Phytochemistry, Vol. 27, No. 2, pp. 323-327, 1988. 0031 9422/88$3.00+0.00
Printed in Great Britain. © 1988 PergamonJournals Ltd.
PURIFICATION AND CHARACTERIZATION OF AVOCADO LIPOXYGENASE
LIONEL MARCUS, DOV PRUSKY~ and BENJAMINJACOBY3~
Department of Fruit and Vegetable Storage, Agricultural Research Organization, The Volcani Center, P.O. Box 6, Bet Dagan 50250,
Israel; :~Department of Agricultural Botany, Faculty of Agriculture, Hebrew University of Jerusalem, P.O.B. 12, Rehovot, 76100,
Israel.
(Revised received 9 July 1987)
Key Word lndex--Persea americana; Lauraceae; lipoxygenase; purification; affinity chromatography
Abstract--Lipoxygenase (E.C. 1.13.1.13) from the avocado cultivar 'Fuerte' was purified to near homogeniety by affinity
chromatography. The enzyme was extracted in potassium phosphate buffer at pH 7.2 in the presence of 2% Triton
X-100. Triton was removed from the homogenate by adsorption on 25.0-350 mesh activated charcoal. Lipoxygenase
was partially purified (seven-fold) by 66% acetone precipitation from a 20% acetone supernatant. The precipitate was
dissolved in a potassium phosphate buffer at pH 7.2 and loaded on an affinity chromatography column. This single-step
chromatographic purification yielded a single lipoxygenase activity peak. The total activity yield of the purification
procedure was ca 65% and the degree of enrichment ca 35-fold. The M, determined by gel filtration and by
electrophoresis, was 74 000. Optimum enzyme activity was found at 36 ° and pH 7.1. The energy of activation, amino
acid composition, isoelectric point, kinetic parameters and inhibitory effect of epicatechin were studied. The enzyme
was found to obey Michaelis-Menten kinetics. The Km of the avocado lipoxygenase for linoleate was 7.2 × 10-2 mM
and the VmaxWaS432 #mol/hr/mg. Epicatechin acted as a competitive inhibitor with a Ks of 9.0 × 10-s mM.
INTRODUCTION
In the presence of molecular oxygen, lipoxygenase (linol-
eate: O: oxidoreductase E.C. 1.13.1.13.) catalyses the
oxidation of C18-unsaturated fatty acids with a cis,cis-1,4-
pentadiene group to give hydroperoxide [1]. Several
conventional and non-conventional methods for the
isolation of this enzyme from a variety of plant and animal
tissues have been described [2-7]. Affinity chromato-
graphy purification was reported by Grossman et al. [8]
for soybean lipoxygenase, and recently by Cohen et al. [9]
for that from pea. Attention has been drawn recently to
avocado lipoxygenase because of its suggested physio-
logical role in the breakdown of the antifungal 1-acetoxy-
2-hydroxy-4-oxo-heneicosa-12,15-diene involved in the
resistance of unripe avocado fruit to fungal infection
[10-12]. The present paper describes conditions for the
purification of avocado lipoxygenase by affinity chroma-
tography and the catalytic properties of this enzyme.
RESULTS
Cold acetone (4o ) was added to the crude enzyme
preparation to a concentration of 20% v/v and the
precipitate was removed after 1 hr by centrifugation at
20000 g for 10 min. The supernatant was adjusted with
Contribution from the Agricultural Research Organization,
The Volcani Center, Bet Dagan, Israel. No. 1941-E, 1987 series.
tAuthor to whom correspondence should be addressed.
additional acetone to 66% v/v and after 2 hr, again
centrifuged at 20000 g for 10 rain. The precipitate was
dissolved in 200ml of 20mM potassium phosphate
buffer, pH 7.2, and the insoluble fraction separated by
centrifugation. A 10 ml aliquot of the supernatant was
applied to the column of cross-linked Sepharose-
hexamethyl-linoleate (10 x 0.7 cm) and eluted with the
same buffer at a flow rate of 18 ml/hr. The eluent was
collected in 6 ml fractions. Protein from the column was
monitored by absorbance at 280 nm.
The elution profile of the crude enzyme preparation
from the cross-linked linoleate column showed that most
of the inactive protein was eluted with the buffer (Fig. 1).
The lipoxygenase activity remained tightly bound to the
column even after prolonged washing with the buffer. The
bound enzyme was released from the linoleate column
with a linear gradient of sodium chloride, 0.0-0.5 M, in
the same buffer. Elution commenced when the NaCI
concentration approached 0.2 M. Fractions containing
lipoxygenase activity were pooled, dialysed against
20 mM potassium phosphate buffer, pH 7.2, for 24 hr,
lyophilized and redissolved in 2 ml of distilled water. This
solution was used as a purified enzyme source for further
study.
The yields, degrees of enrichment and specific activity
of the avocado lipoxygenase for a typical purification
procedure are summarized in Table 1. In this experiment
the specific activity of the purified enzyme was 18.35 pmol
O2/min/mg protein (0.3 × 10-6 Kat/mg) and the enrich-
ment was 35-fold. The purity of the enzyme was evidenced
by the single protein band obtained by SDS gel electro-
phoresis. Cross-linking with ethylene diamine was also
explored, but purification by this column was not ade-
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