As published in the Boston Globe, Thursday, December 27, 2007.

John R. Pappenheimer, 92; taught physiology at Harvard

By Bryan Marquard, Globe Staff  |  December 27, 2007

No part of the body was too tiny or too prominent to escape the attention of John R. Pappenheimer.

At some point, students who pore through a physiology textbook will review Dr. Pappenheimer's explanation of what can pass through the walls of capillaries, the smallest blood vessels. And they might study factor S, a peptide he helped identify that plays a key neurological role in sleep.

"His understanding of the whole body was something that allowed him to do the kinds of experiments that other people mostly had not perceived of," said Woody Hastings, a biology professor at Harvard University. "He was a true biologist, not just a specialist."

Dr. Pappenheimer, the last of three intellectually gifted siblings who taught at Harvard and Harvard Medical School, died of respiratory failure on Dec. 9 in Cadbury Commons at Cambridge. He was 92 and until not long ago still rode his bicycle, adding to a lifetime of pedaling he tallied in the tens of thousands of miles.

"Apart from his work, he had a passion for music, and quartets in particular," said his daughter, Rosamond Zimmerman of Lexington. "He played the cello and was playing right up to the very end. In fact, even when he was in rehab, he had a date with these two Longy [School of Music] students to play some Schubert."

Since childhood, Dr. Pappenheimer had pushed himself to excel intellectually, musically, and physically, often at the behest of his father, a pathology professor at Columbia University who introduced his children to a life of discipline and competitiveness. As a boy, Dr. Pappenheimer began accompanying his father to lectures and to the lab at Columbia.

Dr. Pappenheimer followed his older brother, Alwin, and sister, Anne, from the suburbs of New York City to Cambridge, where all three attended Harvard or Radcliffe. Graduating in 1936, he headed to Cambridge University in England and received a doctorate from Clare College.

Part of his work included studying physiology full time for a year, he said decades later in a speech to the American Physiological Society, which he served as president in the 1960s. "The course consisted of one lecture each day followed by reading, reading, and more reading . . . I have been eternally grateful for this period of intensive study," he said. "It provided a framework for subsequent research and teaching, and made possible the enjoyment of continued reading in fields outside one's own narrow interests."

Returning to the United States in 1940, Dr. Pappenheimer taught at Columbia's College of Physicians and Surgeons, then devoted three years to research at the University of Pennsylvania.

He went back to Harvard in 1946 and seven years later the American Heart Association designated him a career investigator. That meant the organization covered his salary, his daughter said, freeing him to teach and conduct research wherever he wanted. He stayed at Harvard and in 1969 was appointed the George Higginson professor of physiology at Harvard Medical School.

The decision to accept heart association's award prompted a discussion with his father, who "advised him not to take it because it would pollute his own desire to do experiments on his own," said Dr. Pappenheimer's son Will, of Brooklyn, N.Y.

"My father was really from what you would call from a 19th century ethic, of purity of science and purity of motive, although he was definitely a worldly man and definitely competitive," his son said. "He was a very competitive man and therefore good at a variety of things - everything from science to playing tennis. He was a pretty ruthless tennis player."

Dr. Pappenheimer kept one foot in times past through other actions, too. He preferred to get around on a bicycle and didn't like making long-distance phone calls.

"He was parsimonious in quite an English way in that he wanted to do things with paperclips and rubber bands," said Ann Karnovsky, a longtime friend whose late husband, Manfred, collaborated on research with Dr. Pappenheimer. "He wanted to do things as if it were 100 years ago, or even 50 years ago. Those were the times he admired."

Every Christmas Day, she said, he would ride his bicycle to the Karnovskys' house in Cambridge - even if it were snowing - to drop off a loaf of bread he had just baked, and she would send him home with plum pudding.

In the late 1940s, Dr. Pappenheimer was in Western Massachusetts at the Greenwood Music Camp in Cummington when he met Helena Palmer, who is known as Hylie. A talented violinist, she demurred when he suggested they would be a good couple.

"He asked her to marry him and she said, 'I have to think about it,' and went to France for a year," their daughter said.

They married 58 years ago.

After retiring in 1982, Dr. Pappenheimer was given lab space in the Concord Field Station, a Harvard research facility in Bedford, where in recent years he had studied the absorption of sugars and amino acids in the intestine.

And, as always, he continued to create equipment for the lab and home from commonplace objects.

"He was a jury-rigger," his son said. "In the lab that's how he made things. I even saved one of his early valves . . . It's a regulator for oxygen and it's made out of two funnels. He told me that on the inside there's nylon stockings from a women's store. It's a beautiful little object, so I kept it."

In addition to his wife, his son, and daughter, Dr. Pappenheimer leaves another son, Rick Plant of Melbourne, Australia; and five grandchildren.

A service will be announced.