Hormone balance and abiotic stress tolerance in crop plants

Plant hormones play central roles in the ability of plants to adapt to changing environments, by mediating growth, development, nutrient allocation, and source/sink transitions. Although ABA is the most studied stress-responsive hormone, the role of cytokinins, brassinosteroids, and auxins during environmental stress is emerging. Recent evidence indicated that plant hormones are involved in multiple processes. Cross-talk between the different plant hormones results in synergetic or antagonic interactions that play crucial roles in response of plants to abiotic stress. The characterization of the molecular mechanisms regulating hormone synthesis, signaling, and action are facilitating the modification of hormone biosynthetic pathways for the generation of transgenic crop plants with enhanced abiotic stress tolerance.     

Kinetin--a multiactive molecule

Cytokinins are important adenine derivatives that serve as hormones to control many processes in plants. They were discovered as factors that promote cell division in tobacco tissue cultures and have been shown also to regulate several other developmental events. Kinetin which was isolated 50 years ago for the first time as a plant hormone, as well as other cytokinins isopentenyladenine, zeatin and benzylaminopurine induce callus (clusters of dedifferentiated plant cells) to redifferentiate into adventitious buds. Because of some similarities in the biological phenotypes of cancer and callus cells, cytokinins and especially kinetin, affect the differentiation of human cells through a common signal transduction system. Therefore, cytokinins found their way to use in molecular medicine.     

Cytokinins: Biosynthesis metabolism and perception

Summary Cytokinins are essential hormones for plant growth and development. They are also of vital importance for in vitro manipulations of plant cells and tissues. The biological activities and chemistry of cytokinins are well defined but very little is known about their mode of action and it is only recently that cytokinin genes have been identified in plants. This review summarizes the current status of knowledge on cytokinin biosynthesis, metabolism and signal transduction, with an emphasis on genes encoding metabolic enzymes and putative receptors, and genes rapidly induced by cytokinins.     

Phytohormone Conjugates: Nature and Function

Reversible hormone conjugations in plants may represent physiologically and biochemically essential pathways in the regulation of endogenous levels of biologically active pools of phytohormones. Conjugates of auxins, gibberellins, and cytokinins are now widely recognized as serving a storage function for rapid (im)mobilization of these phytohormones, depending on a variety of environmental, developmental, and physiological factors. The significance of conjugates of other phytohormones (abscisic acid, ethylene, jasmonic acid, and salicylic acid) is less well understood. Recent developments in studies on phytohormone conjugation, involving both biochemical and molecular biology approaches, are presented here. The nature and possible functions of the conjugates are discussed. Conjugates of other compounds (e.g., anthranilate-glucosides) are also known (for review, see Hösel, 1981). However, it is not known whether these compounds have a signaling function.     

Strigolactones are a new-defined class of plant hormones which inhibit shoot branching and mediate the interaction of plant-AM fungi and plant-parasitic weeds

Because plants are sessile organisms, the ability to adapt to a wide range of environmental conditions is critical for their survival. As a consequence, plants use hormones to regulate growth, mitigate biotic and abiotic stresses, and to communicate with other organisms. Many plant hormones function plei-otropically in vivo, and often work in tandem with other hormones that are chemically distinct. A newly-defined class of plant hormones, the strigolactones, cooperate with auxins and cytokinins to control shoot branching and the outgrowth of lateral buds. Strigolactones were originally identified as compounds that stimulated the germination of parasitic plant seeds, and were also demonstrated to induce hyphal branching in arbuscular mycorrhizal (AM) fungi. AM fungi form symbioses with higher plant roots and mainly facilitate the absorption of phosphate from the soil. Conforming to the classical definition of a plant hormone, strigolactones are produced in the roots and translocated to the shoots where they inhibit shoot outgrowth and branching. The biosynthesis of this class of compounds is regulated by soil nutrient availability, i.e. the plant will increase its production of strigolactones when the soil phosphate concentration is limited, and decrease production when phosphates are in ample supply. Strigolactones that affect plant shoot branching, AM fungal hyphal branching, and seed ger-mination in parasitic plants facilitate chemical synthesis of similar compounds to control these and other biological processes by exogenous application.     

Role of cytokinins in stress resistance of plants

The facts of both positive and negative influences of cytokinins on stress resistance of plants are known today. Without pretending to a final choice between these points of view, we have made an attempt to analyze the details of the experiments that gave rise to conclusions about the nature of the effect of cytokinins on the resistance to stress-causing influences with a focus on their intensity and duration. The review deals with the data concerning the influence of different adverse factors on the content of endogenous cytokinins and transduction of cytokinin signals, examines the influence on plant resistance of treatment with exogenous hormone, and the effects of genetic modifications causing changes in cytokinin content and signaling. Resistance is considered not only as a mean of plant survival under severe stress but also as an instrument of maintaining growth rate in plants exposed to moderate stress. Literature data and our own results make it possible to conclude that cytokinins play an important role in formation of plant resistance to adverse influences; however, the effect of these hormones depends on stress intensity. Under moderate stress, cytokinins ensure maintenance of plant growth, whereas a drop in cytokinins hampers growth under a strong influence of adverse factors, which is a prerequisite for mobilization of limited resources characteristic of severe stress and ensures preservation of plant viability.     

Plant hormone conjugation: A signal decision

Tight regulation of the auxin hormone indole-3-acetic acid (IAA) is crucial for plant development. Newly discovered IAA antagonists are the amide-linked tryptophan conjugates of IAA and jasmonic acid (JA). JA-Trp and IAA-Trp interfered with root gravitropism in Arabidopsis, and inhibited several responses to exogenously supplied IAA. Relatively low concentrations of the inhibitors occurred in Arabidopsis, but Pisum sativum flowers contained over 300 pmole g⁻¹ FW of JA-Trp. DihydroJA was an even more effective inhibitor than JA-Trp, suggesting that Trp conjugates with other JA derivatives may also be functional. JA-Trp and IAA-Trp add to the list of documented bioactive amide hormone conjugates. The only other example is JA-Ile, the recently discovered jasmonate signal. These examples establish that conjugation not only inactivates hormones, but in some cases creates novel compounds that function in hormone signaling.Key words: jasmonic acid, indole-3-acetic acid, auxin, tryptophan, conjugate, plant hormone, signaling, amino acid, antagonistPlants hold an amazing capacity to auto-regulate their growth and respond to a host of environmental challenges. Since the early discovery of the first plant hormone, indole-3-acetic acid (IAA),¹ science has progressively unveiled ever more complex, and sometimes surprising, ways that plants manipulate hormones to optimize their growth and thwart their opponents. Until recently, the covalent coupling of hormones to sugars, amino acids and peptides was thought to be merely a way to dispose of excess hormone.² The amide linkage of IAA to Asp and Glu does indeed result in IAA catabolism, while IAA-Ala and IAA-Leu are inactive stored forms of IAA.³ But the perception that all hormone conjugates are inactive changed abruptly with the discovery that the isoleucine conjugate of jasmonic acid (JA-Ile) is an active hormonal signal.     

Cytokinin conjugation: recent advances and patterns in plant evolution

Cytokinin (CK) conjugates are important in plant development because they regulate active CK concentrations, CK transport, storage, and irreversible inactivation. While numerous CK conjugates have been identified in higher plants, the biological functions of these compounds, their location within cells and tissues, and the enzymes and genes involved in their regulation are not clearly understood. In this paper, recent advances are reported which have occurred through the study of transgenic plants containing the ipt or rolC genes, the identification of new regulatory enzymes affecting CKs, and the characterization of new CK conjugates. In addition, a survey of the literature is presented which examines the pattern of CK conjugates found in different plant taxa. Based on current knowledge, it appears that green algae, mosses, and ferns contain relatively few CK conjugates of isopentenyl adenine (iP) and zeatin (Z). In contrast, higher land plants, such as gymnosperms and angiosperms, contain a more complex set of CKs, primarily conjugates of Z and dihydrozeatin (DHZ). This suggests that the pattern of CK conjugation has become more complex in parallel with the increasing complexity of higher plants.     

Histones and Plant Hormones: New Evidence for an Interesting Interplay

As any living organism, plants have to respond to a wide variety of biotic and abiotic signals. One of the most important challenges for the plant cell is the response to plant hormones, compounds that regulate almost all aspects of plant development. This process involves the appropriate alteration of chromatin structure and function which can be facilitated by a number of different mechanisms, including histone covalent modifications, incorporation of distinct histone variants, chromatin remodeling and epigenetic regulation of gene expression. On the other hand, distinct chromatin states may in turn influence plant hormone biosynthesis and signaling. This article aims to review the evidence accumulated the last years concerning the effect of the major plant hormones - auxin, gibberellins, cytokinins, ethylene, abscisic acid, brassinosteroids and jasmonic acid - signaling on histone variants and modification pattern and to discuss the interplay between histones and hormones at plant chromatin.     

Cytokinin-induced differentiation of human myeloid leukemia HL-60 cells is associated with the formation of nucleotides, but not with incorporation into DNA or RNA

Cytokinins are important purine derivatives that act as hormones to control many processes in plants. Cytokinins such as isopentenyladenine (IPA), kinetin and benzyladenine were very effective at inducing the granulocytic differentiation of human myeloid leukemia HL-60 cells. The metabolism of cytokinins to their nucleotides was closely associated with cytokinin-induced differentiation and growth inhibition. When the cells were incubated with [14C]-benzyladenine, radioactivity was significantly incorporated into RNA and DNA. However, the radioactive nucleotides in RNA or DNA were adenine nucleotides, not benzyladenine nucleotides, suggesting that cytokinins were not incorporated into RNA and DNA. The benzyladenine nucleotides were not stably released into the medium in intact form. Cytokinins effectively induced a phosphorylated (active) form of mitogen-activated protein kinase (MAPK). MAPK activation was necessary for cytokinin-induced differentiation, because PD98059, an inhibitor of MAPK kinase, suppressed the differentiation induced by cytokinins. These results suggest that cytokinin nucleotides themselves play an important role in inducing the differentiation of HL-60 cells.     

Cytokinin biosynthesis: A black box?

The biosynthetic origin of cytokinins (CKs) in plants has been debated ever since the recognition of CKs as plant hormones. Although several possible biosynthetic pathways have been suggested, none of the data published to date are conclusive. The enzymes involved in CK biosynthesis have not yet been purified and characterized in detail, nor have plant genes encoding CK biosynthetic enzymes been identified.     

植物生长物质与植物抗旱性的关系(综述)

脱落酸、乙烯、多胺和细胞分裂素等是与植物抗旱性关系较为密切的几种植物激素。文章就植物在遭受干旱胁迫时其体内激素变化及喷施植物生长调节剂对植物抗旱性的影响进行概述,为生化调控植物抗旱性提供参考。     

Dependence of root biomass accumulation on the content and metabolism of cytokinins in ethylene-insensitive plants

Diverse functions of ethylene in plants may depend on its ability to interact with other hormones. We studied the participation of ethylene in the regulation of accumulation and metabolism of cytokinins comparing ethylene-insensitive mutant plants of arabidopsis (Arabidopsis thaliana [L.] Heynh., etr1-1) with the plants of original ecotype Columbia (Col-0). Because cytokinins can regulate growth of both leaves and roots, we determined the weights of these organs and the ratio between them. The content of zeatin and its riboside in the roots of etr1-1 plants was two times greater than in Col-0 plants, which could be accounted for by inhibition of conversion of these forms of cytokinins into 9-N-glucosides. In the leaves of mutant plants, expression of IPT3 gene responsible for the synthesis of cytokinins was more intense than in Col-0 plants, which could also contribute to a rise in the content of cytokinins. In this case, the weight of roots in etr1-1 mutants was lower than in the plants of original ecotype. Because high concentrations of cytokinins can inhibit root growth, suppression of accumulation of their biomass in mutant plants may be related to a greater content of cytokinins therein. The obtained results suggest that ethylene can suppress accumulation of cytokinins and, thereby, maintain redistribution of biomass in favor of the roots, which is important for plant adaptation to a shortage of water and ions.     

Effects of Root Cutting on Cytokinin Content in the Shoot Apex Cells of Arabidopsis Plants

The dependence of cytokinin accumulation in the shoot apexes of Arabidopsis plants on the delivery of these hormones from the roots was studied. For the estimation of cytokinin content in the cells, the immunohistochemical localization method using antibodies against zeatin riboside was used. Differential conjugation of free cytokinin bases and their ribosides was used to prevent washing of cytokinins during the dehydration process. Root cutting decreased the immunostaining of zeatin in the cells of the shoot apical meristem, thereby supporting the hypothesis about dependence of cytokinin accumulation in these cells on the hormone delivery from the roots. The level of cytokinins in the cells of the shoot apex decreased under the effect of protonophore, indicating the important role of the secondary-active transmembrane transport process of cytokinins in the maintenance of their level in the cells of the shoot apex.     

N-glucosylation of cytokinins by glycosyltransferases of Arabidopsis thaliana

Cytokinins are plant hormones that can be glucosylated to form O-glucosides and N-glucosides. The glycoconjugates are inactive and are thought to play a role in homeostasis of the hormones. Although O-glucosyltransferases have been identified that recognize cytokinins, the enzymes involved in N-glucosylation have not been identified even though the process has been recognized for many years. This study utilizes a screening strategy in which 105 recombinant glycosyltransferases (UGTs) of Arabidopsis have been analyzed for catalytic activity toward the classical cytokinins: trans-zeatin, dihydrozeatin, N(6)-benzyladenine, N(6)-isopentenyladenine, and kinetin. Five UGTs were identified in the screen. UGT76C1 and UGT76C2 recognized all cytokinins and glucosylated the hormones at the N(7) and N(9) positions. UGT85A1, UGT73C5, and UGT73C1 recognized trans-zeatin and dihydrozeatin, which have an available hydroxyl group for glucosylation and formed the O-glucosides. The biochemical characteristics of the N-glucosyltransferases were analyzed, and highly effective inhibitors of their activities were identified. Constitutive overexpression of UGT76C1 in transgenic Arabidopsis confirmed that the recombinant enzyme functioned in vivo to glucosylate cytokinin applied to the plant. The role of the N-glucosyltransferases in cytokinin metabolism is discussed.     

细胞分裂素对植物衰老的延缓作用

细胞分裂素是一类重要的植物激素,它可在一定程度上延缓植物的衰老。主要从3个方面综述了细胞分裂素与植物衰老之间的关系,即:(1)植物衰老过程中内源细胞分裂素含量变化;(2)外源细胞分裂素的影响;(3)转入与细胞分裂素的合成、降解相关的基因对植物衰老产生的影响。此外,还从细胞分裂素与糖、与脂质氧化反应以及与其它植物激素的关系方面探讨了细胞分裂素在延缓植物衰老中的作用机理。     

The discovery of cytokinin receptors and biosynthesis of cytokinins: A true story

The year 2001 saw an amazing progress in cytokinin studies. Ten years ago, cytokinin receptor genes and genes encoding cytokinin biosynthetic enzymes together with the corresponding proteins were identified in plants. These studies elucidated the molecular mechanism of cytokinin effects on the expression of cytokinin responsive genes and ultimately established the endogenous synthesis of cytokinins in plant cells, justifying their membership among plant hormones. The paper describes in short the edifying and sometimes paradoxical story of these fundamental and captivating discoveries.