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Linkage (allelic interactions) & Epistasis (non-allelic interactions) F1 TtRr TtRr Ttrr Tall/Resist Tall/Susc x Recessive ttrr ttRr Short/Resist ttrr Short/Susc Test Cross Phenotype Genotype Frequency Tall and Resistant TtRr 79 Short and Resistant ttRr 18 Tall and Susceptible Ttrr 22 Short and Susceptible ttrr 81 Linkage Recombination Number of Recombinants Number of Observations 18+22 = 0.2 200 F1 TtRr Self TR 0.4 Tr 0.1 tR 0.1 tr 0.4 Linkage Gametes from female parent Gametes from male parent TR-0.4 Tr-0.1 tR-0.1 tr-0.4 TR – 0.4 TTRR 0.16 TTRr 0.04 TtRR 0.04 TtRr 0.16 Tr – 0.1 TTRr 0.04 TtRR 0.04 TtRr 0.16 TTrr 0.01 TtRr 0.01 Ttrr 0.04 TtRr 0.01 ttRR 0.01 ttRr 0.04 Ttrr 0.04 ttRr 0.04 ttrr 0.16 tR – 0.1 tr – 0.4 16 TTRR:8 TTRr:1 TTrr:8 TtRR:34 TtRr:8 Ttrr:1 ttRR:8 ttRr:16 ttrr Linkage F2 Genotype 50% Recom. 20% Recom. TTRR TTRr TTrr TtRR TtRr Ttrr ttRR ttRr ttrr 6 13 6 12 26 12 6 13 6 16 8 1 8 34 8 1 8 16 Linkage F2 Phenotype 50% Rec. 20% Rec. T_R_ T_rr ttR_ ttrr 56 19 19 6 66 9 9 16 Linkage Independent Assortment ¼ AB Coupling Linkage > ¼ AB Repulsion Linkage < ¼ AB ¼ Ab < ¼ Ab > ¼ Ab ¼ Ab < ¼ aB > ¼ aB ¼ ab > ¼ ab < ¼ ab F2 TTRR TTRr TTrr TtRR TtRr Ttrr ttRR ttRr ttrr 160 80 10 80 340 80 10 80 160 Total F3 F2 TTRR TTRr TTrr TtRR TtRr Ttrr ttRR ttRr ttrr 160 80 10 80 340 80 10 80 160 TTRR 160 20 - 20 54 - - - - 254 TTRr - 40 - - 27 - - - - 67 TTrr - 20 10 - 4 20 - - - 54 TtRR - - - 40 27 - - - - 67 TtRr - - - - 116 - - - - 116 Ttrr - - - - 27 40 - - - 67 ttRR - - - 20 4 - 10 20 - 54 ttRr - - - - 27 - - 40 - 67 Ttrr - - - - 54 20 - 20 160 254 Total Linkage F3 Genotype TTRR TTRr TTrr TtRR TtRr Ttrr ttRR ttRr ttrr 50% Rec. 141 94 141 94 62 94 141 94 141 20% Rec. 254 67 54 67 116 67 54 67 254 Linkage F3 Phenotype 50% Rec. 20% Rec. T_R_ T_rr ttR_ ttrr 391 234 234 141 504 121 121 254 Undesirable Linkage Increased selfing or crossing before selection Three gene test cross AABBCC x aabbcc AaBbCc 1 2 3 Three gene test cross AABBCC x aabbcc AaBbCc x aabbcc 1 2 3 Three gene test cross Phenotype Observed Expected A_B_C_ aabbcc aaB_C_ A_bbcc A_bbC_ aaB_cc A_B_c aabbC_ Total 500 510 50 55 35 38 4 3 1195 149.4 149.4 149.4 149.4 149.4 149.4 149.4 149.4 1195 Three gene test cross Acb aCB ACb acB AcB aCb A C B a c b A C B a c b A C B a c b 55 50 35 38 4 3 Three gene test cross A – B locus A_B_ 500+4 = 504 aabb 510+3 = 513 aaB_ 50 + 38 = 88 A_bb 55 + 35 = 90 Parent types Aa Cc Bb Recombinants % Recombination = 178/1195 x 100 = 14.9% Three gene test cross A – C locus A_C_ 500+35 = 535 Aacc 510+38 = 548 aaC_ 50 + 3 = 53 A_cc 55 + 4 = 59 Parent types Aa Cc Bb Recombinants % Recombination = 112/1195 x 100 = 9.4% Three gene test cross B - C locus B_C_ 500+50 = 550 Bbcc 510+55 = 565 bbC_ 35 + 3 = 38 B_cc 38 + 4 = 42 Parent types Aa Cc Bb Recombinants % Recombination = 80/1195 x 100 = 6.7% Three gene test cross A A 14.9 m 9.4 m C B 6.7 m B Three gene test cross A - B locus Aa Cc Bb NonRecombinant types A_B_C_ 500 aabbcc 510 Recombination events aaB_ 50+39 = 88 A_bb 55+35 = 90 Double cross overs 2 x (3+4) = 14 % Recombination = 192/1195 x 100 = 16.1% Three gene test cross A A 16.1 m 9.4 m C B 6.7 m cM = [-Ln(1-2R)] x 50 B Three gene test cross A A 16.1 m 9.4 m C B 6.7 m B cM = [-Ln(1-2R)] x 50 A A 17.7 cM 10.5 cM C s.e. = [R/(1-R)]/n B 7.0 cM B Epistasis Non-allelic Interactions Epistasis Parent 1 White flower x Parent 2 White flower F1- 100% Blue Flower Hairbells Epistasis Parent 1 White flower Parent 2 White flower x F1- 100% Blue Flower 9 Blue : 7 White Epistasis Parent 1 White flower w1w1W2W2 Parent 2 White flower W1W1w2w2 x F1- 100% Blue Flower W1w1W2w2 Blue flower W1_W2_ 9 Blue : 7 White White flower 3-W1_w2w2 3-w1w1W2_ 1-w1w1w2w2 Epistasis Precursor Enzyme Gene W_ substrate Enzyme Gene M_ Blue Epistasis Parent 1 White flower x Parent 2 Magenta flower F1- 100% Blue Flower Blue Eyed Mary (Collinsia parviflora) Epistasis Parent 1 White flower Parent 2 Magenta flower x F1- 100% Blue Flower 9 Blue : 3 Magenta : 4 White Epistasis Colorless Magenta Gene W_ Blue Gene M_ Epistasis Colorless Magenta Gene W_ Parent 1 White flower wwMM Blue Gene M_ x Parent 2 Magnolia flower WWmm Epistasis Parent 1 White flower wwMM Parent 2 Magnolia flower WWmm x F1- 100% Blue Flower WwMm 9 Blue W_M_ : 3 Magnolia W_mm : 4 White wwM_ & wwmm F2 phenotype A_B_ A_bb aaB_ aabb Genetic explanation 9 3 3 1 Complete dominance: A dominant to a, B dominant to b. 9 3 4 0 Recessive epistasis: aa epistatic to B and b. 12 0 3 1 Dominant epistasis: A epistatic to B or b. 0 Dominant & recessive epistasis: A epistatic to B and b; bb epistatic to A and a; A_ and bb identical phenotypes 0 Duplicate recessive epistasis: aa epistasis to B and b; bb epistatic to A and aa 1 Duplicate dominant epistasis: A epistasis to B and bb; B epistasis to A and aa. 13 9 15 0 0 0 3 7 0 Autotetraploid Segregation & Chi-square