Meiotic recombination ensures accurate chromosome segregation and promotes genetic diversity by generating crossovers between homologous chromosomes. While essential in most sexually reproducing organisms, recombination is variably regulated and can be absent in some lineages, a condition known as achiasmy. However, obligate achiasmy in both sexes of a sexual species has not been previously documented. Here, we investigate the beak-sedge Rhynchospora tenuis, a holocentric plant with the lowest known chromosome number among flowering plants (n = 2) and inverted meiosis. Using chromosome-scale genome assemblies from nine accessions, molecular cytogenetics, immunocytochemistry, high-throughput single-gamete sequencing and whole-genome sequencing of controlled crosses, we show that R. tenuis undergoes obligate, genome-wide achiasmy in both male and female meiosis. Despite normal early meiotic axis formation, synapsis fails, crossovers are not detected cytologically or genetically, and univalents persist at metaphase I. Extensive haplotype-specific accumulation of transposable elements (TEs) generates segregation distortion (e.g. meiotic drive), favouring the transmission of larger, TE-rich chromosomes. Remarkably, sexual reproduction is retained with fertilisation producing viable seeds only when translocation-compatible gametes meet, indicating strong post-meiotic selection that eliminates incompatible homozygous combinations. As a result, all surviving offspring are genetically identical to the maternal genotype, effectively restoring heterozygosity each generation and mimicking clonal reproduction. We propose that the combined effects of recombination loss, low chromosome number, holocentricity, inverted meiosis, and selective transmission of longer chromosomes enable faithful segregation and clonal-like inheritance despite sexual reproduction. These findings challenge the boundary between sex and clonality, revealing a unique evolutionary strategy linking genome architecture, recombination loss, and transmission bias. ### Competing Interest Statement The authors have declared no competing interest.
Hop is an essential ingredient in brewing, providing beer with its characteristic bitterness and aroma. Most modern hop cultivars are hybrids between European and North American hop lineages, but how these ancestries contribute to bitter acid content, the most important trait in hop breeding, remains unclear. Here, we report chromosome-scale, haplotype-resolved assemblies of the hybrid hop cultivar Apollo, assign European and North American ancestry across the genome, and identify varying levels of recombination suppression between chromosomes of either origin. Using this reference, we uncover genetic and chemical diversity in core bittering pathways between European and North American hops. We further show additive effects of beneficial European and North American alleles on bitter acid content, providing a foundation for genomics-assisted hop breeding.
Advancements in plant genome sequencing and assembly have enabled the production of increasingly accurate and contiguous genome sequences. Here, we present the chromosome-level assembly of the durum wheat (Triticum turgidum L. ssp. durum, cv. Svevo) reference genome produced using accurate long-reads, optical mapping and Hi-C. The new assembly (Svevo Rel.2.0) comprises 263 hybrid scaffolds with an N50 value of 112.3 Mb, arranged into 14 contiguous pseudomolecules spanning 10.4 Gb. The Svevo Rel.2.0 genome assembly was annotated using extensive short- and long-read RNA sequencing data obtained from 60 tissue/treatment combinations. The resulting annotation comprises 68 154 high-confidence protein-coding genes, which have been integrated into a comprehensive transcriptome atlas accessible through an eFP browser. Annotation was manually curated for storage protein gene families and for Leucine-Rich Repeat-Containing Receptor genes yielding 3763 LRR-CR loci. The genome assembly's accuracy and completeness were demonstrated by the correct reconstruction of the physical map of Tg1-B (Tenacious glumes 1), a locus controlling the free threshing trait located on chromosome 2B that was not assembled in the previous genome release (Svevo Rel.1.0). A wealth of 6621 QTLs/MTAs from the literature were mapped onto Svevo Rel.2.0 to identify QTL hotspots and trait-specific candidate genes. The ancestry of the durum genome to representative wild emmer populations from North-Eastern and Southern-Levant Fertile Crescent assessed by tracing haplotype transmission patterns revealed a clear mosaic pattern. This new durum reference genome, enhanced with advanced annotation and an expression atlas linked to QTLome data, is the most comprehensive tool available for durum wheat genomics.
To exploit allelic variation in Hordeum vulgare subsp. spontaneum, the Wild Barley Diversity Collection was subjected to paired-end Illumina sequencing at ∼9 × depth and evaluated for several agronomic traits. We discovered 240.2 million single nucleotide polymorphisms (SNPs) after alignment to the Morex V3 assembly and 24.4 million short (1 to 50 bp) insertions and deletions. A genome-wide association study of lemma color identified one marker-trait association (MTA) on chromosome 1H close to HvBlp, the cloned gene controlling black lemma. Four MTAs were identified for seedling stem rust resistance, including 2 novel loci on chromosomes 1H and 6H and one co-locating to the complex RMRL1-RMRL2 locus on 5H. The whole-genome sequence data described herein will facilitate the identification and utilization of new alleles for barley improvement.
Abstract Drought is a major constraint on the productivity of durum wheat across Mediterranean and North African regions. To elucidate the mechanisms underlying drought resilience, we employed a combination of scenario-controlled phenomics and flag leaf transcriptomics across ten durum wheat genotypes. These included the Tunisian landraces Chili and Mahmoudi, seven breeding lines, and the reference cultivar Svevo. The plants were grown to maturity under well-watered or long-term drought conditions in pots and rhizotrons, enabling a comprehensive assessment of growth, yield components, root architecture, physiological traits, and reaction norm plasticity. Drought markedly reduced performance, yet Chili and Mahmoudi consistently maintained superior biomass, grain number and intrinsic water use efficiency (iWUE). This was supported by balanced C/N allocation, strong osmotic adjustment, and the ability to sustain robust root systems under stress, albeit through partly divergent physiological strategies. Transcriptomic profiling revealed highly genotype specific responses, with drought tolerance unrelated to the number of differentially expressed genes. Instead, the landraces displayed distinct regulatory programs involving mainly photosynthesis protection, ABA-related transporters, osmotic adjustment pathways, and stress-responsive transcription factors. These mechanistic insights identify actionable physiological and molecular determinants of drought plasticity and provide high value targets for accelerating the breeding of climate resilient durum wheat. Highlights Integrated phenomics and transcriptomics revealed landrace-specific physiological and molecular mechanisms enabling superior drought resilience and identifying actionable targets for durum wheat improvement.
Improving drought adaptation in wheat requires robust physiological traits that can be genetically dissected and translated into breeding tools. Carbon isotope composition (δ13C) is a stable proxy for intrinsic water-use efficiency (iWUE) and enables the identification of genetic factors underlying drought-stress responses. In this study, δ13C was measured in the Bavarian MAGIC Wheat (BMW) population, and quantitative trait locus (QTL) mapping at the drought-prone Morgenrot field site identified six significant loci on chromosomes 1D, 2B, 4B, 4D, 6A, and 6B. Based on phenotypic extremes at this site, eight genetically diverse lines with contrasting δ13C/iWUE values (four high δ13C and four low δ13C) were selected for detailed characterization. These extreme lines were evaluated under controlled greenhouse conditions using high-throughput phenotyping under both well-watered and drought-stress treatments.Transcriptomic analyses revealed pronounced drought-induced expression changes, with enrichment of genes involved in abscisic acid signaling, stress perception, stomatal regulation, and energy metabolism. To identify robust molecular targets, QTL mapping results were integrated with differential gene expression (DGE) and weighted gene co-expression network analysis in a multi-layer framework. This approach prioritized 18 high-confidence candidate genes located mainly on chromosomes 1D, 4B, 4D, 6A, and 6B. These genes are embedded in regulatory networks dominated by transcription factors such as WRKY and NAC. Collectively, this study provides genetically anchored candidate genes for iWUE and drought adaptation, offering valuable resources for marker-assisted selection in wheat breeding.
Wheat is the most widely cultivated crop in the world, with over 215 million hectares grown annually. The 10+ Wheat Genomes Project recently sequenced and assembled to chromosome-level the genomes of nine wheat cultivars, uncovering genetic diversity and selection within the pan-genome of wheat. Here, we provide a wheat pan-transcriptome with de novo annotation and differential expression analysis for these wheat cultivars across multiple tissues. Using the de novo annotations we identify cultivar-specific genes and define the core and dispensable genomes. Expression analysis across cultivars and tissues reveals conservation in expression between a large core set of homeologous genes, in addition to widespread changes in subgenome homeolog expression bias between cultivars and cultivar-specific expression profiles. We utilise both the newly constructed gene-based wheat pan-genome and pan-transcriptome, demonstrating variation in the prolamin superfamily and immune-reactive proteins across cultivars.
Stomatal guard cells, located at the interface between the leaf and the atmosphere, play a key role in transpiration control and photosynthetic CO2 uptake. Halophytes like Chenopodium quinoa tolerate high soil salinity, but the mechanisms governing guard cell responses to salinity stress in relation to the associated epidermal bladder cells (EBCs) remain unknown. In this study, responses of C. quinoa guard cells under salinity stress and external ABA application were analyzed using RNA profiling and voltage-clamp-based electrophysiology. Under salt stress, guard cell RNA profiles reported the activation of ABA synthesis and signaling pathways. However, unlike EBCs, guard cells exhibited a profoundly attenuated transcriptional response to ABA. Voltage-clamp recordings revealed that under high Na+ concentrations, guard cells' K+-uptake channel activity remained unaffected, while they were impaired in ABA-induced activation of anion channels. As a consequence of a unique guard cell ABA response in salt-adapted plants, stomatal transpiration was reduced and CO2 sensitivity was enhanced. We propose that under salt stress, C. quinoa guard cells rewire their hormone signaling to switch from an ABA-sensitive to a less ABA-responsive mode. This adaptation may reflect the halophyte's ability to perceive salinity as a nonstressful condition, allowing efficient water usage and sustained growth in saline environments.
Oat grain is a traditional human food that is rich in dietary fibre and contributes to improved human health1,2. Interest in the crop has surged in recent years owing to its use as the basis for plant-based milk analogues3. Oat is an allohexaploid with a large, repeat-rich genome that was shaped by subgenome exchanges over evolutionary timescales4. In contrast to many other cereal species, genomic research in oat is still at an early stage, and surveys of structural genome diversity and gene expression variability are scarce. Here we present annotated chromosome-scale sequence assemblies of 33 wild and domesticated oat lines, along with an atlas of gene expression across 6 tissues of different developmental stages in 23 of these lines. We construct an atlas of gene-expression diversity across subgenomes, accessions and tissues. Gene loss in the hexaploid is accompanied by compensatory upregulation of the remaining homeologues, but this process is constrained by subgenome divergence. Chromosomal rearrangements have substantially affected recent oat breeding. A large pericentric inversion associated with early flowering explains distorted segregation on chromosome 7D and a homeologous sequence exchange between chromosomes 2A and 2C in a semi-dwarf mutant has risen to prominence in Australian elite varieties. The oat pangenome will promote the adoption of genomic approaches to understanding the evolution and adaptation of domesticated oats and will accelerate their improvement.
Robinia pseudoacacia L. (black locust) is a nitrogen (N)-fixing legume tree with significant ecological and agricultural importance. Unlike well-studied herbaceous legumes, R. pseudoacacia is a perennial woody species, representing an understudied group of legume trees that establish symbiosis with Mesorhizobium. Understanding its genomic and transcriptional responses to nodulation provides key insights into N fixation in long-lived plants and their role in ecosystem N cycling. We assembled a high-quality 699.6-Mb reference genome and performed transcriptomic analyses comparing inoculated and noninoculated plants. Differential expression and co-expression network analyses revealed organ-specific regulatory pathways, identifying key genes associated with symbiosis, nutrient transport, and stress adaptation. Unlike Medicago truncatula, which predominantly responds to nodulation in roots, R. pseudoacacia exhibited stem-centered transcriptional reprogramming, with the majority of differentially expressed genes located in stems rather than in roots. Co-expression network analysis identified gene modules associated with "leghemoglobins", metal detoxification, and systemic nutrient allocation, highlighting a coordinated long-distance response to N fixation. This study establishes R. pseudoacacia as a genomic model for nodulating trees, providing essential resources for evolutionary, ecological, and applied research. These findings have significant implications for reforestation, phytoremediation, forestry, and sustainable N management, particularly in depleted, degraded, and contaminated soil ecosystems.
Deoxynivalenol (DON) is a toxicologically relevant trichothecene mycotoxin frequently found in cereal products. It is a virulence factor produced by the plant pathogen Fusarium graminearum during cereal crop infections. Investigating plant defense mechanisms is crucial for understanding plant resistance to F. graminearum and identifying new biocatalysts for DON detoxification. Previous studies identified DON-thiol adducts in cereal samples, indicating partial DON detoxification by glutathione transferases (GSTs). DON possesses two electrophilic centers for thiol conjugation, resulting in either epoxide opening at C13 or Michael addition at C10. At present, information on plant GSTs that catalyze these reactions is limited. In this study, Fusarium-inducible wheat GSTs were identified by analyzing the transcriptome of Fusarium-infected wheat heads. Twelve highly induced genes of the tau and phi GST classes were heterologously expressed and purified, biochemically characterized with model substrates, and assayed for activity with DON. Use of LC-MS showed that four of the selected tau class GSTs conjugated DON to GSH by epoxide opening (DON-13-GSH) and/or the reversible Michael addition reaction (DON-10-GSH). The crystal structure of a wheat GST (herein designated "TaGST-10") in complex with DON-13-GSH was solved at a resolution of 2.3 Å and provided insights into the binding of DON at the active site of tau class GSTs. Our results corroborate the hypothesis that enzyme-catalyzed, GSH-mediated DON detoxification may be involved in plant response to Fusarium infection.
The commercial value of economically significant fruits, including date palm fruit (dates), is influenced by various factors, such as biochemical composition and morphological features like size, shape, and visual appearance, which are key determinants of their quality and market value. Dates are typically consumed at the dry stage (Tamar), during which they exhibit a wide range of physical characteristics, such as color, length, weight, and skin appearance. Understanding the genetic basis of these traits is crucial for improving crop quality and breeding new cultivars. In this study, we integrated a genome dataset from highly diverse date cultivars with phenotypes of dry fruit such as length, width, area, and weight, identifying multiple significant genetic loci (SNPs) associated with these traits. We also identified candidate genes located near the associated SNPs that are involved in biological processes such as cell differentiation, proliferation, growth, and the regulation of signalling pathways for growth regulators like auxin and abscisic acid, as observed in other plants. Gene expression analysis reveals that many of these genes are highly expressed in the early stage of fruit development when the fruit attains its maximum size and weight. These findings will enhance our understanding of genetic determinants of fruit size particularly at the commercially important Tamar stage.
Genome sequence assemblies form a durable and precise framework supporting nearly all areas of biological research, including evolutionary biology, taxonomy and conservation science, pathogen population diversity, crop domestication, and biochemistry. In the early days of plant genomics, resources were limited to a handful of tractable genomes, leading to a tension between focus on discovering mechanisms in experimental species such as Arabidopsis thaliana (Arabidopsis) and on trait analyses in crop species. This tension arose from challenges in translating knowledge of gene function across the large evolutionary distances between Arabidopsis and diverse crop species in the absence of comparative genome support. For some time, these clashing interests influenced funding priorities in plant science that limited both the acquisition of knowledge of mechanisms in Arabidopsis and the timely development of the capacity of crop science to incorporate emerging knowledge of genes and their mechanisms. In this review we show how advances in genomics analysis technologies are revealing a high degree of conservation of molecular mechanisms between evolutionarily distant plant species. This progress is bridging the model-species-to-crop barrier, resulting in ever-increasing unification of plant science that is now accelerating progress in understanding mechanisms underlying diverse traits in crops and improving their performance. We lay out some examples of important priorities and outcomes arising from these new opportunities.
Wild plants can contribute valuable genes to their domesticated relatives1. Fertility barriers and a lack of genomic resources have hindered the effective use of crop-wild introgressions. Decades of research into barley's closest wild relative, Hordeum bulbosum, a grass native to the Mediterranean basin and Western Asia, have yet to manifest themselves in the release of a cultivar bearing alien genes2. Here we construct a pangenome of bulbous barley comprising 10 phased genome sequence assemblies amounting to 32 distinct haplotypes. Autotetraploid cytotypes, among which the donors of resistance-conferring introgressions are found, arose at least twice, and are connected among each other and to diploid forms through gene flow. The differential amplification of transposable elements after barley and H. bulbosum diverged from each other is responsible for genome size differences between them. We illustrate the translational value of our resource by mapping non-host resistance to a viral pathogen to a structurally diverse multigene cluster that has been implicated in diverse immune responses in wheat and barley.
Date palm (Phoenix dactylifera L.) is an important crop in arid regions and it is well adapted to desert ecosystems. To understand its remarkable ability to grow and yield in water-limited environments, we conducted experiments in which water was withheld for up to 4 weeks. In response to drought, root, rather than leaf, osmotic strength increased, with organic solutes such as sugars and amino acids contributing more to the osmolyte increase than minerals. Consistently, carbon and amino acid metabolism was acclimated toward biosynthesis at both the transcriptional and translational levels. In leaves, a remodeling of membrane systems was observed, suggesting changes in thylakoid lipid composition which, together with the restructuring of the photosynthetic apparatus, indicated an acclimation preventing oxidative damage. Thus, xerophilic date palm avoids oxidative damage under drought by combined prevention and rapid detoxification of oxygen radicals. Although minerals were expected to serve as cheap key osmotics, date palm also relies on organic osmolytes for osmotic adjustment in the roots during early drought acclimation. The diversion of these resources away from growth is consistent with the date palm strategy of generally slow growth in harsh environments and clearly indicates a trade-off between growth and stress-related physiological responses.
Duckweeds are among the smallest and fastest-growing flowering plants. In a new study that combines experimental data with phylogenomic comparisons across the clade, the authors explore how changes in gene content, epigenetic pathways, and their interplay shaped the body plan, aquatic lifestyle, and clonal growth habit of this plant family.
A pan-transcriptome describes the transcriptional and post-transcriptional consequences of genome diversity from multiple individuals within a species. We developed a barley pan-transcriptome using 20 inbred genotypes representing domesticated barley diversity by generating and analyzing short- and long-read RNA-sequencing datasets from multiple tissues. To overcome single reference bias in transcript quantification, we constructed genotype-specific reference transcript datasets (RTDs) and integrated these into a linear pan-genome framework to create a pan-RTD, allowing transcript categorization as core, shell or cloud. Focusing on the core (expressed in all genotypes), we observed significant transcript abundance variation among tissues and between genotypes driven partly by RNA processing, gene copy number, structural rearrangements and conservation of promotor motifs. Network analyses revealed conserved co-expression module::tissue correlations and frequent functional diversification. To complement the pan-transcriptome, we constructed a comprehensive cultivar (cv.) Morex gene-expression atlas and illustrate how these combined datasets can be used to guide biological inquiry.
A pan-transcriptome describes the transcriptional and post-transcriptional consequences of genome diversity from multiple individuals within a species, revealing an assortment of functions that drive biological outcomes. We developed a barley pan-transcriptome using twenty inbred genotypes representing domesticated* barley diversity by generating and analysing extensive short- and long-read RNA sequencing datasets from multiple tissues. To overcome single reference bias and facilitate downstream analyses we constructed genotype-specific reference transcript datasets (RTDs) and integrated these into a linear pan-genome framework to create a single pan-RTD. Categorising transcripts based upon presence or absence across genotypes defined them as core (expressed in all), shell (absent in one or more) or cloud (expressed in only one). Focusing on the core we observed significant transcript abundance variation among tissues and between genotypes. We show that drivers of transcript abundance variation in this category include RNA processing, gene copy number, large structural rearrangements and degree of conservation of promotor motifs. We reveal conserved patterns of co-expression module-tissue correlations encompassing distinct biological functions, as well as frequent functional diversification. We complement the pan-transcriptome by integrating extensive and diverse replicated public RNA-seq datasets from the reference cultivar (cv.) Morex into a comprehensive gene-expression atlas
The generation of antibody-drug conjugates with optimal functionality depends on many parameters. These include binder epitope, antibody format, linker composition, conjugation site(s), drug-to-antibody ratio, and conjugation method. The production of matrices that cover all possible parameters is a major challenge in identifying optimal antibody-drug conjugates. To address this bottleneck, we adapted our Format Chain Exchange technology (FORCE), originally established for bispecific antibodies, toward the generation of binder-format-payload matrices (pair-FORCE). Antibody derivatives with exchange-enabled Fc-heterodimers are combined with payload-conjugated Fc donors, and subsequent chain-exchange transfers payloads to antibody derivatives in different formats. The resulting binder-format-conjugate matrices can be generated with cytotoxic payloads, dyes, haptens, and large molecules, resulting in versatile tools for ADC screening campaigns. We show the relevance of pair-FORCE for identifying optimal HER2-targeting antibody-drug conjugates. Analysis of this matrix reveals that the notion of format-defines-function applies not only to bispecific antibodies, but also to antibody-drug conjugates. Designing optimal antibody-drug conjugates (ADCs) involves screening many complex parameters. Here, the authors present a payload-coupled chain-exchange technology for efficient ADC matrix production, demonstrating its power in designing ADCs targeting HER2.