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Abstract Flavonols are a subclass of flavonoids widely found in plants and typically exist in glycosylated forms, decorated with various sugars at different positions on the flavonol aglycone. The composition and abundance of flavonol glycosides vary across species and among tissues within a species. Although flavonols are collectively known for their antioxidant activity, the specific physiological functions of individual flavonol structures remain poorly understood. Here, we show that 2 flavonol glycosides, kaempferol 3-O-glucosyl(1 → 2)galactoside (K2) and quercetin 3-O-glucosyl(1 → 2)galactoside (Q2), predominantly accumulate in the pollen of Solanaceae plants. K2 is evolutionarily conserved across Solanaceae, while Q2 has been lost in species such as tomato (Solanum lycopersicum). Our transcriptome profiling and biochemical analysis revealed SlUGT78D-B (78-B) as a pollen-specific flavonol 3-O-galactosyltransferase responsible for K2 production in tomato. Disruption of 78-B abolished K2 accumulation, leading to defective pollen tube growth in our in vitro assays. Supplementation with kaempferol 3-O-galactoside (K2 precursor) restores pollen tube growth, whereas quercetin 3-O-galactoside (Q2 precursor) or flavonol aglycones do not, suggesting distinct roles for individual flavonol structures. We further show that 3 key amino acid residues of 78-B dictate its sugar specificity, favoring galactosylation over glucosylation. Substitution of any one of these residues enables 78-B to acquire glucosyltransferase activity. However, 78-B remains evolutionarily constrained from gaining this activity, suggesting selective pressure to maintain flavonol galactoside accumulation in pollen. These findings indicate that individual flavonol glycosides can have specific physiological roles beyond enhancing solubility and stability. 

ABSTRACT The Mediator complex is a multisubunit transcription coregulator that transfers regulatory signals from different transcription factors to RNA polymerase II (Pol II) to control Pol II‐dependent transcription in eukaryotes. Studies on Arabidopsis Mediator subunits have revealed their unique or overlapping functions in various aspects of plant growth, stress adaptation and metabolite homeostasis. Therefore, the utilization of the plant Mediator complex for crop improvement has been of great interest. Advances in genome editing and sequencing techniques have expedited the characterization of Mediator subunits in economically important crops such as tomato, rice, wheat, soybean, sugarcane, pea, chickpea, rapeseed and hop. In this review, we summarize recent progress in understanding the molecular mechanisms of how the Mediator complex regulates crop growth, development and adaptation to environmental stress. We also discuss the conserved and diverse functions of the Mediator complex in different plant species. In addition, we propose several future research directions to deepen our understanding of the important roles of Mediator subunits and their interacting proteins, which would provide promising targets for genetic modification to develop new cultivars with desirable agronomic traits. 

Abstract Aldoximes are amino acid derivatives that serve as intermediates for numerous specialized metabolites including cyanogenic glycosides, glucosinolates, and auxins. Aldoxime formation is mainly catalyzed by cytochrome P450 monooxygenases of the 79 family (CYP79s) that can have broad or narrow substrate specificity. Except for SbCYP79A1, aldoxime biosynthetic enzymes in the cereal sorghum (Sorghum bicolor) have not been characterized. This study identified nine CYP79-encoding genes in the genome of sorghum. A phylogenetic analysis of CYP79 showed that SbCYP79A61 formed a subclade with maize ZmCYP79A61, previously characterized to be involved in aldoxime biosynthesis. Functional characterization of this sorghum enzyme using transient expression in Nicotiana benthamiana and stable overexpression in Arabidopsis thaliana revealed that SbCYP79A61 catalyzes the production of phenylacetaldoxime (PAOx) from phenylalanine but, unlike the maize enzyme, displays no detectable activity against tryptophan. Additionally, targeted metabolite analysis after stable isotope feeding assays revealed that PAOx can serve as a precursor of phenylacetic acid (PAA) in sorghum and identified benzyl cyanide as an intermediate of PAOx-derived PAA biosynthesis in both sorghum and maize. Taken together, our results demonstrate that SbCYP79A61 produces PAOx in sorghum and may serve in the biosynthesis of other nitrogen-containing phenylalanine-derived metabolites involved in mediating biotic and abiotic stresses. 

SUMMARY Phenylpropanoids are specialized metabolites derived from phenylalanine. Glucosinolates are defense compounds derived mainly from methionine and tryptophan in Arabidopsis. It was previously shown that the phenylpropanoid pathway and glucosinolate production are metabolically linked. The accumulation of indole‐3‐acetaldoxime (IAOx), the precursor of tryptophan‐derived glucosinolates, represses phenylpropanoid biosynthesis through accelerated degradation of phenylalanine ammonia lyase (PAL). As PAL functions at the entry point of the phenylpropanoid pathway, which produces indispensable specialized metabolites such as lignin, aldoxime‐mediated phenylpropanoid repression is detrimental to plant survival. Although methionine‐derived glucosinolates in Arabidopsis are abundant, any impact of aliphatic aldoximes (AAOx) derived from aliphatic amino acids such as methionine on phenylpropanoid production remains unclear. Here, we investigate the impact of AAOx accumulation on phenylpropanoid production using Arabidopsis aldoxime mutants,ref2andref5. REF2 and REF5 metabolize aldoximes to respective nitrile oxides redundantly, but with different substrate specificities.ref2andref5mutants have decreased phenylpropanoid contents due to the accumulation of aldoximes. As REF2 and REF5 have high substrate specificity toward AAOx and IAOx, respectively, it was assumed thatref2accumulates AAOx, not IAOx. Our study indicates thatref2accumulates both AAOx and IAOx. Removing IAOx partially restored phenylpropanoid content inref2, but not to the wild‐type level. However, when AAOx biosynthesis was silenced, phenylpropanoid production and PAL activity inref2were completely restored, suggesting an inhibitory effect of AAOx on phenylpropanoid production. Further feeding studies revealed that the abnormal growth phenotype commonly observed in Arabidopsis mutants lacking AAOx production is a consequence of methionine accumulation. 

Auxins are a class of plant hormones playing crucial roles in a plant’s growth, development, and stress responses. Phenylacetic acid (PAA) is a phenylalanine-derived natural auxin found widely in plants. Although the auxin activity of PAA in plants was identified several decades ago, PAA homeostasis and its function remain poorly understood, whereas indole-3-acetic acid (IAA), the most potent auxin, has been used for most auxin studies. Recent studies have revealed unique features of PAA distinctive from IAA, and the enzymes and intermediates of the PAA biosynthesis pathway have been identified. Here, we summarize the occurrence and function of PAA in plants and highlight the recent progress made in PAA homeostasis, emphasizing PAA biosynthesis and crosstalk between IAA and PAA homeostasis.