He was born in Nafplio, the first capital of Greece and his paternal family home. During World War II, he accompanied his parents to Egypt, South Africa, and finally the United States (U.S.), where his father held ministerial posts in the Greek Government in exile. The family stayed in the U.S. until 1944, when, following the liberation of Greece after the defeat of Nazi Germany, they returned permanently to Greece. 
After his secondary education at the 8th Gymnasium of Athens, he graduated from the Medical School of the National and Kapodistrian University of Athens in 1956. After completion of Army service he joined the team of Prof. Peter Karlson at the Institut für Physiologische Chemie of the Ludwig Maximilian University of Munich. He completed his doctoral degree in biochemistry in 1962 from the Medical School of the Ludwig Maximilian University of Munich.