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M系列調(diào)制葉綠素?zé)晒獬上裣到y(tǒng)IMAGING-PAM
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CI-340手持式光合儀；CI-203手持式激光葉面積儀；CI-202葉面積儀；CI-110冠層分析儀；CI-600根系生長(zhǎng)監(jiān)測(cè)儀
傳統(tǒng)的光纖型調(diào)制熒光儀（如pam-2100、mini-pam等）只能測(cè)量葉片上一個(gè)點(diǎn)的光合活性。利用一個(gè)點(diǎn)的數(shù)據(jù)代表一個(gè)葉片，利用一個(gè)葉片代表一個(gè)植株，進(jìn)而代表一個(gè)群體（如森林、大田作物等），這種方法的誤差是比較大的。
從1980年代末期開(kāi)始，科研人員就期望能通過(guò)成像型熒光儀來(lái)測(cè)量全葉片光合活性，并進(jìn)行了不懈的嘗試，但受技術(shù)上的限制，所設(shè)計(jì)的儀器無(wú)法商品化或商品化了但得不到大家認(rèn)可。其中一個(gè)很重要的原因就是能夠發(fā)出飽和脈沖水平強(qiáng)光的二極管（led）尚未面世。要利用調(diào)制熒光技術(shù)測(cè)量全葉片水平的光合作用，首先要保證葉片上任何一點(diǎn)所接受到的光強(qiáng)必須是*相同的。調(diào)制熒光技術(shù)要求光源必須能發(fā)出很強(qiáng)的飽和脈沖光。鹵素?zé)裟馨l(fā)出很強(qiáng)的光，但其光場(chǎng)非常不均勻，根本不能用于成像！裝在一個(gè)平面上的led陣列發(fā)出的光很均勻，但在2000年前，能發(fā)出*光合有效輻射（par）的led根本沒(méi)有面世！
2000年，能發(fā)出*par的藍(lán)光led面世。2001年，zui的調(diào)制熒光儀制造商德國(guó)walz公司設(shè)計(jì)制造了真正的*臺(tái)多功能調(diào)制熒光成像系統(tǒng)imaging-pam。imaging-pam采用*發(fā)光led作為光源，保證葉片表明受光均勻且光強(qiáng)足夠強(qiáng)；imaging-pam采用ccd作為檢測(cè)器，能檢測(cè)葉片上每個(gè)像素的光合作用；imaging-pam秉承了walz公司pam系列熒光儀的一貫優(yōu)點(diǎn)，功能強(qiáng)大，測(cè)量參數(shù)多，操作極其簡(jiǎn)單，一面世就受到植物學(xué)家的青睞，迅速占領(lǐng)市場(chǎng)。
2005年，walz又推出了m系列imaging-pam，一個(gè)主機(jī)可以連接不同的探頭（microscopy-，micro-，mini-和maxi-探頭），分別在130×150 um、3.5×4.5 mm、24×32 mm或10×13 cm的面積上測(cè)量熒光成像?，F(xiàn)在，只需一個(gè)主機(jī)連接不同的探頭，即可滿足從單細(xì)胞到全葉片，從分子生物學(xué)到生態(tài)學(xué)研究的全面需要。
m系列imaging-pam不同版本的比較
maxi-版 | mini-版 | micro-版 | microscopy-版 |
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成像面積10×13 cm | 成像面積24×32 mm | 成像面積3.5×4.5 mm | 成像面積130×150 um |
放大1.5倍 | 放大6倍 | 放大45倍 | 放大130-1300倍 |
功 能
＊ 一個(gè)主機(jī)連接不同的探頭可滿足從單細(xì)胞到全葉片、從分子生物學(xué)到生態(tài)學(xué)的不同需求
＊ 全葉片光合作用分析（熒光成像），可測(cè)熒光誘導(dǎo)曲線并進(jìn)行淬滅分析
＊ 可測(cè)快速光響應(yīng)曲線（120 s內(nèi)完成，比光合放氧和氣體交換等技術(shù)快得多）
＊ 葉片光合作用的橫向異質(zhì)性檢測(cè)
＊ *相同的條件下同時(shí)測(cè)量多個(gè)樣品（植物、地衣、苔蘚、微藻等）
＊ 遺傳育種、突變株篩選的強(qiáng)大工具
＊ 不同的測(cè)量面積，不同的分辨率
＊ 可利用多孔板（如96孔板）做多個(gè)微藻樣品的同時(shí)成像
＊ 脅迫損傷的早期檢測(cè)
＊ 不連接顯微鏡即可測(cè)量綠色熒光蛋白（gfp）熒光
＊ 可測(cè)量葉片吸光系數(shù)
測(cè)量參數(shù)
＊ 以上所有參數(shù)均可成像
＊ 吸光系數(shù)abs和新參數(shù)ql、y(npq)和y(no)的成像是imaging-pam*的
＊ 生態(tài)毒理學(xué)研究中，選一個(gè)參考點(diǎn)，可以直接求出其它處理（如農(nóng)藥）的受抑制程度inh.
各種熒光參數(shù)的成像是將0.0（黑色）至1.0（紫色）的數(shù)值轉(zhuǎn)換成顏色來(lái)顯示的。
應(yīng)用范圍
| ＊ 環(huán)境科學(xué) ＊ 水生生物學(xué) ＊ 海洋與湖沼學(xué) ＊ 生態(tài)毒理學(xué) ＊ 園藝學(xué) ＊ 農(nóng)業(yè)科學(xué) ＊ 林學(xué) | ＊ 環(huán)境科學(xué) ＊ 水生生物學(xué) ＊ 海洋與湖沼學(xué) ＊ 生態(tài)毒理學(xué) ＊ 園藝學(xué) ＊ 農(nóng)業(yè)科學(xué) ＊ 林學(xué) |
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m系列imaging-pam不同版本介紹
maxi-版 |
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測(cè)量盆栽植物 | 測(cè)量離體葉片 | 測(cè)量微藻樣品 | 新增鏡頭可調(diào)放大倍數(shù) |
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y(npq) | ps/50 | f | 96個(gè)微藻樣品成像 |
mini-版 |
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由于mini-探頭的便攜式設(shè)計(jì)，使其特別適合野外應(yīng)用。由于mini-探頭的成像面積僅為maxi-探頭的1/16，因而前者發(fā)出的zui大光強(qiáng)更大，但耗電卻小得多。mini-探頭可以安裝在光合儀gfs-3000的葉室3010-s上，同步測(cè)量全葉片氣體交換和熒光成像，并且其光源可由gfs-3000控制，達(dá)到真正的同步測(cè)量。
mini-探頭采用1/3”數(shù)碼相機(jī)ccd（640×480象素）和f1.2/f=12mm物鏡。其設(shè)計(jì)目的為測(cè)量固定距離下的熒光成像。
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與光合儀gfs-3000連用 | 長(zhǎng)時(shí)間測(cè)量可裝在三角架上 | fm | qn |
micro-版
微探頭，成像面積3.5×4.5 mm
調(diào)制熒光成像系統(tǒng)的micro-探頭是一個(gè)極便攜的探頭，采用整合式cosmicar-pentax cctv物鏡（f1.4/f=16mm），直接安裝在數(shù)碼ccd（1/3” chip, 640×480像素）上。
micro-探頭只配備一個(gè)luxeon led（藍(lán)光，450 nm）和一個(gè)特制雙色分光鏡，類似于落射熒光顯微鏡。
盡管成像面積只有3.5×4.5 mm，但45倍的放大率卻允許對(duì)葉片熒光成像的異質(zhì)性分析達(dá)到支脈（minor veins）級(jí)。同時(shí)還可提供一個(gè)特制版本用于測(cè)量gfp的成像。 
micro-探頭還可安裝在標(biāo)準(zhǔn)版imaging-pam（2001年設(shè)計(jì)）主機(jī)上。該探頭提供x-y軸可調(diào)的樣品臺(tái)。其設(shè)計(jì)目的為測(cè)量固定距離下的熒光成像。
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fo | gfp成像 | 微探頭直接安裝在ccd上 |
microscopy-版
顯微探頭，成像面積130×150 um
必須與特制落射熒光顯微鏡（hund或zeiss）結(jié)合使用，該顯微鏡可以提供激發(fā)光并檢測(cè)熒光
imag-max/k（數(shù)碼相機(jī)ccd）[1392×1040象素，4象素組合（binning）技術(shù)]可以提供高靈敏度。
探頭標(biāo)準(zhǔn)配置是一個(gè)*luxeon led（450-480 nm），用于提供測(cè)量光、光化光和飽和脈沖。目前已可提供rgb探頭（紅-綠-藍(lán)-白led光源），它采用了phyto-pam技術(shù)，可以顯微鏡下自動(dòng)對(duì)藍(lán)藻、綠藻、硅/甲藻、紅藻進(jìn)行分類并測(cè)量光合作用。
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分類，紅色為硅藻，綠色為絲狀綠藻 | 光合，fv/fm活性，可區(qū)分細(xì)胞不同部位的活性 |
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51. bonfig kb, schreiber u, gabler a, roitsch t, berger s: infection with virulent and avirulent p. syringae strains differentially affects photosynthesis and sink metabolism in arabidopsis leaves planta 2006;225:1-12.
52. dima e, manetas y, psaras gk: chlorophyll distribution pattern in inner stem tissues: evidence from epifluorescence microscopy and reflectance measurements in 20 woody species trees 2006;20:515-521.
53. escher bi, quayle p, muller r, schreiber u, mueller jf: passive sampling of herbicides combined with effect analysis in algae using a novel high-throughput phytotoxicity assay (maxi-imaging-pam). journal of environmental monitoring 2006;8:456-464.
54. heddad m, norén h, reiser v, dunaeva m, andersson b, adamska i: differential expression and localization of early light-induced proteins in arabidopsis thaliana. plant physiology 2006:in press.
55. hölzl g, witt s, kelly aa, zähringer u, warnecke d, dörmann p, heinz e: functional differences between galactolipids and glucolipids revealed in photosynthesis of higher plants. proc. natl. acad. sci. usa 2006;103:7512-7517.
56. ivanov ag, hendrickson l, krol m, selstam e, öquist g, hurry v, huner npa: digalactosyl-diacylglycerol deficiency impairs the capacity for photosynthetic intersystem electron transport and state transitions in arabidopsis thaliana due to photosystem i acceptor-side limitations. plant cell and physiology 2006;47:1146-1157.
57. kaiser h, grams tee: rapid hydr-opassive opening and subsequent active stomatal closure follow heat-induced electrical signals in mimosa pudica. journal of experimental botany 2006;57:2087-2092.
58. kuster a, altenburger r: development and validation of a new fluorescence-based bioassay for aquatic macrophyte species. chemosphere 2006;67:194-201.
59. 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.
60. nagel ka, schurr u, walter a: dynamics of root growth stimulation in nicotiana tabacum in increasing light intensity. plant cell and environment 2006;29:1936-1945.
61. petit a-n, vaillant n, boulay m, clément c, fontaine f: alteration of photosynthesis in grapevines affected by esca. phytopathology 2006;96:1060-1066.
62. pieruschka r, schurr u, jensen m, wolff wf, jahnke s: lateral diffusion of co2 from shaded to illuminated leaf parts affects photosynthesis inside homobaric leaves. new phytologist 2006;169:779-788.
63. swarbrick pj, schulze-lefert p, scholes jd: metabolic consequences of susceptibility and resistance (race-specific and broad-spectrum) in barley leaves challenged with powdery mildew. plant cell and environment 2006;29:1061-1076.
64. vopel k, hawes i: photosynthetic performance of benthic microbial mats in lake hoare, antarctica. limnology and oceanography 2006;51:1801-1812.
65. 蔡馬, 賀立紅, 梁紅: 2種銀杏葉片葉綠素?zé)晒馓匦缘谋容^. 安徽農(nóng)業(yè)科學(xué) 2006;34:3322-3324.
66. 鞏擎柱, 呂金印, 徐柄成, 李鳳民, 張海波: 水分脅迫和種植方式對(duì)小麥葉綠素?zé)晒鈪?shù)及水分利用效率的影響. 西北農(nóng)林科技大學(xué)學(xué)報(bào) 2006;34:83-87.
67. 郭學(xué)民, 王貴禧, 高榮孚, 梁麗松: 果實(shí)表皮毛的掃描電鏡觀察及其對(duì)果實(shí)表面熒光特性的影響. 內(nèi)蒙古農(nóng)業(yè)大學(xué)學(xué)報(bào) 2006;27:43-47.
68. 賀立紅, 賀立靜, 顧群, 梁紅: 銀杏同一葉片不同部位葉綠素?zé)晒馓匦缘难芯? 北方園藝 2006:27-29.
69. 賀立紅, 賀立靜, 梁紅: 銀杏不同品種葉綠素?zé)晒鈪?shù)的比較. 華南農(nóng)業(yè)大學(xué)學(xué)報(bào) 2006;27:43-46.
70. aldea m, hamilton jg, resti jp, zangerl ar, berenbaum mr, delucia eh: indirect effects of insect herbivory on leaf gas exchange in soybean. plant cell and environment 2005;28:402-411.
71. baek m-h, kim j-h, chung by, kim j-s, lee is: alleviation of salt stress by low dose g-irradiation in rice. biologia plantarum 2005;49:273-276.
72. borisjuk l, nguyen th, neuberger t, rutten t, tschiersch h, claus b, feussner i, webb ag, jakob p, weber h, wobus u, rolletschek h: gradients of lipid storage, photosynthesis and plastid differentiation in developing soybean seeds. new phytologist 2005;163:761-776.
73. gog l, berenbaum mr, delucia eh, zangerl ar: autotoxic effects of essential oils on photosynthesis in parsley, parsnip, and rough lemon. chemoecology 2005;15:115-119.
74. kühl m, chen m, ralph pj, schreiber u, larkum awd: a niche for cyanobacteria containing chlorophyll d. nature 2005;433:820.
75. lautner s, grams tee, matyssek r, fromm j: characteristics of electrical signals in poplar and responses in photosynthesis. plant physiology 2005;138:2200-2209.
76. manetas y, pfanz h: spatial heterogeneity of light penetration through periderm and lenticels and concomitant patchy acclimation of corticular photosynthesis trees 2005;19:409-414.
77. marjanovic z, uwe n, hampp r: mycorrhiza formation enhances adaptive response of hybrid poplar to drought annals of the new york academy of sciences 2005;1048:496-499.
78. podola b, melkonian m: se-lective real-time herbicide monitoring by an array chip biosensor employing diverse microalgae. journal of applied phycology 2005;17:261-271.
79. ralph pj, macinnis-ng cmo, frankart c: fluorescence imaging application: effect of leaf age on seagrass photokinetics. aquatic botany 2005;81:69-84.
80. ralph pj, schreiber u, gademann r, kühl m, larkum awd: coral photobiology studied with a new imaging pulse amplitude modulated fluorometer. journal of phycology 2005;41:335-342.
81. berger s, papadopoulos m, schreiber u, kaiser w, roitsch t: complex regulation of gene expression, photosynthesis and sugar levels by pathogen infection in tomato. physiologia plantarum 2004;122:419-428.
82. hill r, larkum awd, frankart c, kühl m, ralph pj: loss of functional photosystem ii reaction centres in zooxanthellae of corals exposed to bleaching conditions: using fluorescence rise kinetics. photosynthesis research 2004;82:59-72.
83. hill r, schreiber u, gademann r, larkum awd, kühl m, ralph pj: spatial heterogeneity of photosynthesis and the effect of temperature-induced bleaching conditions in three species of corals. marine biology 2004;144:633-640.
84. podola b, nowack ecm, melkonian m: the use of multiple-strain algal sensor chips for the detection and identification of volatile organic compounds. biosensors and bioelectronics 2004;19:1253-1260.
85. koziolek c, grams tee, schreiber u, matyssek r, fromm j: transient knockout of photosynthesis mediated by electrical signals. new phytologist 2003;161:715-722.
86. schreiber u, walz h, kolbowski j: propagation of spatial variations of chlorophyll fluorescence parameters in dandelion leaves induced by spot laser heating. pam news 2003.
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