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Postharvest Biology and Technology of Fruits, Vegetables, and Flowers

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Chapter 15<br />

Polyamines <strong>and</strong> Regulation <strong>of</strong> Ripening<br />

<strong>and</strong> Senescence<br />

Savithri Nambeesan, Avtar K. H<strong>and</strong>a, <strong>and</strong> Autar K. Mattoo<br />

15.1 Introduction<br />

Polyamines (PAs) are small polycationic, biogenic amines that have pr<strong>of</strong>ound effects on<br />

growth, development, <strong>and</strong> senescence in eukaryotic cells (Galston <strong>and</strong> Kaur-Sawhney, 1995;<br />

Cassol <strong>and</strong> Mattoo, 2003; Casero <strong>and</strong> Marton, 2007). Diamine putrescine (Put) is a major<br />

PA in plants <strong>and</strong> a precursor for triamine spermidine (Spd) <strong>and</strong> tetraamine spermine (Spm).<br />

PAs influence many biochemical <strong>and</strong> physiological processes such as cell division, cell<br />

elongation, flowering, fruit set <strong>and</strong> development, fruit ripening, <strong>and</strong> senescence (Evans <strong>and</strong><br />

Malmberg, 1989; Galston <strong>and</strong> Kaur-Sawhney, 1990; Bouchereau et al., 1999). Many <strong>of</strong><br />

these processes have direct implications on various aspects <strong>of</strong> postharvest biology <strong>of</strong> fruit<br />

<strong>and</strong> vegetable crops including quality, storage life, senescence, chilling <strong>and</strong> other stresses,<br />

<strong>and</strong> disease development (Valero et al., 2002a). Significant information about PA action has<br />

emerged from indirect studies employing pharmacological levels <strong>of</strong> different PAs <strong>and</strong> as<br />

well as their biosynthetic inhibitors. In recent years, reverse genetics has begun to provide<br />

direct evidence for involvement <strong>of</strong> PAs in determining fruit quality, ripening, <strong>and</strong> processing<br />

attributes (Tiburcio et al., 1997; Martin-Tanguy, 2001; Mehta et al., 2002; Mattoo et al.,<br />

2006; Srivastava et al., 2007). Molecular aspects <strong>of</strong> PA action have also begun to emerge,<br />

especially from nonplant systems (Igarashi <strong>and</strong> Kashiwagi, 2006; Wallace <strong>and</strong> Niiranen,<br />

2007). PAs have been shown to bind to membranes, nucleic acids, <strong>and</strong> other macromolecules,<br />

<strong>and</strong> have been implicated in stabilizing chromatin conformation, regulating ion channels,<br />

scavenging free radicals, <strong>and</strong> regulating gene expression (Casero <strong>and</strong> Marton, 2007; Srivastava<br />

et al., 2007). Since maintenance <strong>of</strong> membrane stability <strong>and</strong> homeostasis is essential<br />

for many cellular, physiological, <strong>and</strong> biochemical processes, it is likely that PAs can have<br />

a pivotal role in extending postharvest shelf life <strong>of</strong> fruits <strong>and</strong> vegetables. The focus <strong>of</strong> this<br />

chapter is on recent progress made in underst<strong>and</strong>ing the roles <strong>of</strong> PAs in plants with special<br />

emphasis on postharvest biological processes.<br />

15.2 Regulation <strong>of</strong> polyamine metabolism in plants<br />

15.2.1 Polyamine biosynthesis<br />

The cellular concentration <strong>of</strong> different PAs ranges from micromolar to millimolar levels<br />

in plants <strong>and</strong> is highly regulated during growth <strong>and</strong> development (Imai et al., 2004;<br />

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