Table Of ContentCURRENT TOPICS IN PHOTOSYNTHESIS
III
Current topics in photosynthesis
Dedicated to Professor L.N.M. Duysens on the occasion of his retirement
edited by
1. AMESZ, A.l. HOFF and H.l. VAN GORKUM
Department oj Biophysics
State University oj Leiden
Leiden, The Netherlands
Reprinted from Photosynthesis Research, Volume 9, Numbers ]-2
1986 MARTINUS NIJHOFF PUBLISHERS
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Table of contents
Foreword vii
FRENCH, C.S.: Introduction ix
ANTENNAE
SHUBIN, V.V., KARAPETYAN, N.Y. and KRASNOVSKY, A.A.:
Molecular arrangement of pigment-protein complexes of photo-
system I [1]
GOLEN, D., KNOX, R. and BRETON, J.: Optical effects of sodium
dodecyl sulfate treatment of the isolated light harvesting complex
of higher plants [11 ]
SCHERZ, A. and PARSON, W.W.: Interactions of the bacteriochloro-
phylls in antenna bacteriochlorophyll-protein complexes of photo-
synthetic bacteria [19]
VAN DORSSEN, R.I., VASMEL, H. and AMESZ, J.: Pigment organi-
zation and energy transfer in the green photosynthetic bacterium
Chloroflexus aurantiacus. II. The chlorosome [31]
REACTION CENTERS
SETIF, P. and MATHIS, P.: Photosystem-I photochemistry: A new
kinetic phase at low temperature [45]
VAN GORKOM, H.I., MEIBURG, R.F. and de VOS, L.I.: Thermo-
dynamics of the charge recombination in photosystem II [53]
MURATA, N., ARAKI, S., FUJITA, Y., SUZUKI, K., KUWABARA,
T. and MATHIS, P.: Stoichiometric determination of pheophytin
in photosystem II of oxygenic photosynthesis [61 ]
SAYGIN, 0., GERKEN, S., MEYER, B. and WITT, H.T.: Total re
covery of 02 evolution and nanosecond reduction kinetics of
chlorophyll-ail (P-680+) after inhibition of water cleavage with
acetate [69]
CLELAND, R.E., MELIS, A. and NEALE, P.I.: Mechanism of photo
inhibition: Photochemical reaction center inactivation in system
II of chloroplasts [77]
LOUS, E.I. and HOFF, A.I.: Triplet-minus-singlet absorbance differ
ence spectra of reaction centers of Rhodopseudomonas sphaer
oides R-26 in the temperature range 24· -290 K measured by Mag-
neto Optical Difference Spectroscopy (MODS) [87]
ELECTRON TRANSPORT
PADHYE, S.,KAMBARA, T.,HENDRICKSON,D.N. and GOVINDJEE:
Manganese-histidine cluster as the function center of the water
oxidation complex in photosynthesis [101]
v
vi
JOLlOT, P. and JOLlOT, A.: Mechanism of proton-pumping in the
cytochrome b/f complex [111]
GHANOTAKIS, D.F., YOCUM, C.F. and BABCOCK, G.T.: ESR
spectroscopy demonstrates that cytochrome bss9 remains low
potential in Ca2+- reactivated, salt-washed PS II particles [123]
MYERS, J.: Photosynthetic and respiratory electron transport in a
cyanobacterium [133 J
SYBESMA, C., SCHOWANEK, D., SLOOTEN, L. and WALRAVENS,
N.: Anoxygenic photosynthetic hydrogen production and electron
transport in the cyanobacterium Oscillatoria limnetica [147]
VAN DER WAL, H.N., GORTER, P.Y. and VAN GRONDELLE,
R.: Oxidation of cytochrome C2 by photosynthetic reaction centers
of Rhodospirillum rubrum and Rhodopseudomonas sphaeroides in
vivo. Effect of viscosity on the rate of reaction (157]
OVERFIELD, R.E. and WRAIGHT, C.A.: Photooxidation of mito
chondrial cytochrome c by isolated bacterial reaction centers:
Evidence for tight-binding and diffusional pathways. [165]
WYNN, R.M., GAUL, D.F.,WON-KI CHOI, SHAW, R.W. and KNAFF,
D.B.: Isolation of cytochrome bCl ·complexes from the photo
synthetic bacteria Rhodopseudomonas viridis and Rhodospirillum
rubrum (179J
FORSTER, V. and JUNGE, W.: On the action of hydroxylamine,
hydrazine and their derivatives on the water-oxidizing complex [195]
MEMBRANE STRUCTURE AND ION TRANSPORT
VAN KOOTEN, 0., SNEL, J.F.H. and VREDENBERG, W.J.: Photo
synthetic free energy transduction related to the electric potential
changes across the thylakoid membrane [209]
FALUDI-DANIEL, A., MUSTARDY, L.A., VASS, I. and KISS, J.G.:
Energization and ultrastructural pattern of thylakoids formed
under periodic illumination followed by continuous light [227]
BARBER, J. and GOUNARIS, K.: What role does sulpholipid play
within the thylakoid membrane? [237]
EVOLUTION
OLSON, J.M. and PIERSON, BK.: Photosynthesis 3.5 thousand
million years ago [249]
METHODS
SCHREIBER, U.: Detection of rapid induction kinetics with a new
type of high-frequency modulated chlorophyll fluorometer [259]
LAVOREL, J.: A Mop.te-Carlo method for the simulation of kinetic
models [271]
Foreword
Four decades ago, when Lou Duysens was about to start his work on fluo
rescence and energy transfer in photosynthesis that would lead to his thesis
[1], very little was known about the molecular mechanisms of photosyn
thesis, certainly from our present-day point of view. However, this state of
affairs would rapidly change in the ensuing years by the introduction of
modern physical and biochemical techniques. Especially the field of optical
spectroscopy, on which the work of Duysens had such a significant impact,
has proved to be one of the most fruitful techniques in the study of primary
processes and electron transfer reactions in photosynthesis.
Duysens' thesis established the role of energy transfer in photosynthesis
and also showed for the first time the existence in photosynthetic bacteria
of light-induced absorbance changes of what is now known as the primary
electron donor P-870. Subsequent studies by the same method demonstrated
the photo-oxidation of cytochromes, both in bacteria [2] and in algae
[3,4] and of the absorbance changes [3] that were later found to be due to
electro chromic band shifts of antenna pigments. Measurements of cyto
chrome kinetics in light of various wavelengths led to the concept of two
photosystems in green plant photosynthesis [5], whereas a study of the
factors affecting the fluorescence yield of chlorophyll gave the first infor
mation on the electron acceptor Q of photo system II [6].
Research in many laboratories all over the world has led to an under
standing of primary and secondary processes in photosynthesis and of the
structure of the photosynthetic membrane that is vastly more detailed
than could have been envisaged 40 years ago. This is well illustrated by this
special issue on the occasion of Lou Duysens' retirement, which we hope
will give an impression of the present status of research.
As editors of this issue of Photosynthesis Research we are very grateful
to our colleagues - many of which at one time or another worked at the
Leiden laboratory for some months or years - for their contributions.
We are much indebted to the Editors of Photosynthesis Research, Prof.
Govindjee and Dr. R. Marcelle who endorsed our plans, and to Ir. A.C.
Plaizier of Martinus Nijhoff Publishers for his kind help.
The Editors,
J. Amesz
A.J. Hoff
H.J. van Gorkom
References
1. Duysens LNM (1952) Doctoral thesis, University of Utrecht
vii
viii
2. Duysens LNM (1954) Nature 173:692
3. Duysens LNM (1954) Science 120:353-354
4. Duysens LNM (1955) Science 121:210-211
5. Duysens LNM, Amesz J and Kamp BM (1961) Nature 190:510-511
6. Duysens LNM and Sweers HE (1963) In 'Studies on Microalgae and Photosyn
thetic Bacteria', Special Issue Plant Cell Physiol, pp 353-372. Tokyo, University
of Tokyo Press
Introduction
A major event in the history of photosynthesis research was the establish
ment of the Biophysical Research Group in Utrecht with support of the
Rockefeller Foundation about 1935. This group was initiated by Prof. AJ.
Kluyver, the Delft microbiolOgist, and Prof. Ornstein of the Physics Depart
ment at Utrecht to combine expertise in both biology and physics. This
group, directed by Prof. Wassink and later by Prof. Thomas, produced an
abundance of top quality research and also trained very distinguished scien
tists. One of these, Prof. Louis l~.M. Duysens, whose impending retirement
is the occasion for this volume, has indeed more than fulft11ed the hopes of
the Utrecht Group's founders by his distinguished research career and pro
ductive leadership of the Biophysical laboratory at Leiden.
Because Lou Duysens had been thoroughly trained in basic physics he
was able to think clearly in mathematical terms about biological problems
even as a graduate student. Starting with studies of energy transfer between
photosynthetic pigments, Duysens' 1952 Utrecht thesis clearly presented
the basic optics of cell suspensions. The concluding part of that thesis re
ported the ingenious measurement of very small changes of absorption
induced by light or by chemical treatment in photosynthetic bacteria. Such
measurements of absorption changes have become widely used in many
laboratories and have made possible the chemical identification of many
intermediate steps in the process of photosynthesis. One of his discoveries
when he came as a postdoctoral fellow to the Carnegie Institution was to
find an absorption change at 515 nm in Chlorella. The challenge to explain
this electrochromic change and its usefulness in studies of photosynthesis
is attested by the numerous papers that have appeared on this subject since
then. In the red alga Porphyridium cruentum Duysens and Amesz and Kamp
found the 420 nm absorption change of cytochrome-f to indicate its oxi
dation by illumination at 680 nm, light absorbed by chlorophyll, and its
reduction by 562 nm, light absorbed by phycoerythrin. They proposed the
names system 1 and system 2 for these two separately acting groups of
pigments which has become universally used. Studies of chlorophyll fluor
escence, of absorption changes, and of energy transfer in photosynthetic
organisms has been a continuing activity in Duysens laboratory. Photo
synthetic bacteria and red or blue green algae, each with their own pigment
systems, have been used as well as green algae and leaf chloroplasts through
out his career. The general principles of photochemical energy conversion
ix
x
have been elucidated by selection of the most suitable organism for each
investigation. As this volume will show Duysens' efforts have been out
standingly successful in this competitive field of worldwide interest.
Department of Plant Biology C. Stacy French
Carnegie Institution of Washington
Stanford, CA 94305, USA
[l]
Molecular arrangement of pigment-protein complex of
photosystem 1
V.V. SHUBIN, N.V. KARAPETYAN and A.A. KRASNOVSKY
A.N. Bakh Institute of Biochemistry, USSR Academy of Sciences, Moscow 117071,
USSR
(Received 25 June 1985)
Key words: light-harvesting complex (pea); cPU, optical absorption, linear dichroism,
chroism, protein secondary structure
Abstract. The circular dichroism (CD) method was applied to study the molecular
organization of P700, antenna chlorophyll and protein of photo system 1 complexes
(CPl), isolated from chloroplasts under mild treatment with Triton X-100. Analysis
of CD spectra and protein: chlorophyll: P700 ratios for CP1 complexes that were differ
ent in their chlorophyll content indicate that CP1 preparations can be considered as a
mixture of CP1-RC, containing P700 (10-20%), and CPI-LH without P700 (80-90%).
Both types of complexes contain approximately 25 chlorophyll molecules, and the
destruction of their spatial organization with detergents represents a cooperative tran
sition. The rate of chlorophyll destruction in CPI-LH is much higher than that in CPI
RC. In both complexes a 65 kDa polypeptide predominates, whose secondary structure
(typical for 01./13 proteins) is stable to Triton X-100 and does not depends on the chloro
phyll content. Chlorophyll seems to be grouped in clusters (5-7 molecules) in the
hydrophobic cores of 2-3 parallel 01./13 domains of the 65 kDa protein. Only one of the
clusters in CPI-RC includes P700; on P700 photooxidation the change of its inter
action with the nearest pigment environment results in a complicated shape of the light
induced CD spectra.
Abbreviations
PSI, photosystem I; CPI pigment-protein complex of PSI; Chi, chlorophyll
a; CPI-140, CPI with ratio Chl:P700 140; RC, reaction center; LH, light
harvesting pigment; CPI-RC, CPI, containing P700; CPI-LH, CPI without
P700 (containing LH); CD, circular dichroism; SDS, sodium dodecyl sulfate.
Introduction
It is generally accepted that the primary photosynthetic reactions take place
in the reaction centers (RC) of photosystems of green plants and photo
synthetic bacteria. The suggestion [5, 10] that only a part of Chi is photo
chemically active anticipated the discovery of the RC. Bacterial RC can be
isolated in purified state and crystallized, whereas the RC's of green plants con
tain a large amount of antenna Chi. Difficulties in isolation and crystallization
Dedicated to Prof. L.N.M. Duysens on the occasion of his retirement
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