HKT1;5-like cation transporters linked to Na⁺ exclusion loci in wheat, Nax2 and Kna1

Bread wheat (Triticum aestivum) has a greater ability to exclude Na ⁺ from its leaves and is more salt tolerant than durum wheat (Triticum turgidum L. subsp. durum [Desf.]). A novel durum wheat, Line 149, was found to contain a major gene for Na⁺ exclusion, Nax2, which removes Na⁺ from the xylem in the roots and leads to a high K ⁺-to-Na⁺ ratio in the leaves. Nax2 was mapped to the distal region on chromosome 5AL based on linkage to microsatellite markers. The Nax2 locus on 5AL coincides with the locus for a putative Na⁺ transporter, HKT1;5 (HKT8). The Nax2 region on 5AL is homoeologous to the region on chromosome 4DL containing the major Na⁺ exclusion locus in bread wheat, Kna1. A gene member of the HKT1;5 family colocates to the deletion bin containing Kna1 on chromosome 4DL. This work provides evidence that Nax2 and Kna1 are strongly associated with HKT1;5 genes.