An ipt gene under control of the senescence-specific SAG12 promoter from Arabidopsis (P SAG12 -IPT) significantly delayed developmental and postharvest leaf senescence in mature heads of transgenic lettuce (Lactuca sativa L. cv Evola) homozygous for the transgene. Apart from retardation of leaf senescence, mature, 60-d-old plants exhibited normal morphology with no significant differences in head diameter or fresh weight of leaves and roots. Induction of senescence by nitrogen starvation rapidly reduced total nitrogen, nitrate, and growth of transgenic and azygous (control) plants, but chlorophyll was retained in the lower (outer) leaves of transgenic plants. Harvested P SAG12 -IPT heads also retained chlorophyll in their lower leaves. During later development (bolting and preflowering) of transgenic plants, the decrease in chlorophyll, total protein, and Rubisco content in leaves was abolished, resulting in a uniform distribution of these components throughout the plants. Homozygous P SAG12 -IPT lettuce plants showed a slight delay in bolting (4-6 d), a severe delay in flowering (4-8 weeks), and premature senescence of their upper leaves. These changes correlated with significantly elevated concentrations of cytokinin and hexoses in the upper leaves of transgenic plants during later stages of development, implicating a relationship between cytokinin and hexose concentrations in senescence.Leaf senescence is a type of programmed cell death characterized by loss of chlorophyll, lipids, total protein, and RNA (Smart, 1994; Gan and Amasino, 1997). This process is believed to be an evolutionarily acquired, active genetic trait that contributes to plant fitness, for example, by remobilizing nutrients from vegetative tissues to reproductive organs (Oh et al., 1997). The biological importance and potential for improvement of crop characteristics, particularly plant productivity and postharvest storage, have prompted extensive physiological, molecular, and genetic analyses of leaf senescence (Nam, 1997). Senescence can be induced by environmental stress, such as low light intensity, nutrient deficiency, pathogen attack, drought, waterlogging, and detachment from the plant. Endogenous factors, including leaf age and reproductive development, also trigger senescence (Smart, 1994; Gan and Amasino, 1997).Plant growth regulators, including auxins, gibberellins, ethylene, abscisic acid, and cytokinins, are believed to play major roles in regulating senescence. Attention has focused on cytokinins that are key components of plant senescence (Van Staden et al., 1988;Singh et al., 1992;Gan and Amasino, 1996, 1997; Buchanan-Wollaston, 1997;Nam, 1997). These compounds have been implicated in several aspects of plant development and are thought to be synthesized mainly in the roots and transported to the shoots via the xylem (Gan and Amasino, 1996). Three main approaches have been used to study the effect of cytokinins in plant senecence, namely exogenous application of cytokinin solutions, measurement of endogenous cytokinins duri...
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