Abstract
This paper deals with how Govindjee taught the Z-Scheme of electron transport in oxygenic photosynthesis at Ravenshaw University, Cuttack, Odisha, India, in 2014, in a unique and highly effective fashion—using students to act as molecules, representing the entire electron transport chain from water to nicotinamide adenine dinucleotide phosphate (NADP+). It culminated in a show by B.Sc. students in the garden of the Department of Botany, Ravenshaw University. The first author (PKM) personally acted as Ferredoxin NADP Reductase (FNR) catalyzing the reduction of NADP+ to NADPH, taking electrons from reduced ferredoxin at the end of Photosystem I. On the other hand, the Q-cycle was played by M.Sc. students, who acted as molecules running this ingenious cycle that produces extra protons. An interesting event was when a student, acting as a herbicide, who was dressed like a devil (fierce looking, in black clothes with a sword; “Yamaraj: The God of Death”, as he called himself), stopped all reactions by throwing out QB, the second plastoquinone molecule of Photosystem II, and that too aggressively, taking its position instead. The second author was the major organizer of the Z-scheme show. We provide here a basic background on the process, a bit on Govindjee’s teaching, and some selected pictures from the drama played in March, 2014 at Ravenshaw University. Here, we also recognize the teacher Govindjee for his ingenious and fun-filled teaching methods that touched the hearts and the souls of the students as well as the teachers of Ravenshaw University. He was rated as one of the most-admired teachers of plant biology at our university.
References
Benson AA (2002) Following the path of carbon in photosynthesis: a personal story. Photosynth Res 73:29–49
Blankenship RE (2014) Molecular mechansisms of photosynthesis. Blackwell-John Wiley, UK
Blankenship RE, Madigan T, Bauer CE (eds) (1995) Anoxygenic photosynthetic bacteria. Advances in Photosynthesis and Respiration, vol 2. Springer, Dordrecht
Crofts AR, Holland JT, Kolling DRJ, Victoria D, Dikanov SA, Gilbreth R, Lhee S, Kuras R, Kraus MG (2008) The Q-cycle reviewed: how well does a monomeric mechanism of the bc1 complex account for the function of a dimeric complex. Biochim Biophys Acta 1777:1001–1019
Demmig-Adams B, Garab G, Adams WW III, Govindjee (eds) (2014) Non-photochemical quenching and energy dissipation in plants, algae and cyanobacteria. Advances in Photosynthesis and Respiration,vol 40. Springer, Dordrecht
Duysens LNM (1989) The discovery of the two photosynthetic systems: a personal account. Photosynth Res 21:61–79
Duysens LNM, Amesz J, Kamp BM (1961) Two photochemical systems in photosynthesis. Nature 190:510–511
Eaton-Rye JJ, Govindjee (1988a) Electron transfer through the quinone acceptor complex of photosystem II in bicarbonate-depleted spinach thylakoid membranes as a function of actinic flash number and frequency. Biochim Biophys Acta 935: 237–247
Eaton-Rye JJ, Govindjee (1988b) Electron transfer through the quinone acceptor complex of photosystem II after one or two actinic flashes in bicarbonate-depleted spinach thylakoid membranes. Biochim Biophys Acta 935:248–257
Emerson R, Lewis CM (1943) The dependence of the quantum yield of Chlorella photosynthesis on wavelength of light. Am J Bot 30:165–178
Emerson R, Chalmers R, Cederstrand CN (1957) Some factors influencing the long-wave limit of photosynthesis. Proc Natl Acad Sci USA 43:133–143
Govindjee (1999a) On the requirement of minimum number of four versus eight quanta of light for the evolution of one molecule of oxygen in photosynthesis: a historical note. Photosynth Res 59:249–254
Govindjee (1999b) Carotenoids in photosynthesis: an historical perspective. In: Frank HA, Young AJ, Britton G, Cogdell RJ (eds) The photochemistry of carotenoids. Kluwer Academic Publishers, Dordrecht, pp 1–19
Govindjee (2000) Milestones in photosynthesis research. In: Younis M, Pathre U, Mohanty P (eds) Probing photosynthesis: mechanism, regulation and adaptation. Taylor & Francis, London, pp 9–39
Govindjee (2008) Teaching photosynthesis: some thoughts. In: Allen JF, Gantt E, Golbeck JH, Osmond B (eds) Photosynthesis: energy from the sun. Springer, Dordrecht, pp 1619–1624
Govindjee, Björn LO (2012) Dissecting oxygenic photosynthesis: the evolution of the “Z”-scheme for thylakoid reactions. In: Itoh S, Mohanty P, Guruprasad KN (eds) Photosynthesis: Overviews on recent progress and future perspective. IK Publishers, New Delhi, pp 1–27
Govindjee, Govindjee R(ajni) (1974) Primary events in photosynthesis. Sci Amer 231:68–82
Govindjee, Rabinowitch E (1960) Two forms of chlorophyll a in vivo with distinct photochemical functions. Science 132:355–356
Govindjee, Veit W (2010) The Z-scheme of electron transport in photosynthesis: http://www.life.illinois.edu/govindjee/Z-Scheme.html (accessed continuously since August 2010)
Govindjee R(ajni), Rabinowitch EI, Govindjee (1968) Maximum quantum yield and action spectrum of photosynthesis and fluorescence in Chlorella. Biochim Biophys Acta 162:539–544
Govindjee, Pulles MPJ, Govindjee R(ajni), van Gorkom HJ, Duysens LNM (1976) Inhibition of the reoxidation of the secondary electron acceptor of photosystem II by bicarbonate depletion. Biochim Biophys Acta 449:602–605
Govindjee, Amesz J, Fork DC (eds) (1986) Light Emission by Plant and Bacteria. Acadamic Press, Orlando
Govindjee, Beatty JT, Gest H, Allen JF (eds) (2005) Discoveries in photosynthesis. Advances in Photosynthesis and Respiration, vol 20. Springer, Dordrecht
Grabolle M, Dau H (2005) Energetics of primary and secondary electron transfer in photosystem II membrane particles of spinach revisited on basis of recombination-fluorescence measurements. Biochim Biophys Acta 1708:209–218
Hill R, Bendall F (1960) Function of the two cytochrome components in chloroplasts: a working hypothesis. Nature 186:136–137
Hill J, Govindjee (2014) The controversy over the minimum quantum requirement for oxygen evolution. Photosynth Res. doi:10.1007/s11120-014-0014-8
Ke B (2001) Photobiochemistry and photobiophysics. Advances in Photosynthesis and Respiration, Vol 10, Springer, Dordrecht
Khanna R, Govindjee, Wydrzynski T (1977) Site of bicarbonate effect in Hill reaction: evidence from the use of artificial electron acceptors and donors. Biochim Biophys Acta 462:208–214
Khanna R, Wagner R, Junge W, Govindjee (1980) Effects of CO2-depletion on proton uptake and release in thylakoid membranes. FEBS Lett 121:222–224
Khanna R, Pfister K, Keresztes A, van Rensen JJS, Govindjee (1981) Evidence for a close spatial location of the binding sites of and for photosystem II inhibitors. Biochim Biophys Acta 634:105–116
Kok B (1959) Light induced absorption changes in photosynthetic organisms. II. A split beam difference spectrophotometer. Plant Physiol l34:184–192
Koroidova S, Shevela D, Shutova T, Samuelsson G, Messinger J (2014) Mobile hydrogen carbonate acts as proton acceptor in photosynthetic water oxidation. Proc Natl Acad Sci USA 111:6299–6304
Losada M, Whatley FR, Arnon DI (1961) Separation of two light reactions in non-cyclic photo-phosphorylation of green plants. Nature 190:606–610
Marcus R (1993) Electron transfer reactions in chemistry—theory and experiment (Nobel lecture). Angewandte Chemie-Int Ed 32:1111–1121
Moser CC, Keske JM, Warncke K, Farid RS, Dutton PL (1992) Nature of biological electron transfer. Nature 355:796–802
Nickelsen K, Govindjee (2011) The maximum quantum yield controversy: Otto Warburg and the ‘‘Midwest-Gang.’’ Bern studies in the history and philosophy of science, University of Bern, Bern, Switzerland
Papageorgiou GC, Govindjee (eds)(2004) Chlorophyll a fluorescence: A signature of photosynthesis. Kluwer Academic Publishers (now Springer), Dordrecht
Prakash JSS, Tiwari S (2013) Prasanna Mohanty (1934–2013): a pioneer and a loving teacher. Physiol Mol Biol Plants 19:301–305
Rabinowitch EI (1945) Photosynthesis and related processes, vol I. Interscience Publishers, NY
Rabinowitch EI (1956) Photosynthesis and related processes: vol 2, part 2 and addenda to vol 1 and vol 2, part 1. Interscience, NY
Rabinowitch E, Govindjee (1961) Different forms of chlorophyll a in vivo and their photochemical function. In: McElroy WD, Glass B (eds) A symposium on light and life. The Johns Hopkins Press, Baltimore, pp 378–386
Rabinowitch E, Govindjee (1969) Photosynthesis. John Wiley & Sons, NY
Shevela D, Eaton-Rye JJ, Shen JR, Govindjee (2012) Photosystem II and unique role of bicarbonate: a historical perspective. Biochim Biophys Acta 1817:1134–1151
Shevela D, Pishchalinikov R, Eichacker LA, Govindjee (2013) Oxygenic photosynthesis in cyanobacteria. In: Srivastava A, Rai AN, Neilan BA (eds) Stress biology of cyanobacteria : Molecular mechanism to cellular responses. CRC Press Boca Raton, FL, pp 3–40
Shinkarev VP, Wraight CA (1993) Kinetic factors in the bicycle model of oxygen evolution by photosystem II. Photosynth Res 38:315–321
Taiz L, Zeiger E (2010) Plant Physiology, 5th edition, Sinauer Associate
Tiwari S, Tripathy BC, Jajoo A, Das AB, Murata N, Sane PV, Govindjee (2014) Prasanna K. Mohanty (1934–2013): A great photosynthetiker and a wonderful human being who touched the hearts of many. Photosynth Res (in press)
Trissl HW, Gao Y, Wulf K (1993) Theoretical fluorescence induction curve derived from coupled differential equations describing the primary photochemistry of photosystem II by an excitation-radical pair equilibrium. Biophys J 64:974–988
Umena Y, Kawakami K, Shen JR, Kamiya N (2011) Crystal structure of oxygen-evolving photosystem II at a resolution of 1.9 Å. Nature 473:55–60
Van Rensen JJS, Xu C, Govindjee (1999) Role of bicarbonate in the photosystem II, the water-plastoquinone oxido-reductase of plant photosynthesis. Physiol Plant 105:585–592
Verne J (1875) L’IleMysterieuse. Pierre-Jules Hetzel, France
Warburg O, Krippahl G, Lehman A (1969) Chlorophyll catalysis and Einstein’s law of photochemical equivalence in photosynthesis. Am J Bot 56:961–971
Wydrzynski T, Govindjee (1975) New site of bicarbonate effect in photosystem II of photosynthesis: evidence from chlorophyll fluorescence transients in spinach-chloroplasts. Biochim Biophys Acta 387:403–408
Acknowledgments
We thank the Vice Chancellor Professor Baishnab Charan Tripathy for inviting Professor Govindjee to Ravenshaw University. We are thankful to Dr. Soumendra Kumar Naik, Dr. Padan Kumar Jena, all the faculty members of the Department of Botany, and all the B.Sc. & M.Sc. students for their cooperation. Special mention is made of Ankita Prusty and Prakash Prasad for their hard and friendly work and assistance during Govindjee’s stay. We are highly thankful to Tina (Trinath Barik) for the photographs and the video. We thank Sripadma Debata for her help in the demonstration of fluorescence to High School students attending the INSPIRE program; Sasmita (one name only) for showing students how to access electronic journals, available through Ravenshaw University library, and Y. Chandrakala (for help in several demonstations), and many others of the department, who made the stay of Govindjee at Ravenshaw comfortable and, more importantly, encouraged us to prepare this news report for the purpose of spreading education around the world. We are grateful to the following for their valuable comments that led to an improvement of this manuscript: Robert Blankenship, Julian Eaton-Rye, and Baishnab Charan Tripathy. Finally, we thank Govindjee for providing us a great deal of information that is included here as well as for reading this text and approving its publication in its present form.
Author information
Authors and Affiliations
Corresponding author
Electronic supplementary material
Below is the link to the electronic supplementary material.
Appendices
Appendix 1
Demonstration of Chlorophyll a fluorescence
The exciting power of chlorophyll a fluorescence as a signature of photosynthesis was shown by Govindjee to all the higher secondary school level students in a special lecture he was invited to give at Ravenshaw University; this lecture was coupled with practical demonstration, which was attended by ~160 students, and many teachers, from different colleges of the state of Odisha. Without taking the help of any sophisticated equipment Govindjee demonstrated how the tiny (2–10 % of the absorbed light; see e.g., Trissl et al. 1993) chlorophyll a fluorescence can be seen through naked eye in a comparatively dark room. Using leaves, he showed that the red chlorophyll a fluorescence, in a leaf, was not visible at all; and that from extracted chlorophyll was barely visible in the lecture hall. When he excited the leaf with a UV lamp, however, it was brilliant red in extracted chlorophyll (see a photograph of chlorophyll fluorescence in Govindjee and Govindjee 1974). What was interesting for students was to see that as leaves were infused with DCMU, they could observe red fluorescence, from the surface of the leaves with their eyes. This was so because when electron flow was inhibited beyond QA, the first plastoquinone acceptor of PS II, chlorophyll a fluorescence was high since the reaction center of PS II was “closed”, and photochemistry was not possible leading to lowered competition with fluorescence. He also explained how chlorophyll a fluorescence has proved to be an open window of photosynthesis research in general and light reactions in particular due to its intricate connection with a series of interactive and interdependent processes that take place during light-induced electron transport (see chapters in Govindjee et al. 1986; Papageorgiou and Govindjee 2004). While lecturing on fluorescence Govindjee did not forget to mention about the origin of the concept of fluorescence and showed the “celestial blue” light under UV light from tonic water. Like a young man he searched around the city of Cuttack, with a motorbike (driven by Prakash Prasad, in his class), to procure tonic water for his lecture-demonstration class, which shows his passion and commitment for the subject and deserves appreciation. (Fig. 4).
Appendix 2
-
(A)
Some useful web sites are:
-
(1)
Photosynthesis Web resources by L.Orr and Govindjee is at: http://www.life.illinois.edu/govindjee/Electronic%20Publications/2006/2006_gov_krogmann.pdf
-
(2)
The Photosynthetic Process by J. Whitmarsh and Govindjee is at: http://www.life.illinois.edu/govindjee/paper/gov.html
-
(3)
Milestones in Photosynthesis Research by Govindjee is at: http://www.life.illinois.edu/govindjee/papers/milestones.html
-
(4)
Photosynthesis and Time is at: http://www.life.illinois.edu/cgi-bin/gov/gov.cgi
-
(5)
Introduction of Photosynthesis and its Applications by Wim Vermaas is at http://photoscience.la.asu.edu/photosyn/education/photointro.html
-
(6)
TimeLine of Photosynthesis by Govindjee and D. Krogmann is at: http://www.life.illinois.edu/govindjee/Electronic%20Publications/2006/2006_gov_krogmann.pdf
-
(7)
Govindjee’s book collections are at: http://www.life.illinois.edu/govindjee/g/Books.html
-
(8)
Publications of Govindjee from 1994-2014 can be found at (there are some educational reviews there) http://www.life.illinois.edu/govindjee/recent_papers.html
-
(1)
-
(B)
Other web sites students may want to explore include:
http://www.hansatech instruments.com/forum/uploads/david_walker/down %20hill.pdf
http://en.wikipedia.org/wiki/Photosynthesis
http://en.wikipedia.org/wiki/Light-dependent_reactions
http://icewater.cms.udel.edu/mast634/lectures/12-photosynthesis.pdf
-
(C)
You-Tube presentations can be searched through “Google”; two are mentioned below:
-
(1)
An interview with Govindjee is at: https://www.youtube.com/watch?v=cOzuL0vxEi0
-
(2)
An interview with Pierre Joliot is at: https://www.youtube.com/watch?v=Tz4uIveE2hI
-
(1)
Rights and permissions
About this article
Cite this article
Mohapatra, P.K., Singh, N.R. Teaching the Z-Scheme of electron transport in photosynthesis: a perspective. Photosynth Res 123, 105–114 (2015). https://doi.org/10.1007/s11120-014-0034-4
Received:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s11120-014-0034-4