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New systems for agrochemical delivery in plants will foster precise agricultural practices and provide new tools to study plants and design crop traits, as standard spray methods suffer from elevated loss and limited access to remote plant tissues. Silk-based microneedles can circumvent these limitations by deploying a known amount of payloads directly in plants' deep tissues. However, plant response to microneedles' application and microneedles' efficacy in deploying physiologically relevant biomolecules are unknown. Here, it is shown that gene expression associated with Arabidopsis thaliana wounding response decreases within 24 h post microneedles' application. Additionally, microinjection of gibberellic acid (GA 3 ) in A. thaliana mutant ft-10 provides a more effective and efficient mean than spray to activate GA 3 pathways, accelerating bolting and inhibiting flower formation. Microneedle efficacy in delivering GA 3 is also observed in several monocot and dicot crop species, i.e., tomato (Solanum lycopersicum), lettuce (Lactuca sativa), spinach (Spinacia oleracea), rice (Oryza Sativa), maize (Zea mays), barley (Hordeum vulgare), and soybean (Glycine max). The wide range of plants that can be successfully targeted with microinjectors opens the doors to their use in plant science and agriculture., (© 2022 The Authors. Advanced Materials published by Wiley-VCH GmbH.)

There is some debate as to whether phytohormone metabolites should be classified as primary or secondary metabolites. Phytohormones have profound effects on growth - a typical trait of primary metabolites - yet several of them are formed from secondary metabolite precursors. This is further exacerbated by the blurred distinction between primary and secondary metabolism. What is clearer, however, is that phytohormones display distinctive regulatory mechanisms from other metabolites. Moreover, by contrast to microbial and mammalian systems, the majority of plant metabolite receptors characterized to date are hormone receptors. Here, we provide an overview of the metabolic links between primary metabolism and phytohormone biosynthesis in an attempt to complement recent reviews covering the signaling crosstalk between elements of core metabolism and the phytohormones. In doing so, we cover the biosynthesis of both the classical metabolic phytohormones namely auxins, salicylic acid, jasmonate, ethylene, cytokinins, brassinosteroids, gibberellins and abscisic acid as well as recently described plant growth regulators which have been proposed as novel phytohormones namely strigolactones blumenols, zaxinone and β-cyclocitral as well as melatonin. For each hormone, we describe the primary metabolite precursors which fuel its synthesis, act as conjugates or in the case of 2-oxoglutarate act more directly as a co-substrate in the biosynthesis of gibberellin, auxin and salicylic acid. Furthermore, several amino acids operate as hormone conjugates, such as jasmonate-conjugates. In reviewing the biosynthesis of all the phytohormones simultaneously, the exceptional intricacy of the biochemical interplay that underpins their interaction emerges., (Copyright © 2021 Elsevier GmbH. All rights reserved.)

Arbuscular mycorrhizal (AM) play an important role in improving plant growth and development. The interaction between phytohormones and AM symbiosis is gradually revealed. Here we examined the effect of Brassinosteroids (BR) on AM symbiosis and discussed the synergistic promotion of plant growth by BR and AM symbiosis. The xylophyta Eucalyptus grandis Hill (E. grandis) was inoculated with AM fungi Rhizoglomus irregularis R197198 (R. irregularis) and treated with different concentrations (0, 1, 10, and 100 nM) of 24-epibrassinolide (24-epiBL) for 6 weeks. With the increase of 24-epiBL concentration, E. grandis growth was firstly promoted and then inhibited, but inoculation with AM fungi alleviated this inhibition. 24-epiBL and R. irregularis colonization significantly improved E. grandis growth and antioxidant system response, and the synergistic effect was the best. Compared with the control group, 24-epiBL treatment significantly increased the mycorrhizal colonization and arbuscular abundance of AM fungi R. irregular in E. grandis roots. The expression of AM symbiosis maker genes was significantly increased by 24-epiBL treatment. Both 24-epiBL treatment and AM colonization upregulated gibberellins (GA) synthesis genes, but no inhibition caused by GA levels was found. 24-epiBL is a kind of synthetic highly active BR. Based on the results of 24-epiBL treatment, we hypothesized that BR actively regulates AM symbiosis regulates AM symbiosis without affecting GA-INSENSITIVE DWARF1 (GID1)-DELLA expression. The synergistic treatment of BR and AM symbiosis can significantly promote the growth and development of plants. IMPORTANCE Brassinosteroids (BR) and Arbuscular mycorrhizas (AM) symbiosis play an important role in improving plant growth and development. Previous studies have shown that there is a complex regulatory network between phytohormones and AM symbiosis. However, the interactions of BR-signaling and AM symbiosis are still poorly understood. Our results suggest that BR actively regulates the colonization and development of AM fungi, and AM fungal colonization can alleviate the inhibition of plant growth caused by excessive BR. In addition, BR actively regulates AM symbiosis, but does not primarily mediate gibberellins-DELLA interaction. The synergistic treatment of BR and AM symbiosis can significantly promote the growth and development of plants. The conclusions of this study provide a reference for phytohormones-AM symbiosis interaction.

Plant hormone signaling pathways have diversified during plant evolution. A new study reveals conservation of DELLA functions in growth and environmental stress responses across land plants., (Copyright © 2021 Elsevier Inc. All rights reserved.)

Chemical inhibitors are invaluable tools for investigating protein function in reverse genetic approaches. Their application bears many advantages over mutant generation and characterization. Inhibitors can overcome functional redundancy, their application is not limited to species for which tools of molecular genetics are available and they can be applied to specific tissues or developmental stages, making them highly convenient for addressing biological questions. The use of inhibitors has helped to elucidate hormone biosynthesis and signaling pathways and here we review compounds that were developed for the plant hormones brassinosteroids (BRs). BRs are steroids that have strong growth-promoting capacities, are crucial for all stages of plant development and participate in adaptive growth processes and stress response reactions. In the last two decades, impressive progress has been made in BR inhibitor development and application, which has been instrumental for studying BR modes of activity and identifying and characterizing key players. Both, inhibitors that target biosynthesis, such as brassinazole, and inhibitors that target signaling, such as bikinin, exist and in a comprehensive overview we summarize knowledge and methodology that enabled their design and key findings of their use. In addition, the potential of BR inhibitors for commercial application in plant production is discussed.

Plants are under relentless challenge by pathogenic bacteria, fungi, and oomycetes, for whom they provide a resource of living space and nutrients. Upon detection of pathogens, plants carry out multiple layers of defense response, orchestrated by a tightly organized network of hormones. In this review, we provide an overview of the phytohormones involved in immunity and the ways pathogens manipulate their biosynthesis and signaling pathways. We highlight recent developments, including the discovery of a defense signaling molecule, new insights into hormone biosynthesis, and the increasing importance of signaling hubs at which hormone pathways intersect., (Copyright © 2019 Elsevier Inc. All rights reserved.)

Calcium-dependent protein kinases (CPKs/CDPKs) are Ca 2+ -sensors that decode Ca 2+ signals into specific physiological responses. Research has reported that CDPKs constitute a large multigene family in various plant species, and play diverse roles in plant growth, development, and stress responses. Although numerous CDPKs have been exhaustively studied, and many of them have been found to be involved in plant hormone biosynthesis and response mechanisms, a comprehensive overview of the manner in which CDPKs participate in phytohormone signaling pathways, regulating nearly all aspects of plant growth, has not yet been undertaken. In this article, we reviewed the structure of CDPKs and the mechanism of their subcellular localization. Some CDPKs were elucidated to influence the intracellular localization of their substrates. Since little work has been done on the interaction between CDPKs and cytokinin signaling pathways, or on newly defined phytohormones such as brassinosteroids, strigolactones and salicylic acid, this paper mainly focused on discussing the integral associations between CDPKs and five plant hormones: auxins, gibberellins, ethylene, jasmonates, and abscisic acid. A perspective on future work is provided at the end., Competing Interests: The authors declare no conflict of interest.

Strigolactones (SLs) and gibberellins (GAs) are plant hormones that share some unique aspects of their perception and signalling pathways. Recent discoveries indicate that these two phytohormones may act together in processes of plant development and that SL biosynthesis is regulated by GAs., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Plant hormones are for a long time known to act as chemical messengers in the regulation of physiological processes during a plant's life cycle, from germination to senescence. Furthermore, plant hormones simultaneously coordinate physiological responses to biotic and abiotic stresses. To study the hormonal regulation of physiological processes, three main approaches have been used (1) exogenous application of hormones, (2) correlative studies through measurements of endogenous hormone levels, and (3) use of transgenic and/or mutant plants altered in hormone metabolism or signaling. A plant hormone profiling method is useful to unravel cross talk between hormones and help unravel the hormonal regulation of physiological processes in studies using any of the aforementioned approaches. However, hormone profiling is still particularly challenging due to their very low abundance in plant tissues. In this chapter, a sensitive, rapid, and accurate method to quantify all the five "classic" classes of plant hormones plus other plant growth regulators, such as jasmonates, salicylic acid, melatonin, and brassinosteroids is described. The method includes a fast and simple extraction procedure without time consuming steps as purification or derivatization, followed by optimized ultrahigh-performance liquid chromatography coupled to electrospray ionization-tandem mass spectrometry (UHPLC-MS/MS) analysis. This protocol facilitates the high-throughput analysis of hormone profiling and is applicable to different plant tissues.

The genus Penicillium is one of the most versatile "mycofactories", comprising some species able to produce gibberellins, bioactive compounds that can modulate plant growth and development. Although plants have the ability to synthesize gibberellins, their levels are lower when plants are under salinity stress. It has been recognized that detrimental abiotic conditions, such as saline stress, have negative effects on plants, being the availability of bioactive gibberellins a critical factor for their growth under this conditions. This review summarizes the interplay existing between endophytic Penicillium strains and plant host interactions, with focus on bioactive gibberellins production as a fungal response that allows plants to overcome salinity stress., (Copyright © 2015 Elsevier GmbH. All rights reserved.)

Plant phenotypic plasticity is controlled by diverse hormone pathways, which integrate and convey information from multiple developmental and environmental signals. Moreover, in plants many processes such as growth, development, and defense are regulated in similar ways by multiple hormones. Among them, gibberellins (GAs) are phytohormones with pleiotropic actions, regulating various growth processes throughout the plant life cycle. Previous work has revealed extensive interplay between GAs and other hormones, but the molecular mechanism became apparent only recently. Molecular and physiological studies have demonstrated that DELLA proteins, considered as master negative regulators of GA signaling, integrate multiple hormone signaling pathways through physical interactions with transcription factors or regulatory proteins from different families. In this review, we summarize the latest progress in GA signaling and its direct crosstalk with the main phytohormone signaling, emphasizing the multifaceted role of DELLA proteins with key components of major hormone signaling pathways., (Copyright © 2016 The Author. Published by Elsevier Inc. All rights reserved.)

We now have the various genomics tools for monocot (Oryza sativa) and a dicot (Arabidopsis thaliana) plant. Plant is not only a very important agricultural resource but also a model organism for biological research. It is important that the interaction between ABA and GA is investigated for controlling the transition from embryogenesis to germination in seeds using genomics tools. These studies have investigated the relationship between dormancy and germination using genomics tools. Genomics tools identified genes that had never before been annotated as ABA- or GA-responsive genes in plant, detected new interactions between genes responsive to the two hormones, comprehensively characterized cis-elements of hormone-responsive genes, and characterized cis-elements of rice and Arabidopsis. In these research, ABA- and GA-regulated genes have been classified as functional proteins (proteins that probably function in stress or PR tolerance) and regulatory proteins (protein factors involved in further regulation of signal transduction). Comparison between ABA and/or GA-responsive genes in rice and those in Arabidopsis has shown that the cis-element has specificity in each species. cis-Elements for the dehydration-stress response have been specified in Arabidopsis but not in rice. cis-Elements for protein storage are remarkably richer in the upstream regions of the rice gene than in those of Arabidopsis.

Cell culture of Taxus spp. represents a potential alternative source of taxol and related taxanes used in cancer chemotherapy. We have analyzed the effect of different culture media components on growth and production of taxol in developing callus cultures of T. cuspidata. Several sequential modifications were made to the basal B5 medium, which included addition and/or variation in the concentration of sucrose, B5 organic supplements, gibberellic acid, 36 combinations of 2,4-D/kinetin ratios, media salts and organic supplements, phenylalanine, casein hydrolysate and medium pH. The experiments were conducted during a 55 day-growth period followed by taxane extraction and analysis. Significant increases in taxol yield and growth over basal medium grown calli were observed with some of the modified media.

Gibberellic acid promotes doubling, early flowering, and fruiting directly in Impatiens. It also lengthens the stem, probably through a neutralization of auxin inhibitors. Such a mechanism would leave the growth-promoting auxins unchecked to produce the increased elongation characteristic of gibberellin application. It seems that application of additional auxin produces a rapid build-up of the auxin inhibitor(s) in the plant.

The subcellular distribution of the enzymes alpha-amylase, protease and ribonuclease in wheat aleurone layers after treatment with gibberellic acid was determined by differential centrifugation. Of the alpha-amylase 56% was precipitable from cell homogenates, indicating that it is a particulate enzyme. Similar results were recorded with protease. Particulate alpha-amylase showed distinct structural latency, and membrane-rupturing mechanical or chemical treatments were required to release the enzyme in an active form; the results were completely analogous to results with lysosomal enzymes found in animal tissues. The identification of the hormonally induced enzymes as lysosomal suggests that the hormonal mechanism may be more closely associated with extracellular enzyme synthesis rather than with nucleic acid metabolism.

Gibberellic acid production from various substrates was studied in 43 strains of Fusarium, among which F. moniliforme strain IOC-3326 was selected as the best producer. Experiments were carried out in shaker flasks and pilot plant fermentors. The results indicate that the best substrate for gibberellic acid production with this strain is composed of the following: glucose, 20 g; corn steep liquor, 25 g; ammonium nitrate, 2.6 g; monopotassium phosphate, 0.5 g; potassium sulfate, 0.2 g; and water, 1000 ml. Glucose, ammonium nitrate, and corn steep liquor were found to be critical. With this medium, maximal yields of 1196 mg per liter in shaker flasks and 997 mg per liter in fermentors were produced.

Plant hormones are signalling compounds that regulate crucial aspects of growth, development and environmental stress responses. Abiotic stresses, such as drought, salinity, heat, cold and flooding, have profound effects on plant growth and survival. Adaptation and tolerance to such stresses require sophisticated sensing, signalling and stress response mechanisms. In this Review, we discuss recent advances in understanding how diverse plant hormones control abiotic stress responses in plants and highlight points of hormonal crosstalk during abiotic stress signalling. Control mechanisms and stress responses mediated by plant hormones including abscisic acid, auxin, brassinosteroids, cytokinins, ethylene and gibberellins are discussed. We discuss new insights into osmotic stress sensing and signalling mechanisms, hormonal control of gene regulation and plant development during stress, hormone-regulated submergence tolerance and stomatal movements. We further explore how innovative imaging approaches are providing insights into single-cell and tissue hormone dynamics. Understanding stress tolerance mechanisms opens new opportunities for agricultural applications.

Seed germination is a fundamental process in the plant life cycle and is regulated by functionally opposing internal and external inputs. Here we explored the role of a negative regulator of photomorphogenesis, a B-box-containing protein (BBX19), as a molecular link between the inhibitory action of the phytohormone abscisic acid (ABA) and the promoting role of light in germination. We show that seeds of BBX19-overexpressing lines, in contrast to those of BBX19 RNA interference lines, display ABA hypersensitivity, albeit independently of elongated hypocotyl 5 (HY5). Moreover, we establish that BBX19 functions neither via perturbation of GA signaling, the ABA antagonistic phytohormone, nor through interference with the DELLA protein germination repressors. Rather, BBX19 functions as an inducer of ABA INSENSITIVE5 (ABI5) by binding to the light-responsive GT1 motifs in the gene promoter. In summary, we identify BBX19 as a regulatory checkpoint, directing diverse developmental processes and tailoring adaptive responses to distinct endogenous and exogenous signals.

The activation of the meristem identity gene VERNALIZATION1 (VRN1) is a critical regulatory point in wheat (Triticum spp.) flowering. In photoperiod-sensitive wheat varieties, VRN1 is expressed only under long days (LDs), but mutants carrying deletions in a regulatory element in its promoter show VRN1 transcription and early spike development under short days (SDs). However, complete spike development is delayed until plants are transferred to LDs, indicating the existence of an additional regulatory mechanism dependent on LDs. We show here that exogenous gibberellin (GA) application accelerates spike development under SDs, but only in wheat lines expressing VRN1. The simultaneous presence of GA and VRN1 results in the up-regulation of the floral meristem identity genes SUPPRESSOR OF OVEREXPRESSION OF CONSTANS1-1 and LEAFY, whereas inhibition of GA biosynthesis with paclobutrazol precludes the LD induction of these two genes. The inductive role of GA on wheat flowering is further supported by the up-regulation of GA biosynthetic genes in the apices of plants transferred from SDs to LDs and in photoperiod-insensitive and transgenic wheat plants with increased FLOWERING LOCUS T transcription under SDs. The up-regulation of GA biosynthetic genes was not observed in the leaves of the same genetic stocks. Based on these observations, we propose a model in which FLOWERING LOCUS T is up-regulated in the leaves under LDs and is then transported to the shoot apical meristem, where it simultaneously induces the expression of VRN1 and GA biosynthetic genes, which are both required for the up-regulation of the early floral meristem genes SUPPRESSOR OF OVEREXPRESSION OF CONSTANS1-1 and LEAFY and the timely development of the wheat spike.

Background: In Citrus spp., gibberellic acid (GA) has been proposed to improve different processes related to crop cycle and yield. Accordingly, many studies have been published about how GA affects flowering and fruiting. Nevertheless, some such evidence is contradictory and the use of GA applications by farmers are still confusing and lack the expected results. Purpose: This review aims to collate, present, analyze and synthesize the most relevant empirical evidence to answer the following questions: (i) how does gibberellic acid act on flowering and fruiting of citrus trees?; (ii) why is all this knowledge sometimes not correctly used by farmers to solve yield problems relating to flowering and fruit set? Methods: An extensive literature search to obtain a large number of records about the topic was done. Searches were done in five databases: WoS, Scopus, Google Academics, PubMed and Scielo. The search string used was "Gibberellic acid" AND "Citrus". Records were classified into 11 groups according to the development process they referred to and initial data extraction was done. Records related with flowering and fruit set were drawn, and full texts were screened. Fifty-eight full text records were selected for the final data extraction. Results: Selected studies were published from 1959 to 2017 and were published mainly in Spain, USA, Brazil and Japan. Twelve species were studied, and Citrus sinensis, C. reticulata and C. unshiu were the principal ones. Most publications with pre-flowering treatments agreed that GA decreases flowering, while only 3 out of 18 did not observe any effect. In most of these studies, the effect on fruit set and yield was not evaluated. Studies with treatments at full bloom or some weeks later mostly reported increased fruit set. However, these increases did not imply higher yields. The results on yield were highly erratic as we found increases, decreases, no effects or variable effects. Conclusions: Despite some limitations, the action of GA related to cell division and growth, stimulating the sink ability of the organ and discouraging its abscission, has been clearly established through reviewed studies. GA applications before flowering counteract the floral induction caused by stress reducing flowering. However, on adult trees under field conditions, reducing flowering by applying GA would be difficult because it would be necessary to previously estimate the natural floral induction of trees. During flowering and fruit set, many problems may arise that limit production. Only when the problem is lack of fruit set stimulus can GA applications improve yields. However, much evidence suggests that the main factor-limiting yield would be carbohydrate availability rather than GA levels. GA applications increased fruit set (often transiently), but this increase did not mean improved yields. [ABSTRACT FROM AUTHOR]

Seed germination is considered the beginning of the spermatophyte lifecycle, and it is a crucial stage in determining subsequent plant growth and development. Although many previous studies have found that melatonin can promote seed germination, the role of melatonin in cotton germination remains unexamined. The main objective of this study is the characterization of potential promotional effects of melatonin (at doses of 0, 10, 20, 50, 100 and 200 μM) on cotton seed germination. This experiment demonstrated that low concentrations of melatonin can promote germination, while high concentrations failed to promote germination and even inhibited germination. Together, these results indicate that a 20 μM melatonin treatment optimally promotes cotton seed germination. Compared with the control, germination potential (GP), germination rate (GR), and final fresh weight (FW) increased by 16.67%, 12.30%, and 4.81%, respectively. Although low concentrations of melatonin showed some improvement in vigor index (VI), germination index (GI), and mean germination time (MGT), these effects were not statistically significant. Antioxidant enzyme activity during seed germination was most prominent under the 20 μM melatonin treatment. Superoxide dismutase (SOD) and peroxidase (POD) activities were significantly increased by 10.37–59.73% and 17.79–47.68%, respectively, compared to the melatonin-free control. Malondialdehyde (MDA) content was reduced by 16.73–40.33%. Two important plant hormones in seed germination, abscisic acid (ABA) and gibberellins (GAs), were also studied. As melatonin concentration increased, ABA content in seeds decreased first and then increased, and GA3 content showed a diametrically opposite trend, in which the 20 μM melatonin treatment was optimal. The 20 μM melatonin treatment reduced ABA content in seeds by 42.13–51.68%, while the 20 μM melatonin treatment increased GA3 content in seeds to about 1.7–2.5 times that of seeds germinated without melatonin. This study provides new evidence suggesting that low concentrations of melatonin can promote cotton seed germination by increasing the activity of antioxidant enzymes, thereby reducing the accumulation of MDA and regulating plant hormones. This has clear applications for improving the germination rate of cotton seeds using melatonin. [ABSTRACT FROM AUTHOR]