双通道PAM-100荧光仪 Dual-PAM-100

dual-pam-100: 一套空前强大的测量系统
schreiber教授因发明pam系列调制叶绿素荧光仪而获得首届国际光合作用协会（ispr）创新奖

1985年开始商品化的全世界第一台调制荧光仪pam-100被几代科学家所广泛采用。dual-pam-100相当于两台pam-100的功能。一方面，它继承了pam-100的所有优点，可以进行复杂的叶绿素荧光分析（ps ii活性）；另一方面，它还可以通过测量p700的吸收变化来检测ps i的活性。特别需要强调的是，dual-pam-100可以在完全同步的情况下测量叶绿素荧光和p700吸收变化。此外，通过特殊的激发-检测单元还可以测量叶绿体或微藻的许多重要光合参数，如跨膜质子梯度delta ph（通过9-aa荧光或吖啶黄荧光）、类囊体膜的电势（通过类胡萝卜素的差示吸收，“p515”）和nadp的氧还状态（通过nadph荧光）等。如果需要极高的灵敏度可以通过连接光电倍增管检测器实现。

特点： 1）声誉卓著的pam-101/102/103的升级版 2）专业的dual-pam操作软件 3）自动测量模式下操作极其简单 4）创新的光-电配件使得仪器非常便携 5）整合式光源（红光、蓝光、远红光、单脉冲饱和闪光、多脉冲饱和闪光） 6）多版本可选，用于测量高等植物和微藻 7）利用饱和脉冲法测量ps i活性 8）同步测量ps i和ps ii的量子产量 9）可以分析蓝藻的光合作用 10）可选配用于测量ph、nadph和p515等的附件

功能： 1）ps i和ps ii的光化学量子产量 2）叶绿素荧光淬灭参数 3）系统间电子传递动力学 4）系统间电子载体库的大小（特殊的st/mt方法） 5）围绕ps i的环式电子传递动力学 6）ps i和ps ii能量传递的光响应曲线 7）荧光快速上升相（线性时间和对数时间） 8）测量pq库的还原态 9）测量光系统ii的吸收截面积

与pam-100相比，dual-pam-100的主要特点： 1）dual-pam-100完全由电脑控制，通过专业的windows操作软件dualpam进行。 2）软件dualpam除了基本的系统操作外，还提供许多特定的测量程序。 3）所有必需的光源（激发叶绿素荧光的红光和蓝光、测量p700的近红外光、红色和蓝色的光化光、单脉冲与多脉冲饱和闪光、远红光）均整合在基础系统中，不再需要复杂的电缆连接。 4）采用了专为dual-pam-100设计的许多新光-电配件，使得激发-检测单元和整合式光源非常便携、非常便于安装和拆卸。 5）所有的光源都可通过软件在2.5 μs的时间分辨率下控制。 6）测量光的频率范围非常大（1 hz～400 khz），因此同一个测量光源既可以用于测量fo，也可以用于诱发快速动力学（如荧光快速上升相或闪光弛豫动力学）。 7）用户可将针对特殊实验/样品的仪器设置存储起来，此后可在完全相同的设置下重复实验。 8）叶绿素荧光和p700的信号变化完全同步，并且是用同一个检测器检测，且不会互相干扰。 9）测量蓝藻时注意：用红光激发ps ii的荧光，用蓝光或远红光激发ps i。

dual-pam-100的这些特点开启了基础光合作用研究和应用光合作用研究的新途径。过去，同步测量ps i和ps ii的量子产量需要很强的专业背景和熟练的操作技巧，只有光合作用领域的少数专家会这项技术。现在，即使是初学者，也可迅速掌握同步测量ps i和ps ii活性的技术，不再需要复杂的操作技巧。

同步测量叶绿素荧光（ps ii）和p700(ps i)的诱导曲线

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同步测量叶绿素荧光（ps ii）和p700(ps i)的光响应曲线

专为dual-pam-100设计的windows操作软件dualpam功能强大，允许独立记录叶绿素荧光或p700，或同步记录叶绿素荧光和p700，记录每次打开饱和脉冲时的动力学变化，以线性时间多对数时间记录快速荧光上升动力学（分辨率2.5 us），分析各种荧光参数的变化等等。

p700的测量

以线性时间记录的荧光快速上升动力学

以对数时间记录的荧光快速上升动力学（相当于o-j-i-p相）

kramer的新荧光参数资料，这是一篇2004年发表在photosynthesis research 上的文章，见如下描述:

kramer提出的新参数：
ql，用于代替qp
y（ii）＝y＝（fm’-f）/fm’
y（npq) 表征ps ii处过量光能耗散为热，与光保护有关
y（no） 表征ps ii处过量光能引起的光损伤
y(ii)+y(npq)+y(no)=1

schreiber教授和其学生klughammer博士新设计的dual-pam-100中，借鉴测量叶绿素荧光的方法，新增了几个参数：
y（i） ps i的量子产量或ps i的光合效率
y（nd) 表征ps i处过量光能耗散为热，与光保护有关
y（na） 表征ps i处过量光能引起的光损伤
y(i)+y(nd)+y(na)=1

色素分子处于氧化态和还原态时，或增加/减少亚基后，其吸收峰会有变化。基于此原理，p700吸收变化、p515吸收变化（类胡萝卜素能态）、p505吸收变化（叶黄素循环）等均可通过dual-pam-100测量。主机共用，只是更换激发-检测单元即可。此外，dual-pam-100还可用于测量nadph荧光、9aa荧光（跨膜质子梯度）等。

部分利用dual-pam-100发表的p700文献

1. aronsson h, schöttler ma, kelly aa, sundqvist c, dörmann p, karim s, jarvis p: monogalactosyldiacylglycerol deficiency in arabidopsis affects pigment composition in the prolamellar body and impairs thylakoid membrane energization and photoprotection in leaves. plant physiology 2008;148:580-592.
2. bailey s, melis a, mackey krm, cardol p, finazzi g, dijken gv, berge gm, arrigo k, shrager j, grossman a: alternative photosynthetic electron flow to oxygen in marine synechococcus biochimica et biophysica acta 2008;1777:269-276.
3. ehlert b, schöttler ma, tischendorf g, ludwig-müller j, bock r: the paramutated sulfurea locus of tomato is involved in auxin biosynthesis. journal of experimental biology 2008:in press.
4. ma w, chen l, wei l, wang q: excitation energy transfer between photosystems in the cyanobacterium synechocystis 6803 journal of luminescence 2008;128:546-548.
5. rogalski m, schöttler ma, thiele w, schulze wx, bock r: rpl33, a nonessential plastid-encoded ribosomal protein in tobacco, is required under cold stress conditions. the plant cell 2007;20:2221-2237.
6. schöttler m, flügel c, thiele w, stegemann s, bock r: the plastome-encoded psaj subunit is required for efficient photosystem i excitation, but not for plastocyanin oxidation in tobacco. biochemical journal 2007;403:251-260.
7. volkmer t, schneider d, bernát g, kirchhoff h, wenk s-o, rögner m: ssr2998 of synechocystis sp. pcc 6803 is involved in regulation of cyanobacterial electron transport and associated with the cytochrome b6f complex. journal of biological chemistry 2007;282:3730-3737.
8. lohmann a, schottler ma, brehelin c, kessler f, bock r, cahoon eb, dormann p: deficiency in phylloquinone (vitamin k1) methylation affects prenyl quinone distribution, photosystem i abundance, and anthocyanin accumulation in the arabidopsis atmeng mutant. journal of biological chemistry 2006;281:40461-40472.
9. schöttler ma, flügel c, thiele w, bock r: knock-out of the plastid-encoded petl subunit results in reduced stability and accelerated leaf age-dependent loss of the cytochrome b6f complex. journal of biological chemistry 2006;282:976-985.


部分利用pam-101/102/103发表的p700文献

1. wang p, duan w, takabayashi a, endo t, shikanai t, ye j-y, mi h, 2006. chloroplastic nad(p)h dehydrogenase in tobacco leaves functions in alleviation of oxidative damage caused by temperature stress. plant physiology 141: 465-474.
2. xu d, liu x, zhao j, zhao j, 2005. fesm, a membrane iron-sulfur protein, is required for cyclic electron flow around photosystem i and photoheterotrophic growth of the cyanobacterium synechococcus sp. pcc 7002. plant physiology 138: 1586-1595.
3. pikulenko mm, bulychev aa, 2005. light-triggered action potentials and changes in quantum efficiency of photosystem ii in anthoceros cells russian journal of plant physiology 52: 584-590.
4. peterson rb, 2005. psbs genotype in relation to coordinated function of ps ii and ps i in arabidopsis leaves. photosynthesis research 85: 205-219.
5. okegawa y, tsuyama m, kobayashi y, shikanai t, 2005. the pgr1 mutation in the rieske subunit of the cytochrome b6f complex does not affect pgr5-dependent cyclic electron transport around photosystem i. the journal of biological chemistry 280: 28332-28336.
6. munné-bosch s, shikanai t, asada k, 2005. enhanced ferredoxin-dependent cyclic electron flow around photosystem i and α-tocopherol quinone accumulation in water-stressed ndhb-inactivated tobacco mutants. planta 222: 502-511.
7. munné-bosch s, shikanai t, asada k, 2005. enhanced ferredoxin-dependent cyclic electron flow around photosystem i and -tocopherol quinone accumulation in water-stressed ndhb-inactivated tobacco mutants. planta 222: 502-511.
8. li x-g, bi y-p, zhao s-j, meng q-w, he q-w, zou q, 2005. effects of short-term chilling stress on the photosystems and chloroplast ultrastructure in sweet pepper. agricultural sciences in china 4: 429-435.
9. laisk a, eichelmann h, oja v, rasulov b, padu e, bichele i, pettai h, kull o, 2005. adjustment of leaf photosynthesis to shade in a natural canopy: rate parameters. plant cell and environment 28: 375-388.
10. ivanov b, asada k, kramer dm, edwards g, 2005. characterization of photosynthetic electron transport in bundle sheath cells of maize. i. ascorbate effectively stimulates cyclic electron flow around ps i. planta 220: 572-581.
11. ifuku k, yamamoto y, ono t-a, ishihara s, sato f, 2005. psbp protein, but not psbq protein, is essential for the regulation and stabilization of photosystem ii in higher plants. plant physiology 139: 1175–1184.
12. hirotsu n, makino a, yokota s, mae t, 2005. the photosynthetic properties of rice leaves treated with low temperature and high irradiance plant cell and physiology 46: 1377-1383.
13. gerloff-elias a, spijkerman e, pröschold t, 2005. effect of external ph on the growth, photosynthesis and photosynthetic electron transport of chlamydomonas acidophila negoro, isolated from an extremely acidic lake (ph 2.6). plant cell and environment 28: 1218-1229.
14. georgieva k, maslenkova l, peeva v, markovska y, stefanov d, tuba z, 2005. comparative study on the changes in photosynthetic activity of the homoiochlorophyllous desiccation-tolerant haberlea rhodopensis and desiccation-sensitive spinach leaves during desiccation and rehydration. photosynthesis research 85: 191-203.
15. endo t, kawase d, sato f, 2005. stromal over-reduction by high-light stress as measured by decreases in p700 oxidation by far-red light and its physiological relevance. plant cell and physiology 46: 775-781.
16. egorova ea, drozdova is, bukhov ng, 2005. modulating effect of far-red light on activities of alternative electron transport pathways related to photosystem i russian journal of plant physiology 52: 709-716.
17. dzhibladze tg, polesskaya og, alekhina nd, egorova ea, bukhov ng, 2005. redox states of photosystems i and ii in irradiated leaves of wheat seedlings grown under different conditions of nitrogen nutrition. russian journal of plant physiology 52: 141-146.
18. ducruet j-m, roman m, havaux m, janda t, gallais a, 2005. cyclic electron flow around psi monitored by afterglow luminescence in leaves of maize inbred lines (zea mays l.): correlation with chilling tolerance. planta 221: 567-579.
19. ducruet j-m, roman m, havaux m, janda t, gallais a, 2005. cyclic electron flow around ps i monitored by afterglow luminescence in leaves of maize inbred lines (zea mays l.): correlation with chilling tolerance. planta 221: 567-579.
20. bukhov ng, egorova ea, 2005. nonmonotonic redox conversions of p700 in leaves irradiated with far-red light originate from variable contributions of several alternative electron transport pathways during the induction period russian journal of plant physiology 52: 571-577.
21. bukhov ng, egorova ea, 2005. identification of ferredoxin-dependent cyclic electron transport around photosystem i using the kinetics of dark p700+ reduction. russian journal of plant physiology 52: 283-287.
22. bukhov ng, egorova ea, 2005. evidence for the operation of alternative electron transport routes through photosystem i in intact barley leaves under weak and moderate white light. russian journal of plant physiology 52: 1-6.
23. bukhov ng, dzhibladze tg, egorova ea, 2005. elevated temperatures inhibit ferredoxin-dependent cyclic electron flow around photosystem i russian journal of plant physiology 52: 578-583.
24. zhang s, scheller hv, 2004. photoinhibition of photosystem i at chilling temperature and subsequent recovery in arabidopsis thaliana plant cell and physiology 45: 1595-1602.
25. zhang s, scheller hv, 2004. light-harvesting complex ii binds to several small subunits of photosystem i. the journal of biological chemistry 279: 3180-3187.
26. yabe t, morimoto k, kikuchi s, nishio k, terashima i, nakai m, 2004. the arabidopsis chloroplastic nifu-like protein cnfu, which can act as an iron-sulfur cluster scaffold protein, is required for biogenesis of ferredoxin and photosystem i the plant cell 16: 993-1007.
27. stöckel j, oelmüller r, 2004. a novel protein for photosystem i biogenesis. the journal of biological chemistry 279: 10243-10251.
28. schöttler ma, kirchhoff h, weis e, 2004. the role of plastocyanin in the adjustment of the photosynthetic electron transport to the carbon metabolism in tobacco. plant physiology 136: 4265-4274.
29. mackenzie tdb, burns ra, campbell da, 2004. carbon status constrains light acclimation in the cyanobacterium synechococcus elongatus. plant physiology 136: 3301-3312.
30. bukhov ng, govindachary s, rajagopal s, joly d, carpentier r, 2004. enhanced rates of p700+ dark-reduction in leaves of cucumis sativus l. photoinhibited at chilling temperature. planta 218: 852-861.
31. bukhov ng, govindachary s, egorova ea, carpentier r, 2004. recovery of photosystem i and ii activities during re-hydration of lichen hypogymnia physodes thalli. planta 219: 110-120.
32. amann k, lezhneva l, wanner g, herrmann rg, meurer j, 2004. accumulation of photosystem one1, a member of a novel gene family, is required for accumulation of [4fe-4s] culster-containing chloroplast complexes and antenna proteins. the plant cell 16: 3084-3097.
33. 姚正菊, 叶济宇, 米华玲, 2003. 高温胁迫对烟草叶绿体nadph脱氢酶复合体活性的促进. 植物生理与分子生物学学报 29: 395-400.
34. wang h-w, su j-h, shen y-g, 2003. difference in response of photosynthesis to bisulfite between two wheat genotypes. journal of plant physiology and molecular biology 29: 27-32.
35. wang h-w, mi h, ye j-y, deng y, shen y-k, 2003. low concentrations of nahso3 increase cyclic photophosphorylation and photosynthesis in cyanobacterium synechocystis pcc6803. photosynthesis research 75: 151-159.
36. shikanai t, müller-moulé p, munekage y, niyogi kk, pilonc m, 2003. paa1, a p-type atpase of arabidopsis, functions in copper transport in chloroplasts the plant cell 15: 1333-1346.
37. ryu j-y, suh k-h, chung y-h, park y-m, chow ws, park y-i, 2003. cytochrome c oxidase of the cyanobacterium synechocystis sp. pcc 6803 protects photosynthesis from salt stress. molecules and cells 16: 74-77.
38. rajagopala s, egorova ea, bukhov ng, carpentier r, 2003. quenching of excited states of chlorophyll molecules in submembrane fractions of photosystem i by exogenous quinones. biochimica et biophysica acta 1606: 147-152.
39. rajagopal s, carpentier r, 2003. retardation of photo-induced changes in photosystem i submembrane particles by glycinebetaine and sucrose. photosynthesis research 78: 77-85.
40. oja v, eichelmann h, peterson rb, rasulov b, laisk a, 2003. deciphering the 820 nm signal: redox state of donor side and quantum yield of photosystem i in leaves. photosynthesis research 78: 1-15.
41. maiwald d, dietzmann a, jahns p, pesaresi p, joliot p, joliot a, levin jz, salamini f, leister d, 2003. knock-out of the genes coding for the rieske protein and the atp-synthase δ-subunit of arabidopsis. effects on photosynthesis, thylakoid protein composition, and nuclear chloroplast gene expression. plant physiology 133: 191-220.
42. krause gh, grube e, virgo a, winter k, 2003. sudden exposure to solar uv-b radiation reduces net co2 uptake and photosystem i efficiency in shade-acclimated tropical tree seedlings. plant physiology 131: 745-752.
43. hanson dt, franklin la, samuelsson g, badger mr, 2003. the chlamydomonas reinhardtii cia3 mutant lacking a thylakoid lumen-localized carbonic anhydrase is limited by co2 supply to rubisco and not photosystem ii function in vivo. plant physiology 132: 2267-2275.
44. golding aj, johnson gn, 2003. down-regulation of linear and activation of cyclic electron transport during drought. planta 218: 107-114.
45. egorova ea, bukhov ng, heber u, samson g, carpentier r, 2003. effect of the pool size of stromal reductants on the alternative pathway of electron transfer to photosystem i in chloroplasts of intact leaves. russian journal of plant physiology 50: 431-440.
46. 姚正菊, 米华玲, 叶济宇, 2002. 光诱导叶片p700氧化还原的测量. 植物生理性通讯 38: 54－56.
47. yamasaki t, yamakawa t, yamane y, koike h, satoh k, katoh s, 2002. temperature acclimation of photosynthesis and related changes in photosystem ii electron transport in winter wheat. plant physiology 128: 1087-1097.
48. varotto c, pesaresi p, jahns p, leßnick a, tizzano m, schiavon f, salamini f, leister d, 2002. single and double knockouts of the genes for photosystem i subunits g, k, and h of arabidopsis. effects on photosystem i composition, photosynthetic electron flow, and state transitions. plant physiology 129: 616-624.
49. morgan-kiss rm, ivanov ag, huner npa, 2002. tha antarctic psychrophile, chlamydomonas subcaudata, is deficient in state i-state ii transitions. planta 214: 435-445.
50. miskiewicz e, ivanov ag, huner npa, 2002. stoichiometry of the photosynthetic apparatus and phycobilisome structure of the cyanobacterium plectonema boryanum utex 485 are regulated by both light and temperature. plant physiology 130: 1414-1425.
51. makino a, miyake c, yokota a, 2002. physiological functions of the water–water cycle (mehler reaction) and the cyclic electron flow around psi in rice leaves. plant cell and physiology 43: 1017-1026.
52. laisk a, oja v, rasulov b, rämma h, eichelmann h, kasparova i, pettai h, padu e, vapaavuori e, 2002. a computer-operated routine of gas exchange and optical measurements to diagnose photosynthetic apparatus in leaves. plant cell and environment 25: 923-943.
53. joët t, cournac l, peltier g, havaux m, 2002. cyclic electron flow around photosystem i in c3 plants. in vivo control by the redox state of chloroplasts and involvement of the nadh-dehydrogenase complex plant physiology 128: 760-769.
54. jin m-x, mi h, 2002. thermal sensitivity of the pool size of electrons available to p700+ reduction in intact maize leaves. photosynthetica 40: 437-439.
55. jin m-x, li d-y, mi h, 2002. effects of high temperature on chlorophyll fluorescence induction and the kinetics of far red radiation-induced relaxation of apparent fo in maize leaves. photosynthetica 40: 581-586.
56. jeong sw, choi sm, lee ds, ahn sn, hur y, chow ws, park y-i, 2002. differential susceptibility of photosynthesis to light-chilling stress in rice (oryza sativa l.) depends on the capacity for photochemical dissipation of light. molecules and cells 13: 419-428.
57. bukhov ng, rajagopal s, carpentier r, 2002. characterization of p700 as a photochemical quencher in isolated photosystem i particles using simultaneous measurements of absorbance changes at 830 nm and photoacoustic signal. photosynthesis research 74: 295-302.
58. thomas dj, thomas j, youderian pa, herbert sk, 2001. photoinhibition and light-induced cyclic electron transport in ndhb- and psae- mutants of synechocystis sp. pcc 6803. plant cell and physiology 42: 803-812.
59. peterson r, havir e, 2001. photosynthetic properties of an arabidopsis thaliana mutant possessing a defective psbs gene. planta 214: 142-152.
60. lennartz k, plücken h, seidler a, westhoff p, bechtold n, meierhoff k, 2001. hcf164 encodes a thioredoxin-like protein involved in the biogenesis of the cytochrome b6f complex in arabidopsis. the plant cell 13: 2539-2551.
61. jin m-x, yao z-j, mi h, 2001. multi-phasic kinetics of p700+ dark re-reduction in nicotiana tabacum. photosynthetica 39: 419-425.
62. durchan m, vácha f, krieger-liszkay a, 2001. effects of severe co2 starvation on the photosynthetic electron transport chain in tabacco plants. photosynthesis research 68: 203-213.
63. bukhov n, carpentier r, samson g, 2001. heterogeneity of photosystem i reaction centers in barley leaves as related to the donation from stromal reductants. photosynthesis research 70: 273-279.
64. barth c, krause gh, winter k, 2001. responses of photosystem i compared with photosystem ii to high-light stress in tropical shade and sun leaves. plant cell and environment 24: 163-176.
65. baena-gonzález e, gray jc, tyystjärvi e, aro e-m, mäenpää p, 2001. abnormal regulation of photosynthetic electron transport in a chloroplast ycf9 inactivation mutant. the journal of biological chemistry 276: 20795-20802.
66. allakhverdiev si, kinoshita m, inaba m, suzuki i, murata n, 2001. unsaturated fatty acids in membrane lipids protect the photosynthetic machinery against salt-induced damage in synechococcus. plant physiology 125: 1842-1853.
67. ohkawa h, pakrasi hb, ogawa t, 2000. two types of functionally distinct nad(p)h dehydrogenases in synechocystis sp. strain pcc 6803. the journal of biological chemistry 275: 31630-31634.
68. lunde c, jensen pe, haldrup a, knoetzel j, scheller hv, 2000. the psi-h subunit of photosystem i is essential for state transitions in plant photosynthesis. nature 408: 613-615.
69. jensen pe, gilpin m, knoetzel j, scheller hv, 2000. the psi-k subunit of photosystem i is involved in the interaction between light-harvesting complex i and the photosystem i reaction center core. the journal of biological chemistry 275: 24701-24708.
70. ivanov ag, park y-i, miskiewicz e, raven ja, huner npa, öquist g, 2000. iron stress restricts photosynthetic intersystem electron transport in synechococcus sp. pcc 7942 febs letters 485: 173-177.
71. haldrup a, simpson dj, scheller hv, 2000. down-regulation of the psi-f subunit of photosystem i (psi) in arabidopsis thaliana. the psi-f subunit is essential for photoautotrophic growth and contributes to antenna function. the journal of biological chemistry 275: 31211-31218.
72. naver h, haldrup a, scheller hv, 1999. cosuppression of photosystem i subunit psi-h in arabidopsis thaliana. efficient electron transfer and stability of photosystem i is dependent upon the psi-h subunit. the journal of biological chemistry 274: 10784-10789.
73. manuel n, cornic g, aubert s, choler p, bligny r, heber u, 1999. protection against photoinhibition in the alpine plant geum montanum. oecologia 119: 149-158.
74. endo t, shikanai t, takabayashi a, asada k, sato f, 1999. the role of chloroplastic nad(p)h dehydrogenase in photoprotection. febs letters 457: 5-8.
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