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Dormancy is an adaptation mechanism of plants to environmental stress. Myricaria laxiflora undergoes a long period of flooding stress every year. In order to determine whether this species escapes flooding stress through dormancy, young branches and leaves were collected at different time points before the onset of flooding, and changes in the content/activity of hormones/enzymes that are closely involved in plant growth were monitored. The inducing environmental factors of summer dormancy were identified. The branches and leaves of M. laxiflora showed the following trends as summer flooding approached: (1) gradual increase in the abscisic acid content; (2) gradual decrease in the gibberellin and cytokinin contents; and (3) a continuous decrease in the activities of malate dehydrogenase (MDH), ribulose diphosphate carboxylase (RuBisCo), and glycolate oxidase (GLO). Pearson correlation analysis revealed (1) daylight duration was highly correlated with the hormone content and enzyme activity; (2) the daily mean air temperature (DMAT) was significantly correlated with the cytokinin content. These findings suggest that daylight duration was the main environmental factor leading to changes in the phytohormone content and enzyme activity as well as leading to summer dormancy. M. laxiflora undergoes dormancy before the onset of summer flooding to escape summer flooding stress. Our data indicate that summer flooding does not impede the survival and growth of M. laxiflora.

COMPASS or COMPASS-like is a highly conserved polyprotein complex in eukaryotes that is often involved in methylation of histone H3 lysine 4 (H3K4). However, the biological function of this complex in rice (Oryza sativa) is unclear. Here, we report the identifiction of their functions in growth and development. The osashl1 osashl2 double mutant shows a dwarf and late-flowering phenotype. Lower expression of Ehd1, OsVIL4, and OsMADS51 in the osashl1 osashl2 double mutant background accompanies a delayed vegetative growth phase and photoperiod-sensitive phase compared with that in wild type. Notably, there is less H3K4 mono-, di- and tri-methylation genome-wide in the double mutant, in particular less H3K4 tri-methylation at OsVIL4. Consistent with this result, knockout of OsVIL4 gives rise to a late-flowering phenotype similar to that of the osashl1 osashl2 double mutant, suggesting that OsVIL4 is a target of the COMPASS-like complex. In addition, the expression of key genes in brassinosteroid and gibberellic acid metabolism is altered in the osashl1 osashl2 double mutant, suggesting that the COMPASS-like complex regulates plant growth and development by modulating the levels of these two phytohormones. In summary, we demonstrate that OsASHL1 and OsASHL2 are important for floral transition and plant development.

Nitric oxide (NO) is an endogenous free radical involved in the regulation of a wide array of physio-biochemical phenomena in plants. The biological activity of NO directly depend on its cellular concentration which usually changes under stress conditions, it participates in maintaining cellular redox equilibrium and regulating target checkpoints which control switches among development and stress. It is one of the key players in plant signalling and a plethora of evidence supports its crosstalk with other phytohormones. NO and salicylic acid (SA) cooperation is also of great physiological relevance, where NO modulates the immune response by regulating SA linked target proteins i.e., non-expressor of pathogenesis-related genes (NPR-1 and NPR-2) and Group D bZIP (basic leucine zipper domain transcription factor). Many experimental data suggest a functional cooperative role between NO and SA in mitigating the plant oxidative stress which suggests that these relationships could constitute a metabolic “equilateral triangle”., FJC research work is supported by a European Regional Development Fund-cofinanced grant from the Ministry of Economy and Competitiveness/Science and Innovation (PID2019-10103924GB-I00), the Plan Andaluz de Investigación, Desarrollo e Innovación (PAIDI 2020) (P18-FR-1359) and Junta de Andalucía (group BIO192), Spain.

Tree branches affect the planting density and basal scab, which act as important attributes in the yield and quality of trees. Eucalyptus urophylla is an important pioneer tree with characteristics of strong adaptability, fast growth, short rotation period, and low disease and pest pressures. In this study, we collected ZQUC14 and LDUD26 clones and compared their transcriptomes and metabolomes from mature xylem, phloem, and developing tissues to identify factors that may influence branch development. In total, 32,809 differentially expressed genes (DEGs) and 18 gibberellin (GA) hormones were detected in the five sampled tissues. Searches of the kyoto Encyclopedia of Genes and Genomes pathways identified mainly genes related to diterpenoid biosynthesis, plant MAPK signaling pathways, plant hormone signal transduction, glycerolipid metabolism, peroxisome, phenylpropanoid biosynthesis, ABC transporters, and brassinosteroid biosynthesis. Furthermore, gene expression trend analysis and weighted gene co-expression network analysis revealed 13 genes likely involved in diterpenoid biosynthesis, including five members of the 2OG-Fe(II) oxygenase superfamily, four cytochrome P450 genes, and four novel genes. In GA signal transduction pathways, 24 DEGs were found to positively regulate branch formation. These results provide a comprehensive analysis of branch development based on the transcriptome and metabolome, and help clarify the molecular mechanisms of E. urophylla.

The phytohormone jasmonate plays a pivotal role in various aspects of plant life, including developmental programs and defense against pests and pathogens. A large body of knowledge on jasmonate biosynthesis, signal transduction as well as its functions in diverse plant processes has been gained in the past two decades. In addition, there exists extensive crosstalk between jasmonate pathway and other phytohormone pathways, such as salicylic acid (SA) and gibberellin (GA), in co-regulation of plant immune status, fine-tuning the balance of plant growth and defense, and so on, which were mostly learned from studies in the dicotyledonous model plants Arabidopsis thaliana and tomato but much less in monocot. Interestingly, existing evidence suggests both conservation and functional divergence in terms of core components of jasmonate pathway, its biological functions and signal integration with other phytohormones, between monocot and dicot. In this review, we summarize the current understanding on JA signal initiation, perception and regulation, and highlight the distinctive characteristics in different lineages of plants.

Salicylic acid (SA) is a pivotal hormone required for the development of resistance to many pathogens in plants. As an SA receptor, NPR1(Nonexpressor of Pathogenesis-Related Genes 1) plays a key regulatory role in the plant immune response. The function of NPR1 is dependent on the alteration of its oligomer-to-monomer. Research in recent years has proven that NPRs perceive SA and regulate the expression of downstream defense genes, but the mechanism of NPR1 oligomer-to-monomer conversion remains unclear. In this paper, we mainly studied the oligomerization of NPR1. By mutation experiments on some residues in the BTB domain involved in protein interactions, we found that the residue His80 plays a key role in the oligomerization of NPR1. We also found that NPR1, interacting with zinc ions at a ratio close to 1:1, was independent of the residue His80. These findings may help us to understand the conformational conversion of NPR1.

The plant hormone salicylic acid plays an important role in balancing plant immunity and growth. In this issue of Cell HostMicrobe, Liu et al. (2022) discovered that a long non-coding RNA, lncSABC1, promotes growth in uninfected plants and unleashes defenses when pathogens attack by transcriptionally regulating salicylic acid biosynthesis.

Phosphorus (P) is an essential nutrient for plant growth and reproduction. Plants preferentially absorb P as orthophosphate (Pi), an ion that displays low solubility and that is readily fixed in the soil, making P limitation a condition common to many soils and Pi fertilization an inefficient practice. To cope with Pi limitation, plants have evolved a series of developmental and physiological responses, collectively known as the Pi starvation rescue system (PSR), aimed to improve Pi acquisition and use efficiency (PUE) and protect from Pi-starvation-induced stress. Intensive research has been carried out during the last 20 years to unravel the mechanisms underlying the control of the PSR in plants. Here we review the results of this research effort that have led to the identification and characterization of several core Pi starvation signaling components, including sensors, transcription factors, microRNAs (miRNAs) and miRNA inhibitors, kinases, phosphatases, and components of the proteostasis machinery. We also refer to recent results revealing the existence of intricate signaling interplays between Pi and other nutrients and antagonists, N, Fe, Zn, and As, that have changed the initial single-nutrient-centric view to a more integrated view of nutrient homeostasis. Finally, we discuss advances toward improving PUE and future research priorities.

Optimal plant development requires root uptake of 14 essential mineral elements from the soil. Since the bioavailability of these nutrients underlies large variation in space and time, plants must dynamically adjust their root architecture to optimize nutrient access and acquisition. The information on external nutrient availability and whole-plant demand is translated into cellular signals that often involve phytohormones as intermediates to trigger a systemic or locally restricted developmental response. Timing and extent of such local root responses depend on the overall nutritional status of the plant that is transmitted from shoots to roots in the form of phytohormones or other systemic long-distance signals. The integration of these systemic and local signals then determines cell division or elongation rates in primary and lateral roots, the initiation, emergence, or elongation of lateral roots, as well as the formation of root hairs. Here, we review the cascades of nutrient-related sensing and signaling events that involve hormones and highlight nutrient-hormone relations that coordinate root developmental plasticity in plants.

Abscisic acid (ABA) plays an important role in seed germination, stomatal closure, and seedling growth inhibition in plants. Among downstream genes whose expression levels are regulated by AFA1 (Arabidopsis F-box Protein Hypersensitive to ABA 1), one gene, AtHAD1 upregulated by ABA was selected from Arabidopsis. AtHAD1 was induced by drought and salt stresses as well as by ABA and was found in dry seeds. Its loss-of-function mutants exhibited increased ABA-sensitivity in germination, seedling growth, and stomatal closure. In addition, the mutants displayed a lower water loss rate and higher survival rate under drought stress than the wild-type plants, indicating that a loss of AtHAD1 leads to enhanced drought tolerance. These results show that AtHAD1 has an inhibitory role in ABA response and ABA-mediated drought tolerance. The expression levels of several ABA-responsive genes in athad1 were higher than those in the wild-type under the ABA treatment, suggesting that AtHAD1, as a negative regulator in ABA response, could be associated with the downregulation of the ABA-responsive genes. The phosphatase assay showed that AtHAD1 exhibits phosphatase activity. Monitoring of the subcellular localization of GFP-fused AtHAD1 proteins indicated that AtHAD1 exists in the nucleus and cytoplasm. Overall, this study shows that Arabidopsis HAD1 as an intracellular phosphatase negatively functions in the ABA-mediated cellular responses. This research could serve as a research basis to understand the functional link between ABA signaling and the regulation process of the cellular phosphate level.

The mitogen-activated protein kinase OsMPK1 is involved in abscisic acid (ABA) biosynthesis in rice (Oryza sativa L.). However, the underlying molecular mechanisms of OsMPK1 in regulating ABA biosynthesis are poorly understood. Here, by using yeast two-hybrid assay and firefly luciferase complementary imaging assay, we show that OsMPK1 physically interact with a short-chain dehydrogenase protein OsABA2. However, OsMPK5, a homolog of OsMPK1, does not interact with OsABA2. Further, OsMPK1 can phosphorylate OsABA2S197 in vitro. Phosphorylation at the position of OsABA2S197 does not affect its subcellular localization, but enhances the stability of OsABA2 protein. We also found that OsABA2 has feedback regulation on OsMPK1 kinase activity. Further research reveals that OsMPK1 and OsABA2 coordinately regulate the biosynthesis of ABA, and phosphorylation of OsABA2 at Ser197 by OsMPK1 plays a crucial role in regulating the biosynthesis of ABA. Finally, genetic analysis showed that OsABA2 can enhance the sensitivity of rice to ABA and the tolerance of rice to drought and salt stress.

Salt stress is one of the common environmental threats to crop growth, development, and productivity. Plant growth regulators (PGRs) are natural messengers and are known to play pivotal roles at different stages of the growth and development of plants under various environmental conditions. Keeping in mind the importance of PGRs in stress management, a factorial randomized pot experiment was conducted to evaluate the efficiency of three selected PGRs, namely gibberellic acid (GA), salicylic acid (SA) and triacontanol (Tria) for the amelioration of NaCl stress in mustard. Plants were subjected to four concentrations of NaCl (0, 50, 100 and 150 mM). Two foliar sprays of PGRs (GA, SA and Tria), each at 5 μM were applied to the foliage of plants using a hand sprayer. The increasing levels of NaCl decreased growth, physio-biochemical, histochemical and yield parameters in a dose-dependent manner while increasing activities of antioxidant enzymes, contents of osmolytes and oxidative stress biomarkers linearly with increasing levels of NaCl. The spray of GA, SA and Tria under stressed-free and stressed conditions improved the aforesaid attributes while decreasing the generation of stress biomarkers. Of sprayed PGRs, SA proved to be the best for alleviating the adverse effect of NaCl stress. Furthermore, it provides experimental data for its possible biotechnological applications in mustard crops exposed to high concentrations of salinity and possibly to other environmental stresses which have associated oxidative stress., The authors gratefully acknowledge the Chairperson, Department of Botany and University Sophisticated Instrumentation Facility, Aligarh Muslim University for providing all necessary facilities for carrying out this work. The authors acknowledging the financial support provided by UGC New Delhi (17PHDBT030) at Aligarh Muslim University, and also extend their sincere appreciation to the Researchers Supporting Project number (RSP2023R194), King Saud University, Riyadh, Saudi Arabia.

Chinese hickory (Carya cathayensis Sarg.) is an important nut tree species native to China. Excessive plant height and long juvenile phase has restricted development of its industry. Recently, grafting has been used increasingly in production practice of this species to solve the problems above. Previous studies have proved the importance of auxin during Chinese hickory grafting. However, the function of ATP-binding cassette subfamily B (ABCB) genes during Chinese hickory grafting is less studied. In this study, 23 ABCB genes were identified and characterized in Chinese hickory (CcABCBs). The expression profiles of auxin-related ABCBs among tissues, under auxin-related phytohormone treatments and during grafting were determined. CcABCB proteins were divided into half-size and full-size transporters. Many phytohormone-related cis-acting regulatory elements were detected on the promoters of CcABCB genes. Four CcABCB genes homologous to auxin-related AtABCB1, 6, 19 and 20 in Arabidopsis were selected for expression analysis. The four genes displayed varying expression patterns in different tissues of Chinese hickory. Expressions of the four CcABCB genes were regulated by auxin-related phytohormones to varying degrees. Expression levels of the four genes were significantly changed at different stages of grafting, especially 7 days after grafting, indicating their involvement of auxin homeostasis regulation during grafting. In addition, the expressions of CcABCB1 were regulated by IAA and NPA treatments during grafting in comparison with CK treatment, while expressions of the other 3 CcABCB genes were slightly affected. This study will lay the foundation for understanding the potential regulatory roles of CcABCB genes during Chinese hickory grafting.

Lipoxygenase (LOX, EC 1.13.11.12) is a non-haeme iron-containing dioxygenase family that catalyzes the oxygenation of polyunsaturated fatty acids into bio-functionally fatty acid diverse (oxylipins) and plays vital role in plant growth and development and responses to abiotic and biotic stresses. Though LOX genes have been studied in many plant species, their roles in Brassicaceae species are still unknown. Here, a set of 14, 18, and 33 putative LOX genes were identified in Brassica rapa, Brassica oleracea and Brassica napus (allotetraploid rapeseed), respectively, which could be divided into 9-LOX (LOX1/5), 13-LOX type I (LOX3/4/6), and type II (LOX2) subgroups. There was an expansion of LOX2 orthologous genes in Brassicaceae. Most of the LOX genes are intron rich and conserved in gene structure, and the LOX proteins all have the conserved lipoxygenase and PLAT/LH2 domain. Ka/Ks ratio revealed that the majority of LOXs underwent purifying selection in Brassicaceae. The light-, ABA-, MeJA-related cis-elements and MYB-binding sites in the promoters of BnaLOXs were the most abundant. BnaLOXs displayed different spatiotemporal expression patterns and various abiotic/biotic stress responsive expression patterns. BnaLOX1/5 were slightly or no response to phytohormones and abiotic stresses. BnaLOX3/4/6 predominantly express in roots and were strongly up-regulated by salinity and PEG treatments, and BnaLOX3/4 were the methyl jasmonate (MeJA) and salicylic acid (SA) early response genes and strongly induced by infection of Sclerotinia sclerotiorum; while the BnaLOX2 members predominantly express in stamens, were MeJA and SA continuous response genes and strongly repressed by cold, heat and waterlogging treatments in leaves. Our results are useful for understanding the biological functions of the BnaLOX genes in allotetraploid rapeseed.

Salvia miltiorrhiza Bunge, belonging to Lamiaceae family, is one of the most important Chinese medicinal herbs. The dried roots, also called Danshen in Chinese, are usually used in the formula of Chinese traditional medicine due to the bioactive constituents known as phenolic acids and tanshinones, which are a group of abietane nor-diterpenoid quinone natural products. Cytochrome P450 enzymes (CYPs) usually play crucial roles in terpenoids synthesis, especially in hydroxylation processes. Up to now, several important P450 enzymes, such as CYP76AH1, CYP76AH3, CYP76AK1, CYP71D373, and CYP71D375, have been functionally characterized in the tanshinones biosynthetic pathway. Nevertheless, the tanshinones biosynthesis is a so complex network that more P450 enzymes should be identified and characterized. Here, we report two novel P450 enzymes CYP76AK2 and CYP76AK3 that are involved in tanshinones biosynthetic pathway. These two P450 enzymes were highly homologous to previously reported CYP76AK1 and showed the same expression profile as CYP76AK1. Also, CYP76AK2 and CYP76AK3 could be stimulated by MeJA and SA, resulting in increased expression. We used a triple-target CRISPR/Cas9 system to generate targeted mutagenesis of CYP76AK2 and CYP76AK3 in S. miltiorrhiza. The content of five major tanshinones was significantly reduced in both cyp76ak2 and cyp76ak3 mutants, indicating that the two enzymes might be involved in the biosynthesis of tanshinones. This study would provide a foundation for the catalytic function identification of CYP76AK2 and CYP76AK3, and further enrich the understanding of the network of tanshinones secondary metabolism synthesis as well.

Phytohormones play important roles in controlling leaf size and in the modulation of various stress responses, including drought. In this study, hormone profiling analyses by ultra high-performance liquid chromatography coupled to electrospray ionization tandem mass spectrometry (UHPLC-MS/MS) was performed in leaves collected at three stages of active leaf growth to unravel which phytohormones modulate leaf size in sesame (Sesamum indicum L.) plants, an important oil-rich crop. Furthermore, endogenous contents of phytohormones were measured in parallel to various stress markers in sesame plants exposed to mild water deficit conditions by withholding water in potted plants for one week. Results revealed a major role of cytokinins and auxin in the modulation of leaf growth in sesame plants (which increased by 21.5 and 2.1-fold, respectively, with leaf growth), as well as a putative antagonistic response between jasmonic acid and salicylic acid during leaf development. Furthermore, growth arrest during water deficit stress appeared to be modulated by cytokinins, the endogenous contents of which decreased (by 48%) in parallel with ABA increases (by 59%). Reductions in the contents of the active cytokinin trans-zeatin occurred in parallel with increases in isopentenyladenine contents under drought, which suggests a partial metabolic limitation in cytokinin biosynthesis in leaves upon water deficit stress. These results provide useful information for the hormonal modulation of leaf size and the improvement of leaf growth and production in sesame plants through manipulation of the levels of key regulatory phytohormones.

Cytochrome P450s (CYPs) are a versatile group of enzymes and one of the largest families of proteins, controlling various physiological processes via biosynthetic and detoxification pathways. CYPs perform multiple roles through a critical irreversible enzymatic reaction in which an oxygen atom is inserted within hydrophobic molecules, converting them into the reactive and hydro soluble components. During evolution, plants have acquired significantly more number of CYPs and represent about 1% of the encoded genes . CYPs are highly conserved proteins involved in growth, development and tolerance against biotic and abiotic stresses. Furthermore, CYPs reinforce plants' molecular and chemical defense mechanisms by regulating the biosynthesis of secondary metabolites, enhancing reactive oxygen species (ROS) scavenging and controlling biosynthesis and homeostasis of phytohormones, including abscisic acid (ABA) and jasmonates. Thus, they are the critical targets of metabolic engineering for enhancing plant defense against environmental stresses. Additionally, CYPs are also used as biocatalysts in the fields of pharmacology and phytoremediation. Herein, we highlight the role of CYPs in plant stress tolerance and their applications for human welfare.

Silicon (Si) acts as a beneficial element for plant growth and provides protection against abiotic and biotic stresses. Despite numerous reports on the beneficial role of Si in enhancing plant resistance to fungal pathogens, the underlying mechanisms remain largely unclear. Silicon shows antifungal activity; however, Si-induced improved disease resistance is partly manifested by the formation of Si polymerized mechanical obstruction under the cuticle and in cell walls, which prevents fungal ingress. Moreover, rapid production of defense compounds through secondary metabolic pathways is thought to be a key mechanism of Si-induced chemical defense against fungal pathogens beyond the physical barrier. Besides, improved mineral nutrition assures the healthy status of Si-supplied plants and a healthy plant exhibits better photosynthetic potential, antioxidant capacity and disease resistance. Multiple plant hormones and their crosstalk mediate the Si-induced basal as well as induced resistance; nonetheless, how root uptake of Si systemically modulates resistance to foliar diseases in low Si accumulating plants, needs in-depth investigation. Recent studies also indicate that Si influences effector-triggered immunity by affecting host recognition and/or limiting receptor-effector interactions. Here we review the role of Si in plant response to fungal pathogens. We also discuss and propose potential mechanisms of Si-induced enhanced disease resistance in plants. Finally, we identify some limitations of research approaches in addressing the beneficial roles of Si in biotic stress management.

Heavy metal (HM) accumulation in the agricultural soil and its toxicity is a major threat for plant growth and development. HMs disrupt functional integrity of the plants, induces altered phenological and physiological responses and slashes down qualitative crop yield. Chemical messengers such as phytohormones, plant growth regulators and gasotransmitters play a crucial role in regulating plant growth and development under metal toxicity in plants. Understanding the intricate network of these chemical messengers as well as interactions of genes/metabolites/proteins associated with HM toxicity in plants is necessary for deciphering insights into the regulatory circuit involved in HM tolerance. The present review describes (a) the role of chemical messengers in HM-induced toxicity mitigation, (b) possible crosstalk between phytohormones and other signaling cascades involved in plants HM tolerance and (c) the recent advancements in biotechnological interventions including genetic engineering, genome editing and omics approaches to provide a step ahead in making of improved plant against HM toxicities.

Cold-regulated (COR) genes are considered downstream functional genes in the cold-response pathway. However, we identified a plasma membrane-type, AtCor413pm1, as a regulatory gene for the abscisic acid (ABA) response, and found that ABA induced it predominantly in Arabidopsis roots, vasculature, stipules, and guard cells. Differentially expressed genes combined with qPCR analysis revealed the expressions of three ABA-responsive genes (AtDTX50, AtABR1, and AtCIPK20) were significantly altered in the ABA-treated atcor413pm1 mutant, compared to the wild-type. Furthermore, the ABA-induced transient Ca2+ oscillation in the plasma membrane of atcor413pm1 roots was different from that observed in the wild-type. Our results revealed that AtCor413pm1 might play a role in the cross-talk between the ABA and stress response pathways.

Chichi is a unique biological phenomenon observed in Ginkgo biloba L.. In this study, multi-omics analysis was used to compare basal chichi (C) with roots (R) and stems (S) to explore the regulatory mechanisms of basal chichi ontogenesis. The results showed that compared with roots and stems, the tZ, SA and ABA contents in basal chichi were the highest, and the ratio of IAA/tZ was the lowest. Nucleotides and their derivatives in basal chichi were upregulated, and phenylpropane metabolites were downregulated. Some differentially expressed genes (DEGs) strongly correlated to plant hormones were screened. We speculate that auxin and cytokinin are involved in the morphogenesis of basal chichi and that cytokinin plays a major role. The ontogenesis of basal chichi is closely related to environmental stress, and it may be a coping strategy of G. biloba in the face of environmental stress.

Sweetpotato (Ipomoea batatas [L.] Lam.) stem tips, which contain high concentrations of chlorogenic acid (CGA), are useful as a physiologically functional food to protect against some serious diseases. According to previous studies, exogenous application of phytohormones may be an effective agrotechnical measure to control CGA biosynthesis through the transcriptional regulation of pathway gene expressions. To understand the mechanism of CGA biosynthesis in sweetpotato, we investigated the effects of exogenous phytohormones on CGA metabolism in stem tips of sweetpotato. A significantly elevated CGA content was observed in salicylic acid (SA)-treated sweetpotato stem tips at 72 h, as well as in those subjected to abscisic acid (ABA) or gibberellic acid (GA) treatments. Dynamic expression change of seven enzyme genes involved in sweetpotato CGA biosynthesis were analyzed to determine correlations between transcript levels and CGA accumulation. As revealed by the differential expression of these genes under distinct phytohormone treatments, the regulation of specific pathway genes is a critical determinant of the accumulation of CGA in sweetpotato stem tips. We also found that several hormone-responsive sites, such as those for ABA, GA, SA, and jasmonic acid (JA), were present in the promoter regions of sweetpotato CGA biosynthestic pathway genes. Collectively, phytohormones can regulate the transcription of CGA synthesis-related genes and ultimately affect CGA accumulation in sweetpotato stem tips, whereas the regulatory differences are mirrored by cis-acting elements in the corresponding pathway gene promoters.

The number of organisms that are known to produce cytokinins (CKs) continues to increase. In fact, species from all life kingdoms have now been shown to either produce CKs or at least have the genetic components to make it possible. In vitro growth of microorganisms, plant/animal cells, and tissue cultures often requires nutrient-rich media composed of ingredients with organic origins including: yeast extract, peptone, tryptone, or various plant or animal extracts. These compounds, derived from microbial, plant and animal materials, can be the source of significant levels of exogenous CKs in the culture medium. As CK investigative work continues to expand rapidly, it is of critical importance to draw attention to this complexity; the presence of CKs in growth medium affects CK metabolism of the cultured organism and interferes with the readings of analytical instrumentation used to profile CKs in tested microorganisms or cell cultures.

Leaf shape is highly variable within and among plant species, ranging from slender to oval shaped. This is largely determined by the proximodistal axis of growth. However, little is known about how proximal-distal growth is controlled to determine leaf shape. Here, we show that Arabidopsis leaf and sepal proximodistal growth is tuned by two phytohormones. Two class A AUXIN RESPONSE FACTORs (ARFs), ARF6 and ARF8, activate the transcription of DWARF4, which encodes a key brassinosteroid (BR) biosynthetic enzyme. At the cellular level, the phytohormones promote more directional cell expansion along the proximodistal axis, as well as final cell sizes. BRs promote the demethyl-esterification of cell wall pectins, leading to isotropic in-plane cell wall loosening. Notably, numerical simulation showed that isotropic cell wall loosening could lead to directional cell and organ growth along the proximodistal axis. Taken together, we show that auxin acts through biosynthesis of BRs to determine cell wall mechanics and directional cell growth to generate leaves of variable roundness.

Jasmonate (JA) regulates various aspects of plant growth and development and stress responses, with prominent roles in male reproductive development and defenses against herbivores and necrotrophic pathogens. JASMONATE-ZIM DOMAIN (JAZ) proteins are key regulators in the JA signaling pathway and function to repress the expression of JA-responsive genes. Here, we show that JAZ proteins directly interact with several chromatin-associated Polycomb proteins to mediate repressive chromatin modifications at JA-responsive genes and, thus, their transcriptional repression in Arabidopsis. Genetic analyses revealed that the developmental defects, including anther and pollen abnormalities, resulting from loss or block of JA signaling were partially rescued by loss of Polycomb protein-mediated chromatin silencing (Polycomb repression). We further found that JAZ-mediated transcriptional repression during anther and pollen development requires Polycomb proteins at four key regulatory loci. Analysis of genome-wide occupancy of a Polycomb factor and transcriptome reprogramming in response to JA revealed that Polycomb repression is involved in the repression of various JA-responsive genes. Taken together, our study reveals an important chromatin-based mechanism for JAZ-mediated transcriptional repression and JA signaling in plants.

Carotenoid cleavage oxygenases (CCOs) play crucial roles in plant growth and development, as well as in the response to phytohormonal, biotic and abiotic stresses. However, comprehensive and systematic research on the CCO gene family has not yet been conducted in Saccharum. In this study, 47 SsCCO and 14 ShCCO genes were identified and characterized in Saccharum spontaneum and Saccharum spp. R570 cultivar, respectively. The SsCCOs consisted of 38 SsCCDs and 9 SsNCEDs, while ShCCOs contained 11 ShCCDs and 3 ShNCEDs. The SsCCO family could be divided into 7 groups, while ShCCO family into 5 groups. The genes/proteins contained similar compositions within the same group, and the evolutionary mechanisms differed between S. spontaneum and R570. Gene Ontology annotation implied that CCOs were involved in many physiological and biochemical processes. Additionally, 41 SsCCOs were regulated by 19 miRNA families, and 8 ShCCOs by 9 miRNA families. Cis-regulatory elements analysis suggested that CCO genes functioned in the process of growth and development or under the phytohormonal, biotic and abiotic stresses. qRT-PCR analysis indicated that nine CCO genes from different groups exhibited similar expression patterns under abscisic acid treatment, while more divergent profiles were observed in response to Sporisorium scitamineum and cold stresses. Herein, comparative genomics analysis of the CCO gene family between S. spontaneum and R570 was conducted to investigate its evolution and functions. This is the first report on the CCO gene family in S. spontaneum and R570, thus providing valuable information and facilitating further investigation into its function in the future.

Unfavorable environmental conditions are the critical inimical to the sustainable agriculture. Among various novel strategies designed to protect plants from abiotic stress threats, use of mineral elements as 'stress mitigators' has emerged as the most crucial and interesting aspect. Silicon (Si) is a quasi-essential nutrient that mediates plant growth and development and interacts with plant growth regulators (PGRs) and signaling molecules to combat abiotic stress induced adversities in plants and increase stress tolerance. PGRs are one of the most important chemical messengers that mediate plant growth and development during stressful conditions. However, the individual roles of Si and PGRs have extensively defined but their exquisite crosstalk with each other to mediate plant stress responses is still indiscernible. The present review is an upfront effort to delineate an intricate crosstalk/interaction between Si and PGRs to reduce abiotic stress adversities. The combined effects of interaction of Si with other signaling molecules such as reactive oxygen species (ROS), nitric oxide (NO) and calcium (Ca2+) for the survival of plants under stress and optimal conditions are also discussed.

Much is known about the molecular interactions between positive-strand RNA viruses and dicotyledon plants. However, many important viral pathogens of the monocotyledon rice crop contain negative-strand or double-strand RNA genomes. Recent studies have shown that virus-derived small-interfering RNAs (siRNAs), host microRNAs and phytohormones regulate antiviral responses in rice plants and that rice-infecting RNA viruses encode a diverse repertoire of multifunctional proteins with counter-defensive activities. Moreover, the interactions between viral virulence proteins and host susceptibility factors also shape the virus-rice arms race. This review will focus on these recent advances and discuss strategies and challenges in the translation of discoveries made on molecular virus-rice interactions into practical virus control measures.

Graphical abstract, Highlights • Polyploidization events were the main driving force for GRASs expansion. • Evolutionary analysis reveals that inter-species GRASs functions may be conserved. • GRASs helped plants resist stress by regulating flavonoids pathway. • GRASs regulate flavonoid synthesis by crosstalk with auxin and photosynthetic pathway. • Polyploidized GRASs play key roles in environment adaption, growth and development., Introduction Environmental stress is both a major force of natural selection and a prime factor affecting crop qualities and yields. The impact of the GRAS [gibberellic acid-insensitive (GAI), repressor of GA1–3 mutant (RGA), and scarecrow (SCR)] family on plant development and the potential to resist environmental stress needs much emphasis. Objectives This study aims to investigate the evolution, expansion, and adaptive mechanisms of GRASs of important representative plants during polyploidization. Methods We explored the evolutionary characteristics of GRASs in 15 representative plant species by systematic biological analysis of the genome, transcriptome, metabolite, protein complex map and phenotype. Results The GRAS family was systematically identified from 15 representative plant species of scientific and agricultural importance. The detection of gene duplication types of GRASs in all species showed that the widespread expansion of GRASs in these species was mainly contributed by polyploidization events. Evolutionary analysis reveals that most species experience independent genome-wide duplication (WGD) events and that interspecies GRAS functions may be broadly conserved. Polyploidy-related Chenopodium quinoa GRASs (CqGRASs) and Arabidopsis thaliana GRASs (AtGRASs) formed robust networks with flavonoid pathways by crosstalk with auxin and photosynthetic pathways. Furthermore, Arabidopsis thaliana population transcriptomes and the 1000 Plants (OneKP) project confirmed that GRASs are components of flavonoid biosynthesis, which enables plants to adapt to the environment by promoting flavonoid accumulation. More importantly, the GRASs of important species that may potentially improve important agronomic traits were mapped through TAIR and RARGE-II publicly available phenotypic data. Determining protein interactions and target genes contributes to determining GRAS functions. Conclusion The results of this study suggest that polyploidy-related GRASs in multiple species may be a target for improving plant growth, development, and environmental adaptation.

Upon infection with insect-borne microbial pathogens, plants are exposed to two types of damage simultaneously. Over the past decade, numerous molecular studies have been conducted to understand how plants respond to pathogens or herbivores. However, investigations of host responses typically focus on a single stress and are performed under static laboratory conditions. In this review, we highlight research that sheds light on how plants deploy broad-spectrum mechanisms against both vector-borne pathogens and insect vectors. Among the host genes involved in multistress resistance, many are involved in innate immunity and phytohormone signaling (especially jasmonate and salicylic acid). The potential for genome editing or chemical modulators to fine-tune crop defensive signaling, to develop sustainable methods to control insect-borne diseases, is discussed.

A facile, efficient and reliable method was designed and established to analyze the plant growth regulators (PGRs) in citrus fruit, based on a simplified dispersive solid-phase extraction (d-SPE) using a shaped zirconium-based metal-organic framework (UiO-66-PDMS bead) as the sorbent. In this method, UiO-66-PDMS beads directly extracted the targets from the homogenized and could be easily separated with a tweezer. It avoided the centrifugation or filtration operation required in normal d-SPE, greatly simplifying the d-SPE process. Moreover, the matrix substances were efficiently removed by this d-SPE process. The method showed good linearities (R

Seed germination is a multifaceted process, controlled by many cues, wherein phytohormones play a central role. Despite extensive studies, it remains obscure how hormonal balance and crosstalk between hormones regulate seed germination. Here we highlight new findings showing that crosstalk between jasmonates (JA) and abscisic acid (ABA) delays seed germination.

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.

Carotenoids are an important source of metabolites with regulatory function, which include the plant hormones abscisic acid (ABA) and strigolactones (SLs), and several recently identified growth regulators and signaling molecules. These carotenoid-derivatives originate from oxidative breakdown of double bonds in the carotenoid polyene, a common metabolic process that gives rise to diverse carbonyl cleavage-products known as apocarotenoids. Apocarotenoids exert biologically important functions in all taxa. In plants, they are a major regulator of plant growth, development and response to biotic and abiotic environmental stimuli, and mediate plant's communication with surrounding organisms. In this article, we provide a general overview on the biology of plant apocarotenoids, focusing on ABA, SLs, and recently identified apocarotenoid growth regulators. Following an introduction on carotenoids, we describe plant apocarotenoid biosynthesis, signal transduction, and evolution and summarize their biological functions. Moreover, we discuss the evolution of these intriguing metabolites, which has not been adequately addressed in the literature.

Gibberellins modulate multiple aspects of plant behavior. The molecular mechanism by which these hormones are perceived and how this information is translated into transcriptional changes has been elucidated in vascular plants: gibberellins are perceived by the nuclear receptor GID1, which then interacts with the DELLA nuclear proteins and promote their degradation, resulting in the modification of the activity of transcription factors with which DELLAs interact physically. However, several important questions are still pending: how does a single molecule perform such a vast array of functions along plant development? What property do gibberellins add to plant behavior? A closer look at gibberellin action from an evolutionary perspective can help answer these questions. DELLA proteins are conserved in all land plants, and predate the emergence of a full gibberellin metabolic pathway and the GID1 receptor in the ancestor of vascular plants. The origin of gibberellin signaling is linked to the exaptation by GID1 of the N-terminal domain in DELLA, which already acted as a transcriptional coactivator domain in the ancestral DELLA proteins. At least the ability to control plant growth seems to be encoded already in the ancestral DELLA protein too, suggesting that gibberellins¿ functional diversity is the direct consequence of DELLA protein activity. Finally, comparative network analysis suggests that gibberellin signaling increases the coordination of transcriptional responses, providing a theoretical framework for the role of gibberellins in plant adaptation at the evolutionary scale, which further needs experimental testing.

Forchlorfenuron (CPPU), a plant growth regulator, is widely used in agriculture. However, its long-term exposure effects on humans, especially neonates, remain unclear. Therefore, we investigated the developmental toxicity of prenatal and postnatal gavage administration of CPPU in rats. Pregnant Sprague-Dawley rats were administered 300 mg/kg/day CPPU by gavage from day 6 of gestation to the cessation of nursing. During weaning, rat offspring were administered 0, 30, 100, or 300 mg/kg/day CPPU for 4 weeks, followed by a 4-week CPPU-free recovery period. There were no significant differences in clinical symptoms, body weight, development indicators, serum biochemical parameters, sex hormone levels, sperm motility, relative organ weights, and histopathological changes among the 0–100 mg/kg/day CPPU groups. In the 300 mg/kg/day CPPU group, female rats exhibited decreased body weight, earlier time of vaginal opening (VO) and first estrus time (FE), elevated estradiol and blood urea nitrogen (BUN) levels, and upregulation of estrogen receptor 1 gene expression, whereas male rats only exhibited increases in serum BUN, creatinine, and glucose levels. Most changes were reversed after the recovery period. Furthermore, the endometrial epithelial height was significantly increased in female rats despite the absence of significant changes in uterine wall thickness and endometrial glands. Thus, CPPU may promote estradiol secretion, resulting in altered VO and FE and adverse effects in prepubertal female rats. These findings may be applied for risk assessment following CPPU exposure in humans.

Aegilops tauschii Coss, a notorious wheat field weed, poses a serious threat to wheat in China. Tillers are an important agronomic tool for yield. In this study, a total of 12 Ae. tauschii populations were collected from China to investigate the effect of plant density on tiller occurrence and its phytohormonal regulation. We assayed the growth parameters of Ae. tauschii and the levels of endogenous hormones at different plant densities. The results showed that plant density had a significant effect on the quantity and quality of Ae. tauschii seeds produced per plant. In particular, the tiller and spike numbers per plant were negatively affected by plant density (P 0.0001). The contents of 13 endogenous hormones in the tiller nodes changed in response to plant density. Among them, indole-3-acetic acid (IAA) and gibberellin (GA) positively responded to plant density. However, the reverse result was found for cytokinin (CTK). Interestingly, phylogenetic tree analysis of auxin (AeYUCCA), CK (AeIPT) and GA (AeCPS) biosynthesis related genes found that phylogenies in the Gramineae for the three different genes were various, hinting at horizontal gene transfer. Moreover, the dynamics of the expression of AeYUCCA, AeIPT and AeCPS were roughly consistent with their phytohormone contents during tillering stage. When externally sprayed on plants of Ae. tauschii, 2,4-D isooctyl ester and GA

The relationship of Resistance (R) gene-mediated defense to other forms of resistance in plants is considered, and the natures of the products of dominant and recessive R genes are reviewed. Various factors involved in expressing R gene-mediated resistance are described. These include phytohormones and plant effector molecules: the former regulating different pathways for disease resistance and the latter having direct effects on viral genomes or encoded proteins. Finally, the status of our knowledge concerning the cell-death hypersensitive response and its relationship to the actual resistance response involved in inhibiting virus infection is examined.

Salt stress is one of the major environmental stresses limiting plant growth and productivity. To adapt to salt stress, plants have developed various strategies to integrate exogenous salinity stress signals with endogenous developmental cues to optimize the balance of growth and stress responses. Accumulating evidence indicates that phytohormones, besides controlling plant growth and development under normal conditions, also mediate various environmental stresses, including salt stress, and thus regulate plant growth adaptation. In this review, we mainly discuss and summarize how plant hormones mediate salinity signals to regulate plant growth adaptation. We also highlight how, in response to salt stress, plants build a defense system by orchestrating the synthesis, signaling, and metabolism of various hormones via multiple crosstalks.

Extensins (EXTs) are major protein components in plant cell walls that play crucial roles in higher plants. The function of EXTs has been reported in several plants but is limited in tomato, especially in fruit ripening. In this study, we identified 83 EXTs in tomato, and divided them into seven groups. The gene intron-exon structure and protein-motif composition of SlEXTs were similar within each group but different among groups. SlEXT genes showed different expression patterns in roots, leaves, flowers and fruits, and some SlEXT gene expressions in flowers could be regulated by treatments of auxin, gibberellic acid and jasmonic acid. In particular, SlSEXT8 had higher and increased expression during tomato fruit ripening, and its expression could be induced by ethylene, suggesting SlSEXT8 may be involved in tomato fruit softening. The result provides insights into the function of EXTs, and will facilitate to further study EXT roles in tomato fruit ripening.

High-stress resistance and yield are major goals in crop cultivation, which can be addressed by modifying plant architecture. Significant progress has been made in recent years to understand how plant architecture is controlled under various growth conditions, recognizing the central role phytohormones play in response to environmental stresses. miRNAs, transcription factors, and other associated proteins regulate plant architecture, mainly via the modulation of hormone homeostasis and signaling. To generate crop plants of ideal architecture, we propose simultaneous editing of multiple genes involved in the regulatory networks associated with plant architecture as a feasible strategy. This strategy can help to address the need to increase grain yield and/or stress resistance under the pressures of the ever-increasing world population and climate change.

Phytohormones are interrelated by synergistic or antagonistic crosstalk and play important roles in the regulation of plant growth and development. In order to understand the interaction between phytohormones in the plant physiological network, it is necessary to determine trace levels of multiple phytohormones simultaneously in a complex matrix. Here, we synthesized ionic liquids containing different functional groups and modified the surface of carbon fibers with them. Based on these carbon fibers-ionic liquid (CFs-IL) materials, a solid phase microextraction method was developed to enable the simultaneous extraction of phytohormones. The adsorption specificity of multiple phytohormones was studied by identifying the hydrophobic, electrostatic, and π-π interactions, as well as hydrogen bonds, which favor simultaneous extraction of the relevant acidic, alkaline and neutral phytohormones by improving compatibility. The proposed method, coupled with liquid chromatography-tandem mass spectrometry, was applied to the simultaneous determination of 13 acidic, alkaline and neutral phytohormones in tomato. The limits of quantification were found to be in the range of 0.32–54.05 ng mL−1 and 4.6–185.8 pg mL−1, respectively, when measured by QQQ and Q-TOF. All of the relative recoveries were in the range of 94.40–113.37% with RSDs ≤15.36% (n = 3) for spiked tomato samples. This method is expected to be widely applied to multiple phytohormones analysis for in-depth researches concerning the physiological networks of plants.

Rice kernel smut (RKS), caused by the basidiomycete fungus Tilletia horrida, is one of the most devastating diseases affecting the production of male sterile lines of rice (Oryza sativa) worldwide. However, the molecular mechanisms of resistance to T. horrida have not yet been explored. In the present study, RNA sequencing analysis of rice male sterile lines, that are resistant and susceptible to RKS (Jiangcheng 3A and 9311A, respectively) was conducted after T. horrida infection. Transcriptomic analysis showed that a greater number of differentially expressed gene (DEGs) was observed in Jiangcheng 3A compared with 9311A after T. horrida inoculation. Furthermore, 4, 425 DEGs were uniquely detected in Jiangcheng 3A, and Gene Ontology (GO) and Kyoto Encyclopedia of Genes and Genomes (KEGG) analyses of these DEGs revealed that oxidoreductase activity, peroxidase activity, cutin, suberine and wax biosynthesis, and flavonoid biosynthesis were key pathways for T. horrida resistance. In summary and based on transcriptome analysis, we suggest a preliminary regulatory mechanism for Jiangcheng 3A cultivar resistance response to T. horrida inoculation.

Strigolactones (SLs) control plant development, enhance symbioses, and act as germination stimulants for some of the most destructive species of parasitic weeds, making SLs a potential tool to improve crop productivity and resilience. Field trials demonstrate the potential use of SLs as agrochemicals or genetic targets in breeding programs, with applications in improving drought tolerance, increasing yields, and controlling parasitic weeds. However, for effective translation of SLs into agriculture, understanding and exploiting SL diversity and the development of economically viable sources of SL analogs will be critical. Here we review how manipulation of SL signaling can be used when developing new tools and crop varieties to address some critical challenges, such as nutrient acquisition, resource allocation, stress tolerance, and plant-parasite interactions.

In plants, pluripotent cells in meristems divide to provide cells for the formation of postembryonic tissues. The cambium is the meristem from which the vascular tissue is derived and is the main driver for secondary (radial) growth in dicots. Xylem and phloem are specified on opposing sides of the cambium, and tightly regulated cell divisions ensure their spatial separation. Peptide ligands, phytohormones, and their receptors are central to maintaining this patterning and regulating proliferation. Here, we describe recent advances in our understanding of how these signals are integrated to control vascular development and secondary growth.