Pergamon
Comp.Biochem. Physiol. Vol.107n,No. 4, pp. 585-591,1994
Copyright © 1994 Elsevier Science Ltd
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Regulation of pyruvate kinase (PK) from the ventricle
of the land snail Helix lucorum L. during early
and prolonged estivation and hibernation
Basile Michaelidis and Theologos Pardalidis
Laboratory of Animal Physiology, Department of Zoology, Science School,
University of Thessaloniki, GR-54006 Thessaloniki, Greece
The kinetic properties of pyruvate kinase (PK) from the ventricles of early and prolonged estivated
and hibernated land snails, Helix lucorum, were studied. The kinetic properties of PK from the
ventricles of snails accfimated to cold for 4 days and 2 months were also determined. Short estivation
(4 days) converted the PK to a less active form, possibly via enzyme phosphorylation. After 2 months
of estivation, PK exhibited kinetic properties similar to control enzyme, except that the Vmx increased
about 2-fold, indicating metabolic reorganization. During early and prolonged hibernation, the kinetic
properties of PK did not change significantly. The results indicate that low temperature and low pH
may be the main modulators of enzyme activity in the ventricle of H. iucorum during hibernation.
Key words: Land snail; Estivation; Hibernation; Ventricle; Pyruvate kinase; Glycolysis; Regulation.
Comp. Biochem. Physiol. I07B, 585-591, 1994.
Introduction
Land snails respond to unfavorable conditions
(dehydration, low temperature, food depri-
vation) by entering a state of quiescence
(dormancy). In this state oxygen consumption is
greatly reduced and metabolic rate drops below
the standard resting state (Nopp, 1974).
The dormant period is characterized by
an increase in CO2 content in extracellular
fluids, which results in a drop of extracellular
and intracellular pH (pHi) (Barnhart and
McMahon, 1988). It has been shown that hyper-
capnia strongly depresses oxygen consumption
in Otala lactea and it has been proposed that the
fall of pHi as a result of hypercapnia may be
involved in the depression of metabolic rate
during estivation (Barnhart, 1989; Barnhart and
McMahon, 1987; Rees and Hand, 1990). Regu-
lation of glycolysis via intracellular pH has been
considered elsewhere (Busa and Nuccitelli,
1984) and it is known that a drop in pH i results
Correspondence to: B. Michaelidis, Laboratory of Animal
Physiology, Department of Zoology, Science School,
University of Thessaloniki, GR-54006 Thessaloniki,
Greece, Fax: 206-138.
Received 15 July 1993; accepted 22 September 1993.
in the suppression of the glycolytic rate by
inhibiting the key glycolytic enzyme, phospho-
fructokinase (PFK). Specifically, a drop in pHi
results in a shift of enzyme molecules to a
dissociated form and consequently to loss of
enzyme activity (Carpenter and Hand, 1986a;
Somero and Hand, 1990).
Recent studies, however, have shown that the
depression of the glycolytic rate in the tissues of
land snails during entry into estivation is inde-
pendent of hypercapnia (Rees and Hand, 1991).
More extensive studies have shown that
mechanisms involving the covalent modification
of PK and PFK, possibly via phosphorylation/
dephosphorylation of enzymes, the regulation
of PFK via the potent activator fructose-2,6-bis-
phosphate (F-2,6-P2) and the binding of glyco-
lytic enzymes to subcellular particles, may be
involved in metabolic rate depression in the
tissues of land snails during dormancy (Storey
and Storey, 1990).
Another mechanism which may be involved
in the depression of oxygen consumption
in estivating snails is the low supply of oxygen
to the tissues through circulation. During
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