山东省自然科学基金项目2015ZRB01179;"小麦重要品质性状形成和改良的分子基础"育种专项和山东省种质资源创制课题资助. This study was supported by the Natural Science Foundation of Shandong Province, China2015ZRB01179;the breeding project of "Molecular Foundation for Formation and Improvement of Main Quality Traits of Wheat", and the Shandong Provincial Project for Germplasm Resource Innovation