Alterations in the response of dark-grown seedlings to ethylene (the "triple response") were used to isolate a collection of ethylene-related mutants in Arabidopsis thaliana. Mutants displaying a constitutive response (eto1) were found to produce at least 40 times more ethylene than the wild type. The morphological defects in etiolated eto1-1 seedlings reverted to wild type under conditions in which ethylene biosynthesis or ethylene action were inhibited. Mutants that failed to display the apical hook in the absence of ethylene (his1) exhibited reduced ethylene production. In the presence of exogenous ethylene, hypocotyl and root of etiolated his1-1 seedlings were inhibited in elongation but no apical hook was observed. Mutants that were insensitive to ethylene (ein1 and ein2) produced increased amounts of ethylene, displayed hormone insensitivity in both hypocotyl and root responses, and showed an apical hook. Each of the "triple response" mutants has an effect on the shape of the seedling and on the production of the hormone. These mutants should prove to be useful tools for dissecting the mode of ethylene action in plants.

Exploiting the triple response of Arabidopsis to identify ethylene-related mutants.

Senescence and Postharvest Physiology of Cut Flowers, Part 1

Physiology and Biochemistry of Flower Petal Senescence

We have examined the expression of mRNAs for S-adenosylmethionine synthetase (EC 2.5.1.6), 1-aminocyclopropane-1-carboxylate (ACC) synthase (EC 4.4.1.14), and the ethylene-forming enzyme (EFE) in various floral organs of carnation (Dianthus caryophyllus) during the increase in ethylene biosynthesis associated with petal senescence. The abundance of ACC synthase and EFE mRNAs increased and S-adenosylmethionine synthetase transcripts decreased concomitantly with the ethylene climacteric in senescing petals. The increase in abundance of ACC synthase and EFE mRNAs in aging flowers was prevented by treatment with the ethylene action inhibitor 2,5-norbornadiene. Furthermore, an increase in ACC synthase and EFE transcripts was detected in petals from presenescent flowers within 3 to 6 hours of exposure to 2 microliters per liter of ethylene. The increase in ethylene production by senescing petals was associated with a concomitant increase in ethylene biosynthesis in styles, ovary, and receptacle tissues. In all tissues, this increase was associated with increased activities of ACC synthase and EFE. The increase in EFE activities by all floral organs examined was correlated with increased abundance of EFE transcripts. In contrast, the level of ACC synthase mRNA, as detected by the cDNA probe pCARACC3, did not always reflect enzyme activity. The combined tissues of the pistil exhibited high rates of ACC synthase activity but contained low levels of ACC synthase mRNAs homologous to pCARACC3. In addition, pollinated styles exhibited a rapid increase in ethylene production and ACC synthase activity but did not accumulate detectable levels of ACC synthase mRNA until several hours after the initiation of ethylene production. These results suggest that transcripts for ACC synthase leading to the early postpollination increase in ACC synthase activity and ethylene production are substantially different from the mRNA for the ethylene-responsive gene represented by pCARACC3.

Expression of ethylene biosynthetic pathway transcripts in senescing carnation flowers.

Pollination regulates a syndrome of developmental responses that contributes to successful sexual reproduction in higher plants. Pollination-regulated developmental events collectively prepare the flower for fertilization and embryogenesis while bringing about the loss of floral organs that have completed their function in pollen dispersal and reception. Components of this process include changes in flower pigmentation, senescence and abscission of floral organs, growth and development of the ovary, and, in certain cases, pollination also triggers ovule and female gametophyte development in anticipation of fertilization. Pollination-regulated development is initiated by the primary pollination event at the stigma surface, but because developmental processes occur in distal floral organs, the activity of interorgan signals that amplify and transmit the primary pollination signal to floral organs is implicated. Interorgan signaling and signal amplification involves the regulation of ethylene biosynthetic gene expression and interorgan transport of hormones and their precursors. The coordination of pollination-regulated flower development including gametophyte, embryo, and ovary development; pollination signaling; the molecular regulation of ethylene biosynthesis; and interorgan communication are presented.

Pollination regulation of flower development.

The rise in ethylene production accompanying the respiration climacteric and senescence of cut carnation flowers (Dianthus caryophyllus L. cv. White Sim) was associated with a 30-fold increase in the concentration of 1-aminocyclopropane-1-carboxylic acid (ACC) in the petals (initial content 0.3 nmol/g fresh weight). Pretreatment of the flowers with silver thiosulfate (STS) retarded flower senescence and prevented the increase in ACC concentration in the petals. An increase in ACC in the remaining flower parts, which appeared to precede the increase in the petals, was only partially prevented by the STS pretreatment. Addition of aminoxyacetic acid (2 mM) to the solution in which the flowers were kept completely inhibited accumulation of ACC in all flower parts.

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Changes in 1-aminocyclopropane-1-carboxylic-acid content of cut carnation flowers in relation to their senescence.

Pollination of flowers of standard carnation (Dianthus caryophyllus L. cv. White Sim) with pollen from flowers of miniature carnations (D. caryophyllus L. cv. Exquisite) caused them to wilt irreversibly within 1 to 2 days. Pollination stimulated a sequential increase in ethylene production by stigmas, ovaries, receptacles, and petals of the flowers. The ACC content of the stigmas increased rapidly in the first few hours after pollination. The possibility that subsequent production of ethylene by other parts of the flower is stimulated by translocated ACC is discussed. Ethylene production and ACC content of other parts of the flower reached their maximum 24 h after pollination. The petal tissues contributed the bulk of the ethylene productionper flower thereafter. There appears to be a qualitative difference between the enzyme in the stigmas converting ACC to ethylene and that in other parts of the flower.

Y. Mor

Papers

Changes in 1-aminocyclopropane-1-carboxylic-acid content of cut carnation flowers in relation to their senescence.

Changes in ethylene production and 1-aminocyclopropane-1-carboxylic acid content of pollinated carnation flowers