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William Tinsley Keeton
William Tinsley Keeton (February 3, 1933 – August 17, 1980) was a Biology Professor at Cornell University in Ithaca, New York as well as an author, an Animal Behaviorist, an Ethologist, and a neuroethologist. Keeton was a noted and well-known Biological Science 101 professor beginning in 1958, so much so that his popularity as a professor earned his class the nickname of the “Keeton course”. In addition to his teaching Keeton is known for his work with pigeons and bird orientation and navigation, as he studied pigeon homing behaviors for well over a decade.

Before his work with pigeons, William Keeton studied millipede systematics and was a member of the Department of Entomology. During his early work at Cornell University the Biological Science Departments were reorganized, and as a result William Keeton moved from the Entomology department to the newly created Neurobiology and Behavior department. It was here where he first started his research on pigeon homing, which led him to the discoveries of the effects of the Earth’s magnetic field, the position of the sun, as well as olfactory navigation and visual cues involved in the process that pigeons use to find their way home. Cornell University built Keeton a loft large enough to house two thousand pigeons that were the subjects of Keeton’s experiments on the behaviors and processes involved in pigeon homing. Both students and faculty at Cornell University, as well as other scientists from around the world came and worked alongside William Keeton in his pigeon loft.

William Keeton is also known for his work in writing the biology textbook named Biological Science, that was first published by the W.W. Norton & Company in 1967. It took Keeton approximately five years to write the first edition of the textbook. It went through three editions before his death in 1980. After Keeton's death the textbook was revised for editions 4, 5, and 6 by James L. Gould (and Carol Gould). The textbook was a combination of both botany and zoology. This combination of sciences turned out to be extremely successful in teaching many aspects of biology. The textbook was one of the first that integrated zoology and botany and sought common themes, guided by the process of evolution.

Biography
William Tinsley Keeton was born on February 3, 1933.

William Keeton attended the University of Chicago and received both his Bachelor of Arts and the Bachelor of Sciences degrees. Keeton went to the Virginia Polytechnic Institute to pursue his graduate work on entomology and moved to Cornell University in 1956 in order to continue his research with millipede systematics for his doctorate. He received his doctorate in 1958 and joined the biology faculty at Cornell University as a biology professor in 1958.

Work With Pigeons
William Keeton had always had a fascination with pigeon homing techniques from the time he was a child. When he was nine years old he received his first homing pigeons which he raced and trained with his friends. Many scientists speculated ideas about the devices pigeons were using including the use of the position of the sun, the earth’s magnetic field, and landmark recognition and olfactory navigation. William Keeton tested these various speculations throughout his time at Cornell University.

Magnet Interference

In William Keeton’s 1970 Magnets Interfere with Pigeon Homing paper, William Keeton proved that pigeons were affected by changes in the magnetic field surrounding them, and that pigeons were using the earth’s magnetic field as one way of finding their way home. In this experiment, William Keeton attached magnets to the back of pigeons just before they were released and measured their vanishing point (in what direction they had flown out of sight) and the time it took to find their way home for both the experimental birds and with the control birds (control birds had a piece of brass glued to their back of the same weight as the magnet).

Keeton’s results showed that when the sun was visible, the magnets would not usually prevent the birds from finding their way home, but when the sky was overcast the birds with magnets on their back were much more unsuccessful and slower at finding their way back home than the control birds. Previous scientists had shown that pigeons were not relying entirely on the sun to navigate home as many scientists had speculated, when they found that many pigeons were able to navigate successfully under completely overcast skies. This notion led Keeton to question whether the pigeons were using the earth’s magnetic field to orient themselves and navigate home successfully. Keeton’s experiments with magnets showed that there was a combination of processes being used by the pigeons to navigate home, but the use of the earth’s magnetic field was very important for pigeon orientation and navigation.

Under clear skies with the sun visible, both the birds with magnets and the birds without magnets had little trouble navigating back to the loft, yet at unfamiliar locations with overcast skies the birds with magnets glued to their backs were unable to successfully orient themselves and navigate back to the loft. Keeton speculated that this was occurring because the birds without magnets were able to use the earth’s magnetic fields to orient themselves in the correct direction, whereas the birds with magnets attached to their backs were unable to use the sun, familiar landmarks, or the earth’s magnetic fields to find their way home. This discovery was extremely useful in explaining one of the most interesting questions of bird navigation.

Olfactory Interference

Many scientists hypothesized that pigeons were using olfactory information as part of the process in finding their way back to the loft. In Italy, a study by N.E. Baldaccini, in which the bills of pigeons were applied with a strong odor, showed results that the pigeons had less accurate initial orientation. Baldaccini also performed an experiment in which he reared pigeons in a loft in which the wind was deflected by 45 degrees. Baldaccini's results showed that a deflection in the wind while the pigeons were young proved to have an effect on their initial orientation after being released. Keeton replicated this experiment but found that there was a smaller deflection than the results in Baldaccini's experiment in Italy. Keeton speculated that there may be an effect on initial orientation based off of an olfactory map, but the experiment was too general for proving that this was indeed what was occuring in this experiment.

Awards and Honors
- William Keeton House (a residence hall at Cornell University, recently built and named after William Keeton)

- Liberty Hyde Bailey chair professorship

- Member of the Cornell University Board of Trustees

- Doctor of Science degree from Coe College

- Speaker at the XVII International Ornithological Congress in Berlin (1978)

- “Professor of Merit” award from the class of 1966

Bibliography of Orientation Publications
Journal Articles:

William, Keeton. (1969). Orientation by pigeons: is the sun necessary? Science 165: 922-928.

William, Keeton. (1970). Orientation by pigeons: a reply. Science 168: 153.

William, Keeton & Gobeft, A. (1970). Orientation by untrained pigeons requires the sun. Proc. National Academy of Science. 65: 853-856.

William, Keeton. (1970). Comparative orientational and homing performances of single pigeons and small flocks. Auk 87: 797 799.

William, Keeton. (1970). Do pigeons determine latitudinal displacement from the sun's altitude? Nature 227: 626-627.

William, Keeton. (1970). "Distance effect" in pigeon orientation: an evaluation. Biol. Bull. 139: 510-527.

William, Keeton. (1971). Magnets interfere with pigeon homing. Proc. National Academy of Science. 68: 102-106.

William, Keeton. (1971). Unconventional theories of orientation: a panel discussion. Ann. New York Academy of Science. 188: 331-333, 338-340, 348-349, 351, 353-354.

William, Keeton. (1971). The development and evolution of orientation: a panel discussion. Ann. New York Academy of Science 188: 410-402, 405.

William, Keeton & Alexander, J.R. (1972). The effect of directional training on initial orientation in pigeons. Auk 89: 280-298.

William, Keeton. (1973). Release-site bias as a possible guide to the "map" component in pigeon homing. J. Comp. Physiol. 86: 1-16.

William, Keeton & Kreithen, M.L. (1974). Detection of changes in atmospheric pressure by the homing pigeon, Columba livia. J. Comp. Physiol. 89: 73 82.

William, Keeton & Kreithen, M.L. (1974). Detection of polarized light by the homing pigeon, Columba livia. J. Comp. Physiol. 89: 83-92.

William, Keeton & Kreithen, M.L. (1974). Attempts to condition homing pigeons to magnetic stimuli. J. Comp. Physiol. 91: 355-362.

William, Keeton & Alexander, J.R. (1974). Clock-shifting effect on initial orientation of pigeons. Auk 91: 370-374.

William, Keeton & Larkin, T.S. & Windsor, D. M. (1974). Normal fluctuations in the earth's magnetic field influence pigeon orientation. J. Comp. Physiol. 95: 95-103.

William, Keeton. (1974). The mystery of pigeon homing. Science Amer. 231: 96-107.

William, Keeton. (1974). Pigeon homing: no influence of outward-journey on initial orientation. Monitore Zool. Ital. (N.S.) 8: 227-234.

William, Keeton & Brown, A. I. (1976). Homing behavior of pigeons not disturbed by application of an olfactory stimulus. J. Comp. Physiol. 105: 259-266.

William, Keeton, Wiltschko, W. & Wiltschko, R. (1976). Effects of a "permanent" clock-shift on the orientation of young homing pigeons. Behavioral Ecology and Sociobiology 1: 229-243.

William, Keeton & Larkin, T. (1976. Bar magnets mask the effect of normal magnetic disturbances on pigeon orientation. J. Comp. Physiol. 110: 227-232.

William, Keeton. (1977). Biological sensitivity to magnetic fields in orientational responses. In Brain interactions with weak electric and magnetic fields, Neurosci. Research Prog. Bull. 15: 22-27.

William, Keeton & Schmidt-Koenig, K. (1977. Sun compass utilization by pigeons wearing frosted contact lenses. Auk 94: 143-145.

William, Keeton, Yodlowskia, M.L. & Kreithen, M.L. (1977). Detection of atmospheric infrasound by the homing pigeon. Nature 265: 725-726.

William, Keeton & Edrich, W. (1977). A comparison of homing behavior in feral and homing pigeons. Z. Tierpsychol. 44: 389-401.

William, Keeton. (1977). Migratory behavior. In Encyclopedia of science and technology, 2nd Ed. New York, McGraw-Hill.

William, Keeton, Kreithen, M.L. & Hermayer, K.L. (1977). Orientation by pigeons deprived of olfaction by nasal tubes. J. Comp. Physiol. 114: 289-299.

William, Keeton, Waldvogel, J.A., Benvenuti, S. & Papi, F. (1979). Homing pigeon orientation influenced by deflected winds at home loft. J. Comp. Physiol. 128:297-301.

William, Keeton, Papi, F., Brown, A.I. & Benvenuti, S. (1979). Do American and Italian pigeons rely on different homing mechanisms? J. Comp. Physiol. 128: 303-317.

William, Keeton. (1979). Avian orientation and navigation. Ann. Rev. Physiol. 41: 353-366.

William, Keeton. (1979). Avian orientation and navigation: a brief overview. Brit. Birds, 17: 451-470

William, Keeton & Hermayer, K. L. (1979). Homing behavior of pigeons subjected to bilateral olfactory nerve section. Monitore Zool. Ital. (N.S.) 13: 303-313.

William, Keeton & Waldvogel, J. A., Phillips, J.B. & McCorkle, D.R. (1980). Short-term residence in deflector lofts alters initial orientation of homing pigeons. Behav. Ecol. and Sociobiol. 7:207-211.

Books:

William, Keeton. (1972). Effects of magnets on pigeon homing. Pp. 579-594 in Animal Orientation and Navigation. NASA SP-262. Washington, D.C.

William, Keeton. (1974). The orientational and navigational basis of homing in birds. Pp. 47-132 in Advances in the study of behavior, Vol. 5. New York, Academic Press.

William, Keeton. (1977). Magnetic reception (biology). In Encyclopedia of science and technology, 2nd Ed. New York, McGraw-Hill.

William, Keeton. (1979). Pigeon navigation. Pp. 5-20 in Neural mechanisms of behavior in the pigeon.( A.M. Granda and J. H. Maxwell, Eds.). New York, Plenum Publishing Corp.