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Postharvest Biology and Technology of Fruits, Vegetables, and Flowers

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66 POSTHARVEST BIOLOGY & TECHNOLOGY OF FRUITS, VEGETABLES, & FLOWERS<br />

(Lycopodium cernuum) (Paull <strong>and</strong> Chantrachit, 2001). Jasmonic acid <strong>and</strong> methyl jasmonate,<br />

natural plant growth regulators, are involved in plant defense responses, exhibit direct<br />

antifungal activity as well as increase numerous antifungal compounds in plant tissues<br />

when applied exogenously. They also activate many inducible genes leading to the synthesis<br />

<strong>of</strong> secondary plant products that function as antimicrobial compounds (Meir et al.,<br />

1998).<br />

4.10 Identification <strong>and</strong> classification <strong>of</strong> senescence-related genes<br />

Recent molecular studies have confirmed that senescence <strong>and</strong> ripening are accompanied by<br />

changes in gene expression. Utilizing differential screening <strong>and</strong> subtractive hybridization<br />

techniques, a number <strong>of</strong> cDNAs that are upregulated during senescence have been cloned.<br />

Genes that exhibit enhanced expression during senescence have been cloned from the leaves<br />

<strong>of</strong> Arabidopsis, asparagus, Brassica napus, barley, maize, radish, <strong>and</strong> tomato (Smart, 1994;<br />

Buchanan-Wollaston, 1997; Nam, 1997; Weaver et al., 1997; Quirino et al., 2000). Differential<br />

screening <strong>of</strong> senescence petal cDNA libraries <strong>and</strong> PCR-based differential display<br />

techniques have been utilized to identify genes that are upregulated during senescence <strong>of</strong><br />

carnation <strong>and</strong> daylily flowers (Woodson et al., 1993; Woodson, 1994; Valpuesta et al., 1995;<br />

Guerrero et al., 1998; Panavas et al., 1999).<br />

Most <strong>of</strong> the genes that have been identified as senescence-related are expressed at basal<br />

levels in nonsenescing tissues (green leaves <strong>and</strong> young flowers) <strong>and</strong> increase in abundance<br />

during senescence. A smaller number <strong>of</strong> SR genes are only detectable in senescing tissues<br />

<strong>and</strong> represent senescence-specific genes. An even smaller set <strong>of</strong> genes have been identified<br />

that have high levels <strong>of</strong> expression early in development, decreased expression in young<br />

maturing tissue, <strong>and</strong> increased expression at the onset <strong>of</strong> senescence. Genes that fit within<br />

this class have only been identified in vegetative tissues <strong>and</strong> represent genes that have a<br />

similar role in multiple stages <strong>of</strong> development like germination <strong>and</strong> senescence (Lohman<br />

et al., 1994; Smart, 1994; Buchanan-Wollaston, 1997).<br />

Weaver et al. (1998) shows the patterns <strong>of</strong> expression <strong>of</strong> a selected group <strong>of</strong> SAGs<br />

(senescence-associated genes) during age-related senescence <strong>of</strong> Arabidopsis leaves. Most<br />

<strong>of</strong> these genes exhibit basal levels <strong>of</strong> expression in green nonsenescing tissues. Within this<br />

broad classification, genes are differentially regulated, with some increasing in abundance<br />

gradually as the leaf matures <strong>and</strong> others increasing more abruptly at various stages <strong>of</strong> leaf<br />

development. Only SAG12 <strong>and</strong> SAG13 show senescence-specific expression. Among the<br />

senescence-specific genes, SAG13 is detected before any visible signs <strong>of</strong> leaf senescence<br />

<strong>and</strong>, as such, may be responsible for initiation <strong>of</strong> the senescence process, while SAG12 is<br />

expressed after the leaf is visibly yellowing.<br />

Many <strong>of</strong> the genes that have been identified as senescence-related are identified from<br />

a particular plant organ, <strong>and</strong> it is not known whether they are expressed in other senescing<br />

organs or during other developmental processes. The expression <strong>of</strong> a number <strong>of</strong> SAGs was<br />

investigated in roots, stems, flower buds, <strong>and</strong> mature flowers <strong>of</strong> Arabidopsis (Quirino et al.,<br />

1999). Expression <strong>of</strong> SAG12, SAG13, SAG25, SAG26, <strong>and</strong> SAG29 was not detected in any<br />

nonsenescing tissues but was detected in both senescing flowers <strong>and</strong> leaves, indicating a<br />

common molecular regulation <strong>of</strong> senescence in vegetative <strong>and</strong> floral tissue. Some <strong>of</strong> the<br />

SAGs show low levels <strong>of</strong> expression in multiple tissues with upregulation in senescing

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