Comp. Biochem. Physiol. Vol. 78B, No. 2, pp. 351-353, 1984 0305-0491/8453.00+ 0.00
Printed in Great Britain © 1984 Pergamon Press Ltd
A MODIFIED LOWRY ASSAY TO MEASURE
LEVELS OF PROTEINS LACKING
AROMATIC AMINO ACID RESIDUES
(e.g. METALLOTHIONEINS)
V. W. T. WONG and P. S. RAINBOW
School of Biological Sciences, Queen Mary College, Mile End Road, London E1 4NS, UK
(Tel: 01-980-4811)
(Received 25 November 1983)
Abstract--1. A combination of two modifications to the Lowry protein assay is recommended for
estimation of metallothionein (lacking aromatic amino acid residues and requiring reducing conditions
for isolation).
2. Serum thymic factor (STF), a decapeptide lacking aromatic amino acids, is recommended for use
as a standard after allowance for the effects of short chain length on colour intensity development.
INTRODUCTION
Metallothioneins are a group of low mol. wt metal-
binding proteins which are thought to occupy a
central position in the metabolism of particular trace
metals such as copper, cadmium, zinc and mercury
(Vallee, 1978). They act as ligands for these toxic
metals absorbed by animals exposed to metal pollu-
tion (Noel-Lambot, 1976) and they may play a role
in the regulation of levels of these trace metals in the
organism. Metallothioneins are widespread among
vertebrates and invertebrates, differing only slightly
between species in their amino acid sequences. Char-
acteristically, metallothioneins have a low mol. wt
(about 12,000), are very high in cysteine residues
(about 30~o) and lack the aromatic amino acids
tyrosine and tryptophan (VaUee, 1978). Under ox-
idising conditions, the isolated proteins tend to form
aggregates via disulphide bridges on the cysteine
residues (Minkel et al., 1980), but can be protected by
the use of a reducing agent such as DTT* at concen-
trations of 1 mM (Cleland, 1964).
The present authors are working to isolate and
study metallothionein in the crab Carcinus maenas,
and in the course of this study, we needed to obtain
absolute protein values to relate to metal values. It is
difficult to obtain sufficient amounts of isolated crab
metallothionein for gravimetric or amino acid anal-
ysis and purified metallothionein standards from
other sources are as yet unavailable commercially. A
routine procedure for measuring metallothionein was
therefore required using a common protein as a
standard.
A simple, direct and sensitive procedure for ana-
lysing metallothionein in crude homogenates has not
yet been devised (Olafson and Sim, 1979). Con-
sequently, workers in this field have relied on a
variety of crude and cumbersome techniques: ultra-
*Abbreviations: BSA, bovine serum albumin; CHM,
chymotrypsinogen-A; DTT, dithiothreitol; OVA, oval-
bumin; STF, serum thymic factor.
violet absorption at 250nm based on the molar
extinction coefficient of 6.8ml/mg/cm for metal-
lothionein (Kagi and Vallee, 1961) is too non-specific
to be sufficiently accurate: a modified biuret method
(Nordberg et al. 1972; Piscator, 1962) was found to
be too insensitive for our purposes; dye-binding
methods with reagents such as Amido Black (Pis-
cator, 1962) and Coomassie Blue (Overnell and
Grant, 1981) were unsuitable because of the variable
nature of their responses to different proteins (Pierce
and Suelter, 1977; Van Kley and Hale, 1977); staining
with Schiff's reagent (Overnell and Grant, 1981)
requires further developments; and finally, elaborate
electrochemical techniques (Olafson and Sim, 1979),
fluorescence techniques (Bohlen et al., 1973; Naka-
mura and Pisano, 1976) and determinations based on
amino acid analysis (Hoch and Vallee, 1953; Buhler
and Kagi 1974; Kagi et al., 1974; Overnell, 1982a,b)
were considered to be too time-consuming for routine
application.
Lowry's method for protein assay (Lowry et al.,
1951) is a widely used procedure in routine analysis.
The Lowry assay can be interpreted in terms of two
reactions: (1) the biuret reaction--an initial reaction
whereby copper ions are thought to bind to the
protein backbone, and (2) the Folin phenol
addition--whereby tyrosine and tryptophan residues
enhance the final colour development. Protein mea-
surements using the Lowry method are often made
against a common "standard" protein such as bovine
serum albumin (BSA) and protein values are given in
"BSA units" rather than exact values. The lack of
aromatic amino acid residues in metallothionein will
clearly affect further colour development in com-
parison with BSA. The unmodified Lowry method
was therefore considered unsuitable for our purposes.
Modifications have since been made to the Lowry
method to render it more suitable for our require-
ments. Dorsey et al. (1977) devised a simple yet
effective modification of Lowry's method which ap-
pears to bring different proteins to a similar final
colour development (by the inclusion of a heating
351
C.B,P. 78/2B~D