生长素调控种子的休眠与萌发

植物种子的休眠与萌发,是植物生长发育过程中的关键阶段,也是生命科学领域的研究热点。种子从休眠向萌发的转换是极为复杂的生物学过程,由外界环境因子、体内激素含量及信号传导和若干关键基因协同调控。大量研究表明,植物激素脱落酸(Abscisic acid, ABA)和赤霉素(Gibberellin acid, GA)是调控种子休眠水平,决定种子从休眠转向萌发的主要内源因子。ABA与GA在含量和信号传导两个层次上的精确平衡,确保了植物种子能以休眠状态在逆境中存活,并在适宜的时间启动萌发程序。生长素(Auxin)是经典植物激素之一,其对向性生长和组织分化等生物学过程的调控已有大量研究。但最近有研究证实,生长素对种子休眠有正向调控作用,这表明生长素是继ABA之后的第二个促进种子休眠的植物激素。本文在回顾生长素的发现历程、阐释生长素体内合成途径及信号传导通路的基础上,重点综述了生长素通过与ABA的协同作用调控种子休眠的分子机制,并对未来的研究热点进行了讨论和展望。     

Light Regulation of Gibberellins Metabolism in Seedling Development

Light affects many aspects of plant development, including seed germination, stem elongation, and floral initiation. How photoreceptors control photomorphogenic processes is not yet fully understood. Because phytohormones are chemical regulators of plant development, it may not be surprising that light affects, directly or indirectly, cellular levels and signaling processes of various phytohormones, such as auxin, gibberellins （GA）, cytokinin, ethylene, abscisic acid （ABA）, and brassinosteroids （BR）. Among those phytohormones, light regulation of GA metabolism has probably attracted more attention among photoblologists and it is arguably the most extensively studied plant hormone at present with respect to its role in photomorphogenesis. It has become Increasingly clear that phytochromes and cryptochromes are the major photoreceptors mediating light regulation of GA homeostasis. This short article attempts to examine some recent developments in our understanding of how light and photoreceptors regulate GA blosynthesis and catabolism during seedling development. It is not our intention to carry out a comprehensive review of the field, and readers are referred to recent review articles for a more complete view of this area of study （Kamiya and Garcia-Martinez 1999; Hedden and Phillips 2000; Garcla-Martinez and GI12001; Olszewski et al. 2002; Halliday and Fankhauser 2003; Sun and Gubler 2004）.     

Genetic Dissection of the Relative Roles of Auxin and Gibberellin in the Regulation of Stem Elongation in Intact Light-Grown Peas

Exogenous gibberellin (GA) and auxin (indoleacetic acid [IAA]) strongly stimulated stem elongation in dwarf GA1-deficient le mutants of light-grown pea (Pisum sativum L.): IAA elicited a sharp increase in growth rate after 20 min followed by a slow decline; the GA response had a longer lag (3 h) and growth increased gradually with time. These responses were additive. The effect of GA was mainly in internodes less than 25% expanded, whereas that of IAA was in the older, elongating internodes. IAA stimulated growth by cell extension; GA stimulated growth by an increase in cell length and cell number. Dwarf lkb GA-response-mutant plants elongated poorly in response to GA (accounted for by an increase in cell number) but were very responsive to IAA. GA produced a substantial elongation in lkb plants only in the presence of IAA. Because lkb plants contain low levels of IAA, growth suppression in dwarf lkb mutants seems to be due to a deficiency in endogenous auxin. GA may enhance the auxin induction of cell elongation but cannot promote elongation in the absence of auxin. The effect of GA may, in part, be mediated by auxin. Auxin and GA control separate processes that together contribute to stem elongation. A deficiency in either leads to a dwarfed phenotype.     

Turning on gibberellin and abscisic acid signaling

The phytohormones gibberellin (GA) and abscisic acid (ABA) play essential and often antagonistic roles in regulating plant growth, development and stress responses. The long-awaited identification of receptors for both GA and ABA has shed light upon the initial events that surround the perception of these two phytohormones. The discovery of these receptors also challenges conventional views of plant hormone signaling and raises intriguing questions regarding the nature of GA and ABA perception and the initiation of their signaling pathways. Moreover, recent advances in understanding GA and ABA signaling point to the existence of multiple, non-linear cell- and compartment-specific pathways that regulate genomic and non-genomic responses to these phytohormones.     

Auxin Physiology of Dwarfism in Pisum sativum

Similar levels of diffusible auxin are measured for the apices of both Little Marvel (dwarf) and Alaska (normal) cultivars of the pea when grown in sunlight and darkness. In sunlight, however, diffusible auxin disappears in the subtending internode of the Little Marvel plant but remains at 50 per cent of the level of the apex in the subtending internode of the Alaska plant. The enzyme preparation from the apex of the dwarf plant converts tryptophan and tryptamine to IAA more readily than that from the normal plant. Indoleacetyl aspartate synthetase activity is also higher in the dwarf plant than in the normal plant and the dwarf plant contains four times as much conjugate as the normal plant with or without treatment with gibberellic acid. Gibberellic acid (GA) does not affect the induction of the synthetase enzyme nor the enzymatic formation of indoleacetyl aspartate. The growth induced by GA is the result of an increased synthesis of auxin.     

Effects of Growth Regulators on H+Translocation across the Membranes of Plasma Membrane Vesicles from Potato Tuber Cells

Effects of the growth regulators epibrassinolide-694 (EB), gibberellic acid (GA), and abscisic acid (ABA) on the ATP-dependent translocation of H⁺through the membranes of plasma membrane vesicles of potato (Solanum tuberosumL.) tuber cells were studied. The ATP-dependent accumulation of H⁺in the plasma membrane vesicles from dormant tubers was inhibited by EB and ABA and stimulated by GA. After the break of dormancy, the stimulatory effect of GA increased, the inhibitory effect of ABA decreased, and EB stimulated the accumulation of H⁺in the vesicles. The data suggest that the plasma membrane H⁺ATPase is a target of phytohormones that regulate the dormancy of potato tubers.     

Gibberellin Effect on Tryptophan Metabolism, Auxin Destruction, and Abscission in Coleus

Application of gibberellic acid (GA) to the apical region of the stem enhances ¹⁴CO₂ release from tryptophan-l-¹⁴C in cell free preparations of the apical region. Although GA when applied to the apical region markedly accelerates abscission rates of debladed petioles at the 4th node, the enhancement effect on tryptophan metabolism appears to be restricted to the apical bud region. The increased levels of diffusible auxin in Coleus stems, observed earlier by Muir and Valdovinos (1965), appear to be due to the GA effect on auxin precursor conversion rather than to an altered rate of auxin destruction. GA pre-treatment does not significantly alter destruction rates of auxin in the stem tissue. This is demonstrated by the release of ¹⁴CO₂ from IAA-1-¹⁴C by sections of internode tissue. While a multiple deblading pattern retards abscission of debladed petioles considerably, application of GA to debladed petioles at the basal region of the stem restores the normal rates of abscission at debladed distal nodes. No significant change in the abscission rates at treated nodes is observed. The GA effect on abscission at distal nodes is attributed to the effect of the growth substance on auxin precursor conversion in the apical region. In these experiments, as in the case of plants treated in the apical region with GA, auxin destruction rates in the stem are not altered significantly.     

Exogenous phytohormones and the induction of plant galls by insects

The mechanism of gall induction by insects has remained elusive. Previous studies have met with limited success in attempting to induce galls by injection or application of chemical compounds. To determine whether an exogenous source of phytohormones plays a role in gall induction, we injected cytokinin (CK), auxin (IAA), gibberellic acid (GA), and abscisic acid (ABA) in various concentrations, ratios, and combinations into leaf petioles of Capsicum annuum L. cv. California Wonder (Solanaceae). We found that CK + IAA injections lead to gall-like growth in C. annuum. GA enhanced and ABA inhibited gall growth except in the presence of GA. Isopentenyl adenine (IP) was the most effective type of CK at inducing growth. Our work is consistent with the hypothesis that exogenous CK + IAA produced and supplied by insects leads to gall induction. We hypothesize that insects have obtained the capability to induce galls via acquisition of the biosynthetic pathways to produce IAA and trans-zeatin family CKs through microbial symbiosis or lateral gene transfer.     

多效唑和脱落酸对白兰花内源激素及光合速率的影响

以白兰花为试材,采用土施、喷施多效唑和脱落酸的方式对其植株进行处理,观察株高和开花情况,同时结合主要内源激素含量及光合特性进行分析。结果表明：经处理后,白兰花内源激素IAA、GA3、Z、ZR和ABA含量呈现不同的变化规律,且光合速率也出现明显的变化;经处理后的植株明显矮化,盛花期提前,开花数量大幅度增加,且开花次数也显著增多。其中以土施和喷施多效唑与脱落酸的效果最好,处理后的植株只有正常株高的40%～50%,开花数却达到正常植株的2.3倍。     

Hormonal Regulation of Assimilate Utilization in Barley Leaves in Relation to the Development of Their Source Function

Growth, photosynthesis, utilization of assimilates, and the development of a source function in leaves were studied in relation to changes in concentrations and ratios of phytohormones. Carbon isotope ¹⁴C was used to trace utilization and outflow of photosynthetic products from the leaf. Concentrations of endogenous phytohormones were determined by solid-phase immunoenzyme assay. It was shown that, in juvenile leaves (one-fifth of their final area), which did not attain a high rate of photosynthesis, up to 80% of assimilates were incorporated into structural polysaccharides (cellulose and hemicellulose) one day after feeding with ¹⁴CO₂. During leaf growth and the development of its source function, the synthesis of structural polysaccharides declined to 10%, but the formation of alcohol- and water-soluble compounds (AWSC) grew to 80%. Monosaccharides and oligosaccharides, which could act as transport forms of carbohydrates, constituted 30% and 40% of AWSC, respectively. The percentage of assimilates utilized for protein synthesis decreased with leaf growth. The revealed changes correlate with the concentration and the ratio of free forms of phytohormones at various stages of leaf development. Development of a source function, a decline in cellulose and hemicellulose syntheses, and an increase in AWSC were related to the decrease in ABA and IAA concentrations and the increase in the ABA/IAA ratio. The ABA level can regulate the pathways of photoassimilate utilization in leaves by partitioning carbon flows either to the synthesis of high-molecular-weight compounds (cellulose, hemicellulose, and proteins), used for cell growth in leaves, or to the synthesis of transport forms of carbohydrates. A high ABA level favors the first pathway while low level switches leaf metabolism to the second one.     

Comprehensive transcriptome analysis of phytohormone biosynthesis and signaling genes in microspore/pollen and tapetum of rice

To investigate the involvement of phytohormones during rice microspore/pollen (MS/POL) development, endogenous levels of IAA, gibberellins (GAs), cytokinins (CKs) and abscisic acid (ABA) in the mature anther were analyzed. We also analyzed the global expression profiles of genes related to seven phytohormones, namely auxin, GAs, CKs, brassinosteroids, ethylene, ABA and jasmonic acids, in MS/POL and tapetum (TAP) using a 44K microarray combined with a laser microdissection technique (LM-array analysis). IAA and GA(4) accumulated in a much higher amount in the mature anther compared with the other tissues, while CKs and ABA did not. LM-array analysis revealed that sets of genes required for IAA and GA synthesis were coordinately expressed during the later stages of MS/POL development, suggesting that these genes are responsible for the massive accumulation of IAA and GA(4) in the mature anther. In contrast, genes for GA signaling were preferentially expressed during the early developmental stages of MS/POL and throughout TAP development, while their expression was down-regulated at the later stages of MS/POL development. In the case of auxin signaling genes, such mirror-imaged expression observed in GA synthesis and signaling genes was not observed. IAA receptor genes were mostly expressed during the late stages of MS/POL development, and various sets of AUX/IAA and ARF genes were expressed during the different stages of MS/POL or TAP development. Such cell type-specific expression profiles of phytohormone biosynthesis and signaling genes demonstrate the validity and importance of analyzing the expression of phytohormone-related genes in individual cell types independently of other cells/tissues.     

Control of growth by auxin and its specific neutral inhibitor

Evidence for the existence of a neutral inhibitor specific for auxin has been in the literature for 45 years. The inhibitor is demonstrable by its effect in causing positive curvature of theAvena coleoptile. The growth of the mesocotyl of oat and corn seedlings in darkness and its inhibition by light are determined by the neutral inhibitor as is the phototropic response of the sunflower stem. Production of the inhibitor is promoted by red and fluorescent light. Irradiance at 730 nm promotes auxin production while irradiance at 660 nm promotes production of the inhibitor. The positive curvature induced by 2,3,5-triiodobenzoic acid can be used to quantify the neutral inhibitor. Since the benzoic acid is more effective than iodoacetate in reacting with the sulfhydryl of cysteine, a sulfhydryl group is indicated to bo one reaction site for auxin and to be the basis of polar transport.     

Influence of Gibberellic Acid on Growth and Endogenous Auxin Level in Epicotyl and Hypocotyl Tissues of Normal and Dwarf Bean Plants

Application of gibberellic acid to the apex of dwarf bean plants (cv. Alabaster) stimulated the elongation growth of epicotyl and hypocotyl but showed no significant effect on elongation growth in a normal cultivar (‘Blue Lake’). Gibberellin-treatment of dwarf plants was characterized by about twofold increase in the level of endogenous auxin. Maximum increase in IAA level was observed after 48 h of GA treatment. There was less increase in IAA content in normal bean plants. — Gibberellin treatment to excised epicotyl and hypocotyl sections of either dwarf or normal cultivar showed no effect on elongation growth. However, a considerable increase in the auxin level was observed in the sections of the dwarf cultivar. The maximum effect occurred with only 1 h incubation in basal medium containing gibberellin. — The indolo-α-pyrone spectro-fluoremetric method for IAA determination was used.     

Effect of Growing Conditions on Wheat Hormonal Status and Productivity in Experimental Ecological System

The levels of free and bound forms of IAA, ABA, cytokinins (CK), and gibberellins, as well as growth characteristics and productivity were investigated in two wheat lines. The plants were grown under controlled conditions in an artificial ecosystem that allowed the irradiance, CO₂ concentration, and rhizosphere temperature to be changed. The main difference in the hormonal status of leaves of tall spring wheat, line 232, and dwarf wheat, line 95-3, was the absence of GA₉ gibberellins in the latter. It was found that the light intensity and temperature of rhizosphere insignificantly affected the balance of endogenous phytohormones and HI in wheat. The elevation of CO₂ concentration resulted in a considerable increase in the content of free IAA, an appearance of free GA₉, and a rise in the productivity of wheat, line 232. The concentration of CO₂ was shown to be a major parameter that determined HI in the experimental ecological system.     

Gibberellins and auxin regulate soybean hypocotyl elongation under low light and high-temperature interaction

Soybean is an important oilseed crop grown globally. However, two examples of environmental stresses that drastically regulate soybean growth are low light and high-temperature. Emerging evidence suggests a possible interconnection between these two environmental stimuli. Low light and high-temperature as individual factors have been reported to regulate plant hypocotyl elongation. However, their interactive signal effect on soybean growth and development remains largely unclear. Here, we report that gibberellins (GAs) and auxin are required for soybean hypocotyl elongation under low light and high-temperature interaction. Our analysis indicated that low light and high-temperature interaction enhanced the regulation of soybean hypocotyl elongation and that the endogenous GA₃, GA₇, indole-3-acetic acid (IAA), and indole-3-pyruvate (IPA) contents significantly increased. Again, analysis of the effect of exogenous phytohormones and biosynthesis inhibitors treatments showed that exogenous GA, IAA, and paclobutrazol (PAC), 2, 3, 5,-triiodobenzoic acid (TIBA) treatments significantly regulated soybean seedlings growth under low light and high-temperature interaction. Further qRT-PCR analysis showed that the expression level of GA biosynthesis pathway genes (GmGA3ox1, GmGA3ox2 and GmGA3) and auxin biosynthesis pathway genes (GmYUCCA3, GmYUCCA5 and GmYUCCA7) significantly increased under (i) low light and high-temperature interaction and (ii) exogenous GA and IAA treatments. Altogether, these observations support the hypothesis that gibberellins and auxin regulate soybean hypocotyl elongation under low light and high-temperature stress interaction.     

Phytohormones in fruit development and maturation

Phytohormones are integral to the regulation of fruit development and maturation. This review expands upon current understanding of the relationship between hormone signaling and fruit development, emphasizing fleshy fruit and highlighting recent work in the model crop tomato (Solanum lycopersicum) and additional species. Fruit development comprises fruit set initiation, growth, and maturation and ripening. Fruit set transpires after fertilization and is associated with auxin and gibberellic acid (GA) signaling. Interaction between auxin and GAs, as well as other phytohormones, is mediated by auxin-responsive Aux/IAA and ARF proteins. Fruit growth consists of cell division and expansion, the former shown to be influenced by auxin signaling. While regulation of cell expansion is less thoroughly understood, evidence indicates synergistic regulation via both auxin and GAs, with input from additional hormones. Fruit maturation, a transitional phase that precipitates ripening, occurs when auxin and GA levels subside with a concurrent rise in abscisic acid (ABA) and ethylene. During fruit ripening, ethylene plays a clear role in climacteric fruits, whereas non-climacteric ripening is generally associated with ABA. Recent evidence indicates varying requirements for both hormones within both ripening physiologies, suggesting rebalancing and specification of roles for common regulators rather than reliance upon one. Numerous recent discoveries pertaining to the molecular basis of hormonal activity and crosstalk are discussed, while we also note that many questions remain such as the molecular basis of additional hormonal activities, the role of epigenome changes, and how prior discoveries translate to the plethora of angiosperm species.     

Dynamics of growth and the content of endogenous phytohormones during kidney bean scoto-and photomorphogenesis

The dynamics of growth and the contents of free and bound endogenous IAA, gibberellins (GA), cytokinins (zeatin and its riboside), and ABA in kidney bean plants (Phaseolus vulgaris L., cv. Belozernaya) grown in darkness or in the light was studied. Phytohormones were quantified in 5–15-day-old plants by the ELISA technique. Plant growth and phytohormone content were shown to depend on plant age and the conditions of illumination. During scotomorphogenesis, changes in the biomass and hypocotyl length were highly correlated with the content of GA, whereas during photomorphogeneses, these parameters were correlated with the content of zeatin. In darkness, epicotyl growth displayed a positive correlation with the content of GA, whereas in the light, the correlation was negative. Growth characteristics of the primary leaves were shown to correlate with IAA in darkness and with GA and zeatin in the light. At a low concentration of cytokinins in illuminated leaves, cell divisions occurred, whereas, at the higher cytokinin concentrations, cell expansion occurred. The highest content of GA was characteristic of leaves in the period of growth cessation. ABA accumulated during active leaf and root elongation and biomass increment in the light and during hypocotyl growth in darkness. After plant illumination, the ratio of auxins to cytokinins increased in bean roots and decreased in their epicotyls. Thus, light changed the developmental programs of bean plants, which was manifested in the changed rate and duration of growth of various organs (root, hypocotyl, epicotyl, and leaf). Some mechanisms of light action depended on the contents of IAA, ABA, GA, and cytokinins and the ratios between these phytohormones. Differences between scotonorphogenesis of mono-and dicotyledonous plants are discussed in relation to the levels of phytohormones in them.     

Magnitude and Kinetics of Stem Elongation Induced by Exogenous Indole-3-Acetic Acid in Intact Light-Grown Pea Seedlings

Exogenously applied indole-3-acetic acid (IAA) strongly promoted stem elongation over the long term in intact light-grown seedlings of both dwarf (cv Progress No. 9) and tall (cv Alaska) peas (Pisum sativum L.), with the relative promotion being far greater in dwarf plants. In dwarf seedlings, solutions of IAA (between 10-4 and 10-3 M), when continuously applied to the uppermost two internodes via a cotton wick, increased whole-stem growth by at least 6-fold over the first 24 h. The magnitude of growth promotion correlated with the applied IAA concentration from 10-6 to 10-3 M, particularly over the first 6 h of application. IAA applied only to the apical bud or the uppermost internode of the seedling stimulated a biphasic growth response in the uppermost internode and the immediately lower internode, with the response in the latter being greatly delayed. This demonstrates that exogenous IAA effectively promotes growth as it is transported through intact stems. IAA withdrawal and reapplication at various times enabled the separation of the initial growth response (IGR) and prolonged growth response (PGR) induced by auxin. The IGR was inducible by at least 1 order of magnitude lower IAA concentrations than the PGR, suggesting that the process underlying the IGR is more sensitive to auxin induction. In contrast to the magnitude of the IAA effect in dwarf seedlings, applied IAA only doubled the growth in tall seedlings. These results suggest that endogenous IAA is more growth limiting in dwarf plants than in tall plants, and that auxin promotes stem elongation in the intact plant probably by the same mechanism of action as in isolated stem segments. However, since dwarf plants to which IAA was applied failed to reach the growth rate of tall plants, auxin cannot be the only limiting factor for stem growth in peas.     

The interaction of light irradiance with auxin in regulating growth of <Emphasis Type="Italic">Helianthus annuus</Emphasis> shoots

Plants growing under canopy shade or in near-neighboring proximity of taller vegetation are the receivers of shade light conditions. The effect of light irradiance (photosynthetically active radiation [PAR]), one of the main components of shade light, on the growth of various tissues of sunflower seedlings and the possible role of auxin were investigated. Gradual reductions in PAR irradiance level from near-normal to low and very low result in significant and gradual increases in sunflower hypocotyl growth and endogenous auxin content. Similar reductions in PAR level resulted in significant and gradual decreases in sunflower cotyledon and leaf growth, and endogenous auxin content. Exogenously applied auxin increased hypocotyl elongation under near-normal PAR, where IAA levels are below optimum, but decreased elongation under very low PAR, where IAA levels are already at optimum. These results suggests that auxin acts as positive growth regulator of sunflower hypocotyls subjected to low light irradiance stress. This is further supported by the transfer experiments where seedlings transferred, for example, from near-normal PAR to very low PAR showed increased elongation associated with increased IAA levels. Therefore, it is reasonable to conclude that light irradiance-mediated changes in hypocotyl elongation of young sunflower seedlings are regulated by endogenous auxin levels.