Mutation of the wheat homeobox gene Grain Number Increase 1 increases grain number and grain yield but decreases grain protein content

Inflorescence structure affects final grain yield (GY) in wheat (Triticum aestivum L.). Recent breeding efforts have focused on improving grain number per spike, which is positively correlated with GY. Grain Number Increase 1 (GNI-A1) encodes a homeodomain leucine zipper class I (HD-Zip I) transcription factor that controls the number of grains per spike and GY. However, how this increase in grain number affects grain quality, especially grain protein content (GPC) in wheat, remains elusive. Here we investigated within-spikelet variation in GPC using GNI-A1 near-isogenic lines. Yield trials in two seasons and at two sites demonstrated that lines harboring a reduced-function allele, GNI-A1 (105Y), consistently showed improved GY due to a 27