SELECTION: ESTIMATED BREEDING VALUE Introduction The goal of animal breeder is rapid genetic improvement in traits of economic importance. How can this goal be achieved? In order to achieve this goal, accurate predictions (or evaluations) of animals’ genetic merit for any given traits are required. This information is used by the breeder to rank the animals and cull those with the best evaluations (Van Vleck et al, 1987). Unfortunately, quantitative traits are also influenced by the environment. It is therefore important for a breeder to understand and adjust (remove) the influence of these environmental factors before predicting an animal’s genetic worth for a given trait. Adjusting for environmental factors enables breeders to compare animals of different production levels. Objectives 1. Define the terms breeding value, transmitting ability and producing ability 2. State the difference between producing ability and transmitting ability 3. Discuss the importance of environmental factors in genetic evaluations Estimated Breeding Value Predictions of additive genetic breeding value and future performance are used so frequently in animal breeding that they have been given specific names. The additive genetic value of an animal for a specific trait is referred to as its breeding value (BV) for that trait. The BV estimated from various measures (records) of performance is called estimated breeding value (EBV). The term BV is used to denote additive genetic value because it is mostly the additive values of the individual genes possessed by an animal which are transmitted directly to its offspring. An animal’s additive genetic value, therefore, reflects its value or worth for breeding. Estimated Transmitting Ability In terms of the model of gene action, an animal’s BV for a trait is the sum of the additive effects of the genes it possesses which affect that trait. However, since an animal will transmit half of its genes, its EBV actually represents twice its contributions to any of his offspring. EBV is, therefore, often converted to estimated transmitted ability (ETA). It is simply half of an individual’s EBV. ETAs are referred to by specific names in livestock industries. ETAs for sires computed from the average performance of their progeny are called Predicted Differences (PD) or Predicted Transmitting Ability (PTA) in dairy cattle and Estimated Progeny Differences (EPD) in beef cattle. The term Cow Index is applied to the ETA of a dairy cow, and is computed from its own performance and the average performance of its paternal half-sisters (i.e. the PTA of her sire). A Pedigree Index is an ETA computed for an individual from the performance of his ancestors. ETA of beef sires (EPD) describes particularly well the interpretation of ETA. The word “progeny” indicates that differences among individuals (sire in this case) in ETA will be reflected in the performance of their progeny. ETA measures that part of genetic superiority or inferiority which is transmitted to the progeny. If a group of sires is mated to random samples of females (neither particularly good nor bad) the relative performance of their progeny (if they are managed equally) should correspond to their ETAs. The word difference means that the ETAs are expressed as a difference from some base or “zero” point. The base point represents the population average ETA at some point in time. The choice of a base point or ‘genetic base’ is arbitrary (and may, in fact, be changed) so long as all ETAs are expressed relative to the same point. The practical result of expressing ETAs relative to some average ETA defined as zero is that ETAs maybe positive or negative. Those exceeding the base point are positive, while ETAs less than the base point are negative. The term EPD indicates what the breeder really wants to know – expected performance of an individual’s progeny relative to the progeny of other individuals. Activity: What is EBV and why is it important in livestock improvement? Producing Ability Producing ability is the performance potential of an individual for a repeated trait. A repeated trait is a trait which individuals commonly have more than one performance record. Prediction of future records works in the same way as EBV. Here the breeder is interested in the future performance of an individual in some repeated trait sometimes referred to as true producing ability, of the individual. This gives the best estimate of future performance. The producing ability of an animal is the sum of its total genetic value (G = A+D+I) and any environmental effects which are permanent to all records (PE). Producing ability would therefore indicate exactly the next record, except for the unpredicted temporary environmental effects (TE) which will be associated with that record. Estimated producing abilities, like ETAs, have different specific names in different livestock species. In beef cattle, pigs, and sheep, the term used is Most Probable Producing Ability (MPPA). In dairy cattle, the most commonly used terms are Estimated Producing Ability (EPA) and Estimated Relative Producing Ability (ERPA). Since MPPAs, EPAs and ERPAs are most often used to compare producing abilities of individuals in the same herd; they are generally expressed relative to the herd mean. EPAs are therefore positive (above) or negative (below) relative herd average, and it is necessary to add average EPA to obtain the absolute level of the predicted future record. Producing ability and transmitting ability are clearly related. Producing ability measures A+D+I+PE, while transmitting ability includes only A, and the two measures have relatively different base points. However, individuals having the highest estimated transmitting abilities in a herd often will have the highest estimated producing abilities as well. In the few cases where these measures do not agree, the breeder may use EPA to decide on the fate of the individual, and the Eta to determine the disposition of the individual’s offspring PREDICTION OF BREEDING VALUES Introduction With the understanding that breeding values are needed by animal breeders in order to make accurate selection and culling decisions. Breeders rank animals based on their estimated breeding values, select those with highest estimates as parents of the next generation and cull those with low breeding values. Without the knowledge of breeding value estimates, selection will not be possible. If you are a livestock breeder, you can use this knowledge to improve either one trait at a time (single trait selection) or more than one trait at a time (multiple trait selection) depending on your selection goals. Depending on the amount and source of information available, proper weighting may depend on the heritability (h2) and repeatability (r) of the specific trait. The number of repeated records (n), the genetic relationship (aij) between a relative and the individual, the number of relatives (m) of a specific type and any environmental correlation (C2) due to extra common environmental factors shared by relatives. Table 1: Factors considered in weighting performance records to estimate breeding value and producing ability Observed Performance Individual Estimated Breeding Value Estimating Producing Ability One record n record Single Relatives h2 h2, r, n r r, n One record n records m Relatives h2,aij h2, aij, r,n One record each n records each h2, aij, m, C2 h2, aij, n, r, m, C2 Objectives 1. To estimate an individual’s breeding value from its own records or performance 2. To predict an individual’s breeding value from performance of relatives (progeny, parents, halfsibs) Estimating Breeding Values from individual Performance (own records) Individual performance (one record) Individual performance is simply an animal’s own record. For example, if a dairy cow produces 7000kgs of milk, the yield 7000kgs is the cow’s individual performance. In terms of breeding value estimation there is need to know how much of the yield is due to genetics. In order to do this a heritability value of the trait need to be known. Heritability represents the proportion of differences in performance which is due to differences in the additive effects of individual genes, or due to additive genetic variance. Heritability also represents the proportion of differences in performance which is due to differences in the breeding values of individuals. In practice the heritability has been estimated from performance records which have been adjusted for major environmental sources of variation (age, cow weight, feeding regime, lactation length etc) and expressed as deviations from the average performance in some environmental conditions. It also indicates the proportion of differences in adjusted records which is due to differences in breeding value. An estimate of the individual‘s breeding value is given by; EBV = αΊ€ = h2 (P - µ) =h2 (Individual record – herd average) Where h2 = COV (a,p)/σp2 = σA2/σp2 and indicates the average (for the trait, in the population) number of units change in breeding value per unit change in performance. The denominator and the numerator are the genetic variance and the phenotypic variance respectively. The accuracy with which EBV predicts true value depends on the magnitude of heritability and the definition of ‘accuracy’. Accuracy of EBV is the square root of heritability (h). Thus for a single record, the accuracy (r (a, y)) is given by; πΆππ£ (π,π¦) πΏππΏπ¦ = πΏ2π πΏππΏπ¦ πΏπ = πΏπ¦ = 2 √β = h EXAMPLE: Estimating breeding value from an animal’s own record Question: A Mashona bull at the University farm gains 1.4kg/day on a feeding trial in which average gain is 1.0kg/day. All animals are fed and managed identically, and gains are adjusted for beginning age and weight. If heritability of weight gain is 0.45, what is the animal’s EBV, and the accuracy of that estimate? Solution: Relative to the performance of other individuals under similar conditions, gain of the individual is; (P - µ) = (1.4 – 1.0) = 0.4 kgs/day Therefore EBV = αΊ€ The accuracy of this EBV = h2 (P - µ) = 0.45 (0.4) = 0.18 kgs/day = √h2 = √ (0.45) = 0.67 (67%) Individual performance: n records An estimate of breeding value based on average performance of several records, logically will be more accurate than EBV from a single record. Therefore when several records are available, all should be used to estimate breeding value. The adjusted average performance is weighted by the following coefficient which can be thought of as heritability of average performance or more precisely the regression of breeding value on average performance in n records. The b coefficient is; b = πβ2 1+(π−1)π where ; b is the weight h2 is the heritability n is the number of records in the average performance P r is the repeatability of the trait This b coefficient, which can be thought of as heritability of average performance, is used in estimating breeding value from average performance (mean of n records) in exactly the same way heritability is used with single records. Specifically, EBV = αΊ€ = πβ2 (1+(π−1)π ) (αΉ-µ) In this case P is the average of the n records and it is adjusted for the environmental factors. The accuracy of EBV from the average performance is the square root of the b coefficient. Note that when n = 1, the b coefficient becomes the heritability as shown in the previous case. From the above formulas, one can also conclude that as the number of records increases the accuracy of the estimated breeding values also increases. Table 2: Weights (b coefficients) and accuracy of estimating breeding value (rAP) from average performance 2 h 0.10 0.25 0.35 0.50 n 2 3 4 5 10 2 3 4 5 10 2 3 4 5 10 2 3 4 5 10 b .15 .19 .21 .23 .27 .38 .47 .53 .57 .68 0.30 rA,P .39 .43 .46 .48 .52 .62 .68 .73 .75 .82 b .13 .15 .16 .17 .18 .33 .38 .40 .42 .45 .47 .53 .56 .58 .64 .67 .75 .80 .83 .91 REPEATABILTY 0.50 rA,P .37 .39 .40 .41 .43 .58 .61 .63 .65 .67 .68 .72 .75 .76 .80 .82 .87 .89 .91 .95 NB* h2 = heritability, n = number of records in the average b .12 .13 .14 .14 .15 .30 .33 .34 .35 .36 .42 .46 .47 .49 .51 .61 .65 .68 .69 .73 0.55 rA,P .35 .36 .37 .37 .38 .55 .57 .58 .59 .60 .65 .68 .69 .70 .71 .78 .81 .82 .83 .85