Natural Selection in Galapagos Finches Question: Did natural selection on ground finches occur when the environment changed? Introduction In 1973, biologists Peter and Rosemary Grant began a 30-year long study to find out if they could document natural selection. On Daphne Major, a tiny island in the Galapagos, they caught, weighed, measured, and recorded the diets of hundreds of ground finches, tiny birds about the size of sparrows. The diet of ground finches consists mostly of seeds. Among ground finches, coloration, body size, wing length and the size and length and depth of the beak are inherited traits that vary in the population. The Grants found that finches with large beak depths also have big bodies. (Grant and Grant, 2002) Finch beak depth is illustrated below: Beak Depth Ground Finch In 1977 a severe drought occurred on Daphne Major; instead of receiving the usual 130mm of rain that normally falls, they received only 24mm. Plants withered and seeds of all kinds were scarce. Small soft seeds were quickly eaten by the finches, leaving mainly large, tough seeds that finches normally ignore. Eighty-four percent of the finch population disappeared. Evidence of dead, emaciated birds supported a hypothesis that many of the finches died of starvation. In addition, no individuals from Daphne Major were found on adjacent islands, nor did they return when food supplies returned to normal. The drought ended when a small amount of rain fell on the island in January of 1978. (Grant and Grant, 2002) Hypothesis to be tested: Beak characteristics changed in response to changes in food availability. Modified and Adapted from Delaware Science Coalition Assessments 1 Figures 1, 2, 3, and 4 illustrate data collected by the Grants on Daphne Major. 1976 Finch Population Beak Depth (Total Number in Sample = 751) 80 70 Number of Finches 60 50 40 30 20 10 0 5.8 6.2 6.6 7 7.4 7.8 8.2 8.6 9 9.4 9.8 10.2 10.6 11 11.4 11.8 12.2 12.6 13 13.4 13.8 Beak Depth (mm) Figure 1 = mean beak depth 1978 Finch Population Beak Depth (Total Number in Sample = 89) 80 70 Number of Finches 60 50 40 30 20 10 0 5.8 6.2 6.6 7 7.4 7.8 8.2 8.6 9 9.4 9.8 10.2 10.6 11 11.4 11.8 12.2 12.6 13 13.4 13.8 Beak Depth (mm) = mean beak depth Figure 2 Modified and Adapted from Delaware Science Coalition Assessments 2 Changes in Finch Population Before and After the Drought of 1977 1600 Population Size 1400 1200 1000 800 600 400 200 0 Jan 0 1976 July 6 1976 Jan 12 1977 Jul 18 1977 Jan 24 1978 Jul 30 1978 Jan 36 1979 Figure 3 Changes in Seed Abundance Before and After the Drought of 1977 Seed Abundance (g/m 2) 12 10 8 6 4 2 0 Jan 0 1976 Jul 6 1976 Jan 12 1977 Jul 18 1977 Jan 24 1978 Jul 30 1978 Jan 36 1979 Figure 4 Modified and Adapted from Delaware Science Coalition Assessments 3 Use Information in the Introduction and Figures 1-4 in answering the following questions: 1. Using data in Figure 1, what is the range in beak depth in the finch population in 1976? 5.8 – 13.8 mm Using data in Figure 2, what is the range in beak depth in 1978? 6.8 – 11.4 mm Did variation in beak size increase or decrease after the drought? Decrease 2. How does mean beak depth before the drought compare with mean beak depth after the drought? The mean beak depth before the drought was about 9.2mm. After the drought, it was about 10.0mm. 3. What do the data show about seed abundance before the drought compared with seed abundance after the drought? (Note: Use numerical data in your response.) The seed abundance before the drought ranged from around 7.7 to 10.6 g/m2. During the drought in 1977, the seed abundance decreased to about 2.8 g/m2 but then started to increase to about 3.5 g/m2 after the drought in July 1978. 4. Describe the relationship between the abundance of seeds and the size of the finch population. (Do not explain. If you don’t know the difference between describe and explain, look it up prior to completing this question.) Seed abundance and finch population size are directly proportional. As one increases, the other increases and as one decreases, the other also decreases. 5. What type of natural selection would you say is occurring in the finch population during and after the drought? Explain. Directional selection for a larger beak is occurring in the finch population after the drought because the average beak size is increasing (average phenotype changed in one direction). Modified and Adapted from Delaware Science Coalition Assessments 4 6. Propose an explanation that accounts for the change in the mean depth and the change in variation of the finches’ beaks between 1976 and 1978 using your knowledge of natural selection, and the information in the Introduction and Figures 1-4. A good explanation might look like: The increase in mean beak depth from 9.2 to 10.0 mm and the decrease in the variation of beak depth from a range of 5.8 to 13.8 mm to 6.8 to 11.4 mm can be explained in terms of directional selection for a medium to large beak depth as a result of food availability during the drought in 1977. The introduction states that plants withered and seeds became scarce. Small soft seeds were quickly eaten by the finches leaving mainly large, tough seeds behind. Figure 1 indicates that there was considerable variation in beak depth in the population before the drought, from very small (5.8mm) to very large (13.8mm). Eighty-four percent of the finch population disappeared as a result of the decreased food supply during the drought. The data suggests that finches with smaller beaks are experiencing lower reproductive success during the drought than finches with larger beaks. Finches with medium to very large beak depths were probably able to eat the larger, tougher seeds while finches with smaller beak depths were not able to eat the tough larger seeds. Therefore, many more smaller beak depth finches died than larger beak depth finches since the larger beak depths were able to eat the larger, tougher seeds. In addition, the introduction states that finches with larger beaks also have larger bodies. Since the range in beak depth decreased at both ends (very small and very large), it may indicate that larger body finches with larger beak depths also were at a disadvantage during the drought and were not as fit as finches with medium to large sized bodies and beaks. This could be due to the fact that larger bodies require more food and since food was scarce, very large beak finches (those in the 12-13.8mm range) also starved during the drought. Assuming a genetic component to beak depth, surviving finches which happened to have medium to large beak depths reproduced, passing on the medium to large beak depth genes to the next generation. By 1978, mean beak depth had increased and the range of beak depths had decreased due to the better reproductive success of finches with a medium to large sized beak. Modified and Adapted from Delaware Science Coalition Assessments 5 7. Little rain fell on Daphne Major from 1979 through 1982. At the beginning of 1983 unusually heavy rainfall (1359 mm) occurred on the island. Plants that produced small seeds flourished while bigseed plant populations crashed. With plenty of food available, the size of the finch population increased dramatically over the next several years. a. Predict how the rainy weather of 1983 affected mean beak depth over the next several generations using your knowledge of natural selection. Make your prediction by circling one of the following choices: A. Mean (average) beak depth increased. B. Mean (average) beak depth decreased. C. Mean (average) beak depth remained the same. b. Explain your prediction. Unusually heavy rainfall increased the abundance of small soft seeds and decreased the abundance of larger seeds. This would allow more finches with smaller beak depths to survive and reproduce, decreasing the mean beak depth of the population. 8. It was later discovered in Cliff Tabin’s lab that a gene called Bmp4 involved in cell-cell signaling during beak development is partly responsible for beak depth in ground finches (Abzhanov et al., 2004). Alleles that increase Bmp4 expression increase the depth of the beaks, and were most likely selected for during the drought in 1977 (Abzhanov et al., 2004). In 2001, ground finches were collected from Daphne major and genotypes were analyzed at this particular locus. Phenotype High Bmp4 expression Low Bmp4 expression Low Bmp4 expression Genotype aa Aa AA Number of Individuals 552 336 491 In 2009, allele frequencies were measured again on Daphne major for this particular gene. Phenotype High Bmp4 expression Low Bmp4 expression Low Bmp4 expression Genotype aa Aa AA Number of Individuals 129 493 706 a) If allele frequencies were at Hardy-Weinberg equilibrium, we would expect the allele frequencies in 2001 to be the same as those for 2009 at this particular locus. Determine the allele frequencies for both A and a in 2001 and 2009. Allele Frequency in 2001 Frequency in 2009 A .48 .72 a .52 .28 Modified and Adapted from Delaware Science Coalition Assessments 6 b) Genotypic frequencies can be calculated by multiplying the frequencies of both alleles together. For example, if the frequency of the A allele was .36, the frequency of the AA genotype would be .13. Calculate the Hardy-Weinberg genotypic frequencies in 2001. Remember the sum of all three genotypic frequencies should be 1. f(AA)= .23 f(Aa)= .25*2=.50 f(aa)= .27 c) Calculate the Hardy-Weinberg genotypic frequencies in 2009. f(AA)= .52 f(Aa)= .20*2=.40 f(aa)= .08 Fill in the table with your answers from b) and c). Expected Frequency from Genotype 2001 Observed Frequency in 2009 AA .23 .52 Aa .50 .40 aa .27 .08 d) Assuming Bmp4 is the only gene that controls beak depth, what do these data suggest about the food availability between 2001-2009 on Daphne major? Explain. The data suggests that small soft seeds were more abundant in 2009 compared to 2001. In 2001, the frequency of the ‘a’ allele which causes high Bmp4 expression was much higher, suggesting that there was a selective pressure for increased beak depth (perhaps another drought). In 2009, the frequency of the ‘a’ allele had greatly decreased, suggesting that mean beak depth had decreased due to a selective advantage of low Bmp4 expression (smaller beak depth), or a disadvantage of high Bmp4 expression (larger beak depth). Modified and Adapted from Delaware Science Coalition Assessments 7 References Abzhanov A, Protas M, Grant B R, Grant P R, Tabin C J. Bmp4 and Morphological Variation of Beaks in Darwin’s Finches, Science 2004; 305 (5689): 1462-1465. Grant P, Grant B R. Unpredictable Evolution in a 30-Year Study of Darwin’s Finches, Science 2002; 296 (5568): 707-711. Modified and Adapted from Delaware Science Coalition Assessments 8