Biology subjects

Weigel, D.

Publications and source records attributed to Weigel, D..

At least 19 recordsLinked to original sources

bioRxiv2026

Dynamic co-existence of bacteriophages and their hosts in the Arabidopsis thaliana phyllosphere

Bacterial communities and the bacteriophages infecting them are the basis of every ecosystem, including holobionts. The various ways in which these microorganisms interact with each other in complex communities over the life of the host affects the holobiont fitness. Despite being ubiquitous and environmentally relevant, plant-associated microbial communities remain understudied, especially in the phyllosphere, mainly because of the low abundance of microbes and the complexity of the system. In this work we followed bacteria and phage community dynamics in the phyllosphere over a growing cycle of Arabidopsis thaliana, to understand the ecology and relevance of bacteriophages in complex bacterial communities. We focused on Pseudomonas, a common plant pathogen and commensal, and the phages infecting them, in three setups of increasing complexity: in vitro, controlled experiments in planta and in wild populations of A. thaliana. We found that bacterial communities are resilient to phage infection, and more dynamic than the phages infecting them over the growing season, suggesting that although ubiquitous and abundant, bacteriophages exert selective pressures on leaf bacterial communities only intermittently.

microbiology↗

bioRxiv2026

Biotic-response networks are an important organizer of the transcriptome in wild Arabidopsis thaliana populations

Extensive laboratory experimentation has revealed conserved molecular pathways controlling growth and stress responses in plants, yet how these programs operate in natural settings remains poorly understood. We investigated transcriptome organization in wild populations of Arabidopsis thaliana by sampling plants from 60 natural sites in Europe and North America across two seasons. Transcriptomes varied extensively among individuals and showed largely continuous rather than discrete structure across geography and season. Although disease and microbial colonization were common in the wild, wild transcriptomes did not simply recapitulate canonical laboratory stress signatures. Measured microbial infection, environmental, and phenotypic variables explained only a modest fraction of total expression variation, but infection-associated signals accounted for the largest share of the explainable component. Consistent with this, biotic-response networks defined in controlled laboratory experiments were well conserved in wild transcriptomes, whereas control and abiotic-response networks were substantially reorganized. Together, these results suggest that while core transcriptional modules remain recognizable across environments, regulatory relationships among modules differ markedly between laboratory and natural contexts. Graphical Abstract O_FIG O_LINKSMALLFIG WIDTH=200 HEIGHT=129 SRC="FIGDIR/small/711176v1_ufig1.gif" ALT="Figure 1"> View larger version (28K): org.highwire.dtl.DTLVardef@1c89152org.highwire.dtl.DTLVardef@bcd1acorg.highwire.dtl.DTLVardef@6975a2org.highwire.dtl.DTLVardef@1a9e2fe_HPS_FORMAT_FIGEXP M_FIG C_FIG

genomics↗

bioRxiv2026

Biosynthetic gene clusters in Pseudomonas viridiflava have a fitness cost during Arabidopsis thaliana infection

Specialized or secondary metabolites mediate biotic interactions, including virulence and defense. In plant-pathogenic Pseudomonas, certain specialized metabolites can enhance colonization of plant hosts, yet their broader contribution to plant-microbe interactions and the relative importance of different metabolites remain unclear. Specialized metabolites are products of enzymes encoded in biosynthetic gene clusters (BGCs), whose prediction from genome sequences has become routine but whose functional roles are rarely tested experimentally. Here, we characterize the BGC repertoire of 225 P. viridiflava isolates from Arabidopsis thaliana and assess BGC contributions to fitness in planta and disease severity. The BGC landscape of P. viridiflava was dominated by non-ribosomal peptide synthetase (NRPS) and NRPS-like BGCs, with one-third of families restricted to a single isolate. Transposon mutagenesis coupled with random barcode transposon sequencing (RB-TnSeq) revealed that the majority of BGCs reduce rather than increase fitness during A. thaliana infection, with the magnitude of the fitness cost varying across host genotypes. This cost could be due to exploitation of public goods by cheater mutant strains. In single-isolate plant infections, where public goods are not available, several BGC families were negatively associated with disease severity, which is positively correlated with bacterial growth in this pathosystem, further indicating that BGCs are generally not beneficial in planta. Our findings reveal extensive and largely uncharacterized biosynthetic potential in populations of P. viridiflava and indicate that candidate metabolites are likely not adaptive for direct interactions with the plant, but perhaps for microbe-microbe interactions either in planta or in other ecological niches. IMPORTANCEBacteria living on plant leaves produce a vast array of chemical compounds, called secondary or specialized metabolites, that can mediate their interaction with the plant host or other microorganisms. Some of these compounds are known to directly influence how bacteria interact with plants, but it has been unclear whether this is a general rule. We studied a large collection of closely related leaf-dwelling bacteria that varied in their ability to cause disease, focusing on leaf-associated Pseudomonas viridiflava--a plant pathogen. We found that very few of the gene clusters responsible for making specialized metabolites improved the ability of the bacteria to colonize Arabidopsis thaliana. On the contrary, carrying these gene clusters often reduced bacterial growth and disease severity in plants. Specialized metabolites may instead primarily be important for interacting with other microbes, different host species, or under environmental conditions we did not test. These are questions that remain for future research.

microbiology↗

bioRxiv2026

Recruitment of SERK co-receptors determines signaling specificity within the systemin peptide family

The phytocytokine systemin (SYS1) regulates defense responses of tomato plants against herbivores and necrotrophic pathogens. For more than three decades, systemin signaling was thought to rely solely on SYS1, although the tomato genome encodes the precursors of at least three additional systemin-like peptides (SYS2, SYS3 and SYS4). We show that these recently discovered peptides are processed in a wound-inducible manner and act as genuine wound signals that activate early defense signaling via the receptor SYR1. Although all systemins depend on SYR1 for perception, the new peptides elicit downstream responses distinct from those triggered by SYS1, including sustained ethylene production and extensive transcriptional reprogramming. Signaling specificity is governed by a single C-terminal residue that determines the ability of SYS peptides to recruit SERK co-receptors into the receptor complex. The resulting increase in complex stability underlies the amplified and prolonged signaling responses observed for the new SYS peptides compared to SYS1. Our findings reveal how subtle sequence changes within the systemin peptide family can remodel receptor-co-receptor interactions to generate ligand-specific signaling outputs. This mechanism allows a single receptor to decode related ligands into distinct physiological responses and may represent a broader principle allowing for the diversification of peptide-mediated signaling in plants.

plant biology↗

bioRxiv2026

Natural selection acting on the genetics of host response to commensal bacteria

BackgroundMicrobiota members collectively contribute to individual performance in humans, animals, and plants. This led to the quest for probiotics to improve host health and reproductive performance. Although the efficacy of probiotics is known to be strongly affected by environmental factors, microbe-microbe interactions, and microbial strain identity, the effects of host genotype and the underlying genetic architecture have been overlooked. In addition, the evolutionary causes of such genetic variation are typically not addressed. In this study, we aimed to describe the genetic architecture of the adaptation of the host plant Arabidopsis thaliana to commensal bacterial members of its native microbiota by identifying candidate genes associated with fitness proxies and presenting signatures of natural selection. ResultsA Genome-Wide Association study conducted under field conditions revealed extensive variation within a new mapping population of 162 genotypes of A. thaliana scored for total seed production and its two underlying components, namely fruit number and mean seed number per fruit, in response to 13 commensal strains. In agreement with the strong host genotype x commensal strain identity interactions observed for each reproductive trait, the polygenic genetic architecture was highly flexible among the 13 commensal strains. Candidate genes exhibited a significant enrichment in signatures of both local adaptation and balancing selection. In line with the phenotyped reproductive traits, we identified seven candidate genes with functions specifically and strongly linked to seed germination and fertility. ConclusionsOur findings reveal the importance of genotype-by-genotype interactions when measuring fitness proxies on a wild plant species inoculated with key members of its native microbiota. In addition, this study improves our understanding of the genetic signatures of natural selection acting on native host-microbiota adaptive interactions.

evolutionary biology↗

bioRxiv2026

Multiple redundant mechanisms account for the majority of gene silencing downstream of DNA methylation

DNA methylation is a conserved epigenetic modification crucial for silencing genes and transposable elements (TEs). However, the mechanisms that cause silencing remain unclear, partly because methyl reader protein mutants in both plants and animals show minimal transcriptional changes. To explore the possibility of redundancy among these silencing mechanisms, we generated combinatorial mutants of H1.1, H1.2, ADCP1, MOM1, MBD2, MBD5, and MBD6 lacking key methyl readers and related silencing pathways. We observed massive derepression of genes and TEs at DNA-methylated loci, showing that these pathways account for 73% of silencing compared to DNA methylation-free mutants. We also observed that immune response genes were upregulated, causing an imbalance between growth and defense. Loss of downstream silencing pathways further disrupted 3D genome organization, leading to increased euchromatin-heterochromatin interactions. These findings highlight the cooperative action of multiple downstream mechanisms in DNA methylation-mediated silencing and genome organization.

molecular biology↗

bioRxiv2025

PanGene-O-Meter: Intra-Species Diversity Based on Gene-Content

BackgroundBacterial genome evolution is shaped extensively by horizontal gene transfer, generating gene presence-absence variation that represents a major axis of functional diversity yet is largely invisible to core-genome-based similarity metrics. While genome-wide average nucleotide identity (ANI) remains the most widely used measure of similarity, it considers only the alignable fraction of compared genomes and therefore fails to capture gene-content differences that can have profound phenotypic consequences even among nearly identical strains. ResultsHere we introduce PanGene-O-Meter, a computational framework that quantifies gene-content similarity (GCS) between bacterial genomes by leveraging pan-genome structure and orthology group relationships. Applying PanGene-O-Meter to four large datasets spanning Escherichia coli, Pseudomonas aeruginosa, Staphylococcus aureus, and Helicobacter pylori, we show that GCS captures substantial functional diversity hidden within high-ANI genome pairs, and that gene-content-based trees resolve substructure within sequence types that core-genome analyses cannot. We further introduce GeneContRep, an efficient greedy algorithm for gene-content-based genome dereplication, and validate its utility on 1,000 Klebsiella pneumoniae genomes from two major multidrug-resistant lineages (ST258 and ST307) with matched experimental antimicrobial resistance phenotypes. GeneContRep preserves substantially more clinically relevant diversity than standard ANI-based dereplication, recovering carbapenemase variants, capsule locus types, and resistance determinants that are systematically lost under conventional ANI thresholds. ConclusionsPanGene-O-Meter provides an efficient, complementary, and biologically interpretable framework for quantifying intra-species bacterial diversity, with broad applications in comparative genomics, epidemiological surveillance, and the principled selection of representative genomes for functional studies. PanGene-O-Meter is available at: https://github.com/HaimAshk/PanGene-O-Meter.

microbiology↗

bioRxiv2025

Cell-type specific gating of gene regulatory modules as a hallmark of early immune responses in Arabidopsis leaves

In plants, many cell types contribute to immunity, but what division of labor exists among cell types when immunity is activated? We compared, at single-cell resolution, the response of Arabidopsis thaliana leaf cells during pattern-triggered and effector-triggered immunity (PTI/ETI), sampled at 3 and 5 hours after infection with Pseudomonas syringae DC3000. Core defense modules were broadly shared across cell clusters, but their activation varied in timing and intensity, with key immune receptors also showing cell type-specific expression dynamics. We identified distinct mesophyll cell populations based on their resilience patterns: after the initial response, some cells continue to express defense genes at high levels during both PTI and ETI, while others quickly reinitiate growth-related gene expression programs, but only during PTI. Gene regulatory network inference revealed WRKY-regulated modules enriched in cells sensing effectors, while SA biosynthesis regulators were activated in complementary clusters. Finally, we used the cue1-6 mutant to demonstrate that core immune responses are robust despite altered leaf architecture. In addition, we uncovered cryptic defense pathways, including sucrose-responsive modules, in this mutant. By capturing early immune responses at high resolution, our study reveals cell type-specific coordination of plant immunity and provides a framework for decoding immune signaling networks.

plant biology↗

bioRxiv2025

Seasonality, land use, and diversity shape microbiome-pathogen interactions in wild populations of Arabidopsis thaliana

The microbiome often protects plants against pathogens, but most findings are limited to controlled experiments in the lab. In the context of wild populations, one key challenge is to understand sources of variation that impact the commensal microbiome, which in turn shapes the degree of protection. Here, we surveyed both disease symptoms and microbiomes from wild populations of Arabidopsis thaliana over four consecutive seasons (fall/spring) across three different land use types. Land use types varied in the extent of anthropogenic influences and included forest meadows, human-impacted fields adjacent to agriculture or municipal parks, and highly disturbed habitats near railroad tracks. By building an integrative map of abiotic and biotic variables, we find that a key predictor of disease was biodiversity across ecological scales. Plant communities with higher diversity were associated with reduced disease burden in A. thaliana populations but also increased diversity within the microbiome of A. thaliana. This increased microbial diversity was additionally associated with less disease in A. thaliana. However, the diversity-microbiome-disease relationships were all sensitive to season and further modulated by land use. Taken together, our work highlights the importance of anthropogenic change reshaping species interactions across ecological scales to impact disease risk in wild plant populations.

ecology↗

bioRxiv2025

The mutational dynamics of the Arabidopsis centromeres

Centromeres are specialized chromosomal regions essential for sister chromatid cohesion and spindle attachment during cell division. Although many centromeres consist of highly variable tandem repeat arrays, the mutational processes driving this variability remain poorly understood. Here, using replicated genome assemblies of Arabidopsis thaliana mutation accumulation (MA) lines, we define the centromeric mutation spectrum. We find that point mutations occur at an almost tenfold higher rates than in chromosome arms, largely driven by non-allelic gene conversion between closely linked repeat units. Large kilobase-scale indels are also frequent and consistently preserve the tandem repeat array by adding or removing only complete repeat units. Analysis of MA lines deficient in the helicase RTEL1 supports the involvement of homology-directed DNA repair in these mutational processes. Moreover, simulations of sequence turnover using the centromere-specific mutation spectrum recapitulate the emergence of homogenized repeat blocks characteristic of natural centromeres. Together, our results show that centromere evolution is driven by a distinct mutational spectrum shaped by homology-directed DNA repair, providing a quantitative framework for how local mutational processes generate and maintain the large-scale architecture of centromeric DNA.

genomics↗

bioRxiv2025

Minimizing detection bias of somatic mutations in a highly heterozygous oak genome

Somatic mutations are particularly relevant for long-lived organisms. Sources of somatic mutations include imperfect DNA repair, replication errors, and exogenous damage such as ultraviolet radiation. A previous study estimated a surprisingly low number of somatic mutations in a 234-year-old individual of the pedunculate oak (Quercus robur), known as the Napoleon Oak. It has been suggested that the true number of somatic mutations was underestimated due to gaps in the reference genome and too conservative filtering of potential mutations. We therefore generated new high-fidelity long-read data for the Napoleon Oak (n = 12) to produce both a pseudo-haploid genome assembly and a partially phased diploid assembly. The high heterozygosity allowed for complete reconstruction of phased and gapless centromeres for 22 of the 24 chromosomes. On the other hand, the high heterozygosity posed challenges for short-read alignments. Use of only the pseudo-haploid assembly as a reference led to potential misalignments, while use of only the diploid assembly reduced variant detection sensitivity. Since most somatic mutations are layer-specific, the observed frequency is expected relatively low, even where all cells in a single layer contain a specific mutation. To address this challenge, we employed a read assignment strategy, selecting the appropriate reference sequence (pseudo-haploid or diploid) based on alignment score and mapping quality. Ultimately, we identified 198 high-confidence somatic mutations, compared with 17 somatic mutations identified before with the same set of short reads. Our approach thus increased the total estimated annual mutation rate by a factor of five.

genomics↗

bioRxiv2025

Long-read detection of transposable element mobilization in the soma of hypomethylated Arabidopsis thaliana individuals

BackgroundBecause transposable elements (TEs) can cause heritable genetic changes, past work on TE mobility in Arabidopsis thaliana has mostly focused on new TE insertions in the germline of hypomethylated plants. It is, however, well-known that TEs can also be active in the soma, although the high-confidence detection of somatic events has been challenging. Here, we leveraged the high accuracy of PacBio HiFi long reads to evaluate the somatic mobility of TEs in individuals of an A. thaliana non-reference strain lacking activity of METHYLTRANSFERASE1 (MET1), a major component of the DNA methylation maintenance machinery. Most somatically mobile families coincided with those found in germline studies of hypomethylated genotypes, although the exact TE copies differed. We also discovered mobile elements that had been missed by standard TE annotation methods. Somatic TE activity was variable among individual plants, but also within TE families. Finally, our approach pointed to the possible involvement of alternative transposition as a cause for somatic hypermutability in a region that contains two closely spaced VANDAL21 elements. We conclude that long-read sequencing can reveal widespread TE transposition in the soma of A. thaliana hypomethylated mutants. Assessing somatic instead of germline mobilization is a fast and reliable method to investigate different aspects of TE mobility at the single plant level.

genomics↗

bioRxiv2025

The structure of mitochondrial genomes is associated with geography in Arabidopsis thaliana

Chloroplasts and mitochondria are the primary sites for photosynthesis and respiration, each harboring its own unique genome. Although the organellar genomes are considerably smaller compared to the nuclear genome, they are nonetheless essential for survival of the organism. A common feature of many chloroplast and mitochondrial genomes is the presence of large repeated sequences longer than 1 kb. These can be either in inverted or direct orientation, and recombination between them leads to structural heteroplasmy. To understand the intraspecific evolution of organellar genomes, we assembled chloroplast and mitochondrial genomes of 143 A. thaliana accessions from PacBio HiFi sequencing data. We find large repeats to be associated with heteroplasmy and structural variation. Our extensive genome annotation identifies novel open reading frames (ORFs) in those accessions that lost large repeats, potentially introduced via horizontal gene transfer, illuminating additional paths for diversification of plant organelles. The loss of large repeats correlates with geography and phenotypes, pointing to their adaptive importance. The assembled and annotated organellar genomes constitute a rich source for future functional studies of the interaction between the three genomes of a plant.

evolutionary biology↗

bioRxiv2025

Natural variation modifies centromere proximal meiotic crossover frequency and segregation distortion in Arabidopsis thaliana

BackgroundCentromeres mediate chromosome segregation during cell division. In plants, centromeres are loaded with CENH3-variant nucleosomes, which direct kinetochore formation and spindle microtubule interaction. Plant centromeres are frequently composed of megabase-scale satellite repeat arrays, or retrotransposon nests. In monocentric genomes, such as Arabidopsis thaliana, extended regions of pericentromeric heterochromatin surround the CENH3-occupied satellite arrays. A zone of suppressed meiotic crossover recombination contains the centromere and extends into the pericentromeres. Here, we explore how Arabidopsis natural genetic variation influences centromere-proximal crossover frequency, and segregation distortion through meiosis, when homologous centromeres are structurally heterozygous. ResultsWe used fluorescent crossover reporters to survey the effect of natural variation on centromere-proximal recombination in twelve F1 hybrids, capturing Arabidopsis Eurasian and relict diversity. The majority of F1 hybrids showed either elevated or suppressed centromere-proximal crossovers (49 of 60), relative to inbreds. We relate hybrid crossover frequencies to patterns of centromeric structural variation, and in a subset of accessions, to epigenetic patterns of CENH3 loading and DNA methylation. The fluorescent reporters also allow segregation distortion through meiosis to be compared between inbred and hybrid strains. We observed a minority of hybrids (18 of 60) with distorted segregation through meiosis compared to inbreds, with and without simultaneous change to centromere-proximal crossover frequency. ConclusionsWe reveal a complex relationship between centromere structural variation, epigenetic state, crossover recombination, and segregation distortion. We propose a model for how Arabidopsis centromere structural heterozygosity may cause segregation distortion during female meiosis.

genetics↗

bioRxiv2024

The 1001G+ project: A curated collection of Arabidopsis thaliana long-read genome assemblies to advance plant research

Arabidopsis thaliana was the first plant for which a high-quality genome sequence became available. The publication of the first reference genome sequence almost 25 years ago was already accompanied by genome-wide data on sequence polymorphisms in another accession, or naturally occurring strain. Since then, inventories of genome-wide diversity have been generated at increasingly precise levels. High-density genotype data for A. thaliana, including those from the 1001 Genomes Project, were key to demonstrating the enormous power of GWAS in inbred populations of wild plants, and the comparison of intraspecific polymorphism with interspecific divergence has illuminated many aspects of plant genome evolution. Over the past decade, an increasing number of nearly complete genome sequences have been published for many more accessions. Here, we highlight the diversity of a curated collection of previously published and so far unpublished genome sequences assembled using different types of long reads, including PacBio Continuous Long Reads (CLR), PacBio High Fidelity (HiFi) reads, and Oxford Nanopore Technologies (ONT) reads. This 1001 Genomes Plus (1001G+) resource is being made available at http://1001genomes.org. We invite colleagues with yet unpublished genome assemblies from A. thaliana accessions to contribute to this effort.

genomics↗

bioRxiv2024

Gfa2bin enables graph-based GWAS by converting genome graphs to pan-genomic genotypes

Variation graphs offer superior representation of genomic diversity compared to traditional linear reference genomes, capturing complex features that are otherwise inaccessible to analysis. It seems self-evident that integrating these graphs with genome-wide association studies (GWAS) should enable more comprehensive understanding of genetic landscapes, potentially uncovering novel associations between genetic variations and traits. This approach takes full advantage of rich genomic information, thereby providing deeper insights into the genetic base of complex traits. Our tool, gfa2bin, offers multiple methods to (i) genotype variation graphs and (ii) convert the genotypes to well-established data formats for genome-wide association studies (GWAS). We demonstrate that variation graphs are feasible alternatives to traditional linear references for GWAS. Our case study using Arabidopsis thaliana and 1,695 traits shows that our approach complements SNP-based approaches, often identifying additional associations, with all associations having on average higher significance compared to SNP-based approaches. gfa2bin is implemented in Rust. Commented source code is available under MIT license at https://github.com/MoinSebi/gfa2bin. Examples of how to run gfa2bin are provided in the documentation. We added several Python scripts and a Snakemake pipeline for easy processing of our tool using larger data sets. In addition, we recommend using packing (https://github.com/MoinSebi/packing) for reduced storage and preprocessing (normalization) of sequence-to-graph alignments coverage.

bioinformatics↗

bioRxiv2024

Arabidopsis PRMT5 Buffers Pre-mRNA Splicing and Development Against Genetic Variation in Donor Splice Sites

Genetic variation at splice site signals significantly influences alternative splicing, leading to transcriptomic and proteomic diversity that enhances phenotypic plasticity and adaptation. However, novel splice variants can negatively impact gene expression and developmental stability. Canalization--the ability of an organism to maintain a consistent phenotype despite genetic or environmental variations--helps balance the effects of genetic variation on development and evolution. Protein arginine methyltransferase 5 (PRMT5) is a key splicing regulator in plants and animals. Most splicing changes in prmt5 mutants are linked to weak donor splice sites, suggesting that PRMT5 may buffer splicing against genetic variation. We examined PRMT5s effects on splicing and development in two genetically divergent Arabidopsis thaliana accessions with different single nucleotide polymorphisms (SNPs) affecting donor splice sites. While PRMT5 inactivation similarly affected splicing in both backgrounds, it significantly increased splicing and phenotypic differences between the accessions. Our findings suggest that PRMT5 contributes to canalization, mitigating the impact of splice site polymorphisms and facilitating the evolution of adaptive splicing patterns.

molecular biology↗

bioRxiv2024

The spliceosomal component GAMETOPHYTIC FACTOR 1 (GFA1) regulates a key photoperiodic switch

Plants have evolved sophisticated mechanisms to perceive and interpret daytime in order to flower at an optimal time point. Here we show that the spliceosomal component GAMETOPHYTIC FACTOR 1 (GFA1) constitutes a previously unrecognized key photoperiodic switch, that is essential for flowering in long days. We show that gfa1 hypomorphic (gfa1hyp) plants fail to initiate flowering in long-days (LD), which correlates with ectopic activation of the short-day (SD) flowering repressor ARABIDOPSIS THALIANA CENTRORADIALIS (ATC). Accordingly, flowering is restored upon inactivation of ATC in gfa1hyp mutants. A novel tissue-specific in-planta splice assay and comprehensive RNAseq profiling of gfa1hyp mutants indicate that GFA1 mediated pre-mRNA splicing is substrate specific, as previously suggested for GFA1 orthologs. Furthermore, we show that gfa1hyp mutants accumulate nonsense transcripts of the photoreceptor components PHYB and RRC1, suggesting inappropriate photoreceptor signaling as a potential cause for the ectopic activation of the SD characteristic profile in gfa1hyp. In fact, known downstream targets of the phytochrome system such as RS31, SR34a, SRp30 accumulate reduced amounts of light-dependent splice isoforms. Together, our data reveal a link between spliceosome composition and long-day flowering, based on complex transcriptional readouts in response to day length.

developmental biology↗