Evolution (L567) 2010
Department of Biology, Indiana University


Graduate evolution (L567).--This is a graduate course in evolution, focusing on the theoretical underpinnings.  Beginning in 2010, I will focus the course more on Evolutionary Ecology and Behavioral Ecology.  The goal will be to complement Dr. Montooth's graduate course in Evolutionary Genetics.  Topics will include life-history theory, evolution of sex, sexual selection, evolution of social behavior, evolution of virulence, and the evolutionary ecology of speciation and macroevolution. I also plan to cover some of the basic theory for Population Genetics and Quantitative Genetics.  Graduate standing or permission of the instructor is required. Advanced undergraduates in Biology are welcome. 

The course will be offered again during the Fall semester of 2012.  Typically, the class meetings are part lecture and part discussion in an informal atmosphere. 


Course syllabus for 2010

Grading Policy


Study questions for  2008

Study questions for first exam, 2010 (pdf) <--updated 6 October 2010

Study questions for second exam, 2010
<--updated 26 November 2010


<>Readings and exercises in support of lecture material

Lecture 1: Early history of evolutionary thought: the Darwinian revolution
(see books by Michael Ruse and E. Mayr)

Overheads for Lecture1
First-day Quiz
mean, variance, covariance, & regression (online handout)
Optional reading: Peters RH: Tautology in ecology and evolution

Lecture 2: The origins of theoretical population genetics (see book by Povine)

Overheads for Lecture2


Lecture 3: Basic Population Genetics.  Quantitative Genetics: Breeder's equation. 

Overheads for Lecture3

Optional (but highly recommended) computer exercise in population genetics.  Click on Pop. Gen. Simulation Instructions to download the worksheet.   After you fill out the worksheet, you can compare your answers to  Answers to PopGen exercise.  Make sure you run through the exercise above before looking at the answers.

Optional: solution for qhat (mathematica notebook) in the mutation-selection balance example.  Here is the solution as html

Note: Excel Population-Genetic exercise #1 due 14 Sept 2010.  There will be "individual differences" in formatting, but hand in 3 pages that have the info contained in this example.

Optional: critique of the Breeder's equation (pdf).



Lecture 4: Selection differential vs the selection gradient: towards a more general formulation. 

Overheads for Lecture4


Lecture 5: Additive genetic variance

Overheads for Lecture5

Optional: solution for VA and VD (mathematica notebook) in the mutation-selection balance example.  Here is the solution as html

Optional: Excel spreadsheet calculation of BreedingValues



Lecture 6 (16 Sept): Frequency-dependent selection and game theory.
Overheads for Lecture6


Note: Excel Population/Quantitative-Genetic exercise #2 due 21 Sept 2010.  There will be "individual differences" in formatting, but hand in 3 pages (or more) that have the info contained in this example.

For today's meeting, read: Maynard Smith's chapter on the basic game-theory model (highly recommended)


Read for lecture 7, Tuesday, 21 Sept 2010
Davies, N.B. 1978. Animal Behaviour 26:138-147


Lecture 7 (21 Sept 2010): Mixed vs. Conditional Strategies

Overheads for Lecture7



Lecture 8 (23 Sept 2010): Logic of animal conflict (discuss Davies 1978); RockPaperScissors

Overheads for Lecture8 Part1 (pdf)
Overheads for Lecture8 Part2 (pdf)
Lively's excel model for hawk-dove game H-D game

Optional: update on the Davies study (pdf).

Optional: Rock Paper Scissors paper (Nature 1996)

Lecture 9 (28 Sept 2010): Games against the field: Fisherian sex ratio

Overheads for Lecture9 (pdf)


For help with calculus see S.P. Thompson, "Calculus made easy" Third edition (QA303.T45)



Lecture10 (30 Sept 2010): Local mate competition

Overheads for Lecture10 (pdf)
LMC_overheads.  Color overheads for the LMC model (3.1 mb).

Suggested reading: Hamilton (1967) extraordinary sex ratios

Optional (but recommended).  Fisher's sex ratio: Graph the CSS:  Mathematica notebookpdf file.

 Exercise 3, due Oct. 7th: create an excel worksheet that calculates and graphs fitness against allocation to male function, ai, as a function of the number of mates.  Include on the graph both the resident's allocation and the ESS.  Here is an example

New: highly recommended.   Links to the local mate competition model (Mathematica fileand (pdf file).


Required reading for lecture 11: Salmon paper by Einum&Fleming (2000) Nature 405:565-567.


Lecture11 (5 October 2010): Life-history theory: size-number compromises
Overheads for Lecture11

Highly recommended: Links to the size-number model (Mathematica file) and (pdf file)

Optional, but highly recommended: Steve Stearns paper on Geometric Mean Fitness


Optional: experimental evolution of bet hedging (pdf).

Bet hedging excel worksheet. 


Lecture11(cont.) (7 October 2010): Discussion of Einum&Fleming.  Temporal environmental variation and geometric mean fitness.

(no new overheads)

-----------------------End of material for first exam---------------

Lecture12(12 October 2010): Cost of sex; the Muller's ratchet and Kondrashov's hatchet.

Overheads for Lecture12
the ratchet and the hatchet overheads

FIRST EXAM (14 October 2010)

Lecture13(19 October2010): The ecological hypotheses for sex.

Overheads for Lecture13


More:  Additional color overheads from lecture 13.

Read Burt & Bell 1987 for class on Thursday 21 October.


Lecture14 (21 October 2010): Testing the ecological hypotheses (Burt and Bell), Difficulties on theory, Pluralism. 
Overheads contained in "overheads for lecture 13" above.

Optional reading: Pluralistic hypotheses The ratchet and the red queen in Nature 1994


Epistasis, Linkage Disequiblium, and Recombination
Overheads for Lecture14

Lecture15, Linkage disequilbrium in sexual selection
Overheads for Lecture15
Suggested reading  Kirkpatrick (1982) in Evolution on the evolution of female choice.

Exercise 4, due Nov. 2nd: create an excel worksheet that calculates epistasis for fitness, the generation of linkage disequilibrium, and the effect of recombination on linkage disequilibrium.  Turn in three sheets, each one for different parameter values.  Use parameters different from the ones in my examples.  Here is an exampleHere is another example.


Lecture16 (2 November 2010).  Good genes and the paradox of female choice
Overheads for Lecture16

Optional (but highly suggested) reading: Hamilton and Zuk (1982)

Suggested reading: Sexual selection in plants (it happens): Delph and Ashman 2006


Suggested reading:Wedekind et al 1995: MHC-dependent choice in humans.

Suggested reading. Reusch et al. 2001: MHC-dependent choice in fish.

Suggested reading: interaction-independent sexual selection: Murphey 1998

read for 9 November: 
reproductive mode and speciation Zeh&Zeh 2000


Lecture17 (4 Nov. 2010).  Sexual selection, genetic correlations, and meiotic drive in stalk-eyed flies.

Overheads for Lecture17


Suggested reading: Wilkinson & Reillo 1994. Correlated selection for female preference in stalk-eyed flies. (see also Wilkinson et al 1998, below)

Suggested reading.  Wilkinson et al. 1998. Nature: meiotic drive suppression in stalk-eyed flies (see also Wilkinson & Reillo 1994)

Suggested reading: Wilkinson's overview of stalk-eyed flies. Wilkinson's web page.

Suggested reading: interaction-independent sexual selection: Murphey 1998


Lecture18 (9 Nov. 2010) Sexual conflict, chase-away selection, selection arenas
Overheads (typed: Zeh&Zeh) for Lecture18
Scanned overheads: examples of meiotic drive (not shown for 2010)
Read: reproductive mode and speciation Zeh&Zeh 2000
Read for exam: Rice (1996) paper on conflict between the sexes

Lecture19 (9 Nov. 2010): Social evolution, kin selection.
Overheads for lecture 19
For 11 Nov 2010: Overheads for derivation of the Taylor&Frank method
Taylor & Frank's paper on how to model kin selection (optional)

Lecture 20 (11 Nov. 2010):
Adaptation and integration of evolutionary forces
First, Taylor and Frank's model (see above)

Templeton's paper on integration of evolutionary forces.
Overheads for
lecture 20
3D graphs from malaria lecture showing changes in the adaptive surface with inbreeding (pdf of 3D graphs).
Mathematica file for invasion of S and C alleles.
Lively's excel simulation spread of S and C alleles with and without inbreeding

pdf output for excel simulation model.


Lecture21 (16 Nov. 2010): Group selection, Population structure, Effective population size (Ne), and shifting balance
Overheads for lecture21


Lecture22 (18 Nov. 2010): The ecology of speciation
Overheads for lecture22

scanned overheads 1 of 2 for lecture 22
scanned overheads 2 of 2 for lecture 22

Lecture23 (23 Nov 2010): Macroevolution
Overheads for lecture23
Jablonski's paper on macroevolution in gastropods.
Coyne and Orr's paper on the evolutionary genetics of speciation
Wade's paper on a gene's eye view of speciation


25 Nov. Thanksgiving.

Lecture 24 (30 Nov 2010).  Discussion of Saetre et al. Nature: sexually selected character displacement in a flycatcher. (News and Views on Saetre et al. paper.)
Discussion points for Saetre et al for lecture 22

And possible chalk talk on the evolution of virulence.

-----------------------End of material for second exam---------------


Lecture(guest) (2 Dec. 2010) (guest lecture by Prof. L. Delph: genetic collelations)

Overheads
. correlated trait evolution as pdf file.
suggested reading is support of Delph lecture Arnold, 1994

SECOND EXAM (7 December 2010)


----------------------Updated for 2010 above this line---------------

Read for Tuesday, December 9th: Schemske and Bradshaw PNAS

Lecture24 (9DEC08: Loose ends.  Class discussion of Schemske and Bradshaw)

Lecture 25 (11Dec08): special topic?



C. M. Lively, Dept. of Biology, Indiana University
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