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Hyde Chapter 5 – Solutions 23 5 MODIFICATIONS TO MENDELIAN PATTERNS OF INHERITANCE CHAPTER SUMMARY QUESTIONS 2. In codominance, the heterozygote expresses the phenotypes of both homozygotes. In incomplete dominance, the heterozygote has a phenotype that is intermediate between those of the two homozygotes. 4. Penetrance is the proportion of individuals of a particular genotype that shows the appropriate phenotype; expressivity is the degree to which a trait is expressed. 6. The disease is recessive at the level of the individual but incompletely dominant at the enzymatic level. 8. The IA and IB alleles produce glycosyl transferase enzymes that each add a different substance to the H structure, whereas the i allele produces a functionless protein that adds nothing. Thus, for example, the IAi heterozygote converts all the H structures to the A antigen making IA dominant to i. IAIB heterozygotes convert the H structures to approximately equal numbers of A and B antigens. 10. The complementation test, or cis–trans test, is used to determine if different recessive mutations are allelic. Mutations that are allelic (that is, in the same gene) will produce a mutant phenotype when they are in the trans configuration. Mutations that are non-allelic (that is, in different genes) will produce a wild-type phenotype when they are in the trans configuration. EXERCISES AND PROBLEMS 12. We see three phenotypes in an approximate 1:2:1 ratio (3:7:4). One of the phenotypes (short) is intermediate between long ears and no ears. Therefore, we have incomplete dominance. The cross is between two heterozygotes: L1L2 L1L2, where L1 = long ears and L2 = no ears. 24 Hyde Chapter 5—Solutions 14. Crosses can be IAIA IBIB, IAIA IBi, IAi IBIB, or IAi IBi. IAIA IBIB IAIA IBi IAIB All type AB 1/2 IAIB:1/2 IAi 1/2 AB:1/2 A IAi IBIB IAi IBi 1/2 IAIB:1/2 IBi 1/2 AB:1/2 B 1/4 IAIB:1/4 IAi:1/4 IBi :1/4 ii 1/4 AB:1/4 A:1/4 B:1/4 O 16. Constraints are set by the length of the gene (number of mutable sites) and the different possible phenotypic effects that can result from alteration of the amino acid sequence of the protein product of the gene. 18. The cross is Cc T1T2 (purple, medium) Cc T1T2 (purple, medium) This problem can be worked out using the branched-line approach. To calculate the probability of each class produced, we can independently calculate the probability of each trait and apply the product rule. 1/4 CC 2/4 Cc 1/4 T1T1 1/16 CC T1T1 purple, tall 2/4 T1T2 2/16 CC T1T2 purple, medium 1/4 T2T2 1/16 CC T2T2 purple, dwarf 1/4 T1T1 2/16 Cc T1T1 purple, tall 2/4 T1T2 4/16 Cc T1T2 purple, medium 1/4 T2T2 2/16 Cc T2T2 purple, dwarf Hyde Chapter 5 – Solutions 1/4 cc 25 1/4 T1T1 1/16 cc T1T1 white, tall 2/4 T1T2 2/16 cc T1T2 white, medium 1/4 T2T2 1/16 cc T2T2 white, dwarf Therefore, the phenotypic ratio is 3/16 C– T1T1 purple, tall 6/16 C– T1T2 purple, medium 3/16 C– T2T2 purple, dwarf white, tall 1/16 cc T1T1 2/16 cc T1T2 white, medium 1/16 cc T2T2 white, dwarf 20. a. If gene A is on an autosome, there will be 10 different genotypes and five different phenotypes. Genotypes A1A1; A1A2; A1A3; A1A4 A2A2; A2A3; A2A4 A3A3 A3A4 A4A4 Phenotypes A1 A2 A3 A3A4 A4 b. If gene A is on the X chromosome, the alleles can now be represented as XA1, XA2, XA3, and XA4. There will be more genotypes and phenotypes, because there is a different sex chromosome composition between males and females. Genotypes XA1XA1; XA1XA2; XA1XA3; XA1XA4 XA2XA2; XA2XA3; XA2XA4 XA3XA3 XA3XA4 XA4XA4 XA1Y XA2Y XA3Y XA4Y Phenotypes A1 females A2 females A3 females A3A4 females A4 females A1 males A2 males A3 males A4 males 26 Hyde Chapter 5—Solutions 22. The 119:32:9 ratio is very close to a 12:3:1 ratio indicating two loci with dominant epistasis. For example: P1 red (AABB) white (aabb) F1 red (AaBb) self F2 119 (A–B– + A–bb) 32 (aaB–) 9 (aabb) CRCW SLSS CRCW SLSS (purple, ovoid) (purple, ovoid) This problem can be worked out using the branched-line approach. To calculate the probability of each class produced, we can independently calculate the probability of each trait and apply the product rule. 24. The cross is 1/4 CRCR 2/4 CRCW 1/4 CWCW 1/4 SLSL 1/16 CRCR SLSL red, long 2/4 SLSS 2/16 CRCR SLSS red, ovoid 1/4 SSSS 1/16 CRCR SSSS red, spherical 1/4 SLSL 2/16 CRCW SLSL purple, long 2/4 SLSS 4/16 CRCW SLSS purple, ovoid 1/4 SSSS 2/16 CRCW SSSS purple, spherical 1/4 SLSL 1/16 CWCW SLSL white, long 2/4 SLSS 2/16 CWCW SLSS white, ovoid 1/4 SSSS 1/16 CWCW SSSS white, spherical 26. The F1 indicates lazy is dominant; therefore, a worker must be a recessive homozygote. If one gene is involved, the cross of the F1 female worker male (Ww ww) should produce 1 worker:1 lazy, for this is a testcross. This result is not seen, so we must have more than one gene involved. Perhaps a worker can result Hyde Chapter 5 – Solutions 27 from more than one gene. Let A–B– = lazy, and A–bb, aaB–, or aabb = workers. The original worker is aabb, and the cross is aabb AABB AaBb (lazy) aabb (worker) 1/4 A–B–:1/4 A–bb:1/4 aaB–:1/4 aabb lazy worker worker worker The observed results (3 workers:1 lazy) are consistent with this explanation. If AaBb AaBb 9/16 A–B– lazy 3/16 A–bb 3/16 aaB– 1/16 aabb workers Thus, the cross of two F1 lazy individuals will produce 9 lazy:7 hard workers. 28. In each cross, we see that all ratios could result from a single gene (3:1, 1:2:1, and 1:1). We can easily explain the red and silver if we assume that red is dominant over silver. Thus, the first cross could be RR rr or Rr rr. Similarly, the second cross could be RR R– or Rr Rr. But we cannot explain any of the white progeny with only two alleles. Since all ratios are single gene ratios, we must propose a third allele that, when homozygous, produces white. Red must be dominant to white. Note that a given diploid individual can have only two alleles. To list all the possible genotypes and their phenotypes, let R = red and r = nonred. RR, RrS, Rrw = red (R is dominant to all other alleles), rSrS = silver, and rwrw = white. What is the phenotype of an rSrw individual? Assume silver is dominant; that is, assume rS is dominant to rw. Thus, rSrw = silver. Now go back to the results. By our model, rwrw is white. Therefore, in order to get white progeny, each parent must have at least one rw allele to contribute. The cross of red by silver to produce white offspring must be Rrw rSrw 1/4 RrS:1/4 Rrw :1/4 rSrw :1/4 rwrw (red) (red) (silver) (white) 30. The cross is CBCY Ii (green) CBCY Ii (green) 28 Hyde Chapter 5—Solutions 3/4 I– 3/16 CBCB I– blue 1/4 ii 1/16 CBCB ii colorless 3/4 I– 6/16 CBCY I– green 1/4 ii 2/16 CBCY ii colorless 3/4 I– 3/16 CYCY I– yellow 1/4 ii 1/16 CYCY ii colorless 1/4 CBCB 2/4 CBCY 1/4 CYCY Thus, the phenotypic ratio is 6 green:3 blue:3 yellow:4 colorless (white). 32. The first two crosses indicate that wild type is dominant to both oranges, and the fourth indicates that orange-2 is dominant to pink. The fifth cross produces four phenotypes, indicating we are dealing with at least two genes. The presence of two genes is also suggested by the cross of orange-1 orange-2. If these two traits were allelic, all the progeny should have been orange. The F1 pink produces progeny that resemble those from a testcross of a dihybrid. If A–B = wild type, A–bb = orange-1, aaB– = orange-2, aabb is probably pink. The crosses in question are then AAbb aaBB (orange-1) (orange-2) AaBb (wild type) self 9/16 A–B– 3/16 A–bb 3/16 aaB– 1/16 aabb wild type orange-1 orange-2 pink Thus the F2 ratio is 9 wild type:3 orange-1:3 orange-2:1 pink. 34. Crosses (a) and (b) do not reveal anything about the dominance hierarchy. Each suggests a mating of a heterozygote with a homozygote. Cross (c) yields a higher proportion of mallard than dusky offspring, indicating that mallard is dominant to dusky. Cross (d) yields a higher proportion of restricted than mallard offspring, indicating that restricted is dominant to mallard. Therefore, the dominance hierarchy Hyde Chapter 5 – Solutions 29 is MR > M > MD. The crosses are (a) MDMD MMD; (b) MDMD MRMD; (c) MMD D D R D d MM ; and (d) MM M M . Note: The dusky allele is actually designated m . 36. This problem can be worked out using the branched-line approach. 1/4 BB 3/4 A– 3/4 C– 9/64 A–BBC– 1/4 cc 3/64 A–BBcc 3/4 C– 18/64 A-BbC– 1/4 cc 6/64 A-Bbcc 3/4 C– 9/64 A–bbC– 1/4 cc 3/64 A–bbcc 3/4 C– 3/64 aaBBC– 1/4 cc 1/64 aaBBcc 3/4 C– 6/64 aaBbC– 1/4 cc 2/64 aaBbcc 3/4 C– 3/64 aabbC– 1/4 cc 1/64 aabbcc 2/4 Bb 1/4 bb 1/4 BB 1/4 aa 2/4 Bb 1/4 bb Because allele B is recessive for lethality, all individuals with genotype BB will not survive. So 16/64 (9/64 + 3/64 + 3/64 + 1/64) individuals die before birth. This leaves 48 surviving offspring. Therefore, the expected phenotypic ratio in the surviving offspring is 18 A–BbC–:6 A–Bbcc:9 A–bbC–:3 A–bbcc:6 aaBbC–: 2 aaBbcc:3 aabbC–:1 aabbcc. 38. a. The genotypic frequencies can be obtained from a Punnett square. Gametes AB AB AABB aB AaBB Ab AABb AaBb aB AaBB aaBB ab AaBb aaBb Therefore, 1/8 AABB:1/8 AABb:2/8 AaBB:2/8 AaBb:1/8 aaBB:1/8 aaBb. The phenotypic frequencies can be derived from the genotypic frequencies. Therefore, 6/8 horned:2/8 hornless or a 3:1 ratio. 30 Hyde Chapter 5—Solutions b. Two hornless animals can produce offspring with horns. The animals will have to be recessive for different genes. For example, if the cross is AAbb aaBB, the offspring will all be AaBb and will all have horns. 40. Because the two genes are independently assorting, we can consider each trait separately. First, joy: the offspring are in an approximate ratio of 1 laughing:1 smiling (41:38). Therefore, one of the parents must be laughing (J1J1) and the other smiling (J1J2). Now, movement: the offspring are in an approximate ratio of 1 runner:2 joggers:1 walker (19:39:21). Thus, both parents must be joggers (M1M2). Combining the two traits, we can determine that the one parent is a laughing jogger and the other is a smiling one! 42. a. To be colorless, individuals must be aabb; the alleles at locus C do not matter. They could be either aabbC– or aabbcc. Therefore the probability is given by (1/4 aa)(1/4 bb)(1/1 C– or cc) = 1/16. b. For red probabilities, determine genotypes that will be red, calculate probability for each genotype, and add. A–bbC– 3/4 1/4 3/4 = 9/64 A–bbcc 3/4 1/4 1/4 = 3/64 A–B–cc 3/4 3/4 1/4 = 9/64 aaB–C– 1/4 3/4 3/4 = 9/64 aaB–cc 1/4 3/4 1/4 = 3/64 33/64 Alternatively, calculate the frequency of blacks as 3/4 A– 3/4 B– 3/4 C– = 27/64. We calculated 1/16 or 4/64 colorless, so 1 – (27/64 + 4/64) = 33/64 = frequency of red. 44. Using the dominance hierarchy, we can determine the potential genotypes for every phenotype. Thus, a gray rabbit can have one of four genotypes (CC, Ccch, Cch, and Cc); a chinchilla rabbit, three (cchcch, cchch, and cchc); a Himalayan rabbit, two (chch and chc); and albino rabbits only one (cc). This order of dominance allows recessive alleles to be “hidden” in various heterozygotes. Crosses (a) and (b) suggest that all three gray rabbits are heterozygous. Cross (c), gray-1 chinchilla-1, resulted in three different genotypes, including Himalayan rabbits. Both parents must thus be heterozygous, and at least one of them must carry the Himalayan allele, ch. Therefore, gray-1 = Cch or Cc and chinchilla-1 = cchch or cchc. (Note: Gray-1 and chinchilla-1 cannot both carry the albino allele at the same time.) We can now go back and analyze cross (a): gray-1 gray-2. It can be represented as Cch (or Cc) C–. The offspring include chinchillas, and so the gray-2 rabbit must be heterozygous for the chinchilla allele, cch. Therefore, gray-2 = Ccch. Let’s move on to cross (b): gray-2 gray-3. It can be represented as Ccch (or Cc) C–. The offspring include chinchilla, and so the gray-3 rabbit cannot be homozygous (CC). However, we cannot tell for sure what the other allele is. Indeed, it can be any of the other three alleles: cch, ch, or c. And finally, cross (d): chinchilla-2 Himalayan, or cch– ch–. The Hyde Chapter 5 – Solutions 31 offspring include albinos, and so both rabbits must be heterozygous for albino allele, c. Therefore, chinchilla-2 = cchc, and Himalayan = chc. The following table summarizes the results. Rabbit Gray-1 Gray-2 Gray-3 Chinchilla-1 Chinchilla-2 Himalayan Genotype Cch or Cc (only if chinchilla-1 is cchch) Ccch Ccch, Cch, or Cc cchch or cchc (only if gray-1 is Cch) cchc chc CHAPTER INTEGRATION PROBLEM a. The true breeding gray parent is AABBCC. Each of the albino parents must be homozygous recessive (cc) at the C locus. Because the A, B, and C genes are independently assorting, each mating can be treated as a series of monohybrid or dihybrid crosses. Cross 1: The F2 offspring are in a ratio of 3 gray:1 albino. The presence of no other color indicates that the F1 rats were homozygous for the A and B genes and have genotype AABBCc. Otherwise, other colored types would have appeared in the offspring. Therefore, the albino-1 parent must have been AABBcc. Cross 2: Without considering the C locus, the F2 offspring are in a ratio of 9 gray: 3 yellow, which can be simplified to a 3:1 ratio. Since yellow offspring have the genotype A–bb, the preceding ratio indicates that the F1 rats were heterozygous for the B gene and homozygous for the A gene (genotype would be AABbCc). Therefore, the albino-2 parent must have been AAbbcc. Cross 3: Without considering the C locus, the F2 offspring are in a ratio of 9 gray: 3 black, which can be simplified to a 3:1 ratio. Since black offspring have the genotype aaB–, the preceding ratio indicates that the F1 rats were heterozygous for the A gene and homozygous for the B gene (genotype would be AaBBCc). Therefore, the albino-2 parent must have been aaBBcc. b. Cross 1: The ratio of colored (gray, black, yellow, and cream) to albino rats is 242:83 or approximately 3:1. This indicates a mating between two heterozygotes for the C locus. The colors are in an approximately 9:3:3:1 ratio, indicating that the parents must also be heterozygous for the A and B genes. Cross 2: The ratio of albino to colored (black and cream) rats is 103:99 or approximately 1:1. This indicates a mating between a heterozygote, Cc, and a homozygote, cc. The absence of gray and yellow offspring means that the parents cannot have any A alleles. The approximately 3 black:1 cream ratio means that the parents were heterozygous at the B locus. 32 Hyde Chapter 5—Solutions Cross 3: There are no albino rats, meaning that the parents cannot produce the cc genotype. The colors are in an approximately 9:3:3:1 ratio, indicating that the parents must be heterozygous for the A and B genes. Summary: Parental Phenotypes Gray gray Black albino Gray gray Cross 1 Cross 2 Cross 3 c. Parental Genotypes AaBbCc AaBbCc aaBbCc aaBbcc AaBbCC AaBbCC, or AaBbCC AaBbCc The parental genotypes can be denoted as A–B–C– (gray) A–bbC– (yellow). The offspring include an albino (cc), and a cream-colored cat (aabb); therefore, both parents must be heterozygous for the A and C genes, and the gray parent should also be heterozygous for the B locus. The cross can be diagrammed as such: AaBbCc AabbCc AbC Abc abC abc ABC AABbCC (gray) AABbCc (gray) AaBbCC (gray) AaBbCc (gray) ABc AABbCc (gray) AABbcc (albino) AaBbCc (gray) AaBbcc (albino) AbC AAbbCC (yellow) AAbbCc (yellow) AabbCC (yellow) AabbCc (yellow) Abc AAbbCc (yellow) AAbbcc (albino) AabbCc (yellow) Aabbcc (albino) aBC AaBbCC (gray) AaBbCc (gray) aaBbCC (black) aaBbCc (black) aBc AaBbCc (gray) AaBbcc (albino) aaBbCc (black) aaBbcc (albino) abC AabbCC (yellow) AabbCc (yellow) aabbCC (cream) aabbCc (cream) abc AabbCc (yellow) Aabbcc (albino) aabbCc (cream) Aabbcc (albino) The phenotypic ratio is 9/32 gray:9/32 yellow:8/32 albino:3/32 black:3/32 cream. Hyde Chapter 5 – Solutions 33 d. The polka-dot trait is dominant in males and recessive in females. This is characteristic of a sex-influenced trait. These traits are controlled by autosomal genes, so the dotted gene locus would be expected to be found on an autosome. Therefore, it will assort independently (Mendel’s second law) from X-linked genes, which have loci on the X chromosome. e. Secondary spermatocytes are “future” sperm. Because secondary spermatocytes are haploid, they would have only one allele per gene. The male has a genotype of AabbCcDd. However, one has to consider the X and Y chromosomes as well. Therefore, the genotype can be designated as AabbCcDdXY. The total number of secondary spermatocytes is 24 = 16 (there are four “heterozygous” positions). The genotypes can be listed systematically as such: AbCDX, AbCDY, AbCdX, AbCdY, AbcDX, AbcDY, AbcdX, AbcdY, abCDX, abCDY, abCdX, abCdY, abcDX, abcDY, abcdX, and abcdY. f. Here again, the cross can be treated as a series of monohybrid crosses. (1) An albino rat has to be cc. The probability is simply 1/4. (2) The genotype of a yellow-colored rat is A–bbC–. The genotype of a polka-dotted male rat is D–XY. Therefore, the overall probability is 1/2 1/2 3/4 3/4 1/2 = 9/128. (3) The genotype of a gray-colored rat is A–B–C–. The dot phenotype depends on the genotype at the D locus and the sex of the offspring. The genotype of a male rat with no dots is ddXY, whereas that of a female can be DdXX or ddXX. We can calculate each of the three possibilities, and then add them all (sum rule) to get to the overall probability. P(gray male with no dots) = 1/2 1/2 3/4 1/4 1/2 = 3/128. P(gray female with no dots) = 1/2 1/2 3/4 3/4 1/2 = 9/128. Therefore, the P(gray rat with no dots) = 3/128 + 9/128 = 12/128 = 3/32. (4) The dot phenotype depends on the genotype at the D locus and the sex of the offspring. The genotypes at the A, B, and C loci are irrelevant. A male rat is polkadotted if he has the genotype D–XY, whereas a polka-dotted female has to be DDXX. We can calculate each of these two probabilities, and then add them up (sum rule) to get to the overall probability. P(polka-dotted rat) = (3/4 1/2) + (1/4 1/2) = 4/8 = 1/2.