Because protruding eyes are well-established comorbidity commonly associated with Graves’ disease, an autoimmune disease leading to hyperthyroidism31, and since GLIS1 and GLIS3 family members have been implicated in several thyroid gland-associated diseases25,27,32,33, we examined whether this Glis1-KO phenotype was related to the development of Graves’ disease that is characterized by high circulating levels of T3/T4 and low TSH. Here, GLIS3 is linked to Graves disease.