It's time to flower: the genetic control of flowering time

In plants, successful sexual reproduction and the ensuing development of seeds and fruits depend on flowering at the right time. This involves coordinating flowering with the appropriate season and with the developmental history of the plant. Genetic and molecular analysis in the small cruciform weed, Arabidopsis, has revealed distinct but linked pathways that are responsible for detecting the major seasonal cues of day length and cold temperature, as well as other local environmental and internal signals. The balance of signals from these pathways is integrated by a common set of genes to determine when flowering occurs. Excitingly, it has been discovered that many of these same genes regulate flowering in other plants, such as rice. This review focuses on recent advances in how three of the signalling pathways (the day-length, vernalisation and autonomous pathways) function to control 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.     

Long-distance regulation of flowering time

One of the great mysteries of plant science appears to have been resolved with the discovery that the protein FT can act as a phloem-mobile florigen hormone. The collective evidence from several laboratories, many from studies on photoperiod response, indicates that FT and its homologues are universal signalling molecules for flowering plants. Duplication and divergence of FT-like proteins reveals an increased complexity of function in certain taxonomic groups including grasses and legumes. There are additional components of long-distance flowering time control, such as a role for gibberellins in some species but probably not others. Cytokinins and sugars are further putative signals. Vernalization processes and responses are generally considered to occur in shoot meristems, but systemic responses to cold have been reported several times. Finally, there is increasing evidence that FT does not act purely to switch on flowering, but in addition, has broader roles in seasonal developmental switches such as bud dormancy and tuberization, and in the regulation of meristem determinacy and compound leaf development. This review seeks to highlight recent progress in systemic floral signalling, and to indicate areas in need of further research.     

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.     

Problems of Growth and Development in the Studies by M.Kh. Chailakhyan

Main vistas of Academician M.Kh. Chailakhyan's inquiry into the problems of plant growth and development are reviewed with the emphasis on the input of his research in elucidating the mechanisms of flowering, tuberization, sex expression, and the integrity of plant organism. The pioneer studies conducted by Chailakhyan and his colleagues on agricultural applications of phytohormones, their synthetic analogs, and other growth regulators are considered. Highlighted is the role of theoretical papers written by Chailakhyan, his lectures, and his brave fight against pseudoscientific trends in the Russian science is stressed.     

Integrating hormone signaling and patterning mechanisms in plant development

Plant growth and development are driven by the bustling integration of a vast number of signals, among which plant hormones dominate. Understanding the role of hormones in particular developmental events requires their integration with developmental regulators known to be specific to those events. Using the increasing number of tools that can be utilized to probe hormone biosynthesis, transport and response, several recent studies have taken such an integrative approach, and in so doing have contributed to a clearer picture of precisely how hormones control plant development.     

Photoreceptors and signals in the photoperiodic control of development

Many plant species are sensitive to changes in the seasons, especially with regard to their reproductive behaviour. Sexual or vegetative reproductive structures are often only formed at times of the year when days are sufficiently long, or short. Plants perceive daylength in the leaves, but reproductive changes occur in shoot apices in response to the movement of signals throughout the plant. There is good evidence that phytochrome A is an essential component of the daylength-sensing mechanism in long-day plants, and preliminary evidence suggests a potential interaction between phytochrome C and daylength sensitivity in short-day plants. The sensitivity of reproductive processes to photoperiodic control is directly altered by photoreceptor action, particularly in the case of phytochrome B, which has a major influence on whether flowering or tuberization occurs under non-inductive conditions in both long- and short-day plants, but is not involved in daylength measurement. The signals which move from leaves to the sites of reproductive development are not known, but there is good evidence that gibberellins may be important and some preliminary indication that brassinosteroids might also be involved in photoperiodic signalling.     

The ups and downs of signalling between root and shoot

It is becoming increasingly apparent that the long-distance signalling associated with many developmental processes is complex and that novel hormone-like signals may play substantial roles. The past decades have seen several substances (e.g. brassinosteroids, systemin and other polypeptides, mevalonic and jasmonic acids, polyamines, oligosaccharides, flavonoids, and quinones) vie for a place among the classical plant hormones (e.g. Spaink, 1996). Recent microinjection and grafting studies have also shown that RNA may act as a long-distance signal (Jorgensen et al ., 1998; Xoconostle-Cázares et al ., 1999). In this issue, Hannah et al . describe long-distance signalling and the regulation of root–shoot partitioning in dwarf lethal or dosage-dependent lethal ( DL ) mutants of common bean (Shii et al ., 1980, 1981), and present evidence indicating that substances in addition to classical plant hormones (e.g. cytokinins) may be involved.
As in the report by Hannah et al ., much of the evidence for roles of unidentified long-distance signals in the control of plant development is indirect. The possibility that a small number of long-distance signals might control a multitude of developmental processes arises through the potential for differences in tissue sensitivity, fluctuations in hormone levels and differences in the nature of responses of different tissues to the same hormone. Consequently, particular hormones may influence numerous processes seemingly simultaneously, yet independently. Even so, long-distance signalling is involved in processes as diverse as root–shoot balance, senescence, branching, flowering, nodulation, stress responses and nutrient uptake. Through comparison of even a few different developmental processes, progress can be made to reveal the true complexity of plant development. Using this approach it is also clear that many unknown signals may be involved.     

Cross-talk in plant hormone signalling: what Arabidopsis mutants are telling us

Genetic screens have been extremely useful in identifying genes involved in hormone signal transduction. However, although these screens were originally designed to identify specific components involved in early hormone signalling, mutations in these genes often confer changes in sensitivity to more than one hormone at the whole-plant level. Moreover, a variety of hormone response genes has been identified through screens that were originally designed to uncover regulators of sugar metabolism. Together, these facts indicate that the linear representation of the hormone signalling pathways controlling a specific aspect of plant growth and development is not sufficient, and that hormones interact with each other and with a variety of developmental and metabolic signals. Following the advent of arabidopsis molecular genetics we are beginning to understand some of the mechanisms by which a hormone is transduced into a cellular response. In this Botanical Briefing we review a subset of genes in arabidopsis that influence hormonal cross-talk, with emphasis on the gibberellin, abscisic acid and ethylene pathways. In some cases it appears that modulation of hormone sensitivity can cause changes in the synthesis of an unrelated hormone, while in other cases a hormone response gene defines a node of interaction between two response pathways. It also appears that a variety of hormones may converge to regulate the turnover of important regulators involved in growth and development. Examples are cited of the recent use of suppressor and enhancer analysis to identify new nodes of interaction between these pathways.     

Key players associated with tuberization in potato: potential candidates for genetic engineering

Tuberization in potato (Solanum tuberosum L.) is a complex biological phenomenon which is affected by several environmental cues, genetic factors and plant nutrition. Understanding the regulation of tuber induction is essential to devise strategies to improve tuber yield and quality. It is well established that short-day photoperiods promote tuberization, whereas long days and high-temperatures inhibit or delay tuberization. Worldwide research on this complex biological process has yielded information on the important bio-molecules (proteins, RNAs, plant growth regulators) associated with the tuberization process in potato. Key proteins involved in the regulation of tuberization include StSP6A, POTH1, StBEL5, StPHYB, StCONSTANS, Sucrose transporter StSUT4, StSP5G, etc. Biomolecules that become transported from “source to sink” have also been suggested to be important signaling candidates regulating the tuberization process in potatos. Four molecules, namely StSP6A protein, StBEL5 RNA, miR172 and GAs, have been found to be the main candidates acting as mobile signals for tuberization. These biomolecules can be manipulated (overexpressed/inhibited) for improving the tuberization in commercial varieties/cultivars of potato. In this review, information about the genes/proteins and their mechanism of action associated with the tuberization process is discussed.     

Epigenetic and physiological effects of gibberellin inhibitors and chemical pruners on the floral transition of azalea

The ability to control the timing of flowering is a key strategy in planning the production of ornamental species such as azaleas; however, it requires a thorough understanding of floral transition. DNA methylation is involved in controlling the functional state of chromatin and gene expression during floral induction pathways in response to environmental and developmental signals. Plant hormone signalling is also known to regulate suites of morphogenic processes in plants and its role in flowering-time control is starting to emerge as a key controlling step. This work investigates if the gibberellin (GA) inhibitors and chemical pinching applied in improvement of azalea flowering alter the dynamics of DNA methylation or the levels of polyamines (PAs), GAs and cytokinins (CKs) during floral transition, and whether these changes could be related to the effects observed on flowering ability. DNA methylation during floral transition and endogenous content of PAs, GAs and CKs were analysed after the application of GA synthesis inhibitors (daminozide, paclobutrazol and chlormequat chloride) and a chemical pruner (fatty acids). The application of GA biosynthesis inhibitors caused alterations in levels of PAs, GAs and CKs and in global DNA methylation levels during floral transition; also, these changes in plant growth regulators and DNA methylation were correlated with flower development. DNA methylation, PA, GA and CK levels can be used as predictive markers of plant floral capacity in azalea.     

Light and temperature signal crosstalk in plant development

Light and temperature are two of the most important environmental stimuli regulating plant development. Recent advances have suggested considerable interaction between these signalling pathways at the molecular level. Studies of both flowering and germination have shown the phytochrome family of plant photoreceptors to display altered functional hierarchies at different growth temperatures. The existence of common signalling components in both light and temperature sensing has additionally been proposed. More recently, light quality signals have been shown to regulate plant-freezing tolerance in an ambient temperature-dependent manner. Together, these data suggest that complex crosstalk between light-signalling and temperature-signalling pathways is fundamental to the growth and development of plants in natural environments.     

Modulation of environmental responses of plants by circadian clocks

Circadian clocks are signalling networks that enhance an organism's relationship with the rhythmic environment. The plant circadian clock modulates a wide range of physiological and biochemical events, such as stomatal and organ movements, photosynthesis and induction of flowering. Environmental signals regulate the phase and period of the plant circadian clock, which results in an approximate synchronization of clock outputs with external events. One of the consequences of circadian control is that stimuli of the same strength applied at different times of the day can result in responses of different intensities. This is known as 'gating'. Gating of a signal may allow plants to better process and react to the wide range and intensities of environmental signals to which they are constantly subjected. Light signalling, stomatal movements and low-temperature responses are examples of signalling pathways that are gated by the circadian clock. In this review, we describe the many levels at which the circadian clock interacts with responses to the environment. We discuss how environmental rhythms of temperature and light intensity entrain the circadian clock, how photoperiodism may be regulated by the relationship between environmental rhythms and the phasing of clock outputs, and how gating modulates the sensitivity of the clock and other responses to environmental and physiological signals. Finally, we describe evidence that the circadian clock can increase plant fitness.