ELSEVIER Plant Science 99 (1994) I I 1-124
plan ience
Review article
Phosphoenolpyruvate carboxylase: structure,
evolution
regulation and
Loic Lepiniec *a, Jean Vidal a, Raymond Chollet b, Pierre Gadal a, Claude Cr6tinT a
aLaboratoire de Physiologie Vdgdtale Moldculaire, URA CNRS D 1128, Universitd de Paris-Sud, Bdtiment 430, 91405 Orsay
Cedex, France
bDepartment of Biochemistry, University of Nebraska-Lincoln, Lincoln, NE 68583-0718, USA
Received 29 December 1993; revision received 29 March 1994; accepted 18 April 1994
Abstract
Plant phosphoenolpyruvate carboxylase (EC 4.1.1.31; PEPC) is encoded by a small multigene family in which the
expression of each member is controlled individually by exogenous (light, environmental) and/or endogenous (hor-
monal and developmental) stimuli. The involvement of putative trans-acting factors and consensus cis-elements of
promoters in the specific transcriptional regulation of the PEPC genes is discussed. At the post-translational level, the
regulatory strategy of the plant enzyme is mainly to offset the negative effect of the feedback inhibitor, L-malate, the
end-product of the oxaloacetate reduction. All plant PEPC-forms are under positive and negative allosteric control
by metabolite effectors and possess a consensus phosphorylation site containing a target serine residue near their N-
terminus (e.g. Ser8 in C4 PEPC from sorghum). In C 4 and Crassulacean acid metabolism (CAM) plants, a complex
signal-transduction chain activates ~a Ca2+-independent protein-serine kinase responsible for regulatory phosphoryla-
tion of PEPC. A more thorough understanding of the functional and regulatory properties of the bacterial and C 4
enzymes has emerged by exploiting recombinant proteins and site-directed mutagenesis. In these newly opened areas,
PEPC offers one of the best characterized paradigms of plant signaling. Finally, some emerging ideas on the evolution
and phylogenetic relationships of the various PEPC isoforms are presented.
Keywords: Phosphoenolpyruvate carboxylase; Gene family; Gene expression; Evolution; Mutagenesis; Phosphory-
lation
* Corresponding author, Laboratorium Genetika, Ledegan-
ckstraat 35, Rijksuniversiteit Gent, B-9000 Gent, Belgium.
Tel.: 32 (0) 9264 5192; Fax.: 32 (0) 9264 5349.
tWe would like to dedicate this review to the memory of Dr.
Claude Cr6tin, untimely deceased in August 1993.
1. Introductory remarks
Phosphoenolpyruvate carboxylase (PEPC)
catalyses the 'irreversible' /3-carboxylation of
phosphoenolpyruvate (PEP) in the presence of
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