Arthur Hasler, one of the leaders in the study of freshwater ecology, died on Friday, March 23, 2001, at age 93.

According to John Magnuson of the University of Wisconsin, Madison — where Mr. Hasler was professor emeritus of limnology — the former Utahn's research answered one of the mysteries of nature, how salmon migrate precisely to their home waters to spawn.

The idea for the study happened when Mr. Hasler visited a mountain stream near his Utah hometown and noticed how the smells of native plants seemed to rekindle childhood memories. His research in the late 1940s showed that olfactory imprinting, an ingrained sense of smell, let a salmon journey thousands of miles to spawn in the precise stream where it was hatched.

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Mr. Hasler was born in 1908 in Lehi. He graduated from Brigham Young University, Provo, in 1932, and earned his Ph.D. from the University of Wisconsin at Madison. He was on the Wisconsin faculty for 41 years and made enduring contributions to lake research.

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