The work was conducted under clean-room conditions, with extensive decontamination of the hair samples before analysis to reduce the risk of contamination from researchers or the environment. The age of the material, the way it had been stored, and the inherent limitations of working with hair rather than harder biological tissues all constrained what could be concluded. The researchers were explicit throughout about the difference between what the data confirmed and what remained uncertain.

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## What the Genome Revealed: Health and Its Limits

The genome that Begg’s team reconstructed from Beethoven’s authenticated hair allowed them to ask several questions that had been debated by historians, physicians, and biographers for nearly two centuries.

The most pressing was the cause of his death.

Beethoven died on March 26, 1827, at the age of 56. In the months before his death he had been gravely ill. His abdomen had swollen with fluid—a condition called ascites—that his doctors drained multiple times through surgical punctures. His skin had taken on the yellow tinge associated with jaundice. He was in considerable pain, and he spent his final weeks receiving visitors while confined to his bed, reportedly still receiving letters and scores until near the end.

The cause of his terminal illness had been debated for almost as long as it had been over. Cirrhosis of the liver had long been the leading diagnosis, consistent with the autopsy findings that were recorded at the time and with the accumulation of abdominal fluid. But the mechanism—why his liver had failed—was less certain. Heavy alcohol consumption was widely attributed to him; one close friend is reported to have said that Beethoven drank at least a liter of wine with lunch each day in his later years, and other accounts supported the picture of a man who drank heavily and consistently throughout his adult life. But the relationship between alcohol and liver disease involves both quantity of consumption and individual genetic susceptibility, and those vary considerably from person to person.

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