FT, a mobile developmental signal in plants

Plants synchronise their flowering with the seasons to maximise reproductive fitness. While plants sense environmental conditions largely through the leaves, the developmental decision to flower occurs in the shoot apex, requiring the transmission of flowering information, sometimes over quite long distances. Interestingly, despite the enormous diversity of reproductive strategies and lifestyles of higher plants, a key component of this mobile flowering signal, or florigen, is contributed by a highly conserved gene: FLOWERING LOCUS T (FT). The FT gene encodes a small globular protein that is able to translocate from the leaves to the shoot apex through the phloem. Plants have evolved a variety of regulatory networks that control FT expression in response to diverse environmental signals, enabling flowering and other developmental responses to be seasonally timed. As well as playing a key role in flowering, recent discoveries indicate FT is also involved in other developmental processes in the plant, including dormancy and bud burst.     

Leaf-produced floral signals

Florigen is the hypothetical leaf-produced signal that induces floral initiation at the shoot apex. The nature of florigen has remained elusive for more than 70 years. But recent progress toward understanding the regulatory network for flowering in Arabidopsis has led to the suggestion that FLOWERING LOCUS T (FT) or its product is the mobile flower-inducing signal that moves from an induced leaf through the phloem to the shoot apex. In the past year, physical and chemical evidence has shown that it is FT protein, and not FT mRNA, that moves from induced leaves to the apical meristem. These results have established that FT is the main, if not the only, component of the universal florigen.     

AGL24 acts in concert with SOC1 and FUL during Arabidopsis floral transition

Arabidopsis plants flower in response to long days (LDs). Exposure of leaves to inductive day lengths activates expression of FLOWERING LOCUS T (FT) protein which moves to the shoot apical meristem (SAM) to induce developmental reprogramming. SUPPRESSOR OF OVEREXPRESSION OF CONSTANS 1 (SOC1) and FRUITFULL (FUL) are induced by FT at the apex. We previously screened the SAM for mRNAs of genes required to promote the floral transition in response to photoperiod, and conducted detailed expression and functional analyses on several putative candidates. Here, we show that expression of AGAMOUS-LIKE 24 (AGL24) is detected at the SAM under SD conditions and increases upon exposure to LDs. Mutations in AGL24 further delay flowering of a soc1 ful double mutant, suggesting that flowering is controlled by AGL24 partly independently of SOC1 and FUL.     

The multifaceted roles of FLOWERING LOCUS T in plant development

One of the key developmental processes in flowering plants is the differentiation of the shoot apical meristem into a floral meristem. This transition is regulated through the integration of environmental and endogenous stimuli, involving a complex, hierarchical signalling network. In arabidopsis, the FLOWERING LOCUS T (FT) protein, a mobile signal recognized as a major component of florigen, has a central position in mediating the onset of flowering. FT-like genes seem to be involved in regulating the floral transition in all angiosperms examined to date. Evidence from molecular evolution studies suggests that the emergence of FT-like genes coincided with the evolution of the flowering plants. Hence, the role of FT in floral promotion is conserved, but appears to be restricted to the angiosperms. Besides flowering, FT-like proteins have also been identified as major regulatory factors in a wide range of developmental processes including fruit set, vegetative growth, stomatal control and tuberization. These multifaceted roles of FT-like proteins have resulted from extensive gene duplication events, which occurred independently in nearly all modern angiosperm lineages, followed by sub- or neo-functionalization. This review assesses the plethora of roles that FT-like genes have acquired during evolution and their implications in plant diversity, adaptation and domestication.     

FTIP1 is an essential regulator required for florigen transport

The capacity to respond to day length, photoperiodism, is crucial for flowering plants to adapt to seasonal change. The photoperiodic control of flowering in plants is mediated by a long-distance mobile floral stimulus called florigen that moves from leaves to the shoot apex. Although the proteins encoded by FLOWERING LOCUS T (FT) in Arabidopsis and its orthologs in other plants are identified as the long-sought florigen, whether their transport is a simple diffusion process or under regulation remains elusive. Here we show that an endoplasmic reticulum (ER) membrane protein, FT-INTERACTING PROTEIN 1 (FTIP1), is an essential regulator required for FT protein transport in Arabidopsis. Loss of function of FTIP1 exhibits late flowering under long days, which is partly due to the compromised FT movement to the shoot apex. FTIP1 and FT share similar mRNA expression patterns and subcellular localization, and they interact specifically in phloem companion cells. FTIP1 is required for FT export from companion cells to sieve elements, thus affecting FT transport through the phloem to the SAM. Our results provide a mechanistic understanding of florigen transport, demonstrating that FT moves in a regulated manner and that FTIP1 mediates FT transport to induce flowering.     

From the model to the crop: genes controlling tuber formation in potato

Photoperiod regulates many different developmental processes, including floral induction in several species and tuber formation in potato. Research in Arabidopsis led to the identification of FLOWERING LOCUS T (FT) as a main component of the florigen or mobile flowering promoting signal produced in the leaves. A similar mobile signal or tuberigen has been reported to induce tuber formation in potato, recent evidence obtained in our laboratory indicates that a potato homolog of FT encodes this signal. Flowering regulators, like CONSTANS and miR172, also play a role in tuberization, although it remains unclear whether these regulators function in identical pathways. Here, we highlight differential regulation of these genes in flowering and tuberization control and discuss on their possible tuberization-related function.     

The quest for florigen: a review of recent progress

The photoperiodic induction of flowering is a systemic process requiring translocation of a floral stimulus from the leaves to the shoot apical meristem. In response to this stimulus, the apical meristem stops producing leaves to initiate floral development; this switch in morphogenesis involves a change in the identity of the primordia initiated and in phyllotaxis. The physiological study of the floral transition has led to the identification of several putative floral signals such as sucrose, cytokinins, gibberellins, and reduced N-compounds that are translocated in the phloem sap from leaves to the shoot apical meristem. On the other hand, the genetic approach developed more recently in Arabidopsis thaliana allowed the discovery of many genes that control flowering time. These genes function in 'cascades' within four promotive pathways, the 'photoperiodic', 'autonomous', 'vernalization', and 'gibberellin' pathways, which all converge on the 'integrator' genes SUPPRESSOR OF OVEREXPRESSION OF CO 1 (SOC1) and FLOWERING LOCUS T (FT). Recently, several studies have highlighted a role for a product of FT as a component of the floral stimulus or 'florigen'. These recent advances and the proposed mode of action of FT are discussed here.