1. Evolutionary Fitness

2. Absolute Fitness Reproductive success: change in proportion of offspring in the next generation Can be determined for genotypes or phenotypes W=No/Np, the number of offspring over the number of parents with the trait W=P*N, where P=probability of offspring survival, N=average number of offspring W/W(mean), where W(mean) is the mean fitness of the population as a whole

3. Competitive Index Knight and Roberston, 1957 CI=p/(1-p), where p is the proportion of the offspring with the trait The relative fitness of two strains is given by W=CI1/CI2=[p/(1-p)]/[q/(1-q)]

4. Alternative Competitive Index Jungen and Hartl, 1979 Just use p, not p/(1-p), as a measure of fitness (ie p=N*/N(total)) Relative fitness, W=p/q

5. Long Term/Equilibrium Tests Sved, 1971 W=(r-h)/r(1-h) where r is the viability of offspring of the strain, h is the equilibrium frequency of the strain (average of the last 9 generations of tests) In test by Hartl, 1981, the longer term value gave different results than the single- generation test

6. What can these tell us? These definitions can be useful, as long as we are not trying to test the theory of natural selection itself (ie, natural selection acts to increase W for a population) For example, used to figure out what characteristics make an organism more fit in a certain environment, or to track changes in the characteristics of a population due to changes in the selective pressures

7. Problem: Circularity These definitions are based on the theory of natural selection. We can’t use them to test the theory that natural selection acts to increase the fitness of a population. A phenotype is “more fit” if proportionally more of its offspring survive Natural selection acts to increase the fitness of populations ?

8. Other Measures Viability alone (measure survival rates of offspring) - not tightly linked to fitness as defined previously Reproductive power: the rate at which organisms transform energy to reproduction/offspring (fitness ~ energy used for reproduction) Model proposed by Brown, Marquet and Taper, 1993, widely discredited now, but work on other models

9. Adaptive Landscapes Allele frequency in a population vs fitness (multidimensional map) Looks at fitness of a population, not of a specific genotype The landscape can change (changes in environment, for example, or in competition, available resources, etc) --> shifting surface, different peaks --> no “end point” Local vs global maxima Natural selection acts to increase the fitness of the population as a whole

10. Phenotype vs. Environment

11. Fitness vs. Allele Frequency

12. The initial condition of the population can affect the direction of its selection (extreme example: an environment favoring bigger wings could cause a population to head for more cells/wing or bigger wing cells, or a combination of the two; less extreme example: could get favored AA or BB genotype if favor red flowers) Importance: local, not global, maxima in fitness

13. Adaptive Landscape

14. Random Drift on Adaptive Landscapes

15. Sources: http://links.jstor.org/sici?sici=0003- 0147(199310)142%3A4%3C573%3AEOBSCO%3E2.0.CO%3B2-Chttp://links.jstor.org/sici?sici=0003- 0147(199310)142%3A4%3C573%3AEOBSCO%3E2.0.CO%3B2-C http://links.jstor.org/sici?sici=0003- 0147(199606)147%3A6%3C1087%3AEDOFYB%3E2.0.CO%3B2-Ohttp://links.jstor.org/sici?sici=0003- 0147(199606)147%3A6%3C1087%3AEDOFYB%3E2.0.CO%3B2-O http://links.jstor.org/sici?sici=0269-8463(200110)15%3A5%3C696%3AEOBSLO%3E2.0.CO%3B2- Ghttp://links.jstor.org/sici?sici=0269-8463(200110)15%3A5%3C696%3AEOBSLO%3E2.0.CO%3B2- G http://links.jstor.org/sici?sici=0031-8248(198206)49%3A2%3C268%3AOTPDOF%3E2.0.CO%3B2- Thttp://links.jstor.org/sici?sici=0031-8248(198206)49%3A2%3C268%3AOTPDOF%3E2.0.CO%3B2- T http://www.genetics.org/cgi/reprint/102/3/455 http://www.genetics.org/cgi/reprint/137/2/509.pdf http://www.jstor.org/view/00318248/ap010196/01a00100/5?frame=noframe&userID=823ac019@sw arthmore.edu/01c0a8346d00501c18850&dpi=3&config=jstorhttp://www.jstor.org/view/00318248/ap010196/01a00100/5?frame=noframe&userID=823ac019@sw arthmore.edu/01c0a8346d00501c18850&dpi=3&config=jstor http://www.springerlink.com/content/x81q31h1m6052651/fulltext.pdf

16. Life: The Science of Biology Introduction to Genetic Analysis http://images.google.com/imgres?imgurl=http://www.uwyo.edu/benkman/landscape.jpg &imgrefurl=http://www.uwyo.edu/benkman/rightsideresearch.html&h=992&w=1302&s z=151&hl=en&start=27&um=1&tbnid=aBv4T9zEahoniM:&tbnh=114&tbnw=150&pre v=/images%3Fq%3Dadaptive%2Blandscape%2Bevolution%26start%3D20%26ndsp%3 D20%26um%3D1%26hl%3Den%26client%3Dsafari%26rls%3Den%26sa%3DN