Principles of Life Hillis • Sadava • Heller • Price Answers to the Analyze the Data, Apply the Concept, and Working with Data Questions Chapter 15: Mechanisms of Evolution Analyze the Data Figure 15.19 A Heterozygote Mating Advantage A. For C. philodice, 43.2% of all viable males are heterozygous, so 56.8% must be homozygous. To get expected numbers of heterozygous and homozygous mating males, we multiple the expected proportions (from all viable males) by the total number of mating males sampled. Therefore, we expect to see (0.432)(50) = 21.6 heterozygous mating males, and (0.568)(50) = 28.4 homozygous mating males. If we repeat the same calculations for C. eurytheme, we expect (under the given assumption) to see (0.478)(59) = 28.2 heterozygous mating males, and (0.522)(50) = 30.8 homozygous mating males. B. Chi-square calculations for C. philodice: Genotype Expected (E) Observed (O) O – E (O – E)2 (O – E)2/E Heterozygotes 21.6 31 9.4 88.36 4.091 Homozygotes 28.4 19 9.4 88.36 3.111 The sum of the last column gives the chi-square test statistic: 7.302. Since this value is greater than the critical value (P = 0.05) of 3.841, the observed results are significantly different from the expectations at P < 0.05. In other words, we can reject the null hypothesis and conclude that the proportions of each genotype (heterozygotes and homozygotes) of mating males are significantly different from the proportions of these genotypes seen among all viable males in C. philodice. Chi-square calculations for C. eurytheme: Genotype Expected (E) Observed (O) O – E (O – E)2 (O – E)2/E Heterozygotes 28.2 45 16.8 282.17 10.005 Homozygotes 30.8 14 16.8 282.17 9.162 The sum of the last column gives the chi-square test statistic: 19.167. Since this value is greater than the critical value (P = 0.05) of 3.841, the observed results are significantly different from the expectations at P < 0.05. In other words, we can reject the null hypothesis and conclude that the proportions of each genotype (heterozygotes and homozygotes) of mating males are significantly different from the proportions of these genotypes seen among all viable males in C. eurytheme. © 2011 Sinauer Associates, Inc. 1 Apply the Concept Concept 15.1 Evolution Can Be Measured by Changes in Allele Frequencies, p. 299 1. Observed allele frequencies: Frequency of a = p = 0.5; frequency of A = q = 0.5 Observed genotype frequencies: Frequency aa = 0.4 Frequency Aa = 0.2 Frequency AA = 0.4 2. The population is not in Hardy–Weinberg equilibrium. There are more homozygotes, and thus fewer heterozygotes, than expected under Hardy–Weinberg equilibrium. Since p = 0.5 and q = 0.5, we would expect heterozygotes to be present at a frequency of 2pq = 0.5 at Hardy–Weinberg equilibrium. The population is likely from two or more different source populations (one with a higher frequency of a, and the other with a higher frequency of A). 3. Allele frequencies: p = 0.5, q = 0.5 (unchanged) Genotype frequencies: Frequency of aa = p2 = 0.25 Frequency of Aa = 2pq = 0.5 Frequency of AA = q2 = 0.25 4. Several of the assumptions of Hardy–Weinberg equilibrium are violated in this example, any one of which could lead to deviations in Hardy–Weinberg expectations. For instance, the population size is not infinite, and in fact it is quite small. Thus, it is subject to random deviations due to chance events. Also, random mating is not possible, especially since the six males can only mate with four possible females. Since the genotype frequencies within each sex differ from the genotype frequencies of the population as a whole, this will have a large effect on the resulting offspring. Further, given that gene flow has occurred in the past (that is how the population was founded), it is likely to occur again in the future. Any gene flow from other populations will affect the allele and genotype frequencies. Finally, the effects from the violated assumptions noted above will be quite large, and the effects of mutation and selection will probably be very small by comparison (at least over just one generation). However, over time new mutations and selection may also influence allele and genotype frequencies in this population. © 2011 Sinauer Associates, Inc. 2 Concept 15.2 Genomes Reveal both Neutral and Selective Processes of Evolution, p. 305 1. Codon numbers 12, 15, and 61 are likely to be evolving under positive selection for change because these three codons have each experienced a large number of nonsynonymous substitutions (which give rise to amino acid replacements), but relatively few synonymous substitutions. 2. Codon number 165 is the most likely of these codons to be drifting neutrally among states, as it has experienced similar rates of synonymous and nonsynonomous substitution. 3. Codon numbers 80, 137, 156, and 226 are likely evolving under purifying selection, as the vast majority of changes at these codons are synonymous substitutions, which do not result in amino acid replacements. Substitutions that result in amino acid changes (nonsynonymous substitutions) undoubtedly occur, but are usually selected against in the population. Working with Data 15.1 Testing for Significant Differences (Figure 15.9) 1. A. The details of this answer will depend on your specific randomization trials. If you conduct enough randomization replications, you should expect to see group differences as large as those seen in this experiment less than 5% of the time. B. P < 0.05 2. A. Large sample sizes provide greater power to discriminate between group means, if the groups are actually different. If the actual difference between group means is small, the experiment will require larger sample sizes to detect the difference. Thus, the power of the experiment to detect group differences increases with increasing sample size. B. Try replicating the randomization test above, but increase the sample size by a factor of two. C. Assume that the new data are similar to the first set of data collected, and write out two cards for each observation. Even though the difference in group means is identical, you should see that it is easier to detect the differences with a larger sample size. © 2011 Sinauer Associates, Inc. 3 15.2 Determining the Paternity of Butterfly Larvae (Figure 15.19) 1. (0.75)16 is approximately 0.01002. Therefore, the investigators would need to analyze at least 16 larvae from each batch of eggs to judge the genotype of the father with 99 percent certainty. An easy way to find this answer is to multiply 0.75 0.75, and then multiply the answer by 0.75, and continue until the result is approximately 0.01. Keep track of the number of times you multiply by 0.75 to find the appropriate sample size. 2. (See Analyze the Data for Figure 15.19 above.) 3. Alternative explanations are directly from Question 4: A possible alternative hypothesis to explain why heterozygous males disproportionately father offspring is that females that mate with those males are more likely to lay eggs than females that mate with homozygous males. The investigators rejected that hypothesis because nearly all females laid fertilized eggs in captivity. Another alternative is that fitness differences were not due to alleles at the PGI locus but were due to genes at other loci on the same chromosome. A third possibility is that heterozygote males, being more vigorous flyers, are more difficult to capture. If so, the investigators might have underestimated the frequency of heterozygotes in the wild. 4. As already noted, the first hypothesis was rejected because nearly all the females laid fertilized eggs in captivity. The second hypothesis received no support because the observed differences in flight performance were consistent (could be accounted for) by the laboratory-measured differences among the PGI alleles. Thus, although additional loci on the same chromosome could contribute to flight performance, the observed variation of PGI alleles is sufficient to explain the observed results. To avoid the third possibility (a sampling bias against heterozygotes), the investigators did not capture butterflies while they were attempting to escape (which could have led to a bias against sampling the better fliers). Instead, they captured unaware individuals as they fed on flowers, searched for mates, oviposited, or interacted with one another. This ensured that the likelihood of capture was unrelated to flying ability of the butterflies. © 2011 Sinauer Associates, Inc. 4