dual-pam-100: 一套*強(qiáng)大的測量系統(tǒng)
schreiber教授因發(fā)明pam系列調(diào)制葉綠素?zé)晒鈨x而獲得首屆光合作用協(xié)會（ispr）創(chuàng)新獎

1985年開始商品化的**臺調(diào)制熒光儀pam-100被幾代科學(xué)家所廣泛采用。dual-pam-100相當(dāng)于兩臺pam-100的功能。一方面，它繼承了pam-100的所有優(yōu)點，可以進(jìn)行復(fù)雜的葉綠素?zé)晒夥治觯╬s ii活性）；另一方面，它還可以通過測量p700的吸收變化來檢測ps i的活性。特別需要強(qiáng)調(diào)的是，dual-pam-100可以在*同步的情況下測量葉綠素?zé)晒夂蚿700吸收變化。此外，通過特殊的激發(fā)-檢測單元還可以測量葉綠體或微藻的許多重要光合參數(shù)，如跨膜質(zhì)子梯度delta ph（通過9-aa熒光或吖啶黃熒光）、類囊體膜的電勢（通過類胡蘿卜素的差示吸收，“p515"）和nadp的氧還狀態(tài)（通過nadph熒光）等。如果需要*的靈敏度可以通過連接光電倍增管檢測器實現(xiàn)。

特點： 1）聲譽(yù)*的pam-101/102/103的升級版 2）專業(yè)的dual-pam操作軟件 3）自動測量模式下操作極其簡單 4）創(chuàng)新的光-電配件使得儀器非常便攜 5）整合式光源（紅光、藍(lán)光、遠(yuǎn)紅光、單脈沖飽和閃光、多脈沖飽和閃光） 6）多版本可選，用于測量高等植物和微藻 7）利用飽和脈沖法測量ps i活性 8）同步測量ps i和ps ii的量子產(chǎn)量 9）可以分析藍(lán)藻的光合作用 10）可選配用于測量ph、nadph和p515等的附件

功能： 1）ps i和ps ii的光化學(xué)量子產(chǎn)量 2）葉綠素?zé)晒獯銣鐓?shù) 3）系統(tǒng)間電子傳遞動力學(xué) 4）系統(tǒng)間電子載體庫的大?。ㄌ厥獾膕t/mt方法） 5）圍繞ps i的環(huán)式電子傳遞動力學(xué) 6）ps i和ps ii能量傳遞的光響應(yīng)曲線 7）熒光快速上升相（線性時間和對數(shù)時間） 8）測量pq庫的還原態(tài) 9）測量光系統(tǒng)ii的吸收截面積

與pam-100相比，dual-pam-100的主要特點： 1）dual-pam-100*由電腦控制，通過專業(yè)的windows操作軟件dualpam進(jìn)行。 2）軟件dualpam除了基本的系統(tǒng)操作外，還提供許多特定的測量程序。 3）所有必需的光源（激發(fā)葉綠素?zé)晒獾募t光和藍(lán)光、測量p700的近紅外光、紅色和藍(lán)色的光化光、單脈沖與多脈沖飽和閃光、遠(yuǎn)紅光）均整合在基礎(chǔ)系統(tǒng)中，不再需要復(fù)雜的電纜連接。 4）采用了專為dual-pam-100設(shè)計的許多新光-電配件，使得激發(fā)-檢測單元和整合式光源非常便攜、非常便于安裝和拆卸。 5）所有的光源都可通過軟件在2.5 μs的時間分辨率下控制。 6）測量光的頻率范圍非常大（1 hz～400 khz），因此同一個測量光源既可以用于測量fo，也可以用于誘發(fā)快速動力學(xué)（如熒光快速上升相或閃光弛豫動力學(xué)）。 7）用戶可將針對特殊實驗/樣品的儀器設(shè)置存儲起來，此后可在*相同的設(shè)置下重復(fù)實驗。 8）葉綠素?zé)晒夂蚿700的信號變化*同步，并且是用同一個檢測器檢測，且不會互相干擾。 9）測量藍(lán)藻時注意：用紅光激發(fā)ps ii的熒光，用藍(lán)光或遠(yuǎn)紅光激發(fā)ps i。

dual-pam-100的這些特點開啟了基礎(chǔ)光合作用研究和應(yīng)用光合作用研究的新途徑。過去，同步測量ps i和ps ii的量子產(chǎn)量需要很強(qiáng)的專業(yè)背景和熟練的操作技巧，只有光合作用領(lǐng)域的少數(shù)專家會這項技術(shù)?，F(xiàn)在，即使是初學(xué)者，也可迅速掌握同步測量ps i和ps ii活性的技術(shù)，不再需要復(fù)雜的操作技巧。

同步測量葉綠素?zé)晒猓╬s ii）和p700(ps i)的誘導(dǎo)曲線

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同步測量葉綠素?zé)晒猓╬s ii）和p700(ps i)的光響應(yīng)曲線

專為dual-pam-100設(shè)計的windows操作軟件dualpam功能強(qiáng)大，允許獨立記錄葉綠素?zé)晒饣騪700，或同步記錄葉綠素?zé)晒夂蚿700，記錄每次打開飽和脈沖時的動力學(xué)變化，以線性時間多對數(shù)時間記錄快速熒光上升動力學(xué)（分辨率2.5 us），分析各種熒光參數(shù)的變化等等。

p700的測量

以線性時間記錄的熒光快速上升動力學(xué)

以對數(shù)時間記錄的熒光快速上升動力學(xué)（相當(dāng)于o-j-i-p相）

kramer的新熒光參數(shù)資料，這是一篇2004年發(fā)表在photosynthesis research 上的文章，見如下描述:

kramer提出的新參數(shù)：
ql，用于代替qp
y（ii）＝y(tǒng)＝（fm’-f）/fm’
y（npq) 表征ps ii處過量光能耗散為熱，與光保護(hù)有關(guān)
y（no） 表征ps ii處過量光能引起的光損傷
y(ii)+y(npq)+y(no)=1

schreiber教授和其學(xué)生klughammer博士新設(shè)計的dual-pam-100中，借鑒測量葉綠素?zé)晒獾姆椒?，新增了幾個參數(shù)：
y（i） ps i的量子產(chǎn)量或ps i的光合效率
y（nd) 表征ps i處過量光能耗散為熱，與光保護(hù)有關(guān)
y（na） 表征ps i處過量光能引起的光損傷
y(i)+y(nd)+y(na)=1

色素分子處于氧化態(tài)和還原態(tài)時，或增加/減少亞基后，其吸收峰會有變化?；诖嗽?，p700吸收變化、p515吸收變化（類胡蘿卜素能態(tài)）、p505吸收變化（葉黃素循環(huán)）等均可通過dual-pam-100測量。主機(jī)共用，只是更換激發(fā)-檢測單元即可。此外，dual-pam-100還可用于測量nadph熒光、9aa熒光（跨膜質(zhì)子梯度）等。

部分利用dual-pam-100發(fā)表的p700文獻(xiàn)

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 , 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發(fā)表的p700文獻(xiàn)

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, 葉濟(jì)宇, 米華玲, 2003. 高溫脅迫對煙草葉綠體nadph脫氫酶復(fù)合體活性的促進(jìn). 植物生理與分子生物學(xué)學(xué)報 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, 米華玲, 葉濟(jì)宇, 2002. 光誘導(dǎo)葉片p700氧化還原的測量. 植物生理性通訊 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.
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