents with activity against anaerobes and staphylococci. Journal of Medicine Chemistry 32: 1062-1069
213.Rosett BT, Sankhala RH, Stickings CE, Taylor MEU, Thomas R (1957) Studies in the biochemistry of micro-organisms. Metabolites of Alternaria tenuis auct: culture filtrate products. Biochemical Journal 67: 390-400
214.Rotem J (1994) The Genus Alternaria. Biology, Epidemiology and Pathogenicity. APS Press, New York.
215.Rusterucci C, Stallaert V, Milat ML, Pugin A, Ricci P, Blein JP (1996) Relationship between active oxygen species, lipid peroxidation, necrosis, and phytoalexin production induced by elicitins in Nicotiana. Plant physiology 111: 885-891
216.Rutherford AW, Krieger-Liszkay A (2001) Herbicide-induced oxidative stress in photosystem II. Trends in Biochemical sciences 26: 648-653
217.Ryan GF (1970) Resistance of common groundsel to simazine and atrazine. Weed Science 18: 614-616
218.Sakamoto M, Tada Y, Nakayashiki H, Tosa Y, Mayama S (2005) Two phases of intercellular reactive oxygen species production during victorin-induced cell death in oats. Journal of General Plant Pathology 71: 387-394
219.Samuilov VD, Lagunova EM, Dzyubinskaya EV, Izyumov DS, Kiselevsky DB, Makarova YV (2002) Involvement of chloroplasts in the programmed death of plant cells. Biochemistry (Moscow) 67: 627-634
220.Samuilov VD, Lagunova EM, Gostimsky SA, Timofeev KN, Gusev MV (2003) Role of chloroplast photosystem II and I in apoptosis of pea guard cells. Biochemistry (Moscow) 68: 912-917
221.Santolini J, Minoletti C, Gomis JM, Sigalat C, Andre F, and Haraux F (2002) An insight into the mechanism of inhibition and reactivation of the F(1)-ATPases by tentoxin. Biochemistry 41: 6008-6018
222Satoh K (1981) Fluorescence induction and activity of ferredoxin-NADP+ reductase in Bryopsis chloroplasts.Biochimica et Biophysica Acta 638:327-333
223.Scandalios JG(2002) The rise of ROS.Trends in Biochemical Sciences 27:483-486
224.Schansker G,Tóth SZ,Strasser RJ(2005) Methylviolegen and dibromothymoquinone treatments of pea leaves reveal the role of photosystem Ⅰ in the Chl α fluorescence rise OJIP.Biochimica et Biophysica Acta 1706:250-261
225.Schansker G,Strasser RJ(2005) Quantification of non-Q_B-reducing centers in leaves using a far-red pre-illumination.Photosynthesis Research 84:145-151
226.Schlegei B,Schmidtke M,Dorfelt H,Kleinwachter P,Grafe U(2001) Terpestacin and L-tenuazonie acid,inducers of pigment and aerial mycelium formation by Fusarium culmorum JP 15.Journal of Basic Microbiology 41:179-183
227.Schnick C,Kortgen N,Groth G(2002) Complete Inhibition of the Tentoxin-resistant F1-ATPase from Escherichia coli by the Phytopathogenic Inhibitor Tentoxin after Substitution of Critical Residues in the alpha - and beta -Subunit.Journal of Biological Chemistry 277:5103-5107
228.Schobert R,Jagusch C,Melanophy C,Mullen G(2004) Synthesis and reactions of polymer-bound Ph3P=C=C=O:a quick route to TeA and other optically pure 5-substituted tetramates.Organic and Biomolecular Chemistry 2:3524-3529
229.Schreiber U(1983) Chlorophyll fluorescence yield changes as a tool in plant physiology,Ⅰ.The measuring system.Photosynthesis Research 4:361-373
230.Schreiber U,Reising H,Neubauer C(1991) Contrasting pH-optima of light-driven O_2- and H_2O_2-reduction in spinach chloroplasts as measured via chlorophyll fluorescence quenching.ZeitschriR fur Naturforschung c-a Journal of Biosciences 46c:635-643
231.Schreiber U,Schliwa U,Bilger W(1986) Continuous recording of photochemical and non-photochemical chlorophyll fluorescence quenching with a new type of modulation fluorometer.Photosynthesis Research 10:51-62
232.Schreiber U,Vidaver W(1974) Chlorophyll fluorescence induction in anaerobic Scenedesmus obliquus.Biochimiea et Biophysica Acta 368:97-112
233.Seguin F,Le Bihan F,Leboulanger C,Berard A(2002) A risk assessment of pollution:induction of atrazine tolerance in phytoplankton communities in freshwater outdoor mesoeosms,using chlorophyll fluorescence as an endpoint.Water Research 36:3227-3236
234.Shanghai Society for Plant Physiology(1985) Modern Plant Physiology Experimental Handbook. Science Technology Press, Shang Hai, pp 5-10
235.Shanghai Institute for Plant Physiology of Chinese Acedamy Science (1999) Modern Plant Physiology Experimental Manual. Science Press, Shanghai, pp 104-106
236.Shao N, Krieger-Liszkay A, Schroda M, Beck CF (2007) A reporter system for the individual detection of hygrogen peroxide and singlet oxygen: its use for the assay of reactive oxygen species produced in vivo. Plant Journal 50: 475-487
237.Shimizu N, Hosogi N, Hyon GS, Jiang S, Inoue K, Park P (2006) Reactive oxygen species (ROS) generation and ROS-induced lipid peroxidation are associated with plasma membrane modifications in host cells in response to AK-toxin I from Alternaria alternate Japanese pear pathotype. Journal of General Plant Pathology 72: 6-15
238.Sinning I, Michel HP, Mathis,. Rutherford AW (1989) Characterization of four herbicide-resistant mutant of Rhodopseudomonas viridis by genetic analysis, electiron paramagretic resonance land optical spectroscopy. Biochemistry 28: 5544-5553
239.Soskic M, Plavsic D (2001) QSAR study of l,8-naphthyridin-4-ones as inhibitors of photosystem II. Journal of Chemical Information and Computer Sciences. 41: 1316-1321
240.Spassieva SD, Markham JE, Hille J (2002) The plant disease resistance gene Asc-1 prevents disruption of sphingolipid metabolism during AAL-toxin-induced programmed cell death. Plant Journal 32: 561-572
241.Spyridaki A, Fritzsch G, Kouimtzoglou E, Baciou L, Ghanotakis D (2000) The natural product capsaicin inhibits photosynthetic electron transport at the reducing side of photosystem II and purple bacterial reaction center: structural details of capsaicin binding. Biochimica et Biophysica Acta 1459: 69-76
242.Srivastava A, Guisse B, Greppin H, Strasser RJ (1997) Regulation of antenna structure and transport in photosystem II of Pisum sativum under elevated temperature probed by fast polyphasic chlorophyll a fluorescence transient: OKJIP. Biochimica et Biophysica Acta 1320: 95-106
243.Srivastava A, Juttner F, Strasser RJ (1998) Action of the allelochemical, fischerellin A, on photosystem II. Biochimica et Biophysica Acta 1364: 326-336
244.Srivastava A, Strasser RJ (1996) Stress and stress management of land plants during a regular day. Journal of Plant Physiology 148: 445-455
245.Srivastava A, Strasser RJ, Govindjee (1999) Greening of peas: parallel measurements of 77 K emission spectra, OJIP chlorophyll a fluorescence transient, period four oscillation of the initial fluorescence level,delayed light emission,and P700*.Photosynthetica 37:365-392
246.Steele JA,Uehytil TF,Durbin RD(1978) The stimulation of coupling factor 1ATPase by tentoxin.Biochimica et Biophysica Aeta 504:136-141
247.Steinback KE,Pfister K,and Arntzen CJ(1981) Trypsin-mediated removal of herbicide binding sites within the photosystem Ⅱ complex.Zeitschrift fur Naturforschung c-a Journal of Biosciences 36c:98-108
248.Stirbet A,Govindjee,Strasser BJ,Strasser RJ(1998) Chlorophyll α fluorescence induction in higher plants:modelling and numerical simulation.Journal of Theoretical Biology 193:131-151
249.Strange RN(2003) Introduction to Plant Pathogens.John Wiley & Son,Chlchester.
250.Strasser BJ(1997) Donor side capacity of photosystem Ⅱ probed by chlorophyll αfluorescence transients.Photosynthesis Research 52:147-155
251.Strasser BJ,Strasser RJ(1995) Measuring fast fluorescence transients to address environmental questions:the JIP-test.In Photosynthesis:from Light to Biophere,Kluwer Academic Publishers Press,the Netherland,pp 977-980
252.Strasser RJ,Govindjee(1992) The Fo and the O-J-I-P fluorescence rise in higher plants and algae.In Regulation of Chloroplast Biogenesis,Plenum Press,New York,pp 423-426
253.Strasser R J,Srivastava A,Govindjee(1995) Polyphasic chlorophyll α fluorescence transient in plants and eyanobateria.Photochemistry and Photobiology 61:32-42
254.Strasser RJ,Srivastava A,Tsimilli-Miehael M(2000) The fluorescence transient as a tool to characterize and screen photosynthetic samples.In Probing Photosynthesis:Mechanism,Regulation and Adaptation,Taylor and Francis,London,pp 445-483
255.Strasser RJ,Stirbet A(2001) Estimation of the energetic connectivity of PS Ⅱ centers in plants using the fluorescence rise O-J-I-P fitting of experimental data to three different PS Ⅱ models.Mathematics and Computers in Simulation 56:451-461
256.Strasser RJ,Tsimilli-Michael M,Srivastava A(2004) Analysis of the chlorophyll αfluorescence transient.In Chlorophyll Fluorescence:A Signature of Photosynthesis,Kluwer Academic Publishers Press,The Netherlands,pp 1-45
257.Strauss A,Krüger GHJ,Strasser RJ,Van Heerden PDR(2006) Ranking od dark chilling tolerance in soybean genotypes probed by the chlorophyll α fluorescence transient O-J-I-P.Environmental and Experimental Botany 56:147-157
258.Suntres Z (2002) Role of antioxidants in paraquat toxicity. Toxicology 180: 65-77
259.Sweat TA, Wolpert T (2007) Thioredoxin h5 is required for victorin sensitivity mediated by a CC-NBS-LRR gene in Arabidopsis. Plant Cell 19: 673-687
260.Tada Y, Hata S, Takata Y, Nakayashiki H, Tosa Y, Mayama S (2001) Induction and signaling of an apoptotic response typified by DNA laddering in the defense response of oats to infection and elicitors. Molecular Plant-Microbe Interactions 14: 477-486
261.Takahashi Y, Utsumi K, Yamamoto Y, Hatano A, Satoh K (1996) Genetic engineering of the processing site of D1 precursor protein of photosystem II reaction center in Chlamydomonas reinhardtii. Plant Cell and Physiology 37: 161-168
262.Tanaka A, Tsuge T (2000) Structural and functional complexity of the genomic region controlling AK-toxin biosynthesis and pathogenicity in the Japanese pear pathotype of Alternaria alternata. Molecular Plant-Microbe Interactions 13: 975-986
263.Tellenbach M, Gerber A, Boschetti A (1983) Herbicide-binding to thylakoid membranes of a DCMU-resistant mutant of Chlamydomonas reinhardtii. FEBS Letters 158: 147-150
264.Tischer W, Strotmann H (1977) Relationship between Inhibitor Binding by Chloroplasts and Inhibition of Photosynthetic Electron Transport. Biochimica et Biophysica Acta 460: 113
265.Toth SZ, Schansker G, Strasser RJ (2005) In intact leaves, the maximum fluorescence level (F_M) is independent of the redox state of the plastoquinone pool: A DCMU-inhibition study. Biochimica Biophysica Acta 1708: 275-282
266.Trebst A (1979) Inhibition of photosynthetic electron flow by phenol and diphenylether herbicides in control and trypsin-treated chloroplasts. Zeitschrift fur Naturforschung c-a Journal of Biosciences 34c: 986-991
267.Trebst A (1986) The topology of the plastoquinone and herbicide binding peptides of photosystem II in the thylakoid membrane. Zeitschrift fur Naturforschung c-a Journal of Biosciences 41C: 240-245
268.Trebst A (1987) The three-dimensional structure of the herbicide binding niche on the reaction center polypeptides of photosystem II. Zeitschrift fur Naturforschung c-a Journal of Biosciences 42c: 742-750
269.Trebst A, Draber W (1979) Structure activity correlations of recent herbicides in photosynthetic reactions. In Geissbuler H. (eds.), Advances in Pesticide Science. Pergamon Press, Oxford, pp 233-234
270.Trebst A,Draber W(1986) Inhibitors of photosystem Ⅱ and the topology of the herbicide and Q_B binding polypeptide in the thylakoid membrane.Photosynthesis Research 10:381-392
271.Tsimilli-Miehael M,Eggenberg P,Biro B,Koves-Pechy K,Voros I,Strasser RJ (2000) Synergistic and antagonistic effects of arbuscular mycorrhizal fungi and Azospirillum and Rhizobium nitrogen-fixers on the photosynthetic activity of alfalfa,probed by the polyphasic chlorophyll α fluorescence transient O-J-I-P.Applied Soil Ecology 15:169-182
272.Tsimilli-Michael M,Kürger GHJ,Strasser RJ(1995) Suboptimality as driving force for adaption:a study about the correlation of excitation light intensity and dynamic fluorescence emission in plants.In Photosynthesis:from Light to Biophere,Kluwer Academic Publishers Press,the Netherland,pp 981-984
273.Tylkowska K,Grabarkiewicz-Szczesna J,Iwanowska H(2003) Production of toxins by Alternaria alternata and A.radicina and their effects on germination of carrot seeds.Seed Science and Technology 31:309-316
274.Van Doom WG,Woltering EJ(2005) Mmany ways to exit? Cell death categories in plants.Trends in Plant Science 10:117-122
275.Van Rensen JJS(1984) Interaction of photosystem Ⅱ herbicides with bicarbonate and formate in their effects on photosynthetic electron flow.Zeitschrift fur Naturforschung c-a Journal of Biosciences 39c:374-377
276.Velthuys BR(1981) Electron- dependent competition between plastoquinone and inhibitors for binding to photosystem Ⅱ.FEBS Letters 126:277-281
277.Vermaas WFJ,Arntzen CJ(1983) Synthetic quiniues influencing herbicide binding and photosystem Ⅱ electron transport;the effects of triazine-resistance on quinone binding properties in thylakoid membranes.Biochimica et Biophysica Acta 725:483-491
278.Vermaas WFJ,Dohnt G,Renger G(1984) Binding and release kinetics of inhibitors of QA-oxidation in thylakoid membranes.Biochimiea et Biophysica Acta 765:74-83
279.Vieente JA,Peixoto F,Lopes ML,Madeira VM(2001) Differential sensitivities of plant and animal mitochondria to the herbicide paraquat.Journal of Biochemical and Molecular Toxicology 15:322-330
280.Viviani F,Little JP,Pallett KE(1998) The mode of action of isoxaflutole Ⅱ.Characterization of the inhibition of carrot 4-hydroxyphenylpyruvate dioxygenase by the diketonitrile derivative of isoxaflutole. Pesticide Biochemistry and Physiology 62: 125-134
281.Voet D, Voet JG, Pratt CW (2006) The process of photosynthesis. Fundamentals of Biochemistry. Life at the molecular level (Second Edition), Chapter 18. John Wiley & Sons, Inc., Asia, pp 590-626
282.Vredenberg WJ, Bulychev A (2003) Photoelectric effects on chlorophyll fluorescence of photosystem II in vivo. Kinetics in the absence and presence of valinomycin. Bioelectrochemistry 60: 87-95
283.Vredenberg WJ (2000) A three-state model for energy trapping and chlorophyll fluorescence in photosystem II incorporating radical pair recombination. Biophysical Journal 79: 26-38
284.Wagner D, Przybyla D, Op den Camp R, Kim C, Landgraf F, Lee KP, Wiirsch M, Laloi C, Nater M, Hideg E, Apel K (2004) The genetic basis of singlet oxygen-induced stress responses of Arabidopsis thaliana. Science 306: 1183-1185
285.Walton JD, Panaccione DG (1993) Host-selective toxins and disease specificity: perspectives and progress. Annual Review of Phytophathology 31: 275-303
286.Wan ZX, Qiang S, Xu SC, Sheng ZG, Dong YF (2001 a) Culture conditions for production of phytotoxinbu Alternaria alternata and plant range of toxicity. Chinese Journal of Biology Control 17: 10-15
287. Wan ZX, Zhu JJ, Qiang S (2001 b) The pathogenic mechanism of toxin of Altenaria alternata(Fr.) Keissler to Eupatorium adenophorum. Journal of Plant Resources and Environment 10: 47-50
288.Wang H, Li J, Bostock RM, Gilchrist DG (1996) Apoptosis: a functional paradigm for programmed plant cell death induced by a host-selective phytotoxin and invoked during development. Plant Cell 8: 375-391
289.Williamson JD, Scandalios JG (1992) Differential response of maize catalases and superoxide dismutases to the photoactivated fungal toxin cercosporin. Plant Journal 2: 351-358
290.Wu WJ, Gao XW (2004). Biological Pesticide and Application. Chemical Industry Press, Beijing, pp 73-153
291.Xiong J, Hutchison RS, Sayre RT, Govindjee (1997) Modification of photosystem II acceptor side function in a D1 mutant (arginine-269-glycine) of Chlamydomonas reinhardtii. Biochimica et Biophysica Acta 1322: 60-76
292.Xiong J,Subramaniam S,Govindjee(1996) Modeling of the D1/D2 proteins and cofactors of the photosystem Ⅱ reaction center:implications for herbicide and bicarbonate binding.Protein Science 5:2054-2073
293.Xiong J,Subramaniam S,Govindjee(1998) A knowledge-based three dimensional model of the photosystem Ⅱ reaction center of Chlamydomonas reinhardtii.Photosynthesis Research 56:229-254
294.Xu Q,Axelrod HL,Abresch EC,Paddock ML,Okamura MY,Feher G(2004)X-my structure determination of three mutants of the bacterial photosynthetic reaction centers from Rb.sphaeroides:altered proton transfer pathways.Structure 12:703-715
295.Yao N,Eisfelfler BJ,Marvin J,Greenberg(2004) The mitochondrion-an organelle commonly involved in programmed cell death in Arabidopsis thaliana.Plant Journal 40:596-610
296.Yao N,Imai S,Tada Y,Nakayashiki H,Tosa Y,Park P,Mayama S(2002) Apoptotic cell death is a common response to pathogen attack in oats.Molecular Plant-Microbe Interactions 15:1000-1007
297.Yao N,Tada Y,Park P,Nakayashiki H,Tosa Y,Mayama S(2001) Novel evidence for apoptotie cell response and differential signals in chromatin condensation and DNA cleavage in victorin-treated oats.Plant Journal 28:13-26
298.Yao N,Tada Y,Sakamoto M,Nakayashiki H,Park P,Tosa Y,Mayama S(2002)Mitochondrial oxidative burst involved in apoptotic response in oats.Plant Journal 30:567-579
299.Yuki H,Kariya K,Hashhnoto Y(1967) Synthesis and anti-tumor activity of tenuazonle acid analogues.Chemical & pharmaceutical bulletin(Tokyo).15:727-729
300.Zhang SP,Weng J,Pan JX,Tu TZ,Yao S,Xu CH(2003) Study on the photo-generation of superoxide radicals in Photosystem Ⅱ with EPR spin trapping techniques.Photosynthesis Research 75:41-48.
301.Zhang YS(1991) Reserch status of the toxins production by plant phathogenie fungi.Acta Mycologiea Sinica 10:169-81
302.Zhou B(2007) Multi key techniques for the commercialization of Alternaria Alternata AAC-Toxin bioherbicide.Doctor Thesis,Nanjing Agricultural University,Nanjing
303.Zhu XG,Govindjee,Baker NR(2005) Chlorophyll α fluorescence induction kinetics in leaves predicted from a model describing each discrete step of excitation energy and electron transfer associated with photosystem Ⅱ.Planta 223:114-133
304.Zong WX, Thompson CB (2006) Necrotic death as a cell fate. Genes and Development 20: 1-15
305.Zouni A, Witt HT, Kern J, Fromme P, Krauss N, Saenger W, Orth P (2001) Crystal structure of photosystem II from Synechococcus elongatus at 3.8 A resolution. Nature 409: 739-743
