Sugar Accumulation in Grape Berries

Topics: Fruit, Tomato, Grape Pages: 9 (7304 words) Published: October 30, 2014
Plant Physiol. (1 996) 1 1 1 : 275-283

Sugar Accumulation in Grape Berries
Cloning of Two Putative Vacuolar lnvertase cDNAs and Their Expression in Grapevine Tissues
Christopher Davies* and Simon

P. Robinson

Cooperative Research Centre for Viticulture, P.O. Box 145, Glen Osmond, South Australia, Australia, and Commonwealth Scientific and industrial Research Organization, Division of Horticulture, G.P.O. Box 350, Adelaide, South Australia 5001, Australia

The growth pattern of developing grape berries (Vifis
vinifera L.) can be described as a double sigmoidal curve
with an initial rapid increase in size followed by a lag
period during which berry volume does not increase. The
lag period is followed by a second phase of growth during
which ripening occurs, and viticulturists use the French
word véraison to describe the inception of berry ripening.
During ripening, the increase in volume is accompanied by
an increase in berry softness, accumulation of hexoses in
the berries, and a decrease in the leve1 of malic and tartaric acids, and in red grape varieties the skin becomes colored
due to the accumulation of anthocyanins (Coombe, 1992).
The accumulation of sugar in the form of Glc and Fru
within the vacuole is one of the main features of the ripening process in grape berries and is a major commercial consideration for the grape grower, winemaker, and dried
fruit producer. Sugar accumulation in climacteric fruit has
received considerable attention, but little is known about

this process in nonclimacteric fruit such as grapes. In
grapevines, SUCproduced as a result of photosynthesis in
the leaf is transported via the phloem to the berry (Swanson and Elshishiny, 1958), where it is cleaved to Glc and Fru, which accumulate in roughly equal amounts (Kliewer,
1965).The accumulation of Fru and Glc commences only at
véraison and continues throughout ripening.
Invertase (P-fructosidase; EC catalyzes the conversion of Suc to its monosaccharide constituents Glc and Fru and is probably responsible for this reaction in grape
berries as ít is in other species that store sugar in this way. In tomato, species that accumulate high levels of Glc and
Fru, for example Lycopersicon esculentum Mill., have high
levels of invertase activity and store little Suc (Yelle et al., 1991; Stommel, 1992). In contrast, accumulators of high
levels of Suc (and low levels of Glc and Fru), such as
Lycopersicon peruvianum and Lycopersicon chmielewskii, have
low levels of vacuolar invertase. Further evidence for a role for vacuolar invertase in controlling the composition of
sugars in tomato fruit has recently been provided by antisense (Ohyama et al., 1995) and breeding experiments (Chetelat et al., 1995).
A number of distinct invertases have been characterized
in plants. Most reports deal with either soluble invertases, which have acidic pI values and acid pH optima and are
located in the vacuole, or with insoluble acid invertases,
which have basic pI values and are bound to the cell wall
(Sturm and Chrispeels, 1990). It is thought that the vacuolar invertases are likely to be important in the regulation of hexose levels in certain tissues, e.g. in fruit tissue (Lingle and Dunlop, 1987), and in the utilization of Suc stored in

the vacuoles (Leigh et al., 1979). The cell-wall forms are
associated with rapidly growing tissues (Eschrich, 1980),
are induced by wounding and pathogenic attack (Sturm
and Chrispeels, 1990), and have been implicated in phloem
unloading and source / sink regulation (Eschrich, 1980;
Roitsch et al., 1995).
Hawker (1969a) found that invertase enzyme activity in
Sultana berries increased immediately after flowering and
that the activity peaked 6 to 7 weeks later, at véraison, when

* Corresponding author; e-mail; fax 618-303-8601.

Abbreviations: "Brix, refractive index measure of total soluble solids; RACE, rapid amplification of cDNA ends....

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