Abstract

BackgroundBread wheat (Triticum aestivum) has a large, complex and hexaploid genome consisting of A, B and D homoeologous chromosome sets. Therefore each wheat gene potentially exists as a trio of A, B and D homoeoloci, each of which may contribute differentially to wheat phenotypes. We describe a novel approach combining wheat cytogenetic resources (chromosome substitution ‘nullisomic-tetrasomic’ lines) with next generation deep sequencing of gene transcripts (RNA-Seq), to directly and accurately identify homoeologue-specific single nucleotide variants and quantify the relative contribution of individual homoeoloci to gene expression.ResultsWe discover, based on a sample comprising ~5-10% of the total wheat gene content, that at least 45% of wheat genes are expressed from all three distinct homoeoloci. Most of these genes show strikingly biased expression patterns in which expression is dominated by a single homoeolocus. The remaining ~55% of wheat genes are expressed from either one or two homoeoloci only, through a combination of extensive transcriptional silencing and homoeolocus loss.ConclusionsWe conclude that wheat is tending towards functional diploidy, through a variety of mechanisms causing single homoeoloci to become the predominant source of gene transcripts. This discovery has profound consequences for wheat breeding and our understanding of wheat evolution.

Highlights

  • Bread wheat (Triticum aestivum) has a large, complex and hexaploid genome consisting of A, B and D homoeologous chromosome sets

  • The nullitetras have historically been used for determining presence of wheat genes on specific homoeologous chromosomes [59,60,61], and we reasoned that, in combination with generation sequencing, they could be used to develop a systematic understanding of the relative contributions of individual homoeoloci to overall wheat gene expression

  • Homoeolocus-specific sequence analysis shows at least 45% of wheat genes are expressed from all three homoeoloci We developed a novel strategy for the direct detection of wheat sequence variants and their assignment to homoeologous chromosomes using nullitetra analysis

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Summary

Introduction

Bread wheat (Triticum aestivum) has a large, complex and hexaploid genome consisting of A, B and D homoeologous chromosome sets. Between 3080% of species are currently polyploids [4], while the rest exist as paleopolyploids [5,6], having undergone a gradual process of “diploidization,” or reversion to a diploid state over evolutionary time. This process involves extensive genomic rearrangements, including the physical loss of a large fraction of duplicate regions, and the accumulation of mutations that distinguish the sequences of the duplicate ‘homoeologous’ copies and contribute to their functional divergence [7,8,9]. Allopolyploid plants often undergo major including Arabidopsis [18], maize [19], Brassica [20] and soybean [21,22]

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