Cookies on this website

We use cookies to ensure that we give you the best experience on our website. If you click 'Accept all cookies' we'll assume that you are happy to receive all cookies and you won't see this message again. If you click 'Reject all non-essential cookies' only necessary cookies providing core functionality such as security, network management, and accessibility will be enabled. Click 'Find out more' for information on how to change your cookie settings.

Price׳s extended covariance selection mathematics is applied to class-structured populations with additional assumptions, to derive the 'genetic Price Equation with class structure'. Each individual belongs to a class, and there may be overlapping generations; the equation is genetic because the trait is restricted to an arbitrary weighted sum of allele frequencies. Two special cases are then considered, a demography-like case corresponding to Fisher's fundamental theorem of natural selection, and a sex-ratio-like case corresponding to Fisher's sex ratio argument: these differ in whether it is natural to assume that the per-capita or the total reproductive values of each class are maintained from the parental to the descendant population. These cases also match the two existing attempts to eliminate from the effects of natural selection those passive changes in allele frequencies that are caused by the class structure, and suggest improvements in one of them. In each case a more specialised Price Equation, and a 'fundamental theorem of natural selection', are proved, which hold out of class-structure equilibrium, showing that passive changes can be eliminated in more than one way, and hinting at the possibility of a more general formulation. Previous class-structured Price Equations and a 'fundamental theorem' are linked to these results. The power of Price's formal approach is vividly illustrated by this lucid conspectus of otherwise self-standing theories with confusing interconnections.

Original publication




Journal article


J Theor Biol

Publication Date





62 - 72


Fundamental theorem of natural selection, Natural selection, Overlapping generations, Reproductive value, Animals, Biological Evolution, Gene Frequency, Genetic Variation, Models, Genetic, Reproduction, Selection, Genetic, Sex Ratio