He was one of the first to identify enhancer elements that regulate transcription at a distance, the first to show that the retinoblastoma tumor suppressor regulates transcription and the first to develop gene therapies for autoimmune disease including an ongoing clinical trial for osteoarthritis. More recently, he was part of a collaborative team that was the first to identify senotherapeutic compounds, able to reduce the senescent cell burden and extend healthspan and lifespan in mouse models, that are in more than 15 clinical trials for age related diseases and conditions.