Insect Bioehem. Vol. 16, No. 3, pp. 441~,47. 1986 0020-1790/86 $3.00+0.00
Printed in Great Britain. All rights reserved Copyright © 1986 Pergamon Press Ltd
CHARACTERIZATION OF CUTICULAR PROTEINS FROM
THE MIGRATORY LOCUST, LOCUSTA MIGRATORIA
SVEND OLAV ANDERSEN, 1 PETER HOJRUP 2 and PETER ROEPSTORFF 2
qnstitute of Biological Chemistry A, Copenhagen University, Universitetsparken 13, DK 2100
Copenhagen and 2Department of Molecular Biology, Odense University, Campusvej 55, DK 5230 Odense,
Denmark
(Received 17 June 1985; revised and accepted 21 August 1985)
Abstract--Proteins were extracted from the still unhardened (teneral) cuticle of the migratory locust,
Locusta migratoria. The proteins are soluble only at extreme pH-values and at low ionic strength, the
solubility increases with decreasing temperature. The unhardened cuticle contains approx. 100 different
proteins according to two-dimensional polyacrylamide gel electrophoresis. The majority of the proteins
are very basic. The basicity and solubility properties of the proteins have necessitated development of
modified electrophoretic procedures. The amino acid composition of the bulk protein shows that alanine,
proline, glycine, valine and tyrosine constitute two thirds of the total amino acid content and that cysteine,
methionine and tryptophan are absent.
The proteins have been extracted from various parts of the cuticle and analysed by two-dimensional
electrophoresis. Characteristic protein compositions were found for cuticle from the different body parts.
Amino acid analyses of these extracts are strikingly similar. The only significant difference is in the
glycine-alanine ratio. Cuticles that are destined to become hard are extremely rich in alanine, whereas
the flexible parts of the cuticle are enriched in glycine. The results indicate that the proteins of locust cuticle
constitute a group of structural proteins different from other known structural proteins.
Key Word Index: Locusta migratoria, cuticle, protein mapping, amino acid composition, hydro-
phobicity, solubility
INTRODUCTION
Insect cuticle is an extracellular layer, consisting
mainly of proteins and chitin, which surrounds the
whole animal (Neville, 1975; Hepburn, 1976). The
cuticle functions both as a skeleton and as a barrier
towards the environment, and it is also responsible
for the coloration of most insects, either by pigments
in the cuticle or by submicroscopic structures giving
rise to physical colours. The ultrastructural or-
ganization of cuticle follows a general plan in all
insect groups: long filaments of chitin are embedded
in a matrix of proteins, and most differences between
cuticles are differences in the proteinaceous matrices
and the way they are modified during maturation.
Cuticles vary considerably in physical properties;
most larval cuticles are very soft and flexible, and in
the living animal they are kept distended by the
internal pressure in the larvae, whereas large parts of
the cuticle of adult insects are quite hard and inex-
tensible, and specialized parts, such as mandibles, can
be extremely hard. Some of these differences are due
to differences in the degree of chemical stabilization
(sclerotization) of the cuticular proteins (Andersen,
1974). The proteins are synthesized in the epidermal
cells and secreted from the apical part of the cells.
Some time after secretion, either before or after the
insect has ecdyzed to the adult stage, the proteins can
be sclerotized by an enzyme-catalyzed, oxidative in-
corporation of phenolic material into the cuticle,
presumably as cross-links between the protein mole-
cules (Andersen, 1979). Sclerotization transforms the
proteins from a soft, soluble material which is easily
digested by proteolytic enzymes into a stiff, insoluble
mass, which is much more resistant towards en-
zymatic proteolysis.
Various types of cuticle differ significantly in amino
acid composition, indicating the presence of different
proteins. As a rule, soft cuticles are characterized by
a high content of polar amino acid residues, and hard
cuticle tends to be rich in hydrophobic amino acids
(Andersen, 1979). The old idea of a single, general
cuticular protein, arthropodin, has long been aban-
doned, and more and more evidence shows that all
types of cuticle contain a large number of proteins
(Willis et al., 1981). The exact pattern of proteins in
a cuticle will presumably be of importance for its
properties, but since only a few samples have been
investigated it is too early to draw many conclusions.
Proteins from larval cuticle of fruitflies, Drosophila
melanogaster (Fristrom et al., 1978), larval cuticle of
fleshflies, Sarcophaga bullata (Lipke et al., 1981), and
cuticle from larvae, pupae and adults of the American
silkmoth, Hyalophora cecropia (Willis et al., 1981),
have been isolated and partly characterized. The total
amino acid sequence has been obtained for several of
the Drosophila larval cuticular proteins (Snyder et al.,
1982); the published information for the other
purified proteins concerns mainly molecular weights,
isoelectric points and amino acid compositions. All
these proteins resemble each other in having rela-
tively low isoelectric points (pIs from 4 to 6), low
molecular weights (8-25 kdal), and in being rich in
polar amino acid residues.
Only the easily extractable proteins have been
characterized, and they account for about half of the
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