Acetyl-CoA: Alcohol Acetyltransferase Activity and Aroma
Formation in Ripening Melon Fruits
Moshe Shalit, Nurit Katzir, Yaakov Tadmor, Olga Larkov, Yosef Burger, Fernond Shalekhet,
Elena Lastochkin, Uzi Ravid, Orit Amar, Menahem Edelstein, Zvi Karchi, and
Efraim Lewinsohn*
Department of Vegetable Crops, Agricultural Research Organization (ARO), Newe Ya’ar Research Center,
P.O. Box 1021, Ramat Yishay 30-095, Israel
Melon varieties (Cucumis melo L.) differ in a range of physical and chemical attributes. Sweetness
and aroma are two of the most important factors in fruit quality and consumer preference. Volatile
acetates are major components of the headspace of ripening cv. Arava fruits, a commercially
important climacteric melon. In contrast, volatile aldehydes and alcohols are most abundant in cv.
Rochet fruits, a nonclimacteric melon. The formation of volatile acetates is catalyzed by alcohol
acetyltransferases (AAT), which utilize acetyl-CoA to acetylate several alcohols. Cell-free extract
derived from Arava ripe melons exhibited substantial levels of AAT activity with a variety of alcohol
substrates, whereas similar extracts derived from Rochet ripe melons had negligible activity. The
levels of AAT activity in unripe Arava melons were also low but steadily increased during ripening.
In contrast, similar extracts from Rochet fruits displayed low AAT activity during all stages of
maturation. In addition, the benzyl- and 2-phenylethyl-dependent AAT activity levels seem well
correlated with the total soluble solid content in Arava fruits.
Keywords: Aroma; volatile acetates; biosynthesis; alcohol acetyltransferase; melons (Cucumis
melo. L)
INTRODUCTION
Contemporary melon cultivars differ in shape, size,
color, surface netting, sweetness, flavor, and storability
(1-3) and can be divided into two groups, climacteric
and nonclimacteric, according to their ripening patterns.
The climacteric mode of ripening is characterized by a
significant increase in the levels of respiration and
release of ethylene, triggering profound and rapid
changes in a range of attributes, such as flavor (sweet-
ness and aroma), as well as in other physical parameters
such as firmness of the fruits and a typical slip area
produced around the peduncle in climacteric melons.
Some of the climacteric varieties of melons are consid-
ered to be highly aromatic [such as Galia, Charentais,
and Ananas (Cucumis melo var. reticulatus)], whereas
the nonclimacteric varieties are normally considered to
be less aromatic, that is, Casaba-type melons such as
Rochet (Cucumis melo var. inodorus).
Two of the most important parameters that determine
the quality and consumer preferences of the fruits are
their sweetness and aroma properties. The aroma of
fruits is ordinarily composed of complex mixtures of
volatile compounds present in the headspace (4). Vola-
tile aldehydes, alcohols, and especially the large quanti-
ties of esters present in their headspace are likely to be
the key contributors to the unique aromas of melons (3,
5-11). Although the aromas of some of the individual
volatile compounds often resemble melon or fruity notes,
the peculiar aroma of melons cannot be related to any
single compound.
Many of the volatile aldehydes present in the head-
space of fruits are formed by degradation of fatty acids
(12), whereas the respective alcohols are derived from
aldehydes by the action of alcohol dehydrogenases or
as a result of degradation of amino acids (13).
Volatile esters, important contributors to the aroma
of many fruits, are formed by esterification of alcohols
and carboxylic acids, normally utilizing a CoA ester as
the acyl donor. The ability to esterify alcohols has been
previously noted in many melon cultivars by incubating
fruit slices with isobutyl alcohol (14). The nonaromatic
inodorus types had low esterification potential, and
other cucurbits such as cucumber (Cucumis sativus L.),
squash (Cucurbita maxima Duch.), and watermelon
(Citrullus lanatus Matsumu.) lacked the ability to
esterify isobutyl alcohol (14). The mechanism of ester
formation has been studied in microorganisms, where
a group of enzymes termed alcohol acetyltransferases
(AAT) have been identified (15, 16). AAT catalyze the
transfer of an acetyl moiety from acetyl-CoA into the
corresponding alcohol, forming an ester and free CoA
(Figure 1).
The first attempt to identify AAT activity in fruits
was carried out in banana slices (17). In this study,
isoamyl acetate was synthesized from isoamyl alcohol
and acetyl-CoA in partially purified cell-free protein
extracts. AAT activities have been also detected in apple
(18), strawberry (19, 20), and melon fruits (14, 21, 22).
In all cases, an alcohol is acetylated; the acyl donor is
acetyl-CoA. The partially purified AAT from oriental
sweet melons mainly esterified isobutyl alcohol to form
isobutyl acetate (21). Other alcohols were also esterified,
but benzyl alcohol, the putative precursor of benzyl
* Author to whom correspondence should be addressed
(fax 972-4-983-6936; telephone 972-4-953-9552; e-mail
twefraim@netvision.net.il).
794 J. Agric. Food Chem. 2001, 49, 794-799
10.1021/jf001075p CCC: $20.00 © 2001 American Chemical Society
Published on Web 01/06/2001