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Postharvest Biology and Technology of Fruits, Vegetables, and Flowers

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

detachment <strong>of</strong> leaves in the dark accelerates senescence <strong>and</strong> serves as a means <strong>of</strong> coordinating<br />

the senescence process between leaves, its use as a model system for natural age-related<br />

senescence should be approached with caution. Following the identification <strong>of</strong> three<br />

cDNAs that showed increased expression in dark-treated leaves, experiments revealed that<br />

only one <strong>of</strong> the three showed enhanced expression during natural senescence (Becker <strong>and</strong><br />

Apel, 1993). Only four <strong>of</strong> seven genes identified from Arabidopsis as dark-induced also<br />

showed enhanced expression in naturally senescing leaves (Park et al., 1998).<br />

There are many common molecular events occurring among senescing tissues. The<br />

degradation <strong>and</strong> remobilization <strong>of</strong> cellular constituents is predominant during senescence<br />

<strong>and</strong>, correspondingly, the activities <strong>of</strong> hydrolytic enzymes <strong>and</strong> their mRNAs increase. A<br />

number <strong>of</strong> SR genes have also been identified that encode products with homology to PR<br />

proteins. While it is not known what the role <strong>of</strong> these proteins is in senescence, it appears<br />

that they may serve a protective role similar to their role during the defense response.<br />

These patterns <strong>of</strong> expression indicate that throughout plant development, common molecular<br />

mechanisms are regulated by the same genes in multiple tissues. A few <strong>of</strong> these genes have<br />

been identified as having leaf, flower, or fruit senescence-specific expression. Of these<br />

genes, many encode different is<strong>of</strong>orms <strong>of</strong> the same enzyme, which may be differentially<br />

regulated within the plant organs. Similar to fruit ripening, the developmental regulation <strong>of</strong><br />

germination <strong>and</strong> senescence also share common molecular mechanisms. This is especially<br />

evident when investigating the expression <strong>of</strong> genes involved in protein <strong>and</strong> lipid degradation<br />

<strong>and</strong> remobilization.<br />

While differential cDNA screening, differential display <strong>and</strong> cDNA subtraction have<br />

identified a number <strong>of</strong> senescence-related genes, the expression <strong>of</strong> most genes has not been<br />

investigated in flowers, leaves, <strong>and</strong> fruits. The use <strong>of</strong> enhancer trap lines in Arabidopsis<br />

has resulted in the identification <strong>of</strong> over 100 lines that have reporter gene expression in<br />

senescing, but not in nonsenescing tissues (He et al., 2001). This technology starts to<br />

reveal the complexity <strong>of</strong> the network <strong>of</strong> senescence-regulated pathways, <strong>and</strong> will allow for<br />

the identification <strong>of</strong> many additional SR genes. The identification <strong>of</strong> senescence-specific<br />

promoter elements (Noh <strong>and</strong> Amasino, 1999a) <strong>and</strong> the generation <strong>of</strong> mutants <strong>and</strong> transgenic<br />

plants will help us to better underst<strong>and</strong> the regulation <strong>of</strong> SR genes during senescence.<br />

DNA microarrays will allow temporal <strong>and</strong> spatial expression patterns to be determined for<br />

hundreds <strong>of</strong> genes involved in senescence. These technologies will lead to an increased<br />

underst<strong>and</strong>ing <strong>of</strong> the initiation <strong>and</strong> execution <strong>of</strong> senescence, which will allow us to increase<br />

vase life <strong>and</strong> horticultural performance <strong>of</strong> ornamentals, increase yield in agronomic crops,<br />

<strong>and</strong> decrease postharvest losses <strong>of</strong> fruits <strong>and</strong> vegetables.<br />

4.12 Sugar status <strong>and</strong> cut flower senescence<br />

Sugar solutions are well known for their ability to improve postharvest quality <strong>and</strong> extend the<br />

vase life <strong>of</strong> cut flowers, although the hypothesis <strong>of</strong> a sole sugar starvation or sugar accumulation<br />

signal in inducing petal senescence has not been validated (van Doorn, 2004). Signals<br />

from the lack <strong>of</strong>, the abundance <strong>of</strong>, or the metabolism <strong>of</strong> sugars (i.e., sugar signals) probably<br />

form part <strong>of</strong> a complex array <strong>of</strong> exogenous <strong>and</strong> endogenous signals that initiate senescence,<br />

<strong>and</strong> there is evidence that in carnation petals sucrose decreases ethylene responsiveness<br />

(Verlinden <strong>and</strong> Garcia, 2004), <strong>and</strong> complex interactions occur between sugar- <strong>and</strong> ethylenesignaling<br />

mechanisms that are tissue dependent (Iordachescu <strong>and</strong> Verlinden, 2005). In

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