Class Of Plant Hormones Research Articles

Structural diversity in the strigolactones.

Strigolactones (SLs) are a class of signalling molecules secreted by the roots of plants into the rhizosphere. On the one hand, they serve as the signal for recruiting arbuscular mycorrhizal fungi which have a symbiotic relationship with plants. On the other hand, they are also host detection signals for the non-symbiotic, pathogenic, root parasitic plants, which use the SLs as germination stimulants. Finally, recently the SLs were discovered to be a new class of plant hormones that regulate processes such as branching/tillering and root architecture. Intriguingly, >25 different SLs have already been discovered that all have the so-called D-ring but otherwise display many differences in structure and functional groups. In this review, we will critically discuss the structural diversity in the SLs. How are they synthesized in plants; how has this structural diversity possibly evolved; what is the biological relevance of this diversity; and what does this imply for the perception of the SLs by receptors in the plant itself and in other organisms? Finally, we conclude that only little is known about the biological significance of this structural diversity, and we will give an outlook on how to elucidate their importance further.

Conversion of carlactone to carlactonoic acid is a conserved function of MAX1 homologs in strigolactone biosynthesis.

Strigolactones (SLs) are a class of plant hormones which regulate shoot branching and function as host recognition signals for symbionts and parasites in the rhizosphere. However, steps in SL biosynthesis after carlactone (CL) formation remain elusive. This study elucidated the common and diverse functions of MAX1 homologs which catalyze CL oxidation. We have reported previously that ArabidopsisMAX1 converts CL to carlactonoic acid (CLA), whereas a rice MAX1 homolog has been shown to catalyze the conversion of CL to 4-deoxyorobanchol (4DO). To determine which reaction is conserved in the plant kingdom, we investigated the enzymatic function of MAX1 homologs in Arabidopsis, rice, maize, tomato, poplar and Selaginella moellendorffii. The conversion of CL to CLA was found to be a common reaction catalyzed by MAX1 homologs, and MAX1s can be classified into three types: A1-type, converting CL to CLA; A2-type, converting CL to 4DO via CLA; and A3-type, converting CL to CLA and 4DO to orobanchol. CLA was detected in root exudates from poplar and Selaginella, but not ubiquitously in other plants examined in this study, suggesting its role as a species-specific signal in the rhizosphere. This study provides new insights into the roles of MAX1 in endogenous and rhizosphere signaling.

Gibberellin reduces the susceptibility of rice, Oryza sativa, to the migratory nematode Hirschmanniella oryzae

Summary Upon pathogen attack, the plant defence response is mediated by a set of connected signal transduction pathways, guided by several classes of plant hormones. In this study, experiments were conducted to observe the role of the plant hormone gibberellic acid in the response of rice to infection by the migratory root-rot nematode Hirschmanniella oryzae. Foliar treatments with gibberellic acid showed a negative effect on H. oryzae infection in the roots. Analyses of mutant rice lines impaired in the production or signalling of gibberellic acid confirmed the effect of the plant hormone on H. oryzae infection. Taken together, the results clearly indicate that gibberellic acid has a positive effect on the capability of the rice plant to fend off an infection by the migratory nematode H. oryzae.

Evidence for species-dependent biosynthetic pathways for converting carlactone to strigolactones in plants.

Strigolactones (SLs), comprising compounds with diverse but related chemical structures, are determinant signals in elicitation of germination in root parasitic Orobanchaceae and in mycorrhization in plants. Further, SLs are a novel class of plant hormones that regulate root and shoot architecture. Dissecting common and divergent biosynthetic pathways of SLs may provide avenues for modulating their production in planta. Biosynthesis of SLs in various SL-producing plant species was inhibited by fluridone, a phytoene desaturase inhibitor. The plausible biosynthetic precursors of SLs were exogenously applied to plants, and their conversion to canonical and non-canonical SLs was analysed using liquid chromatography-tandem mass spectrometry. The conversion of carlactone (CL) to carlactonoic acid (CLA) was a common reaction in all investigated plants. Sorghum converted CLA to 5-deoxystrigol (5-DS) and sorgomol, and 5-DS to sorgomol. One sorgomol-producing cotton cultivar had the same SL profile as sorghum in the feeding experiments. Another cotton cultivar converted CLA to 5-DS, strigol, and strigyl acetate. Further, 5-DS was converted to strigol and strigyl acetate. Moonseed converted CLA to strigol, but not to 5-DS. The plant did not convert 5-DS to strigol, suggesting that 5-DS is not a precursor of strigol in moonseed. Similarly, 4-deoxyorobanchol was not a precursor of orobanchol in cowpea. Further, sunflower converted CLA to methyl carlactonoate and heliolactone. These results indicated that the biosynthetic pathways of hydroxy SLs do not necessarily involve their respective deoxy SL precursors.

Recent advances in molecular basis for strigolactone action.

Strigolactones (SLs) are a very special class of plant hormones, which act as endogenous signals to regulate shoot branching in plants, and also serve as rhizosphere signals to regulate interactions of host plants with heterologous organisms such as symbiotic arbuscular mycorrhizal fungi and parasitic weeds. In this short review, we give a brief description of novel discoveries in SL biosynthesis pathway, and mainly summarize the recent advances in SL perception and signal transduction.

Strigolactones Biosynthesis and Their Role in Abiotic Stress Resilience in Plants: A Critical Review.

Strigolactones (SLs), being a new class of plant hormones, play regulatory roles against abiotic stresses in plants. There are multiple hormonal response pathways, which are adapted by the plants to overcome these stressful environmental constraints to reduce the negative impact on overall crop plant productivity. Genetic modulation of the SLs could also be applied as a potential approach in this regard. However, endogenous plant hormones play central roles in adaptation to changing environmental conditions, by mediating growth, development, nutrient allocation, and source/sink transitions. In addition, the hormonal interactions can fine-tune the plant response and determine plant architecture in response to environmental stimuli such as nutrient deprivation and canopy shade. Considerable advancements and new insights into SLs biosynthesis, signaling and transport has been unleashed since the initial discovery. In this review we present basic overview of SL biosynthesis and perception with a detailed discussion on our present understanding of SLs and their critical role to tolerate environmental constraints. The SLs and abscisic acid interplay during the abiotic stresses is particularly highlighted.Main Conclusion: More than shoot branching Strigolactones have uttermost capacity to harmonize stress resilience.

Evolution of strigolactone receptors by gradual neo-functionalization of KAI2 paralogues

BackgroundStrigolactones (SLs) are a class of plant hormones that control many aspects of plant growth. The SL signalling mechanism is homologous to that of karrikins (KARs), smoke-derived compounds that stimulate seed germination. In angiosperms, the SL receptor is an α/β-hydrolase known as DWARF14 (D14); its close homologue, KARRIKIN INSENSITIVE2 (KAI2), functions as a KAR receptor and likely recognizes an uncharacterized, endogenous signal (‘KL’). Previous phylogenetic analyses have suggested that the KAI2 lineage is ancestral in land plants, and that canonical D14-type SL receptors only arose in seed plants; this is paradoxical, however, as non-vascular plants synthesize and respond to SLs.ResultsWe have used a combination of phylogenetic and structural approaches to re-assess the evolution of the D14/KAI2 family in land plants. We analysed 339 members of the D14/KAI2 family from land plants and charophyte algae. Our phylogenetic analyses show that the divergence between the eu-KAI2 lineage and the DDK (D14/DLK2/KAI2) lineage that includes D14 occurred very early in land plant evolution. We show that eu-KAI2 proteins are highly conserved, and have unique features not found in DDK proteins. Conversely, we show that DDK proteins show considerable sequence and structural variation to each other, and lack clearly definable characteristics. We use homology modelling to show that the earliest members of the DDK lineage structurally resemble KAI2 and that SL receptors in non-seed plants likely do not have D14-like structure. We also show that certain groups of DDK proteins lack the otherwise conserved MORE AXILLARY GROWTH2 (MAX2) interface, and may thus function independently of MAX2, which we show is highly conserved throughout land plant evolution.ConclusionsOur results suggest that D14-like structure is not required for SL perception, and that SL perception has relatively relaxed structural requirements compared to KAI2-mediated signalling. We suggest that SL perception gradually evolved by neo-functionalization within the DDK lineage, and that the transition from KAI2-like to D14-like protein may have been driven by interactions with protein partners, rather than being required for SL perception per se.

Identification and Functional Analysis of Two Cotton Orthologs of MAX2 Which Control Shoot Lateral Branching

Cotton (Gossypium spp.), as the most important fiber and oilseed crop in the world, is extremely important for the industry. However, due to its indeterminate growth habit and complex branching system, massive labor costs are needed for shoot apex removal and branch pruning during cotton production. Therefore, it is very important to explore branch-controlling genes and genetically modify the branch architecture of cotton. Strigolactones (SLs) are a novel class of plant hormone that inhibit the outgrowth of lateral branches. To elucidate the role of SLs in branch development of cotton, we cloned and characterized GhMAX2a and GhMAX2b from tetraploid upland cotton (Gossypium hirsutum), the orthologs of Arabidopsis MAX2, rice D3, and petunia RMS4. GhMAX2a/2b was ubiquitously expressed in all tested tissues of cotton, with relatively higher expression levels in leaves and lateral buds. Subcellular localization assay showed that the GhMAX2-GFP fusion protein localized to the nucleus. Both GhMAX2a and GhMAX2b can fully rescue the dwarfed and highly branched phenotypes of the Arabidopsis max2-1 mutant, indicating that GhMAX2s have conserved functions with that of AtMAX2. The cotton GhMAX2b interacted with Arabidopsis Skp1-like 1 (ASK1) proteins in vitro which was further confirmed in the Arabidopsis protoplasts using the co-immunoprecipitation assay, indicating that GhMAX2b probably functions through forming an SCF E3 complex with Skp and other proteins in the Arabidopsis. These results suggest that the cotton GhMAX2s encode functional MAX2 that can inhibit the shoot lateral branching. Further functional analysis of GhMAX2s in determining cotton branch architecture and yield is underway.

A Taylor-Made Design of Phenoxyfuranone-Type Strigolactone Mimic.

Strigolactones are a class of plant hormones that inhibit axillary bud outgrowth and are released from plant roots to act as a rhizosphere communication signal. The Orobanchaceae parasitic plant Striga hermonthica perceives strigolactone as its germination signal, indicating host presence. After germination, the Striga plant parasitises the host plant and suppresses host growth by draining photosynthetic products, water and other essential nutrients. Because of this way of life, this parasite threatens crop production in sub-Saharan Africa with infestation in crop fields and crop devastation. Crop protection in such areas is among the most concerning problems to be dealt with as immediately as possible. With respect to crop protection from Striga, many strigolactone agonists have been developed and used in research to reveal Striga biology, and have contributed to development of crop protection methods. However, an effective method has yet to be found. In a previous study, we reported debranones as a group of strigolactone mimics that inhibit axillary buds outgrowth with moderate stimulation activity for Striga germination. Debranones would be accessible because they are simply synthesized from commercially available phenols and bromo butenolide. Taking this advantage of debranones for Striga research, we tried to find the debranones stimulating Striga seed germination. To modulate functional selectivity and to enhance germination inducing activity of debranones, we studied structure–activity relationships. We investigated effects of substituent position and functional group on debranone activity and selectivity as a strigolactone mimic. As a result, we improved stimulation activity of debranones for Striga seed germination by chemical modification, and demonstrated the pharmacophore of debranones for selective modulation of distinct strigolactone responses.

Gibberellins Promote Brassinosteroids Action and Both Increase Heterosis for Plant Height in Maize (Zea mays L.).

Brassinosteroids (BRs) and Gibberellins (GAs) are two classes of plant hormones affecting plant height (PHT). Thus, manipulation of BR and GA levels or signaling enables optimization of crop grain and biomass yields. We established backcross (BC) families, selected for increased PHT, in two elite maize inbred backgrounds. Various exotic accessions used in the germplasm enhancement in maize project served as donors. BC1-derived doubled haploid lines in the same two elite maize inbred backgrounds established without selection for plant height were included for comparison. We conducted genome-wide association studies to explore the genetic control of PHT by BR and GA. In addition, we used BR and GA inhibitors to compare the relationship between PHT, BR, and GA in inbred lines and heterozygotes from a physiological and biological perspective. A total of 73 genomic loci were discovered to be associated with PHT, with seven co-localized with GA, and two co-localized with BR candidate genes. PHT determined in field trials was significantly correlated with seedling stage BR and GA inhibitor responses. However, this observation was only true for maize heterozygotes, not for inbred lines. Path analysis results suggest that heterozygosity increases GA levels, which in turn promote BR levels. Thus, at least part of heterosis for PHT in maize can be explained by increased GA and BR levels, and seedling stage hormone inhibitor response is promising to predict heterosis for PHT.

Determination of the best 24-eBL application for the accumulation of antioxidant compounds in ‘Alphonse Lavallée’ grape cultivar by using fuzzy logic modelling

Brassinosteroids (BRs) are a class of polyhydroxysteroids that have been recognized as a “sixth class of plant hormones”. At present, 70 analogs have been identified, and among these, 24-epibrassinolide (24-eBL) is known to have an economic impact on plant metabolism, growth and productivity, and experience high stability under field conditions. Exogenous application of BRs are reported to confer resistance to plants against various abiotic stresses like heat, drought, heavy metals, salt and biotic stress like fungi and even viruses. Apart from resistance to biotic stresses, BRs also affect a broad spectrum of physiological responses like cell expansion, vascular differentiation, reproductive development, seed germination, flowering, fruit set and synthesis of antioxidant compounds in plants. In this study, the aim was to determine the best pre- harvest 24-eBL application for the accumulation of antioxidant compounds in 'Alphonse Lavallee' grape cultivar by using fuzzy logic modelling (FLM). For this aim, 24-eBL was applied to 9 years-old vines with 13 combinations including different application times that are our first input variables and 4 different concentrations (0.2, 0.4, 0.6 and 0.8 ppm) which are second input variables of our fuzzy logic model and control. Total phenolic content, total anthocyanin, trans-resveratrol, β-carotene and ascorbic acid were examined as antioxidant compounds in grapes. One, some or all of these can be used as output variables of our FLM to get the optimum result. By making the data continuous the model can be used for obtaining useful information about untested data so that the best solution, that is the best 24-eBL concentrations and application times, can be attained at some non-measured input variables for each antioxidant compound.

Regulation of Strigolactone Biosynthesis by Gibberellin Signaling.

Strigolactones (SLs) are a class of plant hormones that regulate diverse physiological processes, including shoot branching and root development. They also act as rhizosphere signaling molecules to stimulate the germination of root parasitic weeds and the branching of arbuscular mycorrhizal fungi. Although various types of cross talk between SLs and other hormones have been reported in physiological analyses, the cross talk between gibberellin (GA) and SLs is poorly understood. We screened for chemicals that regulate the level of SLs in rice (Oryza sativa) and identified GA as, to our knowledge, a novel SL-regulating molecule. The regulation of SL biosynthesis by GA is dependent on the GA receptor GID1 and F-box protein GID2. GA treatment also reduced the infection of rice plants by the parasitic plant witchers weed (Striga hermonthica). These data not only demonstrate, to our knowledge, the novel plant hormone cross talk between SL and GA, but also suggest that GA can be used to control parasitic weed infections.

When the story proceeds backward: The discovery of endogenous β-aminobutyric acid as the missing link for a potential new plant hormone

ABSTRACTThe capacity of β-aminobutyric acid (BABA) to induce resistance in plants against biotic and abiotic stresses has been known for more than 50 y. In the beginning reports were mainly descriptive of the phenomenon, but it became clear with the discovery of BABA insensitive mutants in Arabidopsis that there was definitely a genetic basis underlying BABA-induced resistance. The study of these mutants, along with the use of regular hormone mutants, allowed establishing the defense pathways activated upon defense induction. To date it is clear that BABA potentiates the defense pathway more appropriate to counteract the upcoming stress situation, through a phenomenon termed priming. Interestingly, plants possess a receptor for BABA, but up to recently there was a general consensus on the fact that BABA was a xenobiotic molecule. The development of an accurate non-destructive assay for measuring aminobutyric acid isomers in planta and the finding of plant-produced BABA, thus seems to represent the missing link for the discovery of a novel plant hormone. Differences and similarities with some of the classical plant hormones are presented here.

Strigolactone Signaling and Evolution

Strigolactones are a structurally diverse class of plant hormones that control many aspects of shoot and root growth. Strigolactones are also exuded by plants into the rhizosphere, where they promote symbiotic interactions with arbuscular mycorrhizal fungi and germination of root parasitic plants in the Orobanchaceae family. Therefore, understanding how strigolactones are made, transported, and perceived may lead to agricultural innovations as well as a deeper knowledge of how plants function. Substantial progress has been made in these areas over the past decade. In this review, we focus on the molecular mechanisms, core developmental roles, and evolutionary history of strigolactone signaling. We also propose potential translational applications of strigolactone research to agriculture.

Gibberellins regulate iron deficiency-response by influencing iron transport and translocation in rice seedlings (Oryza sativa).

Gibberellins (GAs) are a class of plant hormones with diverse functions. However, there has been little information on the role of GAs in response to plant nutrient deficiency. To evaluate the roles of GAs in regulation of Fe homeostasis, the effects of GA on Fe accumulation and Fe translocation in rice seedlings were investigated using wild-type, a rice mutant ( eui1 ) displaying enhnaced endogenous GA concentrations due to a defect in GA deactivation, and transgenic rice plants overexpressing OsEUI . Exposure to Fe-deficient medium significantly reduced biomass of rice plants. Both exogenous application of GA and an endogenous increase of bioactive GA enhanced Fe-deficiency response by exaggerating foliar chlorosis and reducing growth. Iron deficiency significantly suppressed production of GA 1 and GA 4 , the biologically active GAs in rice. Exogenous application of GA significantly decreased leaf Fe concentration regardless of Fe supply. Iron concentration in shoot of eui1 mutants was lower than that of WT plants under both Fe-sufficient and Fe-deficient conditions. Paclobutrazol, an inhibitor of GA biosynthesis, alleviated Fe-deficiency responses, and overexpression of EUI significantly increased Fe concentration in shoots and roots. Furthermore, both exogenous application of GA and endogenous increase in GA resulting from EUI mutation inhibited Fe translocation within shoots by suppressing OsYSL2 expression, which is involved in Fe transport and translocation. The novel findings provide compelling evidence to support the involvement of GA in mediation of Fe homeostasis in strategy II rice plants by negatively regulating Fe transport and translocation.

Heck functionalization of an asymmetric aza-BODIPY core: synthesis of far-red infrared probes for bioimaging applications.

As part of our ongoing work on the synthesis of a new class of plant hormones named Strigolactones (SLs) and their analogues, we became interested in tracing bioactive molecules with red emitting BODIPY fluorophores in order to unravel signaling and distribution of SLs in plants. In this paper we report on an unprecedented Heck functionalization of azadipyrromethenes (aza-DIPY) which allows for the introduction of suitable functional groups to convert aza-BODIPY in bioconjugate complexes useful for untangling biological processes.

Hormone Profiling in Plant Tissues.

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.

Imaging TCSn::GFP, a Synthetic Cytokinin Reporter, in Arabidopsis thaliana.

Cytokinins are classical plant hormones that control numerous developmental processes throughout the plant life cycle. Cytokinin-responsive cells activate transcription via a phospho-relay signaling network. Type-B nuclear RESPONSE REGULATOR (RR) proteins mediate transcriptional activation as the final step in the signaling cascade. They bind to promoters of immediate-early target genes via a conserved Myb-related DNA-binding domain. To monitor transcriptional activation in response to a cytokinin stimulus, we have constructed a synthetic promoter, TCS (two-component signaling sensor) that harbors the concatemerized binding motifs for activated type-B RR in an optimized configuration. Here, we describe our protocols for imaging TCSn::GFP expression in transgenic Arabidopsis plants. The use of the fluorescent reporter GFP allows the visualization of cytokinin-responding cells by fluorescent microscopy without the need for tissue processing steps, or staining reactions. This method is fast and with a low risk of artifacts. However, since cytokinin signaling integrates various environmental information including light, nutrient status, and biotic and abiotic stress, special care needs to be devoted to the control of growth conditions.

Low-Phosphate Induction of Plastidal Stromules Is Dependent on Strigolactones But Not on the Canonical Strigolactone Signaling Component MAX2

Stromules are highly dynamic protrusions of the plastids in plants. Several factors, such as drought and light conditions, influence the stromule frequency (SF) in a positive or negative way. A relatively recently discovered class of plant hormones are the strigolactones; strigolactones inhibit branching of the shoots and promote beneficial interactions between roots and arbuscular mycorrhizal fungi. Here, we investigate the link between the formation of stromules and strigolactones. This research shows a strong link between strigolactones and the formation of stromules: SF correlates with strigolactone levels in the wild type and strigolactone mutants (max2-1 max3-9), and SF is stimulated by strigolactone GR24 and reduced by strigolactone inhibitor D2.

Q&A: How does jasmonate signaling enable plants to adapt and survive?

Jasmonates (JAs) are a class of plant hormones that play essential roles in response to tissue wounding. They act on gene expression to slow down growth and to redirect metabolism towards producing defense molecules and repairing damage. These responses are systemic and have dramatic impacts on yields, making JAs a very active research area. JAs interact with many other plant hormones and therefore also have essential functions throughout development, notably during plant reproduction, leaf senescence and in response to many biotic and abiotic stresses.