Comp, Biochem. Physiol. Vol. 83A, No. 1, pp. 157 177, 1986 0300-9629/86 $3.00+0.00
Printed in Great Britain © 1986 Pergamon Press Ltd
APPARENT AND REAL VARIABILITY IN THE
PRESENCE AND METAL CONTENTS OF
METALLOTHIONEINS IN THE CRAB CARCINUS
MAENAS INCLUDING THE EFFECTS OF
ISOLATION PROCEDURE AND METAL INDUCTION
V. W. T. WONG and P. S. RAINBOW
School of Biological Sciences, Queen Mary College, University of London, Mile End Road,
London E1 4NS, UK. Telephone: 01-980-4811
(Received 8 May 1985)
Abstract--1. Reducing conditions must be maintained throughout the procedure of isolating metal-
lothioneins from crabs. Dithiothreitol is preferred to 2-mercaptoethanol for long-term protection.
2. Two metallothioneins (I0,100 and 4100 mol. wt, respectively) in the hepatopancreas of the crab
Carcinus maenas showed great variability between individual crabs as to their presence and to their
contents of Zn, Cu and Cd.
3. The 10,100 mol. wt metallothionein was induced in the laboratory by exposure to Cu and Cd, and
variably by Zn-exposure. Laboratory induction did not raise significantly the total metal content of
0.88_ 1.13 g atoms/mol protein of this metallothionein in crabs from the Firth of Clyde, Scotland.
4. The 4100 mol. wt metallothionein was not induced in the laboratory by exposure to Cu, Cd or Zn.
This metallothionein in crabs from the Firth of Clyde. Scotland, contained 0.27 + 0.34 g atoms of total
Cu, Cd and Zn per mole of protein.
INTRODUCTION
Varying numbers of different metallothioneins (MTs)
including various isoforms have been reported in
different crab species. For example, the presence of a
single MT (ca 10-12,000mol. wt) separated by
Sephadex G-75 chromatography, resolving after ion-
exchange and electrophoresis into two isoforms, has
been reported in Scylla serrata (Olafson et al., 1979),
Cancer magister (Olafson et al., 1979) and Cancer
pagurus (Overnell and Trewhella, 1979). More re-
cently, Overnell (1982a,b, 1984a,b) has reported
resolution of such an MT into three isoforms in
Cancer pagurus.
Jennings et al. (1979) and Rainbow and Scott
(1979), however, reported the presence of two MTs of
about 27,000 and 12,000 mol. wt in Carcinus maenas
as separated by Sephadex G-75. Engel and Brouwer
(1984) similarly report metallothionein-like proteins
of about 30,000 and 40,000 tool. wt in addition to one
of about 10,000mol. wt in the lobster Homarus
americanus.
The previous paper (Wong and Rainbow, 1986)
has shown that C. maenas does contain two MTs but
these are of 10,100 and 4100 mol. wt, respectively, as
estimated on Sephadex G-50. It is therefore sur-
prising to see how divergent the results are within and
between different crab species whereas in vertebrate
systems, two MTs have been clearly identified (Kagi
and Nordberg, 1979).
*Abbreviations: DTT, dithiothreitol; 2-M, 2-Mercapto-
ethanol; MT, metallothionein; PMSF, phenylmethyl-
sulphonyl fluoride.
Many studies on MTs have been performed with
molecular-sieve chromatography using Sephadex
G-75. The resolving properties of this particular gel
type, however, are not totally suitable for separating
MTs from other low mol. wt impurities for MTs elute
near the lower limit (5000mol. wt) of the linear
fractionating range. Moreover, many such elutions
have been performed under oxidizing conditions (see
Minkel et al., 1980) causing artifactual changes to
elution profiles. Correspondingly many published
elution profiles of invertebrate metal-binding ligands
show considerable variation attributable to both ox-
idation artefacts and poor resolution.
Minkel et al. (1980) have shown that a reducing
agent such as 2-M* can prevent the formation of
oxidation artefacts. Once the effects of oxidation
have been minimized, it would seem appropriate to
switch to a gel type with a more suitable linear
fractioning range such as Sephadex G-50
(30,000-1500 mol. wt).
By using reducing conditions and Sephadex G-50
we hope to be able to distinguish between apparent
and real variability in the presence and metal contents
of MTs in the common shore crab Carcinus maenas.
MATERIALS AND METHODS
Carcinus maenas (L.) were obtained from the University
Marine Biological Station, Millport, Scotland, and held
immersed in the laboratory aquarium at 6°C for more than
a week in an attempt to reduce physiological variability
caused by entrained tidal rhythms.
Induction experiments: Crabs were exposed individually
for 14 days at 6°C under 12:12 light/dark regime to one of
four media at 33.3 ppt salinity: (i) artificial sea-water as
control [Tropical Marin New containing 2.5 ~zg Zn, 6/~g Cu
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