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Postharvest Biology and Technology of Fruits, Vegetables, and Flowers

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

Progressive stimulus-response mode<br />

Ethylene<br />

Ca<br />

ETR 2<br />

P<br />

L<br />

D<br />

MT<br />

P<br />

L<br />

D<br />

Ca<br />

CTR1<br />

MAPK<br />

cascade<br />

PI kinase<br />

P<br />

L<br />

D<br />

PLD<br />

PA<br />

DG<br />

Gene<br />

expression<br />

PL<br />

FFA<br />

FFOX<br />

H<br />

Ca<br />

X<br />

X<br />

Metabolon<br />

Ca<br />

Nucleus<br />

PI kinase<br />

PLD<br />

PL<br />

FFA<br />

PA<br />

DG<br />

Ca<br />

Ethylene<br />

ETR 1<br />

Transient stimulus-<br />

response mode<br />

Fig. 9.21 A model describing potential events involved in the ethylene signal transduction pathway indicating<br />

the role <strong>of</strong> PLD. Ethylene binding to the receptor is proposed to activate a PI kinase that in turn results in the<br />

generation <strong>of</strong> a voltage difference across the membrane. Voltage-sensitive calcium channels may open in response to<br />

the voltage difference, increasing the cytosolic calcium level. PLD can bind to the membrane at the anionic domains<br />

created by PI-kinase activity or in response to the increased cytosolic calcium levels. In the progressive stimulusresponse<br />

model, the hormonal stimulation continues ultimately inactivating calcium <strong>and</strong> proton pumps (H-X,<br />

Ca-X). Excessive cytosolic calcium would ultimately cause senescence. In the transient stimulus-response model,<br />

cytosolic calcium level is maintained at homeostatic levels. Lipid catabolites are recycled back to phospholipids.<br />

ETR1 <strong>and</strong> ETR2 are designation <strong>of</strong> receptors with potentially distinct function. CTR-1 is a protein kinase that acts<br />

downstream from the receptor. PLD could exist as a metabolon (left bottom) in combination with phosphatidate<br />

phosphatase, calmodulin, <strong>and</strong> lipolytic acyl hydrolase, or even bound to microtubules (MT) (DG, diacylglycerol;<br />

FFA, free fatty acid; FFOX, fatty acid oxidation products; PA, phosphatidic acid; PL, phospholipid).<br />

is promoted by calcium. Under normal conditions <strong>of</strong> growth <strong>and</strong> development, this switch<br />

would be tightly regulated (transient stimulus-response model) due to cross talk with other<br />

signals (Lohrmann <strong>and</strong> Harter, 2002). After the initiation <strong>of</strong> a programmed senescence event<br />

(progressive stimulus-response model), such tight regulation could be lost <strong>and</strong> would eventually<br />

lead to high cytosolic calcium <strong>and</strong> lowered cytosolic pH, as the calcium ATPase <strong>and</strong><br />

proton ATPase are inhibited, respectively (Paliyath <strong>and</strong> Thompson, 1988; Paliyath et al.,

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