William Tinsley Keeton (February 3, 1933 – August 17, 1980) was an American
zoologist
Zoology ()The pronunciation of zoology as is usually regarded as nonstandard, though it is not uncommon. is the branch of biology that studies the animal kingdom, including the structure, embryology, evolution, classification, habits, and d ...
known internationally for his work on
animal behavior
Ethology is the scientific study of animal behaviour, usually with a focus on behaviour under natural conditions, and viewing behaviour as an evolutionarily adaptive trait. Behaviourism as a term also describes the scientific and objectiv ...
, especially
bird migration,
and for his work on
millipede
Millipedes are a group of arthropods that are characterised by having two pairs of jointed legs on most body segments; they are known scientifically as the class Diplopoda, the name derived from this feature. Each double-legged segment is a resu ...
taxonomy
Taxonomy is the practice and science of categorization or classification.
A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
. He was a well-liked professor of biology at
Cornell University
Cornell University is a private statutory land-grant research university based in Ithaca, New York. It is a member of the Ivy League. Founded in 1865 by Ezra Cornell and Andrew Dickson White, Cornell was founded with the intention to tea ...
in
Ithaca, New York
Ithaca is a city in the Finger Lakes region of New York, United States. Situated on the southern shore of Cayuga Lake, Ithaca is the seat of Tompkins County and the largest community in the Ithaca metropolitan statistical area. It is named ...
and author of a widely used introductory textbook, ''Biological Science''.
Biography
William Keeton was born February 3, 1933, in
Roanoke, Virginia
Roanoke ( ) is an independent city in the U.S. state of Virginia. At the 2020 census, the population was 100,011, making it the 8th most populous city in the Commonwealth of Virginia and the largest city in Virginia west of Richmond. It is ...
, and grew up in
Lynchburg. Keeton attended the
University of Chicago
The University of Chicago (UChicago, Chicago, U of C, or UChi) is a private university, private research university in Chicago, Illinois. Its main campus is located in Chicago's Hyde Park, Chicago, Hyde Park neighborhood. The University of Chic ...
and received both his Bachelor of Arts and the Bachelor of Science degrees, working under Dr.
Alfred E. Emerson
Alfred Edwards Emerson, Jr. (December 31, 1896 – October 3, 1976) was an American biologist, Professor of Zoology at the University of Chicago, a noted entomologist and leading authority on termites.
Life and work
Emerson was born in Ithaca ...
. Keeton earned a master's degree at
Virginia Polytechnic Institute
Virginia Tech (formally the Virginia Polytechnic Institute and State University and informally VT, or VPI) is a public land-grant research university with its main campus in Blacksburg, Virginia. It also has educational facilities in six re ...
(Virginia Tech), during which he revised the
millipede
Millipedes are a group of arthropods that are characterised by having two pairs of jointed legs on most body segments; they are known scientifically as the class Diplopoda, the name derived from this feature. Each double-legged segment is a resu ...
genus ''
Brachoria''.
[ During his time at Virginia Tech, Keeton met Barbara Orcutt, whom he married in 1958.][ He moved to ]Cornell University
Cornell University is a private statutory land-grant research university based in Ithaca, New York. It is a member of the Ivy League. Founded in 1865 by Ezra Cornell and Andrew Dickson White, Cornell was founded with the intention to tea ...
in 1956 in order to continue his research with millipede systematics for his doctorate, where he studied under Dr. Howard E. Evans. His doctoral research culminated in a monograph on the family Spirobolidae
Spirobolidae is a family of millipedes in the order Spirobolida. The family consists of several genera with numerous species, and is commonly divided into the subfamilies Spirobolinae and Tylobolinae.
Classification
Subfamily Spirobolinae
* ' ...
. He received his doctorate in 1958 and joined the biology faculty at Cornell University as a biology professor in 1958.
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
Columbidae () is a bird family consisting of doves and pigeons. It is the only family in the order Columbiformes. These are stout-bodied birds with short necks and short slender bills that in some species feature fleshy ceres. They primarily ...
and bird orientation and navigation
Navigation is a field of study that focuses on the process of monitoring and controlling the movement of a craft or vehicle from one place to another.Bowditch, 2003:799. The field of navigation includes four general categories: land navigation, ...
, as he studied pigeon homing behaviors for well over a decade.
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
Earth's magnetic field, also known as the geomagnetic field, is the magnetic field that extends from Earth's interior out into space, where it interacts with the solar wind, a stream of charged particles emanating from the Sun. The magnetic ...
, 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
Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytologist is a scientist who specialises in this field. The term "botany" comes from the Ancient Greek w ...
and zoology
Zoology ()The pronunciation of zoology as is usually regarded as nonstandard, though it is not uncommon. is the branch of biology that studies the animal kingdom, including the structure, embryology, evolution, classification, habits, and ...
. 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.[
Keeton died from ]heart attack
A myocardial infarction (MI), commonly known as a heart attack, occurs when blood flow decreases or stops to the coronary artery of the heart, causing damage to the heart muscle. The most common symptom is chest pain or discomfort which ma ...
on August 17, 1980, due to a failure of a mechanical heart valve. He was 47 years old.
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 held speculative ideas about the techniques that pigeons might be using, including the use of the position of the sun, the earth's magnetic field, landmark recognition and olfactory navigation. William Keeton tested these various speculations throughout his time at Cornell University.
Magnetic 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
Earth's magnetic field, also known as the geomagnetic field, is the magnetic field that extends from Earth's interior out into space, where it interacts with the solar wind, a stream of charged particles emanating from the Sun. The magnetic ...
as one way of finding their way home. In this experiment, William Keeton attached magnets
A magnet is a material or object that produces a magnetic field. This magnetic field is invisible but is responsible for the most notable property of a magnet: a force that pulls on other ferromagnetic materials, such as iron, steel, nickel ...
to the back of pigeons just before they were released and measured their vanishing point
A vanishing point is a point on the image plane of a perspective drawing where the two-dimensional perspective projections of mutually parallel lines in three-dimensional space appear to converge. When the set of parallel lines is perpendicul ...
(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 on an olfactory map, but the experiment was too general for proving that this was indeed what was occurring in this experiment.
Bibliography of orientation publications
Books and chapters
*Keeton, William. (1972). Effects of magnets on pigeon homing. pp. 579–594 in Animal Orientation and Navigation. NASA SP-262. Washington, D.C.
*Keeton, William. (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.
*Keeton, William. (1977). Magnetic reception (biology). In Encyclopedia of science and technology, 2nd Ed. New York, McGraw-Hill.
*Keeton, William. (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.
Work with millipedes
Prior to his work on animal navigation, Keeton studied the systematics and taxonomy
Taxonomy is the practice and science of categorization or classification.
A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
of millipede
Millipedes are a group of arthropods that are characterised by having two pairs of jointed legs on most body segments; they are known scientifically as the class Diplopoda, the name derived from this feature. Each double-legged segment is a resu ...
s. His Master's thesis at Virginia Tech was a revision of the genus '' Brachoria'', a Xystodesmid of the order Polydesmida
Polydesmida (from the Greek ''poly'' "many" and ''desmos'' "bond") is the largest order of millipedes, containing approximately 3,500 species, including all the millipedes reported to produce hydrogen cyanide (HCN).
Description
Members of the o ...
. His doctoral research at Cornell resulted in a monograph of the family Spirobolidae
Spirobolidae is a family of millipedes in the order Spirobolida. The family consists of several genera with numerous species, and is commonly divided into the subfamilies Spirobolinae and Tylobolinae.
Classification
Subfamily Spirobolinae
* ' ...
(order Spirobolida) published in 1960, that garnered praise as bringing order and clarity to "a chaos of unrelated genera replete with poorly-known species".[ He split the family into two subfamilies and reduced the number of species through synonymy - determining that various named species actually belonged to previously described species. He made many field excursions to the ]Appalachian Mountains
The Appalachian Mountains, often called the Appalachians, (french: Appalaches), are a system of mountains in eastern to northeastern North America. The Appalachians first formed roughly 480 million years ago during the Ordovician Period. They ...
with fellow millipede expert Richard L. Hoffman
Richard Lawrence Hoffman (September 25, 1927 – June 10, 2012) was an American zoologist known as an international expert on millipedes, and a leading authority on the natural history of Virginia and the Appalachian Mountains. He was a biology ...
, and also made an extended collecting trip to the Vulcan San Martin of Veracruz, Mexico. He published a total of 13 works on millipedes, in which he named 19 new species, two new genera, and the new families Allopocockiidae and Floridobolidae
''Floridobolus'' is a genus of millipedes commonly known as Florida scrub millipedes containing three described species: ''Floridobolus penneri'', ''F. orini'', and ''F. floydi''; the latter two described in 2014. All three species are endemic t ...
, both of the order Spirobolida.[ He also studied ]development
Development or developing may refer to:
Arts
*Development hell, when a project is stuck in development
*Filmmaking, development phase, including finance and budgeting
*Development (music), the process thematic material is reshaped
* Photograph ...
and morphogenesis
Morphogenesis (from the Greek ''morphê'' shape and ''genesis'' creation, literally "the generation of form") is the biological process that causes a cell, tissue or organism to develop its shape. It is one of three fundamental aspects of deve ...
, and worked with Dr. Thomas Eisner
Thomas Eisner (June 25, 1929 – March 25, 2011) was a German-American entomologist and ecologist, known as the "father of chemical ecology." He was a Jacob Gould Schurman Professor of Chemical Ecology at Cornell University, and Director of the ...
on characterizing the defensive secretions of six species of the order Spirostreptida.
Awards and honors
Keeton's work was recognized by many honors and distinguished positions, including:[
* ]Fellow
A fellow is a concept whose exact meaning depends on context.
In learned or professional societies, it refers to a privileged member who is specially elected in recognition of their work and achievements.
Within the context of higher education ...
of the American Ornithologists' Union
* Liberty Hyde Bailey chair professorship
* Chairman of the Department of Neurobiology and Behavior at Cornell
* Member of the Cornell University Board of Trustees
* Honorary Doctor of Science degree from Coe College
Coe College is a private liberal arts college in Cedar Rapids, Iowa. It was founded in 1851 and is historically affiliated with the Presbyterian Church (USA). The college is a member of the Associated Colleges of the Midwest and the Associatio ...
, Iowa
* Visiting professorship at the Max Planck Institute for Behavioral Physiology
The former Max Planck Institute for Behavioral Physiology was located in Bulldern, Westphalia, Germany, moved to Seewiesen in 1957. It was one of 80 institutes in the Max Planck Society (Max Planck Gesellschaft).
Background
A working group was f ...
, Germany
* Visiting professorship at the University of Konstanz
The University of Konstanz (german: Universität Konstanz) is a university in the city of Konstanz in Baden-Württemberg, Germany. Its main campus was opened on the Gießberg in 1972 after being founded in 1966. The university is Germany's ...
, Germany
* Plenary speaker at the XVII International Ornithological Congress in Berlin (1978)
* "Professor of Merit" award from the Cornell graduating class of 1966
Legacy
Keeton is the namesake of William Keeton House, a residential house of Cornell University that opened in 2008. He is also the namesake of the Keeton Prize, established in 1991 and awarded by faculty to outstanding Cornell undergraduate students.
References
External links
Keeton's 1967 preface to ''Biological Science''
{{DEFAULTSORT:Keeton, William Tinsley
1933 births
1980 deaths
20th-century American zoologists
American ornithologists
Cornell University alumni
Cornell University faculty
Ethologists
Myriapodologists
Writers from Roanoke, Virginia
University of Chicago alumni
Virginia Tech alumni
Scientists from Virginia