In contrast to other cereals, typical barley cultivars have caryopses with adhering hulls at maturity, known as covered (hulled) barley. However, a few barley cultivars are a free-threshing variant called naked (hulless) barley. The covered/naked caryopsis is controlled by a single locus (nud) on chromosome arm 7HL. On the basis of positional cloning, we concluded that an ethylene response factor (ERF) family transcription factor gene controls the covered/naked caryopsis phenotype. This conclusion was validated by (i) fixation of the 17-kb deletion harboring the ERF gene among all 100 naked cultivars studied; (ii) two x-ray-induced nud alleles with a DNA lesion at a different site, each affecting the putative functional motif; and (iii) gene expression strictly localized to the testa. Available results indicate the monophyletic origin of naked barley. The Nud gene has homology to the Arabidopsis WIN1/SHN1 transcription factor gene, whose deduced function is control of a lipid biosynthesis pathway. Staining with a lipophilic dye (Sudan black B) detected a lipid layer on the pericarp epidermis only in covered barley. We infer that, in covered barley, the contact of the caryopsis surface, overlaid with lipids to the inner side of the hull, generates organ adhesion.caryopsis ͉ domestication ͉ epidermis ͉ ethylene response factor ͉ grass B arley (Hordeum vulgare L.) is the world's fourth most important cereal crop behind wheat, rice, and maize. A particular botanical feature of domesticated barley is that most cultivars have covered (hulled) caryopses in which the hull (outer lemma and inner palea) is firmly adherent to the pericarp epidermis at maturity; but a few cultivars are of a free-threshing variant called naked (hulless) barley (Fig. 1). No other Poaceae (grass) family crops show such hull-caryopsis adhesion. Both caryopsis types of barley have agronomic value and are used for different purposes. Covered barley is mainly used as an animal feed and for brewing. The hull of covered barley protects embryos from damage during mechanical harvest, and it also provides a filtration medium in separation of fermentable extract (wort) during malt processing (1). In contrast, naked barley is preferred for human food, because extensive pearling to remove the hull is unnecessary. Now that healthy effects of the soluble-fiber-rich barley products have been officially approved (2, 3), consumers' current interest in nutrition might boost the status of barley as human food.Easy processing of edible part can be a primary character of selection during domestication of a food crop (4, 5). Consequently, the naked caryopsis is considered a key domestication character in barley (5-8). The wild progenitor of barley, H. vulgare subsp. spontaneum, has covered grains. The covered grain is therefore considered adaptive in the wild: the hulls protect the caryopses from various biotic and abiotic stresses, and the awn attached to the distal end of the lemma facilitates seed dispersal and burial (9). According to archeological evidence (4)...