Gibberellins, brassinosteroids and light-regulated development

The regulation of plant development by light requires the action of several well-studied plant hormones. However, the mechanism by which light and hormones affect identical developmental responses remains unclear. Recently, studies of mutants altered in light signal perception or transduction have suggested a role for gibberellins and brassinosteroids in light-regulated development. For instance, mutants in the major light-stable phytochrome from several plant species exhibit altered responsiveness to, or metabolism of, gibberellins. In contrast, mutants that develop as light-grown plants in the absence of light have implicated a role for brassinosteroids in the control of cell elongation, the expression of photoregulated genes, and the promotion of apical dominance, leaf senescence and male fertility. Future studies should help elucidate whether light and hormones independently affect these developmental responses or whether hormones are involved in the sequence of events initiated by excitation of photoreceptors.     

Hormonal input in plant meristems: A balancing act

Plant hormones are a group of chemically diverse molecules that control virtually all aspects of plant development. Classical plant hormones were identified many decades ago in physiology studies that addressed plant growth regulation. In recent years, biochemical and genetic approaches led to the identification of many molecular components that mediate hormone activity, such as hormone receptors and hormone-regulated genes. This has greatly contributed to the understanding of the mechanisms underlying hormone activity and highlighted the intricate crosstalk and integration of hormone signalling and developmental pathways. Here we review and discuss recent findings on how hormones regulate the activity of shoot and root apical meristems.     

Plant hormones: metabolism, signaling and crosstalk

Plants synthesize various hormones in response to environmental cues and developmental signals to ensure their proper growth and development.Elucidation of the molecular mechanisms by which plant hormones control growth and development contributes to our understanding of fundamental plant biology and provides tools to improve crops.Because of their critical roles in plant growth and development, plant hormones have been studied extensively since the early days of plant biology.     

Plant heterotrimeric G protein function: insights from Arabidopsis and rice mutants

Heterotrimeric G proteins have been implicated in a wide range of plant processes. These include responses to hormones, drought, and pathogens, and developmental events such as lateral root formation, hypocotyl elongation, hook opening, leaf expansion, and silique development. Results and concepts emerging from recent phenotypic analyses of G-protein component mutants in Arabidopsis and rice are adding to our understanding of G-protein mechanisms and functions in higher plants.     

Auxin-oxylipin crosstalk: relationship of antagonists

Stephan Pollmann (Corresponding author) Phytohormones regulate a wide array of developmental processes throughout the life cycle of plants. Herein, the various plant hormones may interact additively, synergistically, or antagonistically. By their cooperation they create a delicate regulatory network whose net output largely depends on the action of specific phytohormone combinations rather than on the independent activities of separate hormones. While most classical studies of plant hormonal control have focused mainly on the action of single hormones or on the synergistic interaction of hormones in regulating various developmental processes, recent work is beginning to shed light on the crosstalk of nominally antagonistic plant hormones, such as gibberellins and auxins with oxylipins or abscisic acid. In this review, we summarize our current understanding of how two of the first sight antagonistic plant hormones, i.e. auxins and oxylipins, interact in controlling plant responses and development.     

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.     

Auxin: a master regulator in plant root development

The demand for increased crop productivity and the predicted challenges related to plant survival under adverse environmental conditions have renewed the interest in research in root biology. Various physiological and genetic studies have provided ample evidence in support of the role of plant growth regulators in root development. The biosynthesis and transport of auxin and its signaling play a crucial role in controlling root growth and development. The univocal role of auxin in root development has established it as a master regulator. Other plant hormones, such as cytokinins, brassinosteroids, ethylene, abscisic acid, gibberellins, jasmonic acid, polyamines and strigolactones interact either synergistically or antagonistically with auxin to trigger cascades of events leading to root morphogenesis and development. In recent years, the availability of biological resources, development of modern tools and experimental approaches have led to the advancement of knowledge in root development. Research in the areas of hormone signal perception, understanding network of events involved in hormone action and the transport of plant hormones has added a new dimension to root biology. The present review highlights some of the important conceptual developments in the interplay of auxin and other plant hormones and associated downstream events affecting root development.     

Of light and length: regulation of hypocotyl growth in Arabidopsis

At all stages, plant development results from a complex integration of multiple endogenous and environmental signals. The sedentary nature of plants strongly enhances the impact of the environment on plant development as compared to animal development. The embryonic and postembryonic seedling stem, called the hypocotyl, of the model species Arabidopsis (thale cress) has proved to be an excellent system for studying such signal interplay in the regulation of growth and developmental responses. The extension of the hypocotyl, which is regulated by a network of interacting factors, including light and plant hormones, is such a process. These regulatory factors often reciprocally regulate their biosynthesis and/or signalling. Here we present the current state of knowledge about the regulation of hypocotyl growth by a large repertoire of internal and external cues.     

The Yin-Yang of Hormones: Cytokinin and Auxin Interactions in Plant Development

The phytohormones auxin and cytokinin interact to regulate many plant growth and developmental processes. Elements involved in the biosynthesis, inactivation, transport, perception, and signaling of these hormones have been elucidated, revealing the variety of mechanisms by which signal output from these pathways can be regulated. Recent studies shed light on how these hormones interact with each other to promote and maintain plant growth and development. In this review, we focus on the interaction of auxin and cytokinin in several developmental contexts, including its role in regulating apical meristems, the patterning of the root, the development of the gynoecium and female gametophyte, and organogenesis and phyllotaxy in the shoot.     

Moss systems biology en route: phytohormones in Physcomitrella development

The moss Physcomitrella patens has become a powerful model system in modern plant biology. Highly standardized cell culture techniques, as well as the necessary tools for computational biology, functional genomics and proteomics have been established. Large EST collections are available and the complete moss genome will be released soon. A simple body plan and the small number of different cell types in Physcomitrella facilitate the study of developmental processes. In the filamentous juvenile moss tissue, developmental decisions rely on the differentiation of single cells. Developmental steps are controlled by distinct phytohormones and integration of environmental signals. Especially the phytohormones auxin, cytokinin, and abscisic acid have distinct effects on early moss development. In this article, we review current knowledge about phytohormone influences on early moss development in an attempt to fully unravel the complex regulatory signal transduction networks underlying the developmental decisions of single plant cells in a holistic systems biology approach.     

Hormone interactions during lateral root formation

Lateral root (LR) formation, the production of new roots from parent roots, is a hormone- and environmentally-regulated developmental process in higher plants. Physiological and genetic studies using Arabidopsis thaliana and other plant species have revealed the roles of several plant hormones in LR formation, particularly the role of auxin in LR initiation and primordium development, resulting in much progress toward understanding the mechanisms of auxin-mediated LR formation. However, hormone interactions during LR formation have been relatively underexamined. Recent studies have shown that the plant hormones, cytokinin and abscisic acid negatively regulate LR formation whereas brassinosteroids positively regulate LR formation. On the other hand, ethylene has positive and negative roles during LR formation. This review summarizes recent findings on hormone-regulated LR formation in higher plants, focusing on auxin as a trigger and on the other hormones in LR formation, and discusses the possible interactions among plant hormones in this developmental process.     

Hormone signalling from a developmental context

The influence of hormones on plant growth and development has been clearly documented over the past 50 years. Now, with molecular genetics, the genes that convert changes in hormone levels into a cellular response are beginning to be identified. However, recent studies have demonstrated that the developmental context in which the hormones act plays a large influence on their synthesis and action. In this review, examples are given where known hormone response genes have been shown to have broader developmental roles as well as examples where genes that regulate developmental decisions, such as differentiation and fate, also influence hormone metabolism. The early conclusion of these studies is that an understanding of hormone signal transduction cannot be achieved in the absence of a developmental framework.     

Signs of change: hormone receptors that regulate plant development

Hormonal signalling plays a pivotal role in almost every aspect of plant development, and of high priority has been to identify the receptors that perceive these hormones. In the past seven months, the receptors for the plant hormones auxin, gibberellins and abscisic acid have been identified. These join the receptors that have previously been identified for ethylene, brassinosteroids and cytokinins. This review therefore comes at an exciting time for plant developmental biology, as the new findings shed light on our current understanding of the structure and function of the various hormone receptors, their related signalling pathways and their role in regulating plant development.     

Dawning of a new era: photomorphogenesis as an integrated molecular network

Plant growth is shaped by the dynamic integration of environmental, developmental, and metabolic cues. Information from many of these input pathways feeds into the highly connected network of small molecule phytohormones. Signal transduction components for most plant hormones are known and mapping of hormone interactions within the network is well underway. Recent investigations of seedling photomorphogenesis, using well-established physiological and genetic tools in combination with sophisticated application of newer genomic technologies, provide a systems-level view of early seedling development. Factors, such as light, the circadian clock, and organ-specific developmental programs, profoundly influence the hormone network. The integrative approaches described here clarify the mechanisms of signal integration while revealing the flexibility of such relationships.     

Long-range signalling in plant reproductive development

Animals and plants produce regulatory signals at specific places of their bodies, in order to regulate developmental events which take place at a distance. Plants use this mechanism to adjust their development to the changing environment. Flowering and tuber formation are controlled by signals generated in the leaves that travel throughout the plant to reach their target tissues: the shoot apical meristem for flowering and the underground stolons for tuberization. Although the existence of these long-distance plant messengers was postulated almost seventy years ago, their chemical nature is still not clear. These leaf-derived signals are graft-transmissible and move through the plant vascular system. Presumably they are very similar or even identical for flowering and tuberization and common to most plant species. It is generally accepted that their composition is complex and includes positive and negative regulators. Many different substances, including classical plant hormones and metabolites have been postulated to be components of these mobile signals, but conclusive evidence of this is still lacking. Recent work has positioned these signals within the genetic network that regulates flowering time and suggests roles for specific genes in the generation, transport or response to the signalling molecules. Current knowledge of long-range signalling mechanisms in other physiological and developmental events, together with the finding of common regulators involved in flowering, tuberization and other processes like pathogen and wound responses, should help to establish the biochemical composition of these elusive messenger signals.     

Hormonal regulation of leaf senescence through integration of developmental and stress signals

Leaf senescence is a genetically controlled dismantling programme that enables plants to efficiently remobilise nutrients to new growing sinks. It involves substantial metabolic reprogramming whose timing is affected by developmental and environmental signals. Plant hormones have long been known to affect the timing of leaf senescence, but they also affect plant development and stress responses. It has therefore been difficult to tease apart how the different hormones regulate the onset and progression of leaf senescence, i.e., whether they directly affect leaf senescence or affect it indirectly by altering the developmental programme or by altering plants’ response to stress. Here we review research on hormonal regulation of leaf senescence and propose that hormones affect senescence through differential responses to developmental and environmental signals. We suggest that leaf senescence strictly depends on developmental changes, after which senescence can be induced, depending on the type of hormonal and environmental cues.     

The Physiological,Biochemical and Molecular Roles of Brassinosteroids and Salicylic Acid in Plant Processes and Salt Tolerance

Plant hormones regulate plant growth and development by affecting an array of cellular, physiological, and developmental processes, including, but not limited to, cell division and elongation, stomatal regulation, photosynthesis, transpiration, ion uptake and transport, initiation of leaf, flower and fruit development, and senescence. Environmental factors such as salinity, drought, and extreme temperatures may cause a reduction in plant growth and productivity by altering the endogenous levels of plant hormones, sensitivity to plant hormones, and/or signaling pathways. Molecular and physiological studies have determined that plant hormones and abiotic stresses have interactive effects on a number of basic biochemical and physiological processes, leading to reduced plant growth and development. Various strategies have been considered or employed to maximize plant growth and productivity under environmental stresses such as salt-stress. A fundamental approach is to develop salt-tolerant plants through genetic means. Breeding for salt tolerance, however, is a long-term endeavor with its own complexities and inherent difficulties. The success of this approach depends, among others, on the availability of genetic sources of tolerance and reliable screening techniques, identification and successful transfer of genetic components of tolerance to desired genetic backgrounds, and development of elite breeding lines and cultivars with salt tolerance and other desirable agricultural characteristics. Such extensive processes have delayed development of successful salt-tolerant cultivars in most crop species. An alternative and technically simpler approach is to induce salt tolerance through exogenous application of certain plant growth–regulating compounds. This approach has gained significant interest during the past decade, when a wealth of new knowledge has become available on the beneficial roles of the six classes of plant hormones (auxins, gibberellins, cytokinins, abscisic acid, ethylene, and brassinosteroids) as well as several other plant growth–regulating substances (jasmonates, salicylates, polyamines, triacontanol, ascorbic acid, and tocopherols) on plant stress tolerance. Among these, brassinosteroids (BRs) and salicylic acid (SA) have been studied most extensively. Both BRs and SA are ubiquitous in the plant kingdom, affecting plant growth and development in many different ways, and are known to improve plant stress tolerance. In this article, we review and discuss the current knowledge and possible applications of BRs and SA that could be used to mitigate the harmful effects of salt-stress in plants. We also discuss the roles of exogenous applications of BRs and SA in the regulation of various biochemical and physiological processes leading to improved salt tolerance in plants.     

Signaling Interactions During Nodule Development

Nitrogen fixing bacteria, collectively referred to as rhizobia, are able to trigger the organogenesis of a new organ on legumes, the nodule. The morphogenetic trigger is a Rhizobium-produced lipochitin-oligosaccharide called the Nod factor, which is necessary, and in some legumes sufficient, for triggering nodule development in the absence of the bacterium. Because plant development is substantially influenced by plant hormones, it has been hypothesized that plant hormones (mainly the classical hormones abscisic acid, auxin, cytokinins, ethylene and gibberellic acid) regulate nodule development. In recent years, evidence has shown that Nod factors might act in legumes by changing the internal plant hormone balance, thereby orchestrating the nodule developmental program. In addition, many nonclassical hormonal signals have been found to play a role in nodule development, some of them similar to signals involved in animal development. These compounds include peptide hormones, nitric oxide, reactive oxygen species, jasmonic acid, salicylic acid, uridine, flavonoids and Nod factors themselves. Environmental factors, in particular nitrate, also influence nodule development by affecting the plant hormone status. This review summarizes recent findings on the involvement of classical and nonclassical signals during nodule development with the aim of illustrating the multiple interactions existing between these compounds that have made this area so complicated to analyze.