PBG 650 Advanced Plant Breeding Module 6: Quantitative Genetics – Environmental variance – Heritability – Covariance among relatives More interactions For an individual G=A+D+I P=A+D+I+E For a population P A D I E Two-locus interactions 2 2 2 2 2 I AA AD DD 2 2 2 2 More than two loci…. • • Interlocus interactions are important, but difficult to quantify Many designs for genetic experiments lump dominance and epistatic interactions into one component called “non-additive” genetic variance Genetic variances from a factorial model Source of Variation A locus main effect Degrees of Freedom 2 Variance Component 2A (locusA) D2 (locusA) A linear 1 2A (locusA) A quadratic (deviations) 1 D2 (locusA) B locus main effect 2A (locusB) D2 (locusB) 2 B linear 1 2A (locusB) B quadratic (deviations) 1 D2 (locusB) A x B interaction I2 4 A linear x B linear 1 2AA A linear x B quadratic A quadratic x B linear 1 1 2A D (pooled) A quadratic x B quadratic 1 2 DD Bernardo, Chapt. 5 Environmental variance • covariance would occur if better P=G+E P G E 2CovGE • 2 2 2 • genotype by environment interactions • differences in relative performance of P = G + E + GE 2 P 2 G 2 E genotypes are given better environments randomization should generally remove this effect from genetic experiments in plants 2 GE • genotypes across environments experimentally, GE is part of E DeLacey et al., 1990 – summary of results from many crops and locations For a particular crop, only 10% 70-20-10 rule of variation in phenotype is due E: GE: G to genotype! Repeatability • Multiple observations on the same individuals – May be repetitions in time or space (e.g. multiple fruit on a plant) P Between Within 2 Within 2 Between 2 2 2 variation among observations on the same individual due to temporary environmental effects 2 ( Es = special environmental variance) variation among individuals due to genetic differences and permanent environmental effects 2 ( Eg= general environmental variance) 2 P 2 ( G Falconer & Mackay, pg 136 2 2 Eg ) Es Repeatability 2 P 2 ( G Repeatability 2 2 Eg ) Es r 2 G 2 Eg P2 • Sets an upper limit on heritabilities 2 • Es is easy to measure 2 • To separate G2 and Eg , you must evaluate repeatability of 2 genetically uniform individuals Eg r 2 2 Eg Es Gain from multiple measurements 2 P(n) 2 (G fyi 2 1 2 Eg ) n Es P( n ) 1 r(n 1) 2 P n 2 • Multiple measurements can increase precision and increase heritability (by reducing environmental and phenotypic variation) • Greatest benefits are obtained for measurements that 2 have low repeatability (large Es ) Heritability • • For an individual: For a population: P=A+D+I+E 2 P 2 A 2 D • Broad sense heritability – degree of genetic determination • Narrow sense heritability – extent to which phenotype is determined by genes transmitted from the parents Falconer & Mackay, Chapter 8 2 I 2 E 2 G 2 G H 2 P h2 2 A 2 P “heritability” Narrow sense heritability – another view Breeding Value h2 = the regression of breeding value on phenotypic value 4,5 4,0 3,5 3,0 2,5 2,0 1,5 1,0 0,5 0,0 h2=0.5 +1 +2 h2=0.3 0 2 4 6 8 10 Phenotypic Value h2 is trait specific, population specific, and greatly influenced by the choice of testing environments Narrow sense heritability h 2 2 A 2 P 2 A 2 2 A D 2 E – Can be applied to individuals in a single environment (generally the case in animal breeding) – In plants, it is commonly expressed on a family (plot) basis, which are often replicated within and across environments Heritability in plants - complications • • • • Different mating systems, including varying degrees of selfing Different ploidy levels Annuals, perennials For many crops, measurement of some traits is only meaningful with competition, in a full stand – variables such as yield are measured on a plot basis – other traits are averages of multiple plants/plot – plot size varies from one experiment to the next • • Replicates are evaluated in different microenvironments Genotype x environment interaction is prevalent for many important crop traits Nyquist, 1991; Holland et al., 2003 Heritability in plants - definition • Fraction of the selection differential that is gained when selection is practiced on a defined reference unit (Hanson, 1963) Selection Differential S=s-0 Selection Response R=1-0 Y=bX R=h2S R=Sbyx R/S=h2=byx • Main purpose for estimating heritability is to make predictions about selection response under varying scenarios, in order to design the optimum selection strategy Applications in plant breeding • Selection in a cross-breeding population • Selection among purelines (with or without subsequent recombination) • Selection among clones • Selection among testcross progeny in a hybrid breeding program • Must specify the unit of selection, the selection method, and unit on which the response is measured Heritability of a genotype mean GXE 2 P H 2 2 G 2 G 2 G 2 GL l 2 GL l h Error variance e2 rl 2 e2 rl broad sense heritability 2 G 2 A 2 GL l e2 rl narrow sense heritability or “heritability” Resemblance between Relatives • Covariance between relatives measures degree of genetic resemblance • Variance among groups = covariance within groups Intraclass correlation of phenotypic values B t 2 2 B W 2 Strategy: • Determine expected covariance among relatives from theory, and compare to experimental observations • Estimate genetic variances and heritabilities Falconer & Mackay, Chapt. 9 Covariance between offspring and one parent Cov(X,Y) fi X iYi μX μY Mean Genotypic Value of Offspring Genotype Frequency Genotypic Value Breeding Value A1A1 p2 2q(-qd) 2q q A1A2 2pq (q-p)+2pqd (q - p) (1/2)(q - p) A2A2 q2 -2p(+pd) -2p -p CovOP=p2*2q(-qd)q+2pq[(q-p)+2pqd](1/2)(q - p) +q2[-2p(+pd)](-p) CovOP = pq2 = (1/2)σA2 This result is true for a single offspring and for the mean of any number of offspring Resemblance between offspring and one parent • For parents and offspring, observations occur in pairs • Regression is more useful than the intraclass correlation as a measure of resemblance – does not depend on the number of offspring – does not require parents and offspring to have the same variance b Cov(O, P) Estimate P 2 A 2 P 1 2 2 phenotypic variance of the parental population Resemblance between offspring and mid-parent CovO,MP = pq2 = (1/2)σA2 MP P 2 b • • Cov(O, MP ) MP 2 2 1 2 A A 1 2 2 P 2 P 1 2 2 2 Regression on mid-parent is twice the regression of offspring on a single parent Number of offspring does not affect the covariance or the regression Resemblance among half-sibs Genotype Breeding Frequency Value Mean Genotypic Value of Offspring Freq. x Value2 A1A1 p2 2q q p2q22 A1A2 2pq (q - p) (1/2)(q - p) (1/2)pq(q - p)22 A2A2 q2 -2p -p p2q22 Covariance of half-sibs = variance among half-sib progeny CovHS = pq2[(1/2)(q - p)2+2pq] = pq2[(1/2)(p+q)2] = (1/2)pq2=(1/4)σA2 t CovHS P 2 A 2 P 1 4 2 Resemblance among full-sibs Progeny Genotype of parents Frequency of mating A1A1 A1A2 A2A2 a d -a A1A1 A1A1 p4 1 A1A1 A1A2 4p3q 1/2 A1A1 A2A2 2p2q2 A1A2 A1A2 4p2q2 A1A2 A2A2 4pq3 A2A2 A2A2 q4 1/4 Mean Value of Progeny a 1/2 (1/2)(a+d) 1 d 1/2 1/4 (1/2)d 1/2 1/2 (1/2)(d-a) 1 -a CovFS= σFS2 = p4a2+4p3q[(1/2)(a+d)]2….+q4(-a)2 - 2 = pq[a+d(q-p)]2 + p2q2d2 Resemblance among full-sibs CovFS= σFS2 = p4a2+4p3q[(1/2)(a+d)]2….+q4(-a)2 - 2 = pq[a+d(q-p)]2 + p2q2d2 G 2 pqa d q - p 4 p q d 2 2 CovFS FS 2 t CovFS P 2 1 2 2 A D 2 1 4 2 A D 2 P 1 2 2 1 4 2 2 2 General formula for covariance of relatives • Unilineal relatives – Resemblance involves only • A AA AAA 2 2 2 etc. Bilineal relatives – Potential exist for relatives to have two common alleles that are identical by descent (X1X3, X1X4, X2X3, or X2X4) A X1X2 B X3X4 Resemblance will also involve: D AD AAD 2 C D X1X3 X1X3 2 2 etc. Covariance due to breeding values A B X (Ai Aj) C D Y (Ak Al) Covα P(Ai Ak )Cov( i , k ) P(Ai Al )Cov( i , l ) P(A j Ak )Cov( j , k ) P(A j Al )Cov( j , l ) 4θ XY σ 2 i 2θ XY σ A2 rσ A2 Covariance due to dominance deviations A B C X (Ai Aj) D Y (Ak Al) Cov P(Ai Ak , A j Al )Cov( ij , kl ) P(Ai Al , A j Ak )Cov( ij , kl ) (θ AC θBD θ ADθBC )σ D2 XY σ D2 General formula for covariance of relatives A B C X D Y Cov r2A D2 r = 2XY = ACBD + ADBC Extended to include epistasis: 2 Cov r2A D2 r 22AA r2AD 2DD ... Adjusting coefficients for inbreeding A 2 Relatives Parent-offspring D 2 r = 2XY 1/2 0 1/4 0 Half-sibs Common parent not inbred Common parent inbred (1+F)/4 Full-sibs Parents not inbred Parents inbred 1/2 1/4 (2+FA+FB)/4 (1+FA)(1+FB)/4