Introduction to Biology 174

1. Introduction to Biology 174

2. "Ecological and Evolutionary Physiology" (BIOL 174) Winter 2016 Professor: Dr. Theodore Garland, Jr. Department of Biology Office: 2366 Spieth Hall Phone 827-3524 - better for long questions tgarland@ucr.edu Office Hours: Tuesday 8:30-9:30 and Wednesday 10:30-11:30 A.M. in 2366 Spieth, or by appt.

3. Lecture: Tuesday and Thursday 2:10 - 3:30 A.M. in 2200 Spieth Hall Lectures will be posted on the "Blackboard" website ilearn.ucr.eduafter the lecture is finished. Discussion: All are on Wed. 12:10-1:00, 1:10-2:00, 4:10-5:00, 5:10-6:00 PM Syllabus: Both Lecture and Discussion are posted on the course website. T.A. will go over with you.

4. Teaching Assistant: Mr. Jarren Kay Ph.D. candidate Department of Biology jkay001@ucr.edu Office Hours: Monday 10:30-11:30 PM Thursday 11-12 PM in 2378 Spieth Hall, or by appointment

5. "Ecological and Evolutionary Physiology" (BIOL 174) Winter 2014Catalog Description: Interactions between organisms and their environments, emphasizing coadaptation of physiological, morphological, and behavioral phenotypes. Topics include: allometry and scaling, metabolism and locomotion, heat and water exchange, evolution of endothermy, artificial selection experiments, and phylogenetically based statistical methods.

6. Some other Relevant Courses at UCR: "Animal Physiology" (BIOL 175) "Comparative Biomechanics" (BIOL 176) "Hormones and Behavior" (BIOL 178) "Animal Behavior" (BIOL 160) "Functional Anatomy of the Vertebrates" (BIOL 161 A, B)

7. Required Text: Readings posted on the "Blackboard" website ilearn.ucr.edu Don’t get behind on the reading! Exams will take some material directly from the reading, even if we have not discussed it in class! A copy of the first exam from a previous year is posted on the Blackboard website. Warning:Material covered will not be identical !!!Format may differ !!!

8. Grading Item Points % of Total -------------------------------------------------------------------------------------------------- Pre-Course Survey (5 Jan) 20 8.9 Midterm Exam 1 (28 Jan) 40 17.8 Midterm Exam 2 (25 Feb) 40 17.8 Final Exam 3 (14 Mar) 60 26.7 Discussion Section Attendance 10 4.4 Participation 10 4.4 3 Quizzes (6 Jan, 3 Feb, 2 Mar) 15 (5 pts each) 6.7 Paper Critique (20 Jan) 10 4.4 Heritability Exercise (9 Mar) 20 8.9 ------------------------------------------------------------------------------------------------- Total Points 225 100

9. How grades are determined (of total points): > 90% = A > 80% = B > 70% = C > 60% = D < 60% = F Scale may be moved down ("curved"), depending on distribution of total points at end of course. I cannot tell you in advance if this will happenor by how much. + or - may also be used

10. Regrades: only within one week of when you get it back. We will regrade the whole thing, and points on ANY question may go up or down. Missed assignments: no make-ups are allowed, except for ... Valid excuses, absences: tell us when it happens, or before. Do not wait until the end of the quarter! For medical absences, we need a note from your healthcare provider.

11. Lecture 2: • Historical Develop-ment of Ecological and Evolutionary Physiology

12. Some highlights,not a thorough account ...

13. C. Ladd Prosser: a founder of comparative physiology Edited 1950 volume, in whichhe outlined a broadagenda for comparative physiology:Prosser, C. L., ed. 1950. Comparative animal physiology. W. B. Saunders Co., Philadelphia. ix + 888 pp.

14. Five Big Goals: 1. Simply to describe how different kinds of animals meet their needs. = cataloging biological diversity "Biodiversity" often = how many species But perhaps equally important is how variable are those species, morphologically, physiologically, behaviorally? In other words, functional diversity. Recently, some have criticized this as "stamp collecting" and said that we must now move beyond that phase of simple description.

15. 2. Use physiological information to reconstruct phylogenetic relationships of organisms. In principle, we could use physiological information just like we use morphological information or DNA sequences. In practice, has rarely been done, for 4 reasons: a. physiology doesn't leave many fossil cues b. it can't be measured on museum specimens c. it is difficult to measure as compared with morphology or DNA sequences d. more likely to be adaptive than DNA, so subject to parallel and convergent evolution, which confuses phylogenetic reconstruction.

16. 3. To elucidate how physiology mediates interactions between organisms and their environments. = physiological ecology or ecological physiology

17. 4. To identify "model systems" for studying particular physiological functions. = comparative physiology Examples: a. squid giant axons for nerve transmission b. rattlesnake tail shaker muscles because whole animal can be put in an NMR to measurein vivo changes in metabolites Conley, K. E., and S. L. Lindstedt. 1996. Rattlesnake tail-shaking: minimal cost per twitch in striated muscle. Nature 383:71-73. c. ectothermic poikilotherms in general to study effects of temperature on physiology d. lizards to study effects of fever

18. 5. Vague ... but indicates that "kind of animal" can be used as a sort of "experimental variable." Scientists routinely conduct experiments to test hypotheses. Nature conducts "natural experiments," e.g., putting animals in a desert, and we can use the results of these past experiments to understand how evolution occurs.

19. What's in a name? Noun conveys the main focus of the subdiscipline, adjective conveys the particular flavor. Comparative physiology tends to focus on physiology and how it varies among different organisms. Biomedical physiologists may see "comparative" as anything other than human beings or "the rat." Physiological ecology focuses on ecology, and how physiology affects interactions between organisms and their environment.

20. Physiological ecology has always had an evolutionary component. • Calow, P., ed. 1987. Evolutionary physiological ecology. Cambridge University Press, Cambridge. 239 pp., back cover: • "Physiological ecology is concerned with the way that physiological traits fit organisms for the ecological circumstances in which they live, so there is always, by definition, an implicit evolutionary component to it." • Bennett, A. F., and R. B. Huey. 1990. Studying the evolution of physiological performance. Pages 251-284 in D. J. Futuyma andJ. Antonovics, eds. Oxford surveys in evolutionary biology. Vol. 7. Oxford Univ. Press, Oxford., p. 251: • "The field of physiological ecology ... is ... fundamentally evolutionary to the extent that it considers how organisms came to be the way they are and how they might change in the future."

21. Although Comparative Physiology and Physiological Ecology have always had evolutionary components, they were often an afterthought. Typical mode of operation: make whatever physiological measurement they specialized in on some new, exotic or unusual organism ...

23. … then tack on a paragraph about the adaptive (evolutionary) significance of whatever they found.

24. For Example:compare a temperate and an arctic-inhabiting bird. Any difference that was found would be attributed to the past effects of natural selecting(i.e., an evolutionary adaptation). Possibilities of phylogenetic inheritance or random genetic drift were not considered. Often, a priori hypotheses were not tested. Or, if they were tested, it was not with the rigor that would be found in evolutionary biology(e.g., only two species might be compared).

25. These sorts of inferences have been criticized as "just-so stories."

26. "Just So Stories" by Rudyard Kipling Originally published 1902 http://en.wikipedia.org/wiki/Just_So_Stories First published in 1902, these are origin stories, fantastic accounts of how various phenomena came about. 

27. HOW THE ELEPHANT GOT ITS TRUNK HOW THE WHALE GOT HIS THROAT HOW THE CAMEL GOT HIS HUMP HOW THE RHINOCEROS GOT HIS SKIN HOW THE LEOPARD GOT HIS SPOTS

28. So, the standards of evidence and interpretation used by evolutionary biologists were not being employed by most physiologists who wished to say something about the evolution of their favorite physiological system or organism. In short, many physiologists were practicing evolutionary biology without a license.

29. Evolutionary physiology is a new subdiscipline that incorporates much of Prosser's (1950) five objectives, but with more rigorous evolutionary tools and definitions. • This course will emphasize the methods of ecological and evolutionary physiology and their conceptual background, rather than providing lots of empirical facts concerning how different kinds of organisms make their living or have evolved physiologically. • Also, I will show lots of examples. Try to remember the general point of these examples. • And we will need some definitions …

30. For example, the word "adaptation" has several meanings, with two being common in biology: 1. the genetic, evolutionary process of traits that change across generations because of natural selection and consequent changes in gene frequencies (a property of populations); Bennett, A. F. 1997. Adaptation and the evolution of physiological characters. Pages 3-16 in W. H. Dantzler, ed. Handbook of physiology. Section 13: comparative physiology. Vol. I. Oxford Univ. Press, New York.

31. 2. within-individual changes in response to environmental perturbations = phenotypic plasticity (these changes are not passed on to offspring) a. Acclimatization = in the field b. Acclimation = in the lab Often these changes are reversible (phenotypic flexibility) May occur at any point in the lifecycle, but those occurring early, during "critical periods," may be less reversible ... Piersma, T., and J. A. van Gils. 2011. The flexible phenotype: a body-centred integration of ecology, physiology, and behaviour. Oxford Univ. Press, Oxford, U.K. ix + 238 pp.

32. Example with Laboratory House Mice: Mohammed A. Al-kahtani. 2003. Ph.D. in Zoology at the Univ. of Wisconsin - Madison. "Evolutionary and Phenotypic Plasticity of Mammalian Kidney: Using the Laboratory House Mouse as a Model." Currently faculty atKing Faisal University,Saudi Arabia.

33. Three Goals: Determine how mice "adapt" ontogenetically to reduced water availability. Create a "model" for studying kidney function. Create a "model" for studying desert rodents or other desert mammals. Eventually apply to "conservation physiology" in the context of captive breeding progams for endangered gazelle.

34. Two groups, starting at weaning (21 days of age) FW-J = Free Water-Juveniles WR-J = Water Restricted-Juveniles

35. Method for Restricting Water

36. Daily water consumption in both juvenile groups Values are means + standard deviations

37. Food intake significantly decreased in water-restricted group.

38. Water restriction stunted the growth of water-restricted group.

39. Are such changes reversible, or do they carry over as the animals continue to age?

40. Reversal Experiment

41. Control Groups

42. "Water-Restricted, then Free Water" group drank copiously when reversed to free water and remained significantly higher than Free Water group.

43. Food intake is intimately linked with water consumption in house mice.

44. The effects of early-life water restriction are not completely reversible.

45. The kidneys became larger (relative to body mass) in the water-restricted juvenile group. Note that if you did not account for the difference in body mass, you would conclude that the kidneys of the water-restricted group became smaller!

46. The urine was much more concentrated in the water-restricted juvenile group.

47. The heart became smaller (relative to body mass) in the water-restricted juvenile group.

48. What about trans-generational effects? • This would be a good topic for future research • Any evidence for epigenetic inheritance? • Epigenetic processes: • changes caused by modification of gene expression rather than alteration of the genetic code itself • heritable changes in gene expression that do not involve changes to the underlying DNA sequence • e.g., by DNA methylation Stopped here 6 Jan. 2015

49. The origin of modern"Evolutionary Physiology"

50. 1. Debates concerning the metabolic and thermoregulatory status of dinosaurs and mammal-like reptiles. Were dinosaurs warm blooded? Modern evolutionary physiology arose in the late 1970s and early 1980s. Three main contributing factors: