per molecule. He even declared that it could not store the genetic code because it was chemically far too simple. However, his work was a key basis for the later work that determined the structure of DNA. Levene published over 700 original papers and articles on biochemical structures. Levene died in 1940, before the true significance of DNA became clear.

Levene is known for his tetranucleotide hypothesis which proposed that DNA was made up of equal amounts of adenine, guanine, cytosine,