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Simple Summary  On wheat polyploidisation, the major meiotic gene  ZIP4 , duplicated and diverged, such that tetraploid and hexaploid wheat each carry three and four copies of  ZIP4 , respectively. Surprisingly, this study demonstrates that, in hexaploid wheat, despite the presence of the other three  ZIP4  copies, the duplicated  ZIP4  copy is required to prevent major abnormalities during meiosis. Although there is greater disruption of subsequent male rather than female fertility, the duplicated  ZIP4  copy preserves up to 50% of the grain number. High wheat fertility is important since it is consumed by over 4.5 billion people on the planet, of whom 2.5 billion are dependent on it. This study highlights the potentially extraordinary value of the wheat  ZIP4  duplication, mandating further studies to unravel the complexity of the  ZIP4  phenotype in this global crop.    Abstract  Although most flowering plants are polyploid, little is known of how the meiotic process evolves after polyploidisation to stabilise and preserve fertility. On wheat polyploidisation, the major meiotic gene  ZIP4  on chromosome 3B duplicated onto 5B and diverged ( TaZIP4-B2 ).  TaZIP4-B2  was recently shown to promote homologous pairing, synapsis and crossover, and suppress homoeologous crossover. We therefore suspected that these meiotic stabilising effects could be important for preserving wheat fertility. A CRISPR  Tazip4-B2  mutant was exploited to assess the contribution of the 5B duplicated  ZIP4  copy in maintaining pollen viability and grain setting. Analysis demonstrated abnormalities in 56% of meiocytes in the  Tazip4-B2  mutant, with micronuclei in 50% of tetrads, reduced size in 48% of pollen grains and a near 50% reduction in grain number. Further studies showed that most of the reduced grain number occurred when  Tazip4-B2  mutant plants were pollinated with the less viable  Tazip4-B2  mutant pollen rather than with wild type pollen, suggesting that the stabilising effect of  TaZIP4-B2  on meiosis has a greater consequence in subsequent male, rather than female gametogenesis. These studies reveal the extraordinary value of the wheat chromosome 5B  TaZIP4-B2  duplication to agriculture and human nutrition. Future studies should further investigate the role of  TaZIP4-B2  on female fertility and assess whether different  TaZIP4-B2  alleles exhibit variable effects on meiotic stabilisation and/or resistance to temperature change.

A Duplicated Copy of the Meiotic Gene ZIP4 Preserves up to 50% Pollen Viability and Grain Number in Polyploid Wheat