Abstract

Increasing grain yield is an endless challenge for cereal crop breeding. In barley (Hordeum vulgare), grain number is controlled mainly by Six-rowed spike 1 (Vrs1), which encodes a homeodomain leucine zipper class I transcription factor. However, little is known about the genetic basis of grain size. Here, we show that extreme suppression of lateral florets contributes to enlarged grains in deficiens barley. Through a combination of fine-mapping and resequencing of deficiens mutants, we have identified that a single amino acid substitution at a putative phosphorylation site in VRS1 is responsible for the deficiens phenotype. deficiens mutant alleles confer an increase in grain size, a reduction in plant height, and a significant increase in thousand grain weight in contemporary cultivated germplasm. Haplotype analysis revealed that barley carrying the deficiens allele (Vrs1.t1) originated from two-rowed types carrying the Vrs1.b2 allele, predominantly found in germplasm from northern Africa. In situ hybridization of histone H4, a marker for cell cycle or proliferation, showed weaker expression in the lateral spikelets compared with central spikelets in deficiens Transcriptome analysis revealed that a number of histone superfamily genes were up-regulated in the deficiens mutant, suggesting that enhanced cell proliferation in the central spikelet may contribute to larger grains. Our data suggest that grain yield can be improved by suppressing the development of specific organs that are not positively involved in sink/source relationships.

Highlights

  • Increasing grain yield is an endless challenge for cereal crop breeding

  • To determine whether the reduced Vrs1 transcript level was affected by the upstream regulator Vrs4, we examined Vrs4 expression levels but found no significant difference between the wild type and Deficiens 2 (Def2) (Fig. 4B)

  • Considering contemporary NW European winter cultivars, we identified three single-nucleotide polymorphism (SNP) in the coding sequence of Vrs1 leading to amino acid substitutions, D8G, D26E, S184G, and one SNP in the second intron of this gene (Supplemental Table S3)

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Summary

Introduction

Increasing grain yield is an endless challenge for cereal crop breeding. In barley (Hordeum vulgare), grain number is controlled mainly by Six-rowed spike 1 (Vrs1), which encodes a homeodomain leucine zipper class I transcription factor. Row type is controlled by at least five loci: Six-rowed spike 1 (Vrs1 [syn = HvHox1]), Vrs, Vrs, Vrs, and Intermedium spike-c (Int-c [syn = Vrs5]), and all five genes have been identified (Komatsuda et al, 2007; Ramsay et al, 2011; Koppolu et al, 2013; Bull et al, 2017; van Esse et al, 2017; Youssef et al, 2017) In four of these (Vrs, Vrs, Vrs, and Vrs5), wild-type alleles encode transcription factors: homeodomain-leucine zipper class I (HD-Zip I), SHORT INTERNODES, RAMOSA2, and TEOSINTE BRANCHED1, respectively; in Vrs, the wild-type allele putatively encodes a histone H3K9 demethylase. At least 58 lossof-function mutants have been described; it seems to be a driving force of genetic variation in barley

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