Fruit flavors and aromas
Fruit flavor is a complex trait determined by the mixture and balance of sugars,
acids and a large number of aldehyde, ketone and alcohol volatiles (Buttery,
1993; Baldwin
et al., 2000). Lipoxygenases are key enzymes in fatty acid
metabolism, producing the hexanal aldehyde and alcohols that contribute
substantially to the volatile tomato aromas and flavors. At least three
lipoxygenase genes (
LOX) are expressed during tomato fruit ripening (Ferrie
et al., 1994; Kausch and Handa, 1997; Griffiths
et al., 1999a). The expression of
the
LOX genes in ripening tomato fruit is regulated by both ethylene and
developmental factors. In order to modify the C6 aldehyde and alcohol
composition of tomatoes, antisense transgenic tomatoes were developed using a
conserved region of lipoxygenases to potentially suppress expression of all LOX
activities (Griffiths
et al., 1999b). In ripe fruit of these transgenic plants, the
expression of
tomloxA and
tomloxB, but not
tomloxC, was substantially reduced.
However, no changes in the flavor and aroma constituents resulted, suggesting
that regulation of their biosynthesis may be complex or that
tomloxC expression
was sufficient for these pathways to proceed unimpeded.
Another approach to altering the flavor composition of tomato fruit was
engineered by the
expression of a yeast Δ9 desaturase gene (Wang
et al., 1996),
resulting in changes in the fatty acid content of fruit. Concentrations of palmitoleic acid, 9,12-hexadienoic acid, and linoleic acid increased and
concentrations of palmitic acid and stearic acid were reduced. Concentrations
of flavor compounds derived from these fatty acids were changed and resistance
to powdery mildew was enhanced (Wan
et al., 1998). Another approach to
changing the composition of flavor aldehydes in tomato fruit was taken by
altering the expression of alcohol dehydrogenase
adhII (
2) (Speirs
et al., 1998;
Prestage
et al., 1999). The amount of Z-3-hexanol decreased and the amount of
3-methylbutanal increased in plants with reduced expression of
adhII. Fruit from
plants with increased expression of
adhII had a more intense ‘ripe fruit’ flavor
and had increased amounts of hexanol and Z-3-hexanol.