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Comparative physiology

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Comparative physiology is a subdiscipline of physiology that studies and exploits the diversity
of functional characteristics of various kinds of organisms. It is closely related to evolutionary
physiology and environmental physiology. Many universities offer undergraduate courses that
cover comparative aspects of animal physiology. According to Clifford Ladd Prosser,
"Comparative Physiology is not so much a defined discipline as a viewpoint, a philosophy."[1]

Contents
• 1 History
• 2 Methodology
• 3 Funding
• 4 Societies
• 5 Biographies
• 6 Some journals that publish articles in comparative animal physiology
• 7 Further reading
• 8 See also
• 9 References

History
Originally, physiology focused primarily on human beings, in large part from a desire to improve
medical practices. When physiologists first began comparing different species it was sometimes
out of simple curiosity to understand how organisms work but also stemmed from a desire to
discover basic physiological principles. This use of specific organisms convenient to study
specific questions is known as the Krogh Principle.

Methodology
C. Ladd Prosser,[2] a founder of modern comparative physiology, outlined a broad agenda for
comparative physiology in his 1950 edited volume (see summary and discussion in Garland and
Carter[3]):

1. To describe how different kinds of animals meet their needs.

This amounts to cataloging functional aspects of biological diversity, and has recently
been criticized as "stamp collecting" with the suggestion that the field should move
beyond that initial, exploratory phase.[4]
2. The use of physiological information to reconstruct phylogenetic relationships of organisms.

In principle physiological information could be used just as morphological information or


DNA sequence is used to measure evolutionary divergence of organisms. In practice, this
has rarely been done, for at least four reasons:

• physiology doesn't leave many fossil cues,


• it can't be measured on museum specimens,
• it is difficult to quantify as compared with morphology or DNA sequences, and
• physiology is more likely to be adaptive than DNA, and so subject to parallel and
convergent evolution, which confuses phylogenetic reconstruction.

3. To elucidate how physiology mediates interactions between organisms and their


environments.

This is essentially physiological ecology or ecological physiology.

4. To identify "model systems" for studying particular physiological functions.

Examples of this include using squid giant axons to understand general principles of
nerve transmission, using rattlesnake tail shaker muscles for measurement of in vivo
changes in metabolites (because the whole animal can be put in an NMR machine),[5] and
the use of ectothermic poikilotherms to study effects of temperature on physiology.

5. To use the "kind of animal" as an experimental variable.

"While other branches of physiology use such variables as light, temperature, oxygen
tension, and hormone balance, comparative physiology uses, in addition, species or
animal type as a variable for each function." [6]
25 years later, Prosser put things this way: "I like to think of it as that method in
physiology which uses kind of organism as one experimental variable."[1]

Comparative physiologists often study organisms that live in "extreme" environments (e.g.,
deserts) because they expect to find especially clear examples of evolutionary adaptation.[3] One
example is the study of water balance in desert-inhabiting mammals, which have been found to
exhibit kidney specializations.[7]

Similarly, comparative physiologists have been attracted to "unusual" organisms, such as very
large or small ones. As an example, of the latter, hummingbirds have been studied. As another
example, giraffe have been studied because of their long necks and the expectation that this
would lead to specializations related to the regulation of blood pressure. More generally,
ectothermic vertebrates have been studied to determine how blood acid-base balance and pH
change as body temperature changes.

Funding
In the United States, research in comparative physiology is funded by both the National Institutes
of Health and the National Science Foundation.

Societies
A number of scientific societies feature sections on comparative physiology, including:

• American Physiological Society


• Australian & New Zealand Society for Comparative Physiology & Biochemistry
• Canadian Society of Zoologists
• Japanese Society for Comparative Physiology and Biochemistry
• Society for Integrative and Comparative Biology
• Society for Experimental Biology

Biographies
Knut Schmidt-Nielsen (1915–2007) was a major figure in vertebrate comparative physiology,
serving on the faculty at Duke University for many years and training a large number of students
(obituary). He also authored several books, including an influential text, all known for their
accessible writing style.

Grover C. Stephens (1925–2003) was a well-known invertebrate comparative physiologist,


serving on the faculty of the University of Minnesota until becoming the founding chairman of
the Department of Organismic Biology at the University of California at Irvine in 1964. He was
the mentor for numerous graduate students, many of whom have gone on to further build the
field (obituary). He authored several books and in addition to being an accomplished biologist
was also an accomplished pianist and philosopher.

Some journals that publish articles in comparative animal


physiology
• American Journal of Physiology - Regulatory, Integrative and Comparative Physiology
• Annual Review of Physiology
• Comparative Biochemistry and Physiology
• Integrative and Comparative Biology
• Journal of Comparative Physiology
• Journal of Experimental Biology
• Physiological and Biochemical Zoology

Further reading
• Barrington, E. J. W. 1975. Comparative physiology and the challenge of design. Journal
of Experimental Zoology 194:271-286.
• Clark, A. J. 1927. Comparative physiology of the heart. Cambridge University Press,
London.
• Dantzler, W. H., ed. 1997. Handbook of physiology. Section 13: comparative physiology.
Vol. I. Oxford Univ. Press, New York.
• Dantzler, W. H., ed. 1997. Handbook of physiology. Section 13: comparative physiology.
Vol. II. Oxford Univ. Press, New York. viii + 751-1824 pp.
• Feder, M. E., A. F. Bennett, W. W. Burggren, and R. B. Huey, eds. 1987. New directions
in ecological physiology. Cambridge Univ. Press, New York. 364 pp.
• Garland, T., Jr., and P. A. Carter. 1994. Evolutionary physiology. Annual Review of
Physiology 56:579-621. PDF
• Gibbs, A. G. (1999). "Laboratory selection for the comparative physiologist". Journal of
Experimental Biology. 202 (Pt 20): 2709–2718. PMID 10504307.
• Gilmour, K. M.; Wilson, R. W.; Sloman, K. A. (2005). "The integration of behaviour into
comparative physiology". Physiological and Biochemical Zoology. 78 (5): 669–678.
doi:10.1086/432144. PMID 16047293. S2CID 586358.
• Gordon, M. S., G. A. Bartholomew, A. D. Grinnell, C. B. Jorgensen, and F. N. White.
1982. Animal physiology: principles and adaptations. 4th ed. MacMillan, New York. 635
pages.
• Greenberg, M. J., P. W. Hochachka, and C. P. Mangum, eds. 1975. New directions in
comparative physiology and biochemistry. Journal of Experimental Zoology 194:1-347.
• Hochachka, P. W., and G. N. Somero. 2002. Biochemical adaptation — mechanism and
process in physiological evolution. Oxford University Press. 478 pp.
• Mangum, C. P., and P. W. Hochachka. 1998. New directions in comparative physiology
and biochemistry: mechanisms, adaptations, and evolution. Physiological Zoology
71:471-484.
• Moyes, C. D., and P. M. Schulte. 2006. Principles of animal physiology. Pearson
Benjamin Cummings, San Francisco. 734 pp.
• Prosser, C. L., ed. 1950. Comparative animal physiology. W. B. Saunders Co.,
Philadelphia. ix + 888 pp.
• Randall, D., W. Burggren, and K. French. 2002. Eckert animal physiology: mechanisms
and adaptations. 5th ed. W. H. Freeman and Co., New York. 736 pp. + glossary,
appendices, index.
• Ross, D. M. (1981). "Illusion and reality in comparative physiology". Canadian Journal
of Zoology. 59 (11): 2151–2158. doi:10.1139/z81-291.
• Schmidt-Nielsen, K. 1972. How animals work. Cambridge University Press, Cambridge.
• Schmidt-Nielsen, K. 1984. Scaling: why is animal size so important? Cambridge
University Press, Cambridge. 241 pp.
• Schmidt-Nielsen, K. 1997. Animal physiology: adaptation and environment. 5th ed.
Cambridge University Press, Cambridge. ix + 607 pp.
• Schmidt-Nielsen, K. 1998. The camel's nose: memoirs of a curious scientist. 352 pp. The
Island Press. Review
• Somero, G. N. 2000. Unity in Diversity: A perspective on the methods, contributions, and
future of comparative physiology. Annual Review of Physiology 62:927-937.
• Stephens, G.C.; Schinske, R.A. (1961). "Uptake of amino acids by marine invertebrates".
Limnology and Oceanography. 6 (2): 175–181. Bibcode:1961LimOc...6..175S.
doi:10.4319/lo.1961.6.2.0175.
• Stephens, G.C. (1982). "Recent progress in the study of "Die Ernährung der Wassertiere
und der Stoffhaushalt der Gewasser"". American Zoologist. 22 (3): 611–619.
doi:10.1093/icb/22.3.611.
• Manahan, D.T.; Wright, S.H.; Stephens, G.C.; Rice, M.A. (1982). "Transport of dissolved
amino acids by the mussel, Mytilus edulis: Demonstration of net uptake from seawater by
HPLC analysis". Science. 215 (4537): 1253–1255. doi:10.1126/science.215.4537.1253.
PMID 17757542. S2CID 36756710.
• Swallow, J. G.; Jr; Garland, T. (2005). "Selection experiments as a tool in evolutionary
and comparative physiology: insights into complex traits - An introduction to the
symposium". Integrative and Comparative Biology. 45 (3): 387–390.
doi:10.1093/icb/45.3.387. PMID 21676784.
• Willmer, P., G. Stone, and I. Johnston. 2005. Environmental physiology of animals.
Second edition. Blackwell Science, Oxford, U.K. xiii + 754 pp.

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