Biology 629. Advanced Animal Behavior Fall 2006 (Last modified on September 21, 2006)

Instructor: Alexander (Sasha) Kitaysky 413 Irving I 474-5179 e-mail: [email protected] office hours: by appointment.

Tuesdays – Laboratory section 110 Irving I (Kitaysky’s Lab) Thursdays – Lecture/seminar, 2:30p.m.—5:30p.m., 332 Irving II Final Examination Period: 15 Dec., 8-10 a.m.

General philosophy: in this class you should aid the learning of others. Also, I would appreciate everyone's help from time to time in one mundane task as needed, e.g. copying of materials for reserve readings, logistic support of the research project.

General Course Goals: The assigned readings, field research project and hands-on hormonal analyses are intended to introduce graduate students to the field of mechanistic approaches (specifically, the field approach) in studying animal behavior. The emphases are on hormonal regulation of wild birds behavior. The course is organized into three parts:

1. General principles of behavioral endocrinology (i.e. functions of the endocrine system, structure, secretion, action, and basic analytical techniques) will be surveyed. 2. Physiology of behavior, environmental and comparative aspects. 3. Research project using field endocrinology approach, collection of field samples and use of radio-immunoassay for hormonal analyses.

RECOMMENDED BOOKS Norris, D.O. (1997). Vertebrate Endocrinology. Academic Press, New York. Nelson, R.J. (2000). An introduction to Behavioral Endocrinology. Sinauer Associates, Inc. Massachusetts. Becker, J.B., S.M. Breedlove, and D. Crews, editors. (1992). Behavioral Endocrinology. Massachusetts Institute of Technology.

RECOMMENDED JOURNALS General journals that frequently publish articles in behavioral endocrinology: General and Comparative Endocrinology, and Behavior, Journal of Comparative Physiology, Journal of Experimental Zoology, Animal Behaviour, Animal Ecology, Behavioral Ecology, Behavioral Ecology and Sociobiology, Ecology, Functional Ecology, Proceedings of Royal Society B London, Ecology letters, TREE, PNAS.

Note: If you would like to request academic accommodations due to a disability, please contact Disabled Student Services, (2nd fl. Whitaker) [email protected], 474-7043. If you have a letter from Disabled Student Services indicating you have a disability that requires academic accommodations, please present the letter to us so we can discuss the accommodations you might need for class.

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GRADING Participation in discussions (10% of grade). Before each class period you should read all of the required materials and write down several questions and comments. I expect you to make sure that those questions and comments are covered during the class period. At the end of each class period I will collect the question and comment pages. Generally, I’ll return the pages without comment, but I will provide some suggestions if I feel you should be participating more in class.

Leading discussions (10%). You will be designated as the discussion leader, either alone or with another classmate, of one of the readings and will be the leader of the discussion of that reading. Note: I still expect you to read the other readings and to arrive at class with questions and comments about those readings as well as being ready to lead the discussion of the reading assigned to you.

Independent Research project (60%). 1. Write a short (~5 pp of text and 5-10 references) proposal for your independent project. First draft is due on September 28, and final draft is due on October 5. 2. Conduct sample collection by November 15. 3. Conduct RIAs by December 1. 4. Write a research paper – the first draft is due on December 7, the second (a final) draft is due on December 14. Presentation and Discussion of the obtained results in class. The paper should be formatted for submission to a scientific journal. 5. Prepare a poster - due by December 14. 6. In the time block assigned for the final examination you will make a <20 minute presentation of your research paper, allowing 5 minutes for questions from the class. This session will follow the format of a session at a professional meeting.

RIA analyses (20%). This class is designed as a hands-on analytical techniques venture. You will analyze blood samples you collected for corticosterone (direct assay, mandatory for everyone), CBG (mandatory for a lead author and “suggested” for others), and (mandatory for a lead author).

Tentative course outline (subject to change). Readings are assigned for each lecture (see syllabus). During the course, a discussion group with student oral presentations of recent papers in the field of behavioral endocrinology will follow a cycle of independent research project. Copies of papers will be available in Sasha's office for students to sign out, copy themselves and return.

2 Four major components of the course: Lectures, papers for discussion, RIA, independent projects

Tentative Independent projects: Kalb – chickadees Tawna – chickadees Rebecca – eiders Audrey – shorebirds

Lectures – as assigned

Papers – discussion of 2-3 papers in class (LIST OF PAPERS ATTACHED). Major topics: SEASONALITY OF STRESS RESPONSE HORMONES AND MIGRATORY BEHAVIOR METABOLISM, STRESS AND REPRODUCTION HPA and HPT interactions – MECHANISMS AND BEHAVIORAL CONSEQUENCES

RIAs – training (1 week)

Independent project – 1) proposal draft and presentation 2) final proposal 3) RIA and data analyses 4) paper draft and presentation 5) paper final draft and presentation 6) Poster?

Class Tentative Dates Topic Events 1 31 Aug. Introduction to course 2 1-7 Sept. IACUC training; choosing topics for discussions in class 3 31 Aug. – 7 Sept. The concept of allostasis in biology (read assigned papers) 4 7 Sept. Find up to 5 papers in area of your interest (subject – behavioral endocrinology) 5 14 Sept. Paper review in class (Romero 2002). Goal – identify specific areas in behavioral endocrinology that will be covered in class. 6 21 Sept. Lecture: Endocrine system –major glands, various axis

Discussion of a first draft of proposal in class – Kalb’s presentation Chickadees: seasonal changes in physiology

7 22-29 Sept. 1) Choose topic for your independent project (Audrey, Rebecca, Tawna) 2) LAB 1 – collection of blood samples (captive birds – contingent on IACUC approval), preparation of plasma samples 8 28 Sept. Types of hormones

2 Papers on SEASONALITY OF STRESS RESPONSE 9 1-10 Oct. Radio-immunoassay analysis of plasma samples:

3 Review of general Methodology and laboratory protocols

10 5 Oct. Secretion of hormones 2 Papers on SEASONALITY OF STRESS RESPONSE Discussion of “Eider” proposal – Rebecca’s presentation 13-20 Oct. RIA analyses – training in the lab 11 12 Oct. Transport of hormones from a gland to target tissues (CBG etc); free and bound fractions of hormones

2 papers on METABOLISM, STRESS AND REPRODUCTION

Discussion of “Shorebird” proposal – Audrey’s presentation 12 19 Oct. Types of Receptors: membrane and nuclear receptors

2 papers on METABOLISM, STRESS AND REPRODUCTION 13 26 Oct. Hormone metabolism and excretion

2 papers on METABOLISM, STRESS AND REPRODUCTION

14 3 Nov. Details of the HP- Adrenal, gonadal and thyroid axis

2 papers on HORMONES AND MIGRATION 15 10 Nov. 2 papers on HORMONES AND MIGRATION 16 10-30 Nov. RIA and CBG – ANALYSES OF SAMPLES FOR INDEPENDENT PROJECTS 17 7 Dec. 2 papers on HPA and HPT interactions

FIRST DRAFTS OF RESEARCH PAPERS DUE 18 14 Dec FINAL DRAFTS OF RESEARCH PAPERS and Student Presentations

PAPERS FOR DISCUSSION IN CLASS

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SEASONALITY OF THE STRESS RESPONSE

Romero, L.M. 2002. Seasonal changes in plasma glucocorticoid concentrations in free-living vertebrates. General and Comparative Endocrinology 128: 1-24.

Breuner and Orchinik. 2000. Downstream from corticosterone: seasonality of binding globulins, receptors and behavior in the Avian stress response. Avian Endocrinology. Eds Dawson and Chaturvedi.

Breuner, C.W. and T.P. Hahn. 2003. Integrating stress physiology, environmental change, and behavior in free-living sparrows. Hormones and Behavior 43:115-123.

METABOLISM, STRESS AND REPRODUCTION

Love, O. P., C.W. Breuner, F. Vezina, and T.D. Williams. 2004. Mediation of a corticosterone- induced reproductive conflict. Hormones and Behavior 46:59-65.

Chastel et al. 2003. Pre-breeding energy requirements: thyroid hormone, metabolism and the timing of reproduction in house sparrows. J Avian Biology 34, 298-306.

F. Criscuolo et al. 2006. Body mass and clutch size may modulate prolactin and corticosterone levels in eiders. Physiological and Biochemical Zoology 79:514-521.

Robin,J., L. Boucontet, P. Chillet, and R. Groscolas. 1998. Behavioral changes in fasting emperor penguins; evidence for a 'refeeding signal' linked to a metabolic shift. American Physiological Society: R746-R753.

Meek and Lee. 1994. Lutenizing Hormone and Prolactin in mated female meadow voles housed in long and short days. Biology of reproduction 51: 725-730.

Solomon et al. 2001. Mechanisms of reproductive suppression in female pine voles. Reproduction 122, 297-304.

Kotula-Balak et al. 2003. Complementary approaches demonstrate that cellular aromatization in the bank vole testis is related to photoperiod. European Journal of Histochemistry.

HORMONES AND MIGRATION

Holberton, et al. 1999. Endocrine aspects of physiological condition, weather, and habitat quality in landbird migrants during the non-breeding period. Pages 847-866 in Proceedings of the 22nd International Ornithological Congress, Johannesburg, South Africa.

Holberton, R.L. 1999. Changes in patterns of corticosterone secretion concurrent with migratory fattening in a Neotropical migratory bird. Gen. Comp. Endocrinology 116: 49-58.

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Marra, P.P and R.L. Holberton. 1998. Corticosterone levels as indicators of habitat quality: effects of habitat segregation in a migratory bird during the non-breeding season. Oecologia 116: 284-292.

Mizrahi, et al. 2001. Patterns of corticosterone secretion in migrating semipalmated sandpipers at a major spring stopover site. Auk 118(1):79-91.

HPA AND HPT INTERACTIONS

Helmreich et al. 2005. Relation between the Hypothalamic-pituitary-thyroid axis and HPA axis during repeated stress. 81, 183-192.

Lo et al. 1998. Acute effects of thyroid hormones on the production of adrenal cAMP and corticosterone in male rats. Am J Physiol Endocrinol Metab 274, 238-245.

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