Abstract

Flower development at the shoot apex is initiated in response to environmental cues. Day length is one of the most important of these and is perceived in the leaves. A systemic signal, called the floral stimulus or florigen, is then transmitted from the leaves through the phloem and induces floral development at the shoot apex. Genetic analysis in Arabidopsis identified a pathway of genes required for the initiation of flowering in response to day length. The nuclear zinc-finger protein CONSTANS (CO) plays a central role in this pathway, and in response to long days activates the transcription of FT, which encodes a RAF-kinase-inhibitor-like protein. We show using grafting approaches that CO acts non-cell autonomously to trigger flowering. Although CO is expressed widely, its misexpression from phloem-specific promoters, but not from meristem-specific promoters, is sufficient to induce early flowering and complement the co mutation. The mechanism by which CO triggers flowering from the phloem involves the cell-autonomous activation of FT expression. Genetic approaches indicate that CO activates flowering through both FT-dependent and FT-independent processes, whereas FT acts both in the phloem and the meristem to trigger flowering. We propose that, partly through the activation of FT, CO regulates the synthesis or transport of a systemic flowering signal, thereby positioning this signal within the established hierarchy of regulatory proteins that controls flowering.

Highlights

  • The aerial organs of adult plants are derived from the shoot apical meristem (SAM), a collection of stem cells at the apex of the shoot that are formed during embryonic development

  • Genetic analysis in Arabidopsis identified a pathway of genes required for the initiation of flowering in response to day length

  • The nuclear zinc-finger protein CONSTANS (CO) plays a central role in this pathway, and in response to long days activates the transcription of FT, which encodes a RAF-kinase-inhibitor-like protein

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Summary

Introduction

The aerial organs of adult plants are derived from the shoot apical meristem (SAM), a collection of stem cells at the apex of the shoot that are formed during embryonic development. In Arabidopsis, organ primordia are continuously formed on the flanks of the SAM during post-embryonic development. The transition from vegetative to reproductive development is controlled by environmental signals such as day length or temperature. Each of these environmental signals results in floral development at the SAM, the signals are detected in different organs of the plant. Perception of day length in the leaf suggested that a systemic signal, often called the floral stimulus or florigen, is synthesised in the leaf and transmitted to the SAM where it triggers flower development (Knott, 1934; Zeevaart, 1976). The inducing substance has proven elusive despite extensive attempts to purify it from phloem extracts (Corbesier et al, 1998), and is usually believed to represent a complex mixture of substances (Bernier et al, 1993)

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