Nature Plants

Coumarins are phenylpropanoid-derived specialized metabolites that contribute to plant defence, shape plant-microbe interactions in the rhizosphere, and promote iron acquisition. In Arabidopsis thaliana, a model plant for iron-responsive coumarin metabolism, the enzymatic origin of the catecholic coumarin esculetin has long remained unresolved. Here we identify the first O-demethylation reaction in Arabidopsis specialized metabolism and show that 2-oxoglutarate- and Fe(II)-dependent dioxygenases catalyze scopoletin 6-O-demethylation to form esculetin. We designate these enzymes scopoletin 6-O-demethylases (S6ODs) and validate their activity through biochemical characterization, together with metabolomic profiling and independent loss-of-function mutant lines providing genetic evidence in planta. Disruption of S6OD activity remodels coumarin profiles and alters plant performance under limited iron availability, indicating that esculetin biosynthesis contributes to plant responses under these conditions. Our findings resolve the long-sought missing step in esculetin biosynthesis. It establishes O-demethylation as a previously unrecognized reaction in Arabidopsis specialized metabolism and suggest that 2OGD-mediated O-demethylation is recurrently recruited during evolution of plant metabolism, with implications for metabolic engineering and improvement of iron acquisition traits in crops.

Polyacetylene glycosides exhibit notable pharmacological activities, yet the glycosyltransferases acting on their polyacetylene scaffolds remain unknown. Here we report a telomere-to-telomere genome assembly of Codonopsis pilosula and, guided by spatial metabolomics, characterize three UDP-glycosyltransferases: CpUGT76BG1 and CpUGT76BG2 catalyze the direct glycosylation of lobetyol to lobetyolin, while CpUGT94BY2 performs subsequent sugar-sugar coupling to produce lobetyolinin, with each activity confirmed by in planta overexpression. Structural modeling reveals that CpUGT76BG1 and CpUGT76BG2 employ a deep hydrophobic tunnel to fully encase the linear polyacetylene chain, a binding architecture distinct from the shallow pockets used by canonical plant UGTs for planar aromatic substrates. Ancestral sequence reconstruction across eleven nodes partitions the UGT76 lineage into three functionally distinct evolutionary stages, tracing the trajectory from an ancestral shallow pocket to this specialized deep architecture. These findings establish the key glycosylation steps of polyacetylene glycoside biosynthesis, define a tunnel-based paradigm for non-planar substrate recognition, and reveal how tandem duplication-driven active site remodeling generates metabolic novelty.

A central question in plant biology is whether natural transcriptome variation primarily reflects environmental adaptation or intrinsic genetic programs. Using transcriptomes from 665 natural Arabidopsis thaliana accessions, we show that inherited population structure, not environmental gradients, is the primary organizer of global transcriptome architecture. Genomic population structure uniquely explains 30.0% of transcriptome-wide variation, while environmental variables account for only 2.5%. This architecture is dominated by a single co-expression program (proliferation Module Eigengene: prolifME), enriched for cell cycle regulation and ribosome biogenesis, regulated by a bipartite transcription factor architecture, and genetically encoded at discrete loci with broad trans-regulatory effects. This proliferation program is associated with reduced plant size, biomass, growth rate and water-use efficiency, and these relationships persist under common-garden conditions. Gene contributions to this proliferation program are non-randomly conserved in rice and maize across 150-200 million years of divergence. These findings identify an intrinsic transcriptional proliferation program as a primary state of plant population transcriptomes, conserved across plant species.

Reported CRISPR/SpCas9 editing efficiencies in crops span 0-100%, but no quantitative synthesis has separated taxonomic family from delivery method, ploidy, clustering or publication bias. This meta-analysis estimated pooled per-T0-line editing efficiency across Cucurbitaceae, Brassicaceae, Solanaceae and Poaceae, and tested whether family is an independent moderator after adjustment for delivery and ploidy. A PRISMA 2020 systematic review identified peer-reviewed studies using SpCas9 with extractable per-line T0 edit counts; data were extracted independently by two reviewers, with inter-rater agreement reported. Logit proportions were synthesised with a binomial-normal generalised linear mixed model, and the family-as-moderator hypothesis was tested by a small-sample CR2 cluster-robust F-test on a three-level model with study-level clustering. Publication bias was assessed by Eggers regression and trim-and-fill. Twenty-two studies contributed 172 per-line effect sizes (Cucurbitaceae k=14, Brassicaceae k=20, Solanaceae k=68, Poaceae k=70). Pooled editing efficiency was 61.8% (95% CI 54.5-68.6%) with I{superscript 2}=93.4% ({tau}{superscript 2}=3.21) and a 95% prediction interval of approximately 5-98%. Per-family estimates ranged from 47.8% (Poaceae) to 73.8% (Brassicaceae); the univariate Q test was significant (p=0.0016), but family did not survive cluster-robust adjustment (F=0.73, p=0.63). Intraclass correlation placed 64.4% of variance at the study level, and Solanaceae remained dominated by a single study (58/68 rows). Funnel asymmetry was severe (Egger p<0.0001), and trim-and-fill reduced the bias-adjusted estimate to 45.2% (95% CI 39.0-51.5%). Apparent crop-family differences dissolve once within-study clustering and methodological covariates are accounted for; the bias-adjusted pooled estimate is closer to 45% than to 62%, and reported editing efficiencies reflect study-level factors more than taxonomic family. Key MessageApparent between-family differences in CRISPR/SpCas9 editing efficiency across four crop families reflect within-study clustering and publication bias, not intrinsic biology; family is not an independent moderator after cluster-robust adjustment.

Plasmodesmata are intercellular channels that mediate symplastic communication between plant cells. Molecular transport through these channels is critically regulated by dynamic callose deposition and degradation, yet the secretory mechanisms that deliver regulatory components to plasmodesmata remain poorly understood. Here, we identify and characterize a non-canonical plasmodesmata-associated module of the exocyst, an evolutionarily conserved protein complex involved in secretory vesicle tethering and exocytosis. Exocyst subunits EXO70G1, SEC15A, EXO84C, and SEC10A specifically accumulate at plasmodesmata, whereas the canonical exocyst subunits EXO70A1 and SEC8 do not. Genetic and interaction analyses show that EXO70G1 acts as a landmark for recruiting SEC15A and EXO84C to plasmodesmata, revealing a distinct mode of exocyst targeting at these membrane domains. EXO70G1-dependent exocyst targeting to plasmodesmata depends on phosphoinositides and sphingolipids, consistent with the specialized lipid environment of plasmodesmal membranes. Loss of EXO70G1 results in increased callose accumulation and reduced symplastic transport, and strongly enhances developmental defects of a callose-overproducing mutant. In addition, exo70G1 mutants display enhanced resistance to bacterial pathogen Pseudomonas syringae, linking reduced plasmodesmal permeability to anti-bacterial defense. Cross-species analysis further indicates that plasmodesmata association is a derived feature of the EXO70G clade, present in angiosperms but absent from non-angiosperm EXO70 homologs. Together, our findings show that exocyst diversification in plants has generated a specialized trafficking module - plasmodesmata-associated exocyst - that links vesicle delivery to callose homeostasis at plasmodesmata, thereby regulating intercellular communication, development, and immunity. TeaserA specialized secretion module of the exocyst complex regulates plant cell-to-cell connectivity by controlling callose turnover at plasmodesmata

Plastids house the biology of eukaryotic photosynthesis. The majority of a plastids proteome is imported after cytosolic translation, but a few dozen proteins on average remain organelle-encoded, translated by the plastids own ribosomes. While 1000s of plastid genomes have been sequenced, the availability of less than ten proteomes and only two species with full 70S plastid ribosomal structures limit our understanding of land plant evolution. To address this, we optimized a protocol for the rapid isolation of Marchantia polymorpha chloroplasts that provides a highly enriched and intact organelle fraction from gradient volumes as little as 2 mL. Our approach was successfully applied to six other species, including Chlamydomonas reinhardtii and Nicotiana tabacum. Focusing on M. polymorpha, we determined the proteome of the chloroplast fraction, identifying 1337 nuclear-encoded proteins with a high confidence, where 83% belong to orthologs shared with angiosperms. We further isolated large protein complexes by RNA affinity purification using poly-lysine and provide the high-resolution structures of the 50S subunit of the chloroplast ribosome and RuBisCO from this bryophyte using cryogenic EM and image reconstruction to 2.23 and 2.12 [A] resolution, respectively, highlighting the structural conservation of both complexes. For chloroplasts, our data show that the genome reduction event experienced by the common ancestor of bryophytes has had little impact on the organelles complexity and that they underscore a high level of structural conservation of core components of plastid biology. Our data provide novel resources and methods to explore the functional evolution of plastid proteomes and major macromolecular complexes of cyanobacterial origin.

Drought increasingly constrains global rice productivity, yet how water deficit remodels cis-regulatory activity in plants remains poorly resolved. Here we used precision run-on sequencing (PRO-seq) to profile nascent transcription in rice leaves under well-watered and drought conditions and mapped transcription-initiation regions with the tool dREG, which detects genome-wide peaks of bidirectional transcription displaying active-enhancer behaviour. PRO-seq captured a robust drought response at genes and revealed extensive remodelling of initiation landscapes. We detected 85,764 consensus dREG sites, of which 17,193 changed significantly under drought and were predominantly intergenic. Because plant intergenic space is rich in transposable elements and silencing-associated transcription, we integrated transposable-element overlap and small-RNA loci with chromatin accessibility and DNA methylation to prioritize 2,428 drought-responsive intergenic sites (841 induced and 1,308 repressed) that are accessible, locally hypomethylated, and bidirectionally transcribed - features consistent with enhancer-like elements. Activity at proximal candidates correlated with elevated nascent transcription of nearby genes, and a subset overlapped gene-connected chromatin loop anchors, supporting candidate enhancer-target relationships. Motif enrichment further supported the involvement of drought-responsive regulatory programs, and hundreds of candidates overlapped rice STARR-seq enhancers. Together, these data define a drought-responsive atlas of candidate enhancer-like nascent transcription in rice and provide prioritized cis-regulatory candidates for mechanistic validation and crop improvement.

Plants detect neighbours through a reduced red-to-far-red ratio (R:FR), triggering elongation growth that reduces crop yield. Although Gibberellin (GA) is required for the neighbour-proximity (NP) elongation response, bioactive GA levels do not increase sufficiently to account for elongation magnitude, suggesting GA sensitivity as an additional regulated variable. Here, we show that GID1C, one of three Arabidopsis GA receptors, is the primary GA receptor involved in NP-induced elongation. GID1C protein accumulates selectively in hypocotyls and root tips under low R:FR without an increase in bioactive GA. The gid1c mutant shows a reduced elongation response that exogenous GA treatment cannot rescue. Transcriptome profiling reveals that GID1C controls 86% of the NP-responsive transcriptome, including genes for cell growth, division, and transcriptional regulation. Hub analysis identifies ICE1 as a GID1C-repressed transcriptional brake. ICE1 transcript is suppressed under low R:FR in a GID1C-dependent manner, and a phosphorylation-resistant ICE1 allele blocks NP-induced elongation. Together, these findings establish GA perception as an additional regulatory layer in NP, with subfunctionalisation among GID1 paralogs shaping the response to neighbouring plants.

Across kingdoms, cells fold their membranes into precise shapes closely linked to their functions. In mature land-plant chloroplasts, the photosynthetic membranes have been viewed as strictly lamellar and it is unknown whether they can take on a different structure while remaining functional. Here, we show that mature Arabidopsis thaliana chloroplasts can transform this network into a gyroid-type cubic membrane, which we call the gyrobody. The gyrobody forms reversibly during the night and preserves photosystem II photochemistry. A decrease in stromal side thylakoid surface charge, caused by lower protein phosphorylation, triggers the lamellar-to-gyroid transition which the curvature-inducing lipid MGDG facilitates. This shows that the mature plant thylakoid network is not locked into its lamellar form, revealing unexpected structural flexibility of this system.

Members of the K+ efflux antiporter (KEA) family fulfill key roles in plastids and the endomembrane system. Plants and green algae possess at least one KEA mediating K+/H+ exchange across the plastid inner envelope (IE) membrane. Recently, IE KEAs were shown to be essential for plastid gene expression (PGE), chloroplast development, and photosynthesis. Plants lacking these antiporters exhibit reduced stromal protein synthesis and accumulation of unprocessed rRNA precursors. KEA proteins comprise a conserved monovalent cation/proton antiporter 2 (CPA2) domain and a regulatory K transport and NAD-binding (KTN) domain. IE KEAs are distinguished by an additional ~500-amino-acid N-terminal extension containing a coiled-coil (CC) domain embedded within a largely intrinsically disordered region (IDR). Intrigued by this unusual architecture, we performed phylogenetic analyses, revealing that this N-terminal fusion arose early and has been conserved throughout the green lineage. We then investigated the oligomeric state, native distribution, and function of the N-terminal domain. Using Arabidopsis thaliana, we found that IE KEAs localize to discrete clusters within the inner envelope membrane and assemble into complexes of approximately 600 kDa. Finally, complementary approaches using a functional KEA1 variant lacking the core N-terminal domains (KEA1{Delta}N) indicate that this extension plays a regulatory rather than an essential role. Our findings uncover an evolutionarily ancient regulatory module that shapes the molecular organization and function of IE KEAs, advancing our understanding of plastid ion and pH homeostasis and plastid ribosome integrity.

Ribosome biogenesis is critical for plant development and environmental responses. A large number of ribosomal proteins (RPs) and ribosomal biogenesis factors (RBFs) are required for ribosome biogenesis, many of which remain uncharacterized in plants. We report here the identification of Arabidopsis RBF FAN and its interacting partner FAN-INTERACTING PROTEIN 1 (FIP1). As their human and yeast orthologues, FAN-FIP1 interact. Both FAN and FIP1 participate in the processing of pre-rRNAs. Functional loss of FAN or FIP1 knock-down results in developmental retardation and hypersensitivity to heat stresses. We demonstrate that FAN-FIP1 positively mediates brassinosteroid (BR) signaling by ensuring the translation efficiency of the BR receptor-coding gene BRASSINOSTEROID INSENSITIVE 1 (BRI1) through the presence of its upstream open reading frame (uORF). Importantly, BR signaling positively mediates the processing of pre-rRNAs, which may be critical not only for development but also for heat tolerance.

Arbuscular mycorrhizal fungi (AMF) establish intimate symbiosis with plant roots, yet the cell-type-specific regulatory and metabolic programs governing this interaction remain poorly resolved. Here, we integrate single-nucleus RNA sequencing (snRNA-seq) with spatial metabolomics across a temporal gradient of soybean root colonization (2-8 weeks post inoculation) to construct a high-resolution, multi-omic atlas of AMF symbiosis. Profiling 33,410 nuclei spanning all major root cell types, we uncover dynamic, cell-type-resolved transcriptional reprogramming coupled to spatially localized metabolite accumulation. Early colonization triggers a robust, epidermis-localized immune response alongside cortex-specific epigenetic reprogramming, mediated by RNA-directed DNA methylation machinery, suggesting active suppression of defense in fungal accommodation zones. Spatial metabolomics reveals a biphasic metabolic transition from flavonoid- and terpenoid-rich signaling states to lipid-dominated nutrient exchange, aligned with colonization progression. In parallel, coordinated carbon allocation and lipid biosynthesis pathways were activated in cortex and vascular tissues, supporting fungal dependence on host-derived fatty acids and sugars. Nutrient exchange programs, particularly nitrogen and phosphorus transport, exhibit strong pericycle and phloem specificity, highlighting systemic integration of symbiotic benefits. Through co-expression network analysis, we identify a previously uncharacterized coumarin-centered metabolic switch, governed by GmF6H1-2, that is essential for efficient colonization, as validated by natural loss-of-function variants. Collectively, this study provides a comprehensive, spatially resolved framework linking gene regulation, metabolism, and cell identity, revealing that AMF symbiosis is orchestrated through coordinated immune modulation, metabolic rewiring, and nutrient flux partitioning at single-cell resolution.

Chromosomal variation underlies species evolution, but reconstructing its large-scale dynamics remains challenging, obscuring its adaptive significance. Here, we introduce GouMang, a framework that mines conserved genic section compositions across diverse species to trace karyotype evolution. In grasses, applied to 818 highly varied chromosomes spanning eight subfamilies, GouMang resolved a shared karyotype evolution path of 9-to-18 ({rho} whole genome duplication, {rho}WGD)-to-12 chromosomes, followed by lineage specific rearrangements or WGDs. Genes retained from the early {rho}WGD are linked to cold/light adaptation, supporting a key biomass expansion event that impacted subsequent global ecological pattern and human agricultural civilization, in which K-Pg global cooling and subsequent forest degradation drove early understory grasses to sun plants. Parallel analysis in Brassicaceae reconstructed karyotype evolution as well as {beta}WGD which unlinked to cold/light adaptation, reflecting a divergent biogeographic history compared to grasses. Together, GouMang depicts a widespread plant evolutionary pattern where karyotype constantly diversified with WGDs recurrently fueling adaptation.

Terrestrial plant biomass relies on the formation of vascular bundles (VBs) comprising xylem, cambial, and phloem cells that drive secondary growth and wood formation. How VB formation and organization is coordinated with stem growth remains unclear. Here, we show that the conserved CLAVATA3 peptide (CLV3p)-receptor stem cell signaling pathway represses vascular transcriptional programs in the shoot apical meristem (SAM), thereby preventing ectopic vascular differentiation in the stem center, and quantitatively regulates peripheral VB number. Mutational analysis of conserved CLV3p residues partially uncouples its roles in vascular differentiation from stem cell maintenance. However, overexpression and mutational analysis with key phloem regulators reveals that CLV3p controls vascular and stem cell programs via a DOF (DNA-BINDING WITH ONE FINGER) and SMXL (SUPPRESSOR OF MAX2 1-LIKE) transcriptional module. Our findings identify a novel role for CLV3p signaling in shoot vascular development and establish a regulatory framework integrating stem cell signaling with vascular patterning.

High-quality plant genome assemblies are rapidly increasing, but accurate structural annotation remains reliant on transcript and homology evidence, limiting applications in newly sequenced and non-model species. Here, we present PlantGeneAnn, a plant-optimized, strand-specific genome foundation model for ab initio gene structure annotation. Fine-tuned on only nine high-quality model plant annotations, PlantGeneAnn outperformed a multi-species model trained on 42 species, showing that annotation quality is more important than token volume. On a stringent 13-species benchmark covering rosids, asterids, and monocots, PlantGeneAnn surpassed four state-of-the-art baselines across five evaluation levels, from base-level classification to complete transcript recovery. It achieved higher intron precision and better captured complex gene structures. In zero-shot variant effect prediction, PlantGeneAnn identified cryptic splice donors and premature stop codons in maize and rice, with saturation mutagenesis confirming single-nucleotide, context-dependent sensitivity. It also retained generalizability for epigenomic track prediction, highlighting its value for pan-genomics, crop improvement, and non-model plant research.

Eukaryotic cells are composed of different organelles that communicate with one another through direct contacts, which are necessary for a host of cellular reactions and for responding to different developmental and environmental changes. Plasma membrane (PM) forms extensive contacts with the endoplasmic reticulum (ER) at specific sites named ER-PM contact sites. These contacts play crucial functions in lipid homeostasis, Ca2+ regulation and signaling in all eukaryotes. However, the mechanisms by which plant ER-PM contact site proteins tether to the PM, as well as the dynamics of these contact sites, remain poorly understood. Here, we investigate the importance of phosphoinositides in the establishment and dynamics of ER-PM contact site proteins in plants. We found that phosphatidylinositol-4-phosphate (PI4P), rather than phosphatidylinositol 4,5-bisphosphate (PI(4,5)P2), is required for the association of ER-PM contact site proteins with the PM. Furthermore, we identified a PI4P phosphatase, SUPPRESSOR-OF-ACTIN7 (SAC7), that associates with the ER-PM contact site protein SYNAPTOTAGMIN1 (SYT1) and regulates its dynamic association with the PM. In particular, we found that in growing root hairs, a highly polarized cell type, SAC7 removes SYT1-containing contact sites at the growing tip. Consistently, optogenetic induction of ER-PM tethering reduced root hair elongation within minutes of blue light induction. Altogether, we propose a link between SAC7-mediated regulation of PI4P, dynamic ER-PM contact site establishment and polarized cell growth in plants.

Pigmented rice landraces are prized for their anthocyanin-accumulating pericarp, yet the repeat-rich regulatory alleles that govern pigmentation are poorly resolved in draft assemblies. Here we report a gap-free, telomere-to-telomere (T2T) genome of Mojiang purple rice (ZN65), a glutinous purple-pericarp Hani landrace from Yunnan, China. The 395.1-Mb assembly comprises twelve gap-free chromosomes with all 24 telomeres, all 12 centromeres and the 45S/5S ribosomal DNA arrays resolved (contig N50 32.35 Mb; Merqury QV 53.6; 99.6% BUSCO); we annotated 42,090 protein-coding genes, with transposable elements occupying 56.6% of the genome. Against the japonica reference Nipponbare we identified 1,045,956 single-nucleotide variants, 121 inversions and 449 translocations; ZN65 is larger on every chromosome (~22 Mb cumulative excess), reflecting lineage-specific retrotransposon expansion (26.9 Mb of LTR/Gypsy within 73.3 Mb of ZN65-specific sequence), and falls in the indica group. The flavonoid pathway is complete and copy-number-conserved (100 genes), so pigmentation maps to regulators: OsC1/Kala3 is conserved, whereas Kala4/OsB2, Kala1/OsDFR and Rc each carry transposon-associated structural variation. ZN65 encodes a functional pigmentation-allele complement - most diagnostically a full-length Rc with an intact bHLH domain, unlike the truncated rc allele of white Nipponbare. At Kala4/OsB2, ZN65 carries a long-read-validated retrotransposon architecture, including a ZN65-specific proximal-promoter insertion absent across a seven-genome panel - a candidate, lineage-specific realization of the black-rice gain-of-function mechanism, distinct from the canonical tandem-duplication allele. This T2T resource and its pigmentation-locus haplotypes provide a foundation for the functional study and molecular breeding of pigmented rice.

Shoot regeneration is a powerful model for cell fate reprogramming but how it occurs in nature remains poorly understood because studies in Arabidopsis thaliana conventionally rely on in vitro assays supplemented with exogenous hormones and sugars. In this study, we established the Hormone-autonomY Direct Regeneration Assay (HYDRA) in which removal of the shoot apical meristem (SAM) initiates shoot regeneration from the cotyledon-hypocotyl boundary domain without hormone or sugar supplementation. We show that photosynthesis-derived carbon and the boundary domain are two separable but convergent requirements for shoot regeneration in HYDRA. Carbon availability increases in the boundary domain where it activates cell cycle progression via the RETINOBLASTOMA-RELATED1 (RBR1) pathway. Carbon deprivation blocks regeneration despite induction of SAM marker genes, indicating that carbon-dependent cell cycle activation is a limiting factor for regeneration. In parallel, perturbation of the boundary domain or its regulators reduces regeneration despite sufficient carbon, indicating that boundary domain identity is independently required. Additionally, exogenous carbon supply overcomes the requirement for SAM removal to induce shoot formation, indicating that carbon availability also acts as an initiation cue. Together, this study reveals an inherent capacity for hormone-autonomous shoot regeneration and identifies photosynthesis-derived carbon as a central regulator of this process.

Plants perceive neighboring vegetation through an enrichment of far-red light (shade) in the environment. These changes in light quality trigger molecular and physiological responses aimed at outgrowing competitors, collectively known as the shade avoidance syndrome. In this study, we identify the TARGET OF RAPAMYCIN (TOR) complex 1 (TORC1), a major growth-regulating hub in eukaryotes, as a driver of shade-mediated growth responses in plants. Combining physiology, genetics, biochemistry, and proteomics, we show that TOR activity is rapidly enhanced upon shade perception and is required for proper shade responses, as TOR inhibition severely impairs shade-mediated elongation. Furthermore, we found that the control of shade-mediated elongation by TOR involves auxin-dependent mechanisms, requires efficient translation activity, and is closely linked to epidermal cell elongation capacity. Altogether, our work identifies TOR as a key integrator of light quality signals to control adaptive growth responses. Finally, we further highlight the conservation of shade-mediated TOR activation in tomato, with potential implications for engineering crop cultivars better suited to high-density planting.

The phytohormone auxin is a central coordinator of plant growth and development. Besides its canonical effect on gene transcription1,2, auxin triggers an ultra-rapid calcium ion influx that initiates the root gravitropic response3. The nature of the so-called rapid auxin pathway connecting the AFB1 auxin receptor3,4 and plasma membrane calcium channels remained unknown. Here, we show that auxin induces the direct interaction of the AFB1 receptor with the CNGC14 calcium channel. As the AFB1 receptor is independent of the ubiquitin ligase complex5, the auxin-induced interaction translates into relocalization of the receptor to the plasma membrane. We identify the interaction interface and provide evidence that the docking of the receptor to the channel complex activates Ca2+ influx and triggers growth inhibition. These findings position a calcium channel as an unprecedented component of the AFB1 auxin receptor complex. The ligand-dependent localization shift of a TIR1/AFB family receptor represents a novel paradigm in signal transduction and opens the possibility of unforeseen branches of auxin signaling pathways.