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Trihybrid cross.
Tamarin: Principles of Genetics, Seventh Edition
II. Mendelism and the Chromosomal Theory
2. Mendel s Principles
The McGraw Hill Companies, 2001
Two Mendel s Principles
First white variety AAbb
Second white variety aaBB
Purple AaBb Self
F2 Rose Pea
Ab AABb
aB AaBB
ab AaBb
AABB
AABb
AAbb
AaBb
Aabb
AaBB
AaBb
aaBB
aaBb
AaBb
Aabb
aaBb
aabb
Walnut Figure 2.20
Single
Four types of combs in fowl.
F2 Summary Figure 2.22
Purple : White 9 : 7
Color production in corn.
Rose comb RRpp
Pea comb rrPP
is dominant to a for black. The albino gene (ca), when homozygous, is epistatic to the A gene; the A gene is hypostatic to the gene for albinism.
Walnut comb RrPp F1 F1
Mechanism of Epistasis
In this case, the physiological mechanism of epistasis is known.The pigment melanin is present in both the black and agouti phenotypes. The agouti is a modi ed black hair in which yellow stripes (the pigment phaeomelanin) have been added. Thus, with melanin present, agouti is dominant. Without melanin, we get an albino regardless of the genotype of the agouti gene because both agouti and black depend on melanin. Albinism is the result of one of several defects in the enzymatic pathway for the synthesis of melanin ( g. 2.24). Knowing that epistatic modi cations of the 9:3:3:1 ratio come about through gene interactions at the biochemical level, we can look for a biochemical explanation for the 9:7 ratio in corn kernel color ( g. 2.22). Two possible mechanisms for a 9:7 ratio are shown in gure 2.25. Either a two-step process takes a precursor molecule and turns it into purple pigment, or two precursors that must be converted to nal products then combine to produce purple pigment. The dominant alleles from the two genes control the two steps in the process. Recessive alleles are ineffective. Thus, dominants are necessary for both steps to complete the pathways for a purple pig-
RP RP RRPP Walnut RRPp Walnut RrPP Walnut RrPp Walnut
Rp RRPp Walnut RRpp Rose RrPp Walnut Rrpp Rose
rP RrPP Walnut RrPp Walnut rrPP Pea rrPp Pea
rp RrPp Walnut Rrpp Rose rrPp Pea rrpp Single
F2 Summary Figure 2.21
Walnut : Rose : Pea : Single 9 : 3 : 3 : 1
Independent assortment in the determination of comb type in fowl.
Tamarin: Principles of Genetics, Seventh Edition
II. Mendelism and the Chromosomal Theory
2. Mendel s Principles
The McGraw Hill Companies, 2001
Genotypic Interactions
Black
Albino
aaCC
Black hair
AAc ac a
White hair
Agouti
AaCc a
Black hair with yellow stripes
Ac a AACca
aC AaCC
ac a AaCca
AACC AC
Agouti
Agouti
Agouti
Agouti
AACca Ac a
Agouti
AAcaca
AaCca
Aacaca
Albino
Agouti
Albino
AaCC aC
Agouti
AaCca
aaCC
aaCca
Agouti
Black
Black
AaCca ac a
Agouti
Aacaca
aaCca
aacaca
Albino
Black
Albino
Agouti : Black : Albino 9:3:4 Figure 2.23
Epistasis in the coat color of mice.
ment. Stopping the process at any point prevents the production of purple color. Another example of epistasis occurs in the snapdragon (Antirrhinum majus).There, a gene called nivea has alleles that determine whether any pigment is produced; the nn genotype prevents pigment production, whereas the NN or Nn genotypes permit pigment color genes to express themselves. The eosinea gene controls the production of a red anthocyanin pigment. In the
presence of the N allele of the nivea gene, the genotypes EE or Ee of the eosinea gene produce red owers; the ee genotype produces pink owers. When dihybrids are self-fertilized, red-, pink-, and white- owered plants are produced in a ratio of 9:3:4 ( g. 2.26). The epistatic interaction is the nn genotype masking the expression of alleles at the eosinea gene. In other words, regardless of the genotypes of the eosinea gene (EE, Ee, or ee), the owers will be white if the nivea gene has the nn
Tamarin: Principles of Genetics, Seventh Edition
II. Mendelism and the Chromosomal Theory
2. Mendel s Principles
The McGraw Hill Companies, 2001
Two Mendel s Principles
Phenylalanine
CH2 1 OH
CH NH+
COO
2 Tyrosine 3
Thyroxine P-Hydroxyphenyl pyruvate Homogentisic acid 4
CH NH
COO
OH OH 3,4-Dihydroxyphenylalanine (DOPA) CO2 + H2O Melanins 5 Enzyme defect conditions 1 Phenylketonuria (PKU) CH2 CH NH+
COO
2 Genetic goitrous cretinism 3 Tyrosinosis 4 Alkaptonuria 5 Albinism
In humans, errors in melanin synthesis produce different physical conditions and diseases, depending on which part of the tyrosine (an amino acid) metabolic pathway is disrupted. The broken arrows indicate that there is more than one step in the pathways; the conditions listed occur only in homozygous recessives.
Pathway 1 Colorless precursor
Control by gene A Colorless intermediate
Control by gene B Purple pigment
Pathway 2 Colorless precursor 1 Control by gene A A B Colorless precursor 2 Control by gene B + Purple pigment
Possible metabolic pathways of color production that would yield 9:7 ratios in the F2 generation of a self-fertilized dihybrid.
Tamarin: Principles of Genetics, Seventh Edition
II. Mendelism and the Chromosomal Theory
2. Mendel s Principles
The McGraw Hill Companies, 2001
Biochemical Genetics
combination of alleles. Thus, nivea is epistatic to eosinea, and eosinea is hypostatic to nivea. (We should add that at least seven major colors occur in snapdragons, along with subtle shade differences, all genetically controlled by the interactions of at least seven genes.) Other types of epistatic interactions occur in other organisms. Table 2.4 lists several. We do not know the exact physiological mechanisms in many cases, especially when developmental processes are involved (e.g., size and shape). However, from an analysis of crosses, we can know the number of genes involved and the general nature of their interactions.
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