Chapter 10 – Factors Affecting the Rate of Genetic Change I. Elements of the Key Equation for Genetic Change II. The Key Equation in More Precise Terms III. Trade-Offs Among Elements of the Key Equation IV. Comparing Selection Strategies Using the Key Equation V. The Key Equation in Perspective VI. Male Versus Female Selection Chapter 10 – Factors Affecting the Rate of Genetic Change Learning Objective: To understand the factors (heritability, intensity of selection, phenotypic variation, and generation interval) that affect the predicted amount of genetic progress, and how certain factors can be manipulated by the breeder. I. Elements of the Key Equation for Genetic Change The Key Equation: Four Factors Accuracy of Selection – A measure of the strength of the relationship between true breeding values and their predictions for a trait under selection (largely a function of heritability). Selection Intensity – A measure of how “choosy” breeders are in deciding which individuals are selected. Genetic Variation – Variability of breeding values within a population for a trait under selection. This is the key to genetic progress. Generation Interval – The average amount of time required to replace one generation with the next. II. The Key Equation in More Precise Terms ^ i σ rBV,BV BV ∆BV/t = ________ L where ∆BV/t = predicted rate of genetic change per unit of time ^ rBV,BV = accuracy of selection i = selection intensity σBV = genetic variation L = generation interval ∆BV/t = ^ iσ rBV,BV BV ________ L where ∆BV/t = The predicted change in mean breeding value of a population due to selection. Only BV is passed on from selected parents to offspring. ^ rBV,BV = The strength of the relationship between true breeding values and their predictions for a trait under selection. Accuracy ranges from 0 (no information) to 1 (lots of information). i = selection intensity (phenotypic difference between the mean selection criterion of those individuals selected to be parents and the population mean = the “Standardized Selection Differential”). σBV = genetic variation (the standard deviation of breeding values). L = generation interval (the average age of parents when their selected offspring are born). L = generation interval (the average age of parents when their selected offspring are born). Generally, the larger the size of the species, the longer the generation interval. TABLE 10.2 Common generation intervals (modified) Species Horses Dairy cattle Beef cattle Sheep/Goats Swine Chickens/Rabbits Generation Interval (years) 8 to 12 4 to 6 4 to 6 3 to 5 1.5 to 2 1 to 1.5 Source: http://www.lfba.lv/en/industry/history III. Trade-Offs Among Elements of the Key Equation ∆BV/t = ^ iσ rBV,BV BV ________ L Accuracy vs. Generation Interval – To wait for more progeny records to enhance accuracy would increase the generation interval. A strategy is to choose the better young sires with the highest accuracies and use for only one year. Accuracy vs. Intensity – Likewise, to wait for more progeny records to enhance accuracy for only a few sires would decrease the intensity of selection (i.e., the percentage selected). One solution is to test more sires that have fewer progeny records. Intensity vs. Generation Interval – Being too choosy usually results in a longer generation interval. However, the real trade-off is between selecting females vs. males (e.g., a lower replacement rate dictates that animals remain longer in the population). Intensity vs. Risk – A problem may exist if very few sires are used that have low accuracies, whereby their breeding values turn out to be poorer than what was initially reported. Truncation selection – Only need to know the proportion (p) of offspring saved as replacements. Truncation selection – Only need to know the proportion (p) of offspring saved as replacement. ___ __ __ _ SCS - SC i = _______ σSC Mathematical Proof of the Key Equation: Given the following parameters for a sheep population being selected for yearling weight (YW, lbs): μ = 130 Mean of selected males = 190 and females = 158 lbs h2 = 0.36 σP = 30 lbs L = 3 years 1) R = h2 x SD = 0.36 x (174 – 130) R = 15.84 lbs _ 2) R = h2 i σP = 0.36 x 1.465 x 30 {SD (Selection Differential) = 44 lbs _ {i = “Standardized Selection Differential” _ = i = SD/σP ; 44/30 = 1.465 R = 15.84 lbs 3) ˆ _ rBV, BV i σBV ∆BV/t = __________ L { = h2 x σ2P = σ2BV = 0.36 x 302 = 324 (√ 324 = 18) _ i σBV = √.36 x 1.465 x 18 ______________ ______________________ = √ h2 L 3 = 15.84 ______ 3 = 5.28 lbs/yr IV. Comparing Selection Strategies Using the Key Equation Mathematical proof of i : = “Standardized Selection Differential” Determine the intensity of selection values (i) for each sex. Top 1% (p) of males; i = 2.67 Top 10% (p) of females; i = 1.76 Mean of i = (2.67+1.76)/2 = 2.215 Mean of selected males: μ + iσ 5.0 + 2.67(√4.0) = 10.34 kg Mean of selected females: μ + iσ 5.0 + 1.76(√4.0) = 8.52 kg __ Mean of selected offspring : SCS __ SCS = 10.34 + 8.52 = 9.43 kg 2 ___ __ __ _ SCS - SC i = _______ = 9.43 – 5.0 = 2.215 σSC √4 i = “Standardized Selection Differential” IV. Comparing Selection Strategies Using the Key Equation {∆BV = R, genetic “response” to selection} Mean of i = (2.67+1.76)/2 = 2.215 ∆BV = = New mean = = √.35 x 2.215 x √ 1.4 1.55 kg 5.0 + 1.55 6.55 kg R/t = 1.55/1.5 = 1.034 kg/year {σ2BV = h2 x σ2p} = .35 x 4 = 1.4 σBV = √ 1.4 = 1.1832 IV. Comparing Selection Strategies Using the Key Equation Problem 10.1: Calculate the rate of genetic change in feed conversion in a swine population given the following: Heritability of feed conversion (h2) = .35 Phenotypic standard deviation (σp) = .2 lb/lb Accuracy of male selection (rBVm,BVm) = .8 Accuracy of female selection (rBVf,BVf) = .5 Intensity of male selection (im) = -2.4 Intensity of female selection (if) = -1.5 Generation interval for males (lm) = 1.8 years Generation interval for females (lf) = 1.8 years IV. Comparing Selection Strategies Using the Key Equation (Problem 10.1): ∆BV/t = ^ iσ rBV,BV BV _________ L ^ i +r ^ i)σ (rBVm,BV m BV ,BV BV m f f f = _______________________ Lm + Lf = (.8(-2.4) + .5(-1.5))(.118) = -.088 lb/lb/year 1.8 + 1.8 IV. Comparing Selection Strategies Using the Key Equation Problem 10.6: A rancher runs a closed herd of breeding cattle. He normally keeps and breeds the top 3% of his bull calves based on individual performance for yearling weight (YW). His sires average three years of age when their offspring are born. He is studying two female replacement strategies: a. Saving the top 20% of his heifers based on YW (Lf = 6.2 years) Saving the top 60% based on YW (Lf = 3.2 years; cull more older cows) If h2YW = .5, and σPYW = 60 lb, calculate the expected annual rate of genetic change in YW for each strategy. Strategy 1: ^ i +r ^ (rBVm,BV BVf,BVf if) σBV m m ∆BV/t = _______________________ = (.7071(2.27) + .7071(1.4))(42.4264) = 11.97 lb/yr Lm + Lf 3.0 + 6.2 b. What elements of the key equation is the rancher experimenting with? c. What element appears to be more important? IV. Comparing Selection Strategies Using the Key Equation A rancher runs a closed herd of breeding cattle. He normally keeps and breeds the top 3% of his bull calves based on individual performance for yearling weight (YW). His sires average three years of age when their offspring are born. He is studying two female replacement strategies: a. Saving the top 20% of his heifers based on YW (Lf = 6.2 years) Saving the top 60% based on YW (Lf = 3.2 years ; cull more older cows) If h2YW = .5, and σPYW = 60 lb, calculate the expected rate of genetic change in YW for each strategy. Strategy 2: ^ i +r ^ (rBVm,BV BVf,BVf if) σBV m m ∆BV/t = _______________________ = (.7071(2.27) + .7071(.64))(42.4264) = 14.08 lb/yr Lm + Lf 3.0 + 3.2 b. What elements of the key equation is the rancher experimenting with? if and Lf c. What element appears to be more important? The i affects L (i.e., by culling more cows that are aged or less productive, if increases, which potentially can translate into more genetic progress).