1021-4437/01/4802- $25.00 © 2001 MAIK “Nauka /Interperiodica” 0171
Russian Journal of Plant Physiology, Vol. 48, No. 2, 2001, pp. 171–175. From Fiziologiya Rastenii, Vol. 48, No. 2, 2001, pp. 203–207.
Original English Text Copyright © 2001 by Zervoudakis, Georgiou, Angelopoulos.
INTRODUCTION
Pyruvate kinase (PK; ATP: pyruvate phosphotrans-
ferase, EC 2.7.1.40) catalyses the irreversible conver-
sion of phosphoenolpyruvate (PEP) to pyruvate in the
presence of Mg
2+
and K
+
, with concomitant synthesis
of ATP according to the following reaction:
PEP + ADP pyruvate + ATP.
The enzyme has been isolated from a wide variety
of tissues [1]. The kinetics and regulatory properties of
the plant enzyme have been studied mainly in germi-
nating or developing castor bean endosperm [2–4], in
green algae [5, 6] and less in leaves of higher plants [3,
7]. It is considered to be a key regulatory enzyme in the
final steps of aerobic glycolysis and its activity depends
on the rates of respiration [8], photorespiration [9], and
ammonia assimilation [10, 11]. The existence of two
isoenzymes with different immunological, physical,
and kinetic properties has also been reported [5, 6]. One
isoenzyme was found in the cytosol (PK
c
; cytosolic
pyruvate kinase) and the other (PK
p
; plastid pyruvate
kinase) was found in the plastids of seed endosperms as
well as in the chloroplasts of green algae and leaves
from C
3
plants [5, 7, 12].
Research to this point has been focused mainly on
the PK from tissues of C
3
plants and green algae, and,
for studying PK, the lactate dehydrogenase-coupled
assay [13, 14] has been used. PEPC interferes with this
assay, although it does not present a major problem in
C
3
plant extracts, where PEPC is low as compared to
PK, and can be inactivated by the relatively effective
inhibitor maleate [15]. In C
4
plants, however, PEPC
activity is much higher as compared to PK, and its
interference was eliminated by thermal inactivation of
crude extracts of Zea mays [16]. Based on our findings
that the purified enzyme has the optimum pH 6.2 [17]
and PEPC is inactivated by acidic pH [18], we devel-
oped simple extraction and LDH-coupled PK assay
methods that least modified PK quaternary structure to
provide a way of assessing its physiological properties
in crude leaf extracts from C
4
plants.
MATERIALS AND METHODS
Reagents. Mes, Mops, PVP (10 kD), DTT, β-
NADH, ADP, phosphoenolpyruvate, lactate dehydro-
genase (from rabbit muscle, salt free form), EDTA
were from Sigma (United States), Tris and sea sand
from Merck (Germany). All other reagents used were
of the highest purity.
Plant material. Cynodon dactylon (L.) Pers. plants
were grown from rhizomes in soil pots kept outside
during summer months or kept in a greenhouse during
winter months. Fully expanded mature leaves were
taken during daytime hours for the preparation of crude
extracts. Mature leaves from C
4
species Setaria verti-
cillata (L.) Beauv., Sorghum halepense (L.) Pers., Dig-
itaria sanguinalis (L.) Scop., and Amaranthus sp. were
collected in fields.
Extraction, assay conditions, and temperature effect
on PK activity. For enzyme extraction, 1 g of leaf tissue
from C. dactylon and from other C
4
species was ground
at room temperature in a porcelain mortar with purified
Pyruvate Kinase Activity in Crude Extracts of Leaves
of Cynodon dactylon and Other C
4
Plants
1
G. Zervoudakis*, C. D. Georgiou**, and K. Angelopoulos*
* Department of Biology, Laboratory of Plant Physiology, University of Patras, Patra, 26100 Greece
fax +30(61)997228; e-mail: angelop@upatras.gr
** Department of Biology, Section of Genetics, Cell and Developmental Biology, University of Patras, Patra, 26100 Greece
e-mail: c.georgiou@upatras.gr
Received May 17, 2000
Abstract—A new extraction procedure and an LDH-coupled assay method are presented for the study of pyru-
vate kinase (PK) in leaf crude extracts from Cynodon dactylon (L.) Pers and other C
4
plants. Extraction at pH
6.8 and assay at pH 6.2 facilitated the measuring of PK activity by eliminating phosphoenolpyruvate carboxylase
interference more effectively than the thermal inactivation or chemical inhibition previously used. The method
suggested did not affect the kinetic properties of PK as compared to the purified enzyme from C. dactylon.
Key words: Amaranthus sp. - Cynodon dactylon - Digitaria sanguinalis - Setaria verticillata - Sorghum
halepense - C
4
plants - pyruvate kinase
1
This article was submitted by the authors in English.
Abbreviations: BSA—bovine serum albumin; DTT—dithiothrei-
tol; LDH—lactate dehydrogenase; Mops—3-(N-morpholino)pro-
panesulfonic acid; PEPC—phosphoenolpyruvate carboxylase;
PK—pyruvate kinase; PVP—polyvinylpyrrolidone.