Photosynthesis Research

, Volume 113, Issue 1–3, pp 63–74 | Cite as

Quantifying and monitoring functional photosystem II and the stoichiometry of the two photosystems in leaf segments: approaches and approximations

  • Wah Soon ChowEmail author
  • Da-Yong Fan
  • Riichi Oguchi
  • Husen Jia
  • Pasquale Losciale
  • Youn-Il Park
  • Jie He
  • Gunnar Öquist
  • Yun-Gang Shen
  • Jan M. Anderson


Given its unique function in light-induced water oxidation and its susceptibility to photoinactivation during photosynthesis, photosystem II (PS II) is often the focus of studies of photosynthetic structure and function, particularly in environmental stress conditions. Here we review four approaches for quantifying or monitoring PS II functionality or the stoichiometry of the two photosystems in leaf segments, scrutinizing the approximations in each approach. (1) Chlorophyll fluorescence parameters are convenient to derive, but the information-rich signal suffers from the localized nature of its detection in leaf tissue. (2) The gross O2 yield per single-turnover flash in CO2-enriched air is a more direct measurement of the functional content, assuming that each functional PS II evolves one O2 molecule after four flashes. However, the gross O2 yield per single-turnover flash (multiplied by four) could over-estimate the content of functional PS II if mitochondrial respiration is lower in flash illumination than in darkness. (3) The cumulative delivery of electrons from PS II to P700+ (oxidized primary donor in PS I) after a flash is added to steady background far-red light is a whole-tissue measurement, such that a single linear correlation with functional PS II applies to leaves of all plant species investigated so far. However, the magnitude obtained in a simple analysis (with the signal normalized to the maximum photo-oxidizable P700 signal), which should equal the ratio of PS II to PS I centers, was too small to match the independently-obtained photosystem stoichiometry. Further, an under-estimation of functional PS II content could occur if some electrons were intercepted before reaching PS I. (4) The electrochromic signal from leaf segments appears to reliably quantify the photosystem stoichiometry, either by progressively photoinactivating PS II or suppressing PS I via photo-oxidation of a known fraction of the P700 with steady far-red light. Together, these approaches have the potential for quantitatively probing PS II in vivo in leaf segments, with prospects for application of the latter two approaches in the field.


Chlorophyll fluorescence Electrochromic signal Oxygen evolution P700 Photosystem II PS II/PS I stoichiometry 









Electrochromic signal

Fo, Fm

Chl fluorescence corresponding to open and closed PS II traps in the dark-adapted state, respectively


(F m − F o)/F m


Special Chl pair in the PS I reaction center




Photosystem I, II, respectively

\( \phi \)PS II

Effective photochemical yield of PS II


Primary, secondary quinone acceptor in PS II, respectively


Indicator of the oxidation state of QA


Integrated flash-induced transient flow of electrons from PS II to P700+



The support of this work by an Australian Research Council Grant (DP1093827) awarded to W. S. Chow, a Knowledge Innovation Programme of the Chinese Academy of Sciences grant (KSCX2-EW-J-1) to D.-Y. Fan and JSPS Postdoctoral Fellowships for Research Abroad (21-674) to R. Oguchi is gratefully acknowledged. W.S.Chow is grateful to Barry Osmond for providing scientific refuge and saving a career in photosynthesis research.


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Copyright information

© Springer Science+Business Media B.V. 2012

Authors and Affiliations

  • Wah Soon Chow
    • 1
    Email author
  • Da-Yong Fan
    • 1
    • 2
  • Riichi Oguchi
    • 1
  • Husen Jia
    • 1
  • Pasquale Losciale
    • 3
  • Youn-Il Park
    • 4
  • Jie He
    • 6
  • Gunnar Öquist
    • 5
  • Yun-Gang Shen
    • 7
  • Jan M. Anderson
    • 1
  1. 1.Research School of Biology, College of Medicine, Biology and EnvironmentThe Australian National UniversityCanberraAustralia
  2. 2.State Key Laboratory of Vegetation and Environmental Change, Institute of BotanyThe Chinese Academy of SciencesBeijingChina
  3. 3.Dipartimento Colture ArboreeUniversity of BolognaBolognaItaly
  4. 4.Department of Biological Sciences, College of Biological Science and BiotechnologyChungnam National UniversityDaejeonKorea
  5. 5.Department of Plant Physiology, Umeå Plant Science CentreUmeå UniversityUmeåSweden
  6. 6.Natural Sciences and Science Education Academic Group, National Institute of EducationNanyang Technological UniversitySingaporeSingapore
  7. 7.Institute of Plant Physiology and EcologyThe Chinese Academy of SciencesShanghaiChina

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