Pergamon
PH: S0031-9422(96)00193-8
Phytochemistry, Vol. 43, No. 1, pp. 39-44, 1996
Copyright ~) 1996 Elsevier Science Ltd
Printed in Great Britain. All rights reserved
0031-9422/96 $15.00 + 0.00
MAJOR PROTEINASE HYDROLYSING GLIAD1N DURING WHEAT
GERMINATION
ANDREABOTrAR1,ANTONELLA CAPOCCHI, DEBORA FONTANINI and LUCIANO GALLESCHI*
Department of Botanical Sciences, University of Pisa, Via L. Ghini 5, 56126-Pisa, Italy
(Received in revisedform 30 January 1996)
Key Word Index--Triticum durum; Gramineae; wheat; purification and characterization; pro-
teinases; cysteine proteinase.
Abstract--A proteinase, representing the bulk of the enzyme activity for the hydrolysis of gliadin, was extracted
from endosperms isolated from germinated seeds (four days) and was purified by ion-exchange chromatography
and preparative isoelectric focusing. The optimal pH for gliadin hydrolysis was 4.25. The M r, determined by
sodium dodecyl sulphate-polyacrylamide gel electrophoresis, was 30 000; the isoelectric point was 4.5. The
enzyme activity was totally inhibited by E-64 and cystatin, while inhibitors of other classes of proteinases were
barely effective or ineffective. The activity was stimulated by sulphhydryl compounds. The proteinase hydrolysed
to small peptides the gliadins from durum and soft wheat seeds. Other protein substrates were weakly degraded or
not degraded. The proteinase appears to belong to the cysteine class and to play a key role in the initial
mobilization of the main reserve protein in the starchy endosperm. Copyright © 1996 Elsevier Science Ltd
INTRODUCTION
The mobilization of seed storage proteins represents
one of the most important post-germinative events in
growth and seedling development. In fact, hydrolysis of
these reserves to soluble products supports seedling
growth prior to the development of autotrophy.
In a wheat grain most of the nitrogen is found in the
storage proteins, known as gliadins, which are soluble
in alcohol solution. They are deposited in the starchy
endosperm where they constitute up to 50% of the total
proteins [1]. It is known that cereal storage proteins are
degraded during early seedling growth by endo- and
exopeptidases, although the cooperative role of those
enzymes in protein breakdown still remains to be
elucidated in many of the plant materials studied [2].
Carboxypeptidases and aspartic proteinases are already
present in dry wheat seeds [3-5]. In in vitro experi-
ments they showed a weak hydrolytic action on the
gliadin, which increased by about 25% when both
enzymes were added simultaneously [4]. However, the
in vitro hydrolysis of the gliadin started after the first
day of germination of wheat seeds, in association with
the appearance of proteinase activity against this stor-
age protein [6]. Preliminary studies conducted with
crude extracts from germinated (4 days) wheat seeds in
the presence of proteinase inhibitors showed that about
89% of the proteinase activity responsible for the
increase in gliadin hydrolysis was of cysteine-type (EC
3.4.22) [6]. Although cysteine proteinases have been
*Author to whom correspondence should be addressed.
previously found in germinating wheat seeds [4, 7],
they have not been isolated to a sufficiently high degree
of purification for the study of their characteristics and
physiological role. The present work describes the
isolation and the partial characterization of a cysteine
proteinase that is probably involved in the initial
cleavage of gliadin to polypeptides in germinated
Triticum durum seeds.
RESULTS AND DISCUSSION
Purification of the proteinase
The purification of the proteinase from endosperms
of germinating durum wheat is summarized in Table 1.
Endosperms of 4-day-old seedlings were used as the
starting material for the purification of the enzyme,
since our previous work [6] showed that gliadin mobili-
zation and gliadin activity were most prominent during
this period. The fractionation of the proteins on DEAE-
cellulose allowed us to eliminate most of the con-
taminating proteins. Initial attempts to fractionate the
crude protein extract with the Rotofor cell, either by
using ampholytes with a broad pH range (3-10), or
with a narrow pH range (4-6) were unsuccessful. In
both cases there was considerable precipitation, which
started in the end compartments and then spread inward
during the run. Use of glycerol and pre-focus of the pH
gradient before loading the extract did not alleviate the
problem. On the other hand, protein precipitation did
not happen when samples fractionated on DEAE-cellu-
lose were applied to the Rotofor cell. However, the
39