An important determinant of wheat grain quality is the hardness of the grain. The trait is controlled by a major locus, Ha, on the short arm of chromosome 5D. Purified starch granules from soft-grained wheats have associated with them 15-kDa polypeptides called grain softness proteins (GSPs) or "friabilins." Genes that encode one family of closely related GSP polypeptides-GSP-1 genes-were mapped using chromosome substitution lines to the group 5 chromosomes. An F2 population segregating for hard and soft alleles at the Ha locus on a near-isogenic background was used in a single-seed study of the inheritance of grain softness and of GSP-1 alleles. Grain softness versus grain hardness was inherited in a 3:1 ratio. The wheat endosperm is a triploid tissue formed by fusion of two identical polar cells (from the female parent) with a sperm cell. The properties of the endosperm determine the end use of the wheat grain. Grain hardness or endosperm strength is one such important determinant of wheat end use. The mature wheat endosperm can be considered analogous to a polymer matrix (seed storage protein) in which a filler (starch granules) is dispersed (1). In such a structure, both the hardness of individual components and the degree of adhesion between them will determine overall strength (2). There is no detectable difference between hard and soft wheats in the individual hardness of starch granules or storage protein matrix fragments, but the degree of adhesion between starch granules and the surrounding protein matrix is higher in hard wheats (3-5). This higher adhesion results in both greater compressive strength and ductility for the endosperm as a whole (6, 7). Thus, in soft wheats, the main planes of weakness tend to occur at the starch-protein matrix interfaces and result in separation of the starch granules from the matrix with little damage during milling. As a result, starch granules from soft wheats take up less water when dough is formed and the product is more suited to the commercial production of cakes and biscuits than is dough from hard wheat flour [which is preferred for the baking of breads (8)].The publication costs of this article were defrayed in part by page charge payment. This article must therefore be hereby marked "advertisement" in accordance with 18 U.S.C. §1734 solely to indicate this fact.Although there are various techniques for classifying grains as hard or soft, the phenotypic difference is largely controlled by one or two genetic loci (9, 10). The major gene (Ha) is on the short arm of chromosome 5D (11, 12). Morrison et al. (13,14) have shown that the Ha gene is tightly linked or identical to a gene controlling the level of extractable free polar lipids in the grain, Fpl-1. Genetic markers linked to milling energy in barley, which is analogous to grain hardness, were also recently shown to be located on the homologous chromosome 5H (15).The biochemistry that determines the hardness of the grain is not known. The linkage between Ha and Fpl-l suggests that lipid metabolism or...