Photosynthesis Research 57: 203–213, 1998.
© 1998 Kluwer Academic Publishers. Printed in the Netherlands.
203
Regular paper
Changes in the photosynthetic pigments in bean leaves during the first
photoperiod of greening and the subsequent dark-phase. Comparison
between old (10-d-old) leaves and young (2-d-old) leaves
Benoît Schoefs
1
, Martine Bertrand
2
& Yves Lemoine
3
1
Laboratory of Biomembranes, University of South Bohemia, Branisovska 31, 370 05 Ceske Budejovice, Czech
Republic;
2
National Institute for Marine Sciences and Techniques, Conservatoire National des Arts et M´ etiers, BP
324, 50103 Cherbourg Cedex, France;
3
Laboratory of Plant Cytophysiology and Phycology (SN2), University of
Lille 1, 59655 Villeneuve d’Ascq, France
Received 4 November 1997; accepted in revised form 23 June 1998
Key words: carotenoid, chlorophyll b formation, chlorophyllide a esterification, accumulation of photosynthetic
pigments, HPLC, protochlorophyllide a
Abstract
Chlorophyll and carotenoid variations of 2-d-old and 10-d-old bean leaves (Phaseolus vulgaris var Red Kidney)
were analyzed by HPLC during the first photoperiod of greening (16 h light + 8 h dark). The HPLC method used is
suitable for the separation of cis- and trans-carotenoid isomers, Pchlide a and Chlide a as well as their esters. The
main results are (1) before illumination the composition of the carotenoid pool is similar at the two developmental
stages; (2) non-illuminated 2-d-old leaves are devoid of Pchlide a ester; (3) chlorophyll and carotenoid accumu-
lation in 2-d-old leaves presented a lag phase twice longer than observed in 10-d-old ones; (4) Chlide a seems
directly esterified to Chl a in 2-d-old leaves whereas esterification requires four steps in 10-d-old leaves and, (5)
the kinetics of Chl and carotenoid accumulation are different at the two investigated developmental stages.
Abbreviations: Chl – chlorophyll; Chlide – chlorophyllide, C
X-Y
– Chlide a having absorbance and fluorescence
maxima at X and Y nm at 77 K; DHGG – dihydrogeranylgeraniol; GG – geranylgeraniol; HPLC – High Per-
formance Liquid Chromatography; Pchl – protochlorophyll; Pchlide – protochlorophyllide, Ph – phytol; THGG –
tetrahydrogeranylgeraniol
Introduction
Dark-grown angiosperm leaves synthesize carotenoids
but are devoid of Chl (Morren 1858). In the absence of
light, they accumulate a Chl precursor, namely Pchlide
a, a modified collection of xanthophylls and little or no
carotene. Usually no carotenoid precursor may be ob-
served, except if leaves remain for a very long period
in the dark (e.g. Schoefs et al. 1995).
When dark-grown plants are illuminated, carotenoid
accumulation in leaves is triggered (Goodwin and
Phagpolngarm 1960). In the same time photoactive
Pchlide a is transformed to Chlide a which is then
esterified by GG pyrophosphate which after three suc-
cessive dehydrogenations yielding Chlide a Ph, is
integrated into pigment–protein complexes (i.e. Chl a;
reviewed by Rüdiger 1992; Franck and Schoefs 1996).
Chl a accumulation during greening exhibits a lag
phase of several hours whose length appears to depend
upon the age: the younger the leaf, the shorter the
lag phase (Akoyunoglou and Argyroudi-Akoyunoglou
1966, Sisler and Klein 1963). Chl b formation starts
after that of Chl a. It is generally accepted that it is
synthesized at the expense of Chl(ide) a (reviewed by
Schoefs and Bertrand 1997).
10-d-old dark-grown leaves are not encountered in
nature since prolonged darkness does not occur usu-
ally. In natural conditions, proplastids of young leaves