For more than 300 years, thoroughbred horses have been raised for a single purposeto win at the

racetrack. The origin of these horses can be traced to a small group that was imported to England from North Africa and the Middle East in the 1600s. The population of racing horses remained small until the 1800s, when horse racing became increasing popular; today there are approximately half a million thoroughbred horses worldwide.

Although the mathematical methods for analyzing complex characteristics may seem to be imposing at first, most people can intuitively grasp the underlying logic of quantitative genetics. We all recognize family resemblance: we talk about inheriting our fathers height or our mothers intelligence. Family resemblance lies at the heart of the statistical methods used in quantitative genetics. When genes influence a characteristic, related individuals resemble one another more than unrelated individuals. Closely related individuals (such as siblings) should resemble one another more than distantly related individuals (such as cousins). Comparing individuals with different degrees of relatedness, then, provides information about the extent to which genes influence a characteristic.

This chapter is about the genetic analysis of complex characteristics such as racing speed. We begin by considering the differences between quantitative and qualitative characteristics and why the expression of some characteristics varies continuously. Well see how quantitative characteristics are often influenced by many genes, each of which has a small effect on the phenotype. Next, we will examine statistical procedures for describing and analyzing quantitative characteristics. We will consider the question of how much of phenotypic variation can be attributed to genetic and environmental influences and will conclude looking at the effects of selection on quantitative characteristics. Its important to recognize that the methods of quantitative genetics are not designed to identify individual genes and genotypes. Rather, the focus is on statistical predictions based on groups of individuals.

The statistical methods described for use in analyzing quantitative characteristics can be used both to make predictions about the average phenotype expected in offspring and to estimate the overall contribution of genes to variation in the characteristic. These methods do not, however, allow us to identify and determine the influence of individual genes that affect quantitative characteristics. The genes that control polygenic characteristics are referred to as quantitative trait loci (QTLs). Although quantitative genetics has made important contributions to basic biology and to plant and animal breeding, the inability to identify QTLs and measure their individual effects has severely limited the application of quantitative genetic methods.

In recent years, numerous genetic markers have been identified and mapped with the use of recombinant DNA techniques, making it possible to identify QTLs by linkage analysis. The underlying idea is simple: if the inheritance of a genetic marker is associated consistently with the inheritance of a particular characteristic (such as increased height), then that marker must be linked to a QTL that affects height. The key is to have enough genetic markers so that QTLs can be detected throughout the genome. With the introduction of restriction fragment length polymorphisms and microsatellite variations, variable markers are now available for mapping QTLs in a number of different organisms.

The number of genes affecting a quantitative characteristic can be estimated by locating QTLs with genetic markers and adding up the number of QTLs detected. This method will always be an underestimate, because QTLs that are located close together on the same chromosome will be counted together, and those with small effects are likely to be missed.