Abstract
This article starts by reviewing the differences between various measures of fitness, for example, individual fitness, absolute fitness, relative fitness and geometric mean fitness. Differences in fitness (when measured appropriately) can be used to derive selection equations, which show how natural selection changes the genetic composition of a population through time. Quantitative geneticists derived the secondary theorem of natural selection, which shows how selection on fitness will change other, genetically correlated traits. Fitness landscape models, whether continuous or discrete, can be used to analyse how natural selection will drive a population to the top of a fitness peak. Evolutionary geneticists are currently pursuing several empirical approaches to the study of fitness, including direct fitness assays, microbial experimental evolution and the use of DNA sequence data to infer a history of positive natural selection. The concluding section sketches several major unresolved problems in the experimental study of fitness.