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THE PLANT CELL, Vol 6, Issue 4 521-530, Copyright © 1994 by American Society of Plant Biologists
The Never Ripe Mutation Blocks Ethylene Perception in Tomato
M. B. Lanahan, H. C. Yen, J. J. Giovannoni and H. J. Klee
The Agricultural Group, Monsanto Company, 700 Chesterfield Village Parkway, Chesterfield, Missouri 63198
Seedlings of tomato fruit ripening mutants were screened for their ability
to respond to ethylene. Ethylene induced the triple response in etiolated
hypocotyls of all tomato ripening mutants tested except for one, Never ripe
(Nr). Our results indicated that the lack of ripening in this mutant is
caused by ethylene insensitivity. Segregation analysis indicated that
Nr-associated ethylene insensitivity is a single codominant trait and is
pleiotropic, blocking senescence and abscission of flowers and the
epinastic response of petioles. In normal tomato flowers, petal abscission
and senescence occur 4 to 5 days after the flower opens and precede fruit
expansion. If fertilization does not occur, pedicel abscission occurs 5 to
8 days after petal senescence. If unfertilized, Nr flowers remained
attached to the plant indefinitely, and petals remained viable and turgid
more than four times longer than their normal counterparts. Fruit
development in Nr plants was not preceded by petal senescence; petals and
anthers remained attached until they were physically displaced by the
expanding ovary. Analysis of engineered 1-aminocyclopropane-1-carboxylate
(ACC) synthase-overexpressing plants indicated that they are phenotypic
opposites of Nr plants. Constitutive expression of ACC synthase in tomato
plants resulted in high rates of ethylene production by many tissues of the
plant and induced petiole epinasty and premature senescence and abscission
of flowers, usually before anthesis. There were no obvious effects on
senescence in leaves of ACC synthase overexpressers, suggesting that
although ethylene may be important, it is not sufficient to cause tomato
leaf senescence; other signals are clearly involved.
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