HSF4 and Werner syndrome: Other differentially methylated genes have been associated with genome instability (RECQL5, TERF2, and XRCC3), cataract (EYA1, HSF4, OPA3, PAX6, and SORD), thyroid cancer (NDUFA13 and PAX8), osteoporosis (CALCA and CALCR), diabetes mellitus (AKT2 and TCFL2), and diminished fertility (ESR1), all of which contribute to the WS aging phenotype.