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Schnurbusch, T.

Publications and source records attributed to Schnurbusch, T..

At least 19 recordsLinked to original sources

Spatial metabolomics links polyamine turnover and vascular lipid remodeling to barley spikelet fate

Local metabolic programs can determine whether developing organs maintain growth, differentiate, or degenerate. However, the spatial establishment of such programs during reproductive development remains poorly understood. In cereal inflorescences, this question is closely tied to grain number because the fate of initiated floral primordia is shaped by the developmental position, metabolic support, genotype, and environmental stress. To understand the metabolic logic underlying floral fate in barley (Hordeum vulgare L.), we mapped the spatial distribution of metabolites across the inflorescence in wild-type Bowman and in hvcmf4, a mutant that undergoes premature apical inflorescence degeneration. Amino acid-, carbohydrate-, and chlorophyll-associated metabolites form clear developmental gradients along the inflorescence axis. These gradients are progressively remodeled during developmental pre-anthesis tip degeneration but are both accelerated and spatially distorted in hvcmf4. Within this broader metabolic landscape, two discrete domains stand out. First, spermidine-associated domains specifically marked reproductive meristems, a pattern corroborated by the meristem-enriched expression of polyamine biosynthetic genes. This polyamine-rich meristem state declined as developmental tip degeneration proceeded and was prematurely lost in hvcmf4, where polyamine metabolism appeared to shift toward oxidative breakdown via the polyamine oxidase pathway. Second, lysophosphatidylcholine accumulated specifically at floral attachment and vascular supply zones. The developmental reduction of this bioactive lysophospholipid-associated domain in the apical regions, and its stronger disruption in hvcmf4, coincided with the reprogramming of lipid-remodeling and auxin transport-associated genes, suggesting impaired coordination of lipid signaling, auxin canalization, and vascular support. Together, our study reveals spatial polyamine and lysophospholipid domains that couple meristem maintenance with vascular support, providing a metabolic framework for the fate of developing cereal inflorescences.

plant biology↗

Rate of meristem initiation driven by the MADS-WUS axis contributes to floral survival and inflorescence evolution in grasses

Crop domestication has repeatedly shaped inflorescence architecture to improve floral production, but mechanisms coordinating the rate of floral initiation, maturation and survival remain unclear. Combining morphometry, modelling and molecular genetic analyses, we show that floral production in the indeterminate barley (Hordeum vulgare L.) inflorescence follows an "initiate fast-die young" strategy orchestrated by a main MADS-box gene, SPIKELET INITIATION AND FERTILITY (SIF). SIF accomplishes this duality by coordinately terminating the inflorescence meristem via WUSCHEL and activating the floral meristem via APETALA1 (Vrn-H1). Hereby, the ancestral SIF "slow" allele promotes a timely commitment to floral maturation, whereas the derived "fast" allele permits more floral initiations. Postdomestication selection of SIF alleles thus enables diversified reproductive strategies in barley populations to maintain yield traits in the field. Finally, we show that a lineage-specific SIF duplication contributed to meristem fate transition and inflorescence evolution during Triticeae cold adaptation. Our results establish developmental rate as a key driver of architectural innovation and reproductive success.

plant biology↗

Defining critical drivers of cross-pollination for better hybrid grain set in wheat

Hybrid wheat breeding offers a promising route to enhance grain yield and yield stability through heterosis, yet hybrid grain production remains constrained by limited cross-pollination efficiency due to high rates of autogamy. To achieve cross-pollination in an autogamous species like wheat, pollen must shed outside the floret. This is typically assessed by scoring visual anther extrusion (VAEX), a key floral trait that sets the foundation for cross-pollination. However, VAEX explains only part of the variation in hybrid grain set. To address this, we analyzed floral structures and reproductive processes underlying cross-pollination efficiency in wheat. From 24 elite winter wheat genotypes, we developed traits describing anther extrusion kinetics, pollen release, and floral bract architecture. These traits showed substantial genotypic variation and high heritability. While VAEX alone explained approximately 49% of the variation in hybrid grain set, combined trait analyses explained up to 77%, demonstrating that hybrid grain production is governed by coordinated floral and reproductive trait interactions. Together, our analyses define a hierarchical trait architecture linking floral bract mechanics, anther extrusion dynamics, and pollen shedding to cross-fertilization success. This establishes a systems-level phenotyping framework for improving male parent selection in hybrid wheat breeding. HighlightHigh cross-pollination efficiency in wheat is a multi-factorial process that requires lighter floral bract architecture combined with adequate anther extrusion and pollen release for improving hybrid grain production.

developmental biology↗

Structural and coding variation in PHYTOCHROMES A and C underlies differences in flowering time and shade avoidance in wheat

Plant architecture and developmental timing are influenced by light availability, especially in high-density cropping systems, where canopy shading modifies both light intensity and spectral quality. Despite their ecological and agronomical importance, the genetic basis of these responses in wheat remains poorly understood. Here, using a Recombinant Inbred Lines (RILs) population, we investigated phenological and morphological traits under sunlight and simulated canopy shade. We identify a major QTL on chromosome 5A with light-dependent allelic effects, indicating genotype-by-environment variation in developmental responses. This QTL corresponds to a structural rearrangement, consistent with an inversion in the wild emmer reference genome encompassing PHYTOCHROME C (PHYC-A) and VERNALIZATION-1 (VRN-A1), as well as coding polymorphism in PHYC-A. Analysis of a tetraploid wheat diversity panel further showed that natural variation at PHYC-A and an early stop codon in the BB genome copy of PHYTOCHROME A (PHYA-B) on chromosome 4B are associated with differences in heading time. Functional analysis using TILLING-derived phytochrome mutants confirms distinct and complementary roles for PHYA and PHYC in regulating flowering time, plant height, and leaf elongation under simulated canopy shade. These findings highlight the contribution of phytochrome variation to developmental plasticity under canopy-like light environment, thereby extending model insights to agronomically relevant conditions.

plant biology↗

Homoeo-alleles of wheat GNI2 fuel grain yields across input environments

Grain yield in wheat is frequently driven by pre-anthesis growth and how carbon is directed to reproductive parts, determining the potential size of the grain-producing sink. However, the genetic mechanisms controlling this carbon allocation remain unclear. In this study we discovered a series of loci in tetraploid and hexaploid wheats that, when combined, confer an average grain yield advantage in hexaploid winter wheat of approx. 5-7% under high-input farming while approx. 10-15% under low-input conditions. The responsible gene is GRAIN NUMBER INCREASE 2 (GNI2), ancestral to its duplicate GNI1, and represented by the homoeo-allelic series GNI-A2, GNI-B2, and GNI-D2. GNI1 and all GNI2 copies additively affect floral growth and fertility by modulating reproductive allocation and harvest index. Herewith, we describe a series of grain yield-relevant GNI2 homoeo-alleles with a proven track record for being beneficial in both high- and low-input environments. Their deployment offers a sustainable pathway to raise global grain yields in the future.

plant biology↗

Hypermorphic TaARF4 shapes wheat architecture by repressing SPL-mediated developmental timing

Integrating spatial morphogens with temporal developmental clocks is fundamental to optimizing plant architecture and crop yield, yet their molecular interface remains elusive. Here, we characterize Branched shoot 1 (Bsh1), a semi-dominant wheat mutant exhibiting non-canonical upper-aerial branching and aberrant spike development, caused by a T265I substitution in the DNA-binding domain of the auxin response factor TaARF4-A2. Unlike flanking mutations governing protein stability in basal land plants, this central substitution uniquely converts TaARF4-A2 into an auxin-insensitive hypermorphic repressor. The mutant protein disrupts auxin-cytokinin homeostasis and markedly enhances the native repression of SQUAMOSA PROMOTER BINDING PROTEIN-LIKE (SPL) transcription factors, the core timers of developmental phase transitions. This spatial and temporal uncoupling unleashes upper axillary buds from dormancy, promoting aerial branching and diverting resources from reproductive development. Our findings demonstrate how a conserved spatial morphogen effector was evolutionarily rewired in polyploid wheat to dictate the temporal SPL clock, orchestrating species-specific shoot architectural innovation.

plant biology↗

Pre-anthesis light signaling of sheathed embryonic barley inflorescences defines floral fate

The pre-anthesis embryonic inflorescence greening (PEIG) is a distinctive developmental feature in the members of the Triticeae such as barley, wheat and rye. In barley (Hordeum vulgare L.), floral survival and fertility are major determinants of grain yield, yet the physiological processes supporting early inflorescence development remain poorly understood. Here, we investigated PEIG, a light-dependent chlorophyll accumulation occurring while immature inflorescences are still enclosed by leaf sheaths, and its role in overall inflorescence development. Using chlorophyll autofluorescence imaging and chlorophyll quantification, we found that PEIG in barley initiates at a surprisingly early developmental stage, when developing spikes are still enclosed by leaf sheaths. PEIG first appears in the central rachis and then progressively spreads to the spikelet primordia and other floral organs. Using a non-destructive dark treatment method designed for preventing light exposure to developing inflorescences, we further demonstrated that the inhibition of PEIG does not affect floral initiation but significantly reduces floral survival and pollen viability, particularly in the tip region of the inflorescence, indicating a critical role of light mediated PEIG in floral fate. Finally, analysis of natural variation for PEIG revealed a strong positive correlation between PEIG and floral survival. Our findings establish PEIG as an underappreciated hidden trait that supports floral viability and reproductive success, with implications to enhance grain yield potential in cereals.

plant biology↗

Pan-genome Analysis Reveals Hidden Diversity and Selection Signatures of Auxin Response Factors (ARFs) Associated with Breeding in Barley

Auxin Response Factors (ARFs) play a pivotal role in regulating plant growth and development, yet their evolutionary dynamics and functional divergence remain poorly understood in barley (Hordeum vulgare L.) In this study, we conducted a comprehensive genome-wide analysis of the ARF gene family across a 76-accession barley pan-genome. By integrating gene presence/absence variation (PAV) and copy number variation (CNV), phylogeny, expression profiling, transposable element (TE)-mediated regulation, and selection signatures, we characterized 1,911 ARF-coding genes and their structural, transcriptional, and functional variation at the population level. Phylogenetic analysis revealed lineage-specific expansion and dynamic duplications, particularly within the HvARF13 clade. Co-expression networks and tissue-resolved transcriptomes showed that many HvARFs are preferentially expressed in inflorescence and meristematic tissues. Selective sweep and haplotype analyses identified HvARF3 as a candidate gene under selection during European barley breeding. A favorable haplotype of HvARF3, enriched in European cultivars, was significantly associated with increased grain size and weight, demonstrating the utility of pan-genome-enabled frameworks for accelerating gene-trait association and candidate gene discovery. This study highlights the power of multi-omics integration in decoding gene family complexity and provides valuable insights for functional genomics and trait improvement in cereal crops.

plant biology↗

Scaling law links plant growth variation to grain yield in wheat stands

Growth rate, a fundamental biological trait influencing plant resource use, scales predictably with plant mass, following remarkably consistent allometric power laws shaped by biophysical constraints and natural selection. However, how these laws apply to crop plants shaped by artificial selection, and how they manifest in agronomic traits, remains undefined. Under controlled greenhouse conditions, we quantified the relationship between plant mass and growth rate in 195 European winter wheat cultivars. We uncovered genetic variation in allometry linked to plant size, where increased leaf allocation and faster development elevated allometric exponents. Phenotypic and genetic analyses revealed adaptive strategies, ranging from large, slow-growing genotypes that support reproductive initiation to small, fast-growing genotypes that enhance reproductive effort. A shared genetic basis--associated with Photoperiod response-1 (Ppd-1)--linked growth allometry in the greenhouse to genotype-by-environment interactions for grain yield in field trials. This variation in growth allometry, shaped by breeding in diverse environments, reflects strategies that enhance adaptation to weather conditions. Our findings demonstrate that growth allometry is biologically robust and agronomically important because it scales to wheat yield under diverse, realistic field conditions.

plant biology↗

Imputation integrates single-cell and spatial gene expression data to resolve transcriptional networks in barley shoot meristem development

Grass inflorescences are composite structures, featuring complex sets of meristems as stem cell niches that are initiated in a repetitive manner. Meristems differ in identity and longevity, generate branches or split to form flower meristems that finally produce seeds. Within meristems, distinct cell types are determined by positional information and the regional activity of gene regulatory networks. Understanding these local microenvironments requires precise spatio-temporal information on gene expression profiles, which current technology cannot achieve. Here we investigate transcriptional changes during barley development, from the specification of meristem and organ founder cells to the initiation of distinct floral organs, based on an imputation approach integrating deep single-cell RNA sequencing with spatial gene expression data. The expression profiles of more than 40.000 genes can be analysed at cellular resolution in multiple barley tissues using the web-based graphical interface BARVISTA, which enables precise virtual microdissection to analyse any sub-ensemble of cells. Our study pinpoints previously inaccessible key transcriptional events in founder cells during primordia initiation and specification, characterises complex branching mutant phenotypes by barcoding gene expression profiles, and defines spatio-temporal trajectories during flower development. We thus uncover the genetic basis of complex developmental processes, providing novel opportunities for precisely targeted manipulation of barley traits.

plant biology↗

CLAVATA signalling shapes barley inflorescence architecture by controlling activity and determinacy of shoot apical and rachilla meristems

Grasses exhibit a large variety of diverse inflorescence architectures, from complex branched inflorescences in Oryzeae (rice) to simple spike-type inflorescences in Triticeae (e.g. barley, wheat). Inflorescence architecture depends on shape, longevity and determinacy of meristems that direct growth of the main rachis and lateral branches, but how individual meristem activities are determined and integrated within complex inflorescences is not yet understood. We found that activity of distinct meristems in the barley inflorescence is coordinated by a signalling pathway comprising the receptor like kinase Hordeum vulgare CLAVATA1 (HvCLV1) and the secreted CLAVATA3/ENDOSPERM SURROUNDING REGION (CLE)-family peptide FON2- LIKE CLE PROTEIN1 (HvFCP1). HvFCP1 interacts with HvCLV1 to promote spikelet formation but restricts inflorescence meristem and rachilla meristem proliferation. Hvfcp1 or Hvclv1 mutants generate branched inflorescences with additional rows of spikelets and supernumerary florets. Transcriptome analysis reveals that HvFCP1/HvCLV1 signalling controls inflorescence branching through the regulation of trehalose-6-phosphate synthesis and sugar transport. Our discoveries reveal the potential to engineer barley inflorescence architecture by manipulating regulation of distinct meristem activities.

plant biology↗

Anatomical insights into the vascular lay-out of the barley rachis: implications for transport and spikelet connection

Background and aimsVascular patterning is intimately related to plant form and function. However, morphologic a l studies on the vascular anatomy of cereal crops, and inflorescences in particular, are scarce despite their importance for grain yield determination. Here, using barley (Hordeum vulgare) as a model, we study the vascular anatomy of the spike-type inflorescence. Our goal is to clarify the relationship between rachis (spike axis) vasculature and spike size, the implications for transport capacity and its interaction with the spikelets. MethodsWe employed serial transversal internode sections in multiple barley lines with different spike size, and investigated the internode diameter, vascular area and vein number size along the mature barley rachis. We then modeled the vascular dynamics along the main spike axis, and analyzed their relationship with spike size. Key resultsInternode diameter and total vascular area have a clear positive correlation with spike size whereas vascular number is only weakly correlated. While the lateral periphery of the rachis contains large mature veins of constant diameter the central part is occupied by a staggered array of small immature veins. This underlines the importance of minimizing transport resistance and suggests that transport and distribution of nutrients are spatially separated. Spikelet-derived veins enter the rachis either in the central area, where they often merge with the immature rachis veins, or in the periphery where they do not merge with the large mature veins. An increase in floret fertility through the conversion of a two-rowed barley into an isogenic six-rowed line, as well a decrease in floret fertility due to enhanced pre-anthesis tip degeneration caused by the mutation tip sterile 2.b (tst2.b) significantly affected vein size, but had limited to no effects on vein number or rachis diameter. Comparative analysis of a wild barley accession suggests that the domestication of barley may have favored plants with enhanced rachis transport capacity. ConclusionsThe rachis vasculature is the result of a two-step process involving an initial lay-out followed by size adjustment according to floret fertility/spike size. The functional processes of long distance transport and local supply to spikelets are spatially separated while a vascular continuity between rachis and spikelets appears non-essential.

plant biology↗

Dynamic phytomeric growth contributes to local adaptation in barley

Vascular plants segment their body axis with iterative nodes of lateral branches and internodes. Appropriate node initiation and internode elongation are fundamental to plant fitness and crop yield formation; but how they are spatiotemporally coordinated remains elusive. We show that in barley (Hordeum vulgare L.), selections under domestication have extended the apical meristematic phase to promote node initiation, but constrained subsequent internode elongation. In both vegetative and reproductive axes, internode elongation displays a dynamic proximal - distal gradient, and among subpopulations of domesticated barleys at the global range, node initiation and proximal internode elongation are associated with latitudinal and longitudinal gradients, respectively. Genetic and functional analysis suggest that, in addition to their converging roles in node initiation, flowering time genes are repurposed to specify the dynamic internode elongation. Our study provides an integrated view of barley node initiation and internode elongation, and suggests that plant architecture has to be recognized as dynamic phytomeric units in the context of crop evolution.

plant biology↗

Balancing grain yield trade-offs in 'Miracle-Wheat'

Introducing variations in inflorescence architecture, such as the Miracle-Wheat (Triticum turgidum convar. compositum (L.f.) Filat.) with a branching spike, has relevance for enhancing wheat grain yield. However, in the spike-branching genotypes, the increase in spikelet number is generally not translated into grain yield advantage because of reduced grains per spikelet and grain weight. Here, we investigated if such trade-offs might be a function of source-sink strength by using 385 RILs developed by intercrossing the spike-branching landrace TRI 984 and CIRNO C2008, an elite durum (T. durum L.) cultivar; they were genotyped using the 25K array. Various plant and spike architectural traits, including flag leaf, peduncle and spike senescence rate, were phenotyped under field conditions for two consecutive years. On Chr 5AL, we found a new modifier QTL for spike-branching, branched headt 3 (bht-A3), which was epistatic to the previously known bht-A1 locus. Besides, bht-A3 was associated with more grains per spikelet and a delay in flag leaf senescence rate. Importantly, favourable alleles viz., bht-A3 and grain protein content (gpc-B1) that delayed senescence are required to improve grain number and grain weight in the spike-branching RILs. In summary, achieving a balanced source-sink relationship might minimise grain yield trade-offs in Miracle-Wheat. HIGHLIGHTGenetic interplay between sink number and post-anthesis source activity limits grain yield in the spike-branching Miracle-Wheat.

plant biology↗

Dosage of duplicated and antifunctionalized homeobox proteins influences spikelet development in barley

Illuminating the mechanisms of inflorescence architecture of grain crops that feed our world may strengthen the goal towards sustainable agriculture. Lateral spikelet development of barley (Hordeum vulgare L.) is such an example of a floral architectural trait regulated by VRS1 (Vulgare Row-type Spike 1 or Six-rowed Spike 1, syn. HvHOX1). Its lateral spikelet-specific expression and the quantitative nature of suppressing spikelet development were previously shown in barley. However, the mechanistic function of this gene and its paralog HvHOX2 on spikelet development is still fragmentary.Here, we show that these duplicated transcription factors (TFs) have contrasting nucleotide diversity in various barley genotypes and several Hordeum species. Despite this difference, both proteins retain their basic properties of the homeodomain leucine zipper class I family of TFs. During spikelet development, these genes exhibit similar spatiotemporal expression patterns yet with anticyclic expression levels. A gene co-expression network analysis suggested that both have an ancestral relationship but their functions appear antagonistic to each other, i.e., HvHOX1 suppresses whereas HvHOX2 rather promotes spikelet development. Our transgenic promoter-swap analysis showed that HvHOX2 can restore suppressed lateral spikelets when expression levels are increased; however, at its low endogenous expression level, HvHOX2 appears dispensable for spikelet development. Collectively, this study proposes that the dosage of the two antagonistic TFs, HvHOX1 and HvHOX2, influence spikelet development in barley.

plant biology↗

The barley mutant multiflorus2.b reveals quantitative genetic variation for new spikelet architecture

Understanding the genetic basis of yield forming factors in small grain cereals is of extreme importance, especially in the wake of stagnation of further yield gains in these crops. One such yield forming factor in these cereals is the number of grain-bearing florets produced per spikelet. Wildtype barley (Hordeum vulgare L.) spikelets are determinate structures, the spikelet axis (rachilla) degenerates after producing single floret. In contrast, the rachilla of wheat (Triticum ssp.) spikelets, which are indeterminate, elongates to produce up to 12 florets. In our study, we characterized the barley spikelet determinacy mutant multiflorus2.b (mul2.b) that produced up to three fertile florets on elongated rachillae of lateral spikelets. Apart from the lateral spikelet indeterminacy (LS-IN), we also characterized the supernumerary spikelet phenotype in the central spikelets (CS-SS) of mul2.b. Through our phenotypic and genetic analyses, we identified two major QTLs on chromosomes 2H and 6H, and two minor QTLs on 3H for the LS-IN phenotype. For, the CS-SS phenotype we identified one major QTL on 6H, and a minor QTL on 5H chromosomes. Notably, the 6H QTLs for CS-SS and LS-IN phenotypes co-located with each other, potentially indicating that a single genetic factor might regulate both phenotypes. Thus, our in-depth phenotyping combined with genetic analyses revealed the quantitative nature of the LS-IN and CS-SS phenotypes in mul2.b, paving the way for cloning the genes underlying these QTLs in the future. Key messageSpikelet indeterminacy and supernumerary spikelet phenotypes in barley multiflorus2.b mutant show polygenic inheritance. Genetic analysis of multiflorus2.b revealed major QTLs for spikelet determinacy and supernumerary spikelet phenotypes on 2H and 6H chromosomes.

plant biology↗

A development guide for evaluating the maximum yield potential stage in barley

Determining the grain yield potential contributed by grain number is a step towards advancing cereal crops yield. To achieve this aim, it is pivotal to recognize the maximum yield potential (MYP) of the crop. In barley (Hordeum vulgare L.), the MYP is defined as the maximum spikelet primordia number of a spike. Previous barley studies often assumed the awn primordium (AP) stage as the MYP stage regardless of genotypes and growth conditions. From our spikelet-tracking experiments using the two-rowed cultivar Bowman, we found that the MYP stage can be different from the AP stage. Importantly, we find that the occurrence of inflorescence meristem (IM) deformation and its loss of activity coincided with the MYP stage, indicating the end of further spikelet initiation. Thus, we recommend validating the barley MYP stage with the IMs shape and propose this approach (named Spikelet Stop) for MYP staging. Following this approach, we compared the MYP stage and the MYP in 27 two- and six-rowed barley accessions grown in the greenhouse and field. Our results reveal that the MYP stage can be reached at various developmental stages, which majorly depend on the genotype and growth conditions. Furthermore, we found that two-rowed barleys MYP and the duration reaching the MYP stage may determine their yield potential. Based on our findings, we suggest key steps for the identification of the MYP in barley that can also be applied in a related crop such as wheat. HighlightWe show that the maximum yield potential stage in barley can be different from the awn primordium stage as proposed in earlier studies and it varies depending on the genotype and growth conditions. We suggest key steps to identify maximum yield potential in barley that might apply to related cereals.

plant biology↗

Modes of grain number determination differentiate barley row-types

Gaining knowledge on intrinsic interactions of various yield components is crucial to improve the yield potential in small grain cereals. It is well known in barley that increasing the grain number (GN) preponderantly improves their yield potential; however, the yield components determining GN and their association in barley row-types are less explored. In this study, we assessed different yield components like potential spikelet number (PSN), spikelet survival (SSL), spikelet number (SN), grain set (GS), and grain survival (GSL), as well as their interactions with GN by using a selected panel of two- and six-rowed barley types. Also, to analyze the stability of these interactions, we performed the study in two growth conditions, greenhouse and field. From this study, we found that in two-rowed, GN determination is strongly influenced by PSN rather than SSL and/or GS in both growth conditions. Conversely, in six-rowed, GN is associated with SSL instead of PSN and/or GS. Thus, our study exemplified that increasing GN might be possible by augmenting PSN in two-rowed genotypes, while for six-rowed genotypes, the ability of SSL needs to be improved. We speculate that this disparity of GN determination in barley row-types might be due to the fertility of lateral spikelets. Collectively, this study revealed that the GN of two-rowed largely depends on the developmental trait, PSN, while in six-rowed, it mainly follows the ability of SSL. HighlightIn cereals, understanding the interactions of different yield components that influence the grain number is essential to increase their yield by modulating the components. We show in this study that the grain number of two-rowed barley is predominantly determined by the potential spikelet number while in six-rowed by spikelet survival.

plant biology↗