主要功能
采用板載(zai)芯片(pian) LED 陣列(lie)技術，用 6 種不同波(bo)段(duan)的(de)激發(fa)光作為測量光、光化光、飽(bao)和(he)脈沖、單周(zhou)轉飽(bao)和(he)閃光與(yu)多周(zhou)轉飽(bao)和(he)閃光
具備(bei)比 PAM-2500 高 200 倍(bei)的靈敏(min)度
可用(yong)于很稀的懸浮液（藻液、葉綠體懸浮液）測量(liang)
專用葉夾可(ke)用于高(gao)等植(zhi)物/大型海(hai)藻等葉片狀(zhuang)樣品的測量
標準的(de)(de)PAM測(ce)量功能、復雜的(de)(de)多相熒(ying)光上升動(dong)力學擬合分(fen)析、馳豫動(dong)力學分(fen)析
特別(bie)適合狀態(tai)轉換(huan)研(yan)究(jiu)、“非活性(xing)PSII”（“Inactive PS II”）研(yan)究(jiu)
超快時間分辨率達到 10 ms，由此利用獨特的 O-I1 相（O-J相）擬合分析用于分析PSII反映中心異質性分析，得出 PS II 光合單位的連接性參數（p和J），速率常數（Tau）和兩種不同類型 PS II（Type 1 和Type 2）的光學截面積（Sigma(II)λ）等參數
新增 PSII 有效光強 PAR(II)、經過 PSII 的絕對電子傳遞速率 ETR(II)λ 等全新的光合參數(shu)。
配套的操作軟件，用于復雜的擬合分(fen)析
測量(liang)參(can)數
Fo, Fm, F, Fm', Fv/Fm, Y(II), qP, qN, NPQ, Y(NO), Y(NPQ), ETR, ETR(II)λ, p, J, Tau, Sigma(II)λ, PAR、PAR(II) 等
應用領域
主要用于各種藻(zao)(zao)類的(de)(de)(de)(de)深入光合(he)(he)作用機理研究，用適合(he)(he)的(de)(de)(de)(de)波長、全(quan)新的(de)(de)(de)(de)測(ce)量、全(quan)新的(de)(de)(de)(de)參數進行藍藻(zao)(zao)、綠藻(zao)(zao)、硅藻(zao)(zao)、甲(jia)藻(zao)(zao)、紅藻(zao)(zao)、隱藻(zao)(zao)等的(de)(de)(de)(de)深入研究。如選配高(gao)等植物附件，也可實現對高(gao)等植物葉(xie)片的(de)(de)(de)(de)測(ce)量。
主要技術參數
測量光(guang)：提(ti)供 400、440、480、540、590 和(he) 625 nm 的(de)脈沖調制(zhi)測量光(guang)，20 個強度(du)選(xuan)擇，14 個頻(pin)率選(xuan)擇。
光化光：提供 440、480、540、590、625 nm 和 420-640 nm（白光）連續光化光照，最大光強 4000 μmol m-2 s-1；單周轉飽和閃光的最大強度 200 000 μmol m-2 s-1，持續時間 5-50 μs可調；多周轉飽和閃光強度 10 000 μmol m-2 s-1，1-800 ms可調(diao)。
遠紅光(guang)：725 nm。
信號檢測：PIN-光電二極管，帶特(te)制鎖相放(fang)大(da)器（專(zhuan)利設計），最大(da)時間分辨率 10 μs。
Multi-Color-PAM的功能介紹
光(guang)(guang)(guang)系統 II 的(de)(de)相對電子傳遞(di)速率 rETR 是(shi)(shi)很常(chang)用(yong)(yong)的(de)(de)一個(ge)參數(shu)。rETR = PAR × Y(II) × ETR-factor，其中 ETR-factor 是(shi)(shi)指(zhi)光(guang)(guang)(guang)系統II吸收(shou)的(de)(de)光(guang)(guang)(guang)能占總入射 PAR 的(de)(de)比(bi)例(li)。在大(da)多數(shu)已發表的(de)(de)文獻中，均沒有試(shi)圖去測定 ETR-factor，只是(shi)(shi)簡單地假定跟 “模式葉片(pian)” 相同，即有 50% 的(de)(de) PAR 分(fen)配到光(guang)(guang)(guang)系統 II，84% 的(de)(de) PAR 被(bei)光(guang)(guang)(guang)合色素(su)吸收(shou)。因(yin)此在已有的(de)(de)文獻中，rETR一般(ban)是(shi)(shi)用(yong)(yong)公式 rETR = PAR × Y(II) × 0.84 × 0.5 來計算的(de)(de)。
近期，利用多激發波長調制葉綠素熒光儀 MULTI-COLOR-PAM 可以實現光系統II的絕對電子傳遞速率 ETR(II)λ 的測量。首先需要利用 MULTI-COLOR-PAM 測定某個波長下的光系統II功能性光學截面積 Sigma(II)λ（單位nm2）（其中λ為波長），然后求出光系統II的量子吸收速率 PAR(II) = Sigma(II)λ × L × PAR = 0.6022 × Sigma(II)λ× PAR。其中 L 為阿伏伽德羅常數，系數 0.6022 是將 1 μmol quanta m-2 （即 6.022 × 1017 quanta m-2）轉換為 0.6022 quanta nm-2，PAR(II) 的單位為 quanta/(PSII × s)。接下來就可以計算 ETR(II)λ = PAR(II) × Y(II)/Y(II)max，其中 Y(II)max 是經(jing)過暗適(shi)應(ying)達到穩態后的(de)光系統II的(de)量子產量，也就是 Fv/Fm×ETR(II) 的(de)單位為 electrons/(PSII × s)。
傳(chuan)統的(de)(de)(de)調(diao)制(zhi)葉綠素熒光(guang)(guang)(guang)(guang)(guang)儀一(yi)般只能提供(gong)一(yi)種或(huo)兩種顏(yan)色的(de)(de)(de)光(guang)(guang)(guang)(guang)(guang)源，如發(fa)(fa)(fa)(fa)出(chu)(chu)白光(guang)(guang)(guang)(guang)(guang)的(de)(de)(de)鹵素燈、發(fa)(fa)(fa)(fa)出(chu)(chu)藍(lan)光(guang)(guang)(guang)(guang)(guang)的(de)(de)(de)藍(lan)色 LED 或(huo)發(fa)(fa)(fa)(fa)出(chu)(chu)紅光(guang)(guang)(guang)(guang)(guang)的(de)(de)(de)紅色 LED 等。用(yong)不(bu)同(tong)顏(yan)色的(de)(de)(de)光(guang)(guang)(guang)(guang)(guang)測量的(de)(de)(de)結果可能會有(you)不(bu)同(tong)，如圖 1A 所示，用(yong)藍(lan)光(guang)(guang)(guang)(guang)(guang)（440 nm）和紅光(guang)(guang)(guang)(guang)(guang)（625 nm）測量綠藻小球藻的(de)(de)(de)快速光(guang)(guang)(guang)(guang)(guang)曲線有(you)非常顯(xian)著的(de)(de)(de)差別，藍(lan)光(guang)(guang)(guang)(guang)(guang)照(zhao)射下的(de)(de)(de) rETRmax 顯(xian)著小于紅光(guang)(guang)(guang)(guang)(guang)照(zhao)射下，且(qie)在較強的(de)(de)(de)光(guang)(guang)(guang)(guang)(guang)曲線 rETR 有(you)輕微(wei)下降趨(qu)勢(shi)，這(zhe)說明藍(lan)光(guang)(guang)(guang)(guang)(guang)的(de)(de)(de)更容易(yi)引發(fa)(fa)(fa)(fa)光(guang)(guang)(guang)(guang)(guang)抑制(zhi) (Schreiber, Klughammer et al. 2011, Schreiber, Klughammer et al. 2012)。由(you)此(ci)可以推測，過去文(wen)獻報道的(de)(de)(de)很過實驗(yan)結果，可能會存在由(you)于采(cai)用(yong)的(de)(de)(de)激發(fa)(fa)(fa)(fa)光(guang)(guang)(guang)(guang)(guang)源不(bu)同(tong)而引起(qi)的(de)(de)(de)錯誤(wu)理解。
如上文所述，利用 MULTI-COLOR-PAM，已經可以測量真實電子傳遞速率 ETR(II)λ。如果用 ETR(II)λ 來繪制快速光曲線會出現什么結果呢？圖 1B 是將圖 1A 的結果轉換成絕對電子傳遞速率后得到的結果，可以看出無論是照射藍光還是照射紅光，其絕對電子傳遞速率是一致的。由此證明圖 1A 中結果的差異是由于不同波長下藻細胞的光系統 II 功能性光學截面積 Sigma(II)λ 的大小不同引起的 (Schreiber, Klughammer et al. 2011, Schreiber, Klughammer et al. 2012)。這種利用絕對電子傳遞速率 ETR(II)λ 繪制的快(kuai)速光曲線在未來的科研中可能會發揮越(yue)(yue)來越(yue)(yue)重(zhong)要的作用。
 |  |
| 圖1 利用相對電子傳遞速率（A）和絕對電子傳遞速率（B）分別繪制的快速光曲線（引自Schreiber et al., 2012） | |
| 利用 MULTI-COLOR-PAM 分別以藍光（440 nm）和紅光（625 nm）作為光化光源，測量小球藻（Chlorella sp.）的快速光曲線。 | |
| 圖A中，rETR 的計算采用 0.42 作為 ETR factor。 | |
| 圖B中，藍光和紅光激發下獲得的光系統II功能性光學截面積 Sigma(II)λ 分別為 4.547 和 1.669 nm2，計算絕對電子傳遞速率 ETR(II)440 和 ETR(II)625 的 Fv/Fm 分別為 0.68 和 0.66。 | |
選購(gou)指南
一、懸浮樣品測量基本款
系統組(zu)成(cheng)：通用型主機，標準版檢測單(dan)元，懸(xuan)浮(fu)液的光學單(dan)元，數據(ju)線，工作臺，軟件等(deng)
 |
| 懸浮樣品測量基本款 |
二 、高等植物葉片測量基本款
系(xi)統(tong)組成(cheng)：通(tong)用型(xing)主機，標準版(ban)檢測(ce)單元(yuan)，特制葉片夾，數據線，工(gong)作臺(tai)，軟件等
 |
| 高等植物葉片測量特制葉夾 |
三、其他可選附件
1，ED-101US/T： 控(kong)溫(wen)裝置，安裝在(zai) ED-101US/MD 上(shang)，為懸浮液控(kong)溫(wen)；可外接循環水浴(yu)來控(kong)溫(wen),
2，US-SQS/WB： 球狀微(wei)型(xing)光量(liang)(liang)子探頭，可(ke)插(cha)入(ru)樣(yang)品杯中測(ce)量(liang)(liang) PAR；由主機 DUAL-C 控制。
3，PHYTO-MS：磁力攪(jiao)拌器，連接(jie)到光學(xue)單元 ED-101US/MD 的底部(bu)對懸浮(fu)液進行(xing)攪(jiao)拌。
產地：德國WALZ
參考文(wen)獻
數據來源：光合作用文(wen)獻 Endnote 數據庫，更新(xin)至 2021 年 1 月(yue)，文(wen)獻數量超過 10000 篇(pian)
原始數據(ju)來(lai)源：Google Scholar
1.Grund, M., et al. (2022). "Heterologous Lactate Synthesis in Synechocystis sp. Strain PCC 6803 Causes a Growth Condition-Dependent Carbon Sink Effect." Applied and Environmental Microbiology 0(0): e00063-00022.
2.Xiao, X., et al. (2022). "Effects of three graphene-based materials on the growth and photosynthesis of Brassica napus L." Ecotoxicology and Environmental Safety 234: 113383.
3.Xie, S., et al. (2022). "Enhanced lipid productivity coupled with carbon and nitrogen removal of the diatom Skeletonema costatum cultured in the high CO2 level." Algal Research 61: 102589.
4.Bernát, G., et al. (2021). "Photomorphogenesis in the Picocyanobacterium Cyanobium gracile Includes Increased Phycobilisome Abundance Under Blue Light, Phycobilisome Decoupling Under Near Far-Red Light, and Wavelength-Specific Photoprotective Strategies." Frontiers in Plant Science 12(352).
5.Ma, Z., et al. (2021). "Inhibitory effects of Prorocentrum donghaiense allelochemicals on Sargassum fusiformis zygotes probed by JIP-test based on fast chlorophyll fluorescence kinetics." Marine environmental research 170: 105453.
6.MATTILA, H., et al. (2021). "Differences in susceptibility to photoinhibition do not determine growth rate under moderate light in batch or turbidostat–a study with five green algae." Photosynthetica.
7.Mehmood, S. S., et al. (2021). "Integrated analysis of transcriptomics and proteomics provides insights into the molecular regulation of cold response in Brassica napus." Environmental and Experimental Botany 187: 104480.
8.Michel-Rodriguez, M., et al. (2021). "Underwater light climate and wavelength dependence of microalgae photosynthetic parameters in a temperate sea." PeerJ 9: e12101.
9.Qu, L., et al. (2021). "Elevated pCO2 enhances under light but reduces in darkness the growth rate of a diatom, with implications for the fate of phytoplankton below the photic zone." n/a(n/a).
10.Schansker, G. (2021). "Kinetic characterization of the interaction of NO with the S2 and S3 states of the oxygen-evolving complex of Photosystem II." bioRxiv: 2021.2001.2010.426130.
11.Schreiber, U. and C. Klughammer (2021). "Evidence for variable chlorophyll fluorescence of photosystem I in vivo." Photosynthesis Research.
12.Suka?ová, K., et al. (2021). "Perspective Design of Algae Photobioreactor for Greenhouses—A Comparative Study." energies 14(5): 1338.
13.Terentyev, V. V. (2021). "Loss of carbonic anhydrase in the thylakoid lumen causes unusual moderate-light-induced rearrangement of the chloroplast in Chlamydomonas reinhardtii as a way of photosystem II photoprotection." Plant Physiology and Biochemistry 168: 501-506.
14.Wang, C., et al. (2021). "Harmful algal bloom-forming dinoflagellate Prorocentrum donghaiense inhibits the growth and photosynthesis of seaweed Sargassum fusiformis embryos." Journal of Oceanology
15.Limnology: 1-15.
16.Zavafer, A., et al. (2021). "Normalized chlorophyll fluorescence imaging: A method to determine irradiance and photosynthetically active radiation in phytoplankton cultures." Algal Research 56: 102309.
17.Zav?el, T., et al. (2021). "Monitoring fitness and productivity in cyanobacteria batch cultures." Algal Research 56: 102328.
18.Andrzejczak, O. A., et al. (2020). "The Hypoxic Proteome and Metabolome of Barley (Hordeum vulgare L.) with and without Phytoglobin Priming. ." Int. J. Mol. Sci(21): 1546.
19.Chen, Y., et al. (2020). "Astaxanthin biosynthesis in transgenic Dunaliella salina (Chlorophyceae) enhanced tolerance to high irradiation stress." South African Journal of Botany 133: 132-138.
20.Pavaux, A.-S. (2020). "Chemical Ecology of the toxic dinoflagellate Ostreopsis cf. ovata in N.W. Mediterranean Sea." 學(xue)位論文.
21.Smolova, T., et al. (2020). "Cortical photosynthesis as a physiological marker for grape breeding: methods and approaches." BIO Web of Conferences 25: 02018.
22.Terentyev, V. V., et al. (2020). "The Main Structural and Functional Characteristics of Photosystem-II-Enriched Membranes Isolated from Wild Type and cia3 Mutant Chlamydomonas reinhardtii." Life(10): 63.
23.Zhang, X., et al. (2020). "Photosynthetic Properties of Miscanthus condensatus at Volcanically Devastated Sites on Miyake-jima Island." Plants(9): 1212.
24.Grund, M., et al. (2019). "Electron balancing under different sink conditions reveals positive effects on photon efficiency and metabolic activity of Synechocystis sp. PCC 6803." Biotechnology for Biofuels 12(1): 43.
25.R?kke, G. B., et al. (2019). "Unique photosynthetic electron transport tuning and excitation distribution in heterokont algae." PLoS ONE 14(1): e0209920.
26.Schreiber, U., et al. (2019). "Rapidly reversible chlorophyll fluorescence quenching induced by pulses of supersaturating light in vivo." Photosynthesis Research: 1-16.
27.Wang, W. and Y. Sheng (2019). "Pseudomonas sp. strain WJ04 enhances current generation of Synechocystis sp. PCC6803 in photomicrobial fuel cells." Algal Research 40: 101490.
28.Yanykin, D., et al. (2019). "Hydroxyectoine protects Mn-depleted photosystem II against photoinhibition acting as a source of electrons." Photosynthesis Research: 1-15.
29.Chartrand, K. M., et al. (2018). "Living at the margins–The response of deep-water seagrasses to light and temperature renders them susceptible to acute impacts." Marine environmental research.
30.Goessling, J. W., et al. (2018). "Modulation of the light field related to valve optical properties of raphid diatoms: implications for niche differentiation in the microphytobenthos." MARINE ECOLOGY PROGRESS SERIES 588: 29-42.
31.Khorobrykh, A., et al. (2018). "Photooxidation and photoreduction of exogenous cytochrome c by photosystem II preparations after various modifications of the water-oxidizing complex." Photosynthetica: 1-10.
32.Li, F., et al. (2018). "Diatom performance in a future ocean: interactions between nitrogen limitation, temperature, and CO2-induced seawater acidification." ICES Journal of Marine Science.
33.Lin, L., et al. (2018). "Electrochemical oxidation of Microcystis aeruginosa using a Ti/RuO2 anode: contributions of electrochemically generated chlorines and hydrogen peroxide." Environmental Science Pollution Research.
34.Miao, H., et al. (2018). "Calcification Moderates the Increased Susceptibility to UV Radiation of the Coccolithophorid Gephryocapsa oceanica Grown under Elevated CO 2 Concentration: Evidence Based on Calcified and Non‐Calcified Cells." Photochemistry and Photobiology.
35.Morelle, J. and P. Claquin (2018). "Electron requirements for carbon incorporation along a diel light cycle in three marine diatom species." Photosynthesis Research 137(2): 201-214.
36.Morelle, J., et al. (2018). "Annual Phytoplankton Primary Production Estimation in a Temperate Estuary by Coupling PAM and Carbon Incorporation Methods." Estuaries and Coasts: 1-19.
37.Nikkanen, L., et al. (2018). "Regulation of chloroplast NADH dehydrogenase-like complex by NADPH-dependent thioredoxin system." bioRxiv: 261560.
38.Preuss, M. and G. C. Zuccarello (2018). "Comparative studies of photosynthetic capacity in three pigmented red algal parasites: Chlorophyll a concentrations and PAM fluorometry measurements." Phycological Research 0(0).
39.Sung, M.-G., et al. (2018). "Wavelength shift strategy to enhance lipid productivity of Nannochloropsis gaditana." Biotechnology for Biofuels 11(1): 70.
40.Ternon, E., et al. (2018). "Allelopathic interactions between the benthic toxic dinoflagellate Ostreopsis cf. ovata and a co-occurring diatom." Harmful Algae 75: 35-44.
41.Zav?el, T., et al. (2018). "Effect of carbon limitation on photosynthetic electron transport in Nannochloropsis oculata." Journal of Photochemistry and Photobiology B: Biology 181: 31-43
42.Béchet, Q., et al. (2017). "Modeling the impact of high temperatures on microalgal viability and photosynthetic activity." Biotechnology for Biofuels 10(1): 136.
43.Havurinne, V. and E. Tyystj?rvi (2017). "Action spectrum of photoinhibition in the diatom Phaeodactylum tricornutum." Plant and Cell Physiology: pcx156.
44.Kalaji, M. H., et al. (2017). Chlorophyll Fluorescence: Understanding Crop Performance—Basics and Applications, CRC Press.
45.Lamb, J. J. and M. F. Hohmann-Marriott (2017). "Manganese acquisition is facilitated by PilA in the cyanobacterium Synechocystis sp. PCC 6803." PLoS ONE 12(10): e0184685.
46.R?kke, G., et al. (2017). "The plastoquinone pool of Nannochloropsis oceanica is not completely reduced during bright light pulses." PLoS ONE 12(4): e0175184.
47.Savchenko, T., et al. (2017). "The hydroperoxide lyase branch of the oxylipin pathway protects against photoinhibition of photosynthesis." Planta: 1-14.
48.Shin, W.-S., et al. (2017). "Complementation of a mutation in CpSRP43 causing partial truncation of light-harvesting chlorophyll antenna in Chlorella vulgaris." Scientific Reports 7(1): 17929.
49.Laviale, M., et al. (2016). "The importance of being fast: comparative kinetics of vertical migration and non-photochemical quenching of benthic diatoms under light stress." Marine Biology 163(1): 1-12.
50.Murphy, T. E., et al. (2016). "A radiative transfer modeling approach for accurate interpretation of PAM fluorometry experiments in suspended algal cultures." Biotechnology Progress: n/a-n/a.
51.Shin, W.-S., et al. (2016). "Truncated light-harvesting chlorophyll antenna size in Chlorella vulgaris improves biomass productivity." Journal of Applied Phycology: 1-10.
52.Yanykin, D. V., et al. (2016). "Trehalose protects Mn-depleted photosystem 2 preparations against the donor-side photoinhibition." Journal of Photochemistry and Photobiology B: Biology 164: 236-243.
53.He, J., et al. (2015). "Photoinactivation of Photosystem II in wild-type and chlorophyll b-less barley leaves: which mechanism dominates depends on experimental circumstances." Photosynthesis Research: 1-9.
54.Lin, L., et al. (2015). "Effects of electrolysis by low-amperage electric current on the chlorophyll fluorescence characteristics of Microcystis aeruginosa." Environmental Science and Pollution Research: 1-8.
55.Polishchuk, A., et al. (2015). "Cultivation of Nannochloropsis for eicosapentaenoic acid production in wastewaters of pulp and paper industry." Bioresource Technology 193: 469-476.
56.Tamburic, B., et al. (2015). "Gas Transfer Controls Carbon Limitation During Biomass Production by Marine Microalgae." ChemSusChem.
57.Yanykin, D., et al. (2015). "Trehalose stimulation of photoinduced electron transfer and oxygen photoconsumption in Mn-depleted photosystem 2 membrane fragments." Journal of Photochemistry and Photobiology B: Biology 152: 279-285.
58.Klughammer, C. and U. Schreiber (2014). Apparent PS II absorption cross-section and estimation of mean PAR in optically thin and dense suspensions of Chlorella. Photosynth Res.
59.Szabó, M., K. Parker, et al. (2014). Photosynthetic acclimation of Nannochloropsis oculata investigated by multi-wavelength chlorophyll fluorescence analysis. Bioresource Technology 167: 521-529.
60.Szabó, M., D. Wangpraseurt, et al. (2014). Effective light absorption and absolute electron transport rates in the coral Pocillopora damicornis. Plant Physiology and Biochemistry.
61.Tamburic, B., M. Szabó, et al. (2014). Action spectra of oxygen production and chlorophyll a fluorescence in the green microalga Nannochloropsis oculata. Bioresource Technology 169: 320-327.
62.Hakkila, K., T. Antal, et al. (2014). "Oxidative stress and photoinhibition can be separated in the cyanobacterium Synechocystis sp. PCC 6803." Biochim Biophys Acta 1837(2): 217-225.
63.Reigosa, M., D. Wangpraseurt, et al. (2014). "Spectral Effects on Symbiodinium Photobiology Studied with a Programmable Light Engine." PLoS ONE 9(11): e112809.
64.Schreiber, U. and C. Klughammer (2013). Wavelength-dependent photodamage to Chlorella investigated with a new type of multi-color PAM chlorophyll fluorometer. Photosynthesis Research 114(3): 165-177.
65. Bernát G, Schreiber U, Sendtko E, Stadnichuk IN, Rexroth S, R?gner M, Koenig F (2012) Unique Properties vs. Common Themes: The Atypical Cyanobacterium Gloeobacter violaceus PCC 7421 is Capable of State Transitions and Blue-light Induced Fluorescence Quenching. Plant & Cell Physiology: in press.
66.Schreiber U, Klughammer C, Kolbowski J (2012) Assessment of wavelength-dependent parameters of photosynthetic electron transport with a new type of Multi-Color-PAM chlorophyll fluorometer. Photosynthesis Research: in press.
67.Schreiber U, Klughammer C, Kolbowski J (2011) High-end chlorophyll fluorescence analysis with the MULTI-COLOR-PAM. I. Various light qualities and their applications. PAM Application Notes, 4: 1-19.
