Parental investment and family dynamics: interactions between theory and empirical tests

The pattern of parental investment (PI) seen in nature is a product of the simultaneous resolution of conflicts of interest between the members of a family. How these conflicts are resolved depends upon the mating system, the genetic mechanism, on whether extra PI affects current or future offspring, and the behavioural mechanisms underlying supply and demand of PI. Until recently very little empirical work has been done to underpin these key determinants of conflict resolution. This review examines recent empirical progress in understanding both (1) how conflict is resolved and (2) its evolutionary consequences. How offspring demand interacts with parental supply of resources determines how conflict is resolved. Two extremes are: passive parental choice of competing offspring, relating to offspring control of resource allocation, and active parental choice relating to parental control. Although most previous empirical work has tended to conclude or assume that parents primarily control resource allocation decisions, recent studies explicitly examining predictions from theoretical analyses have shown that offspring control of resource allocation is more important than previously realised. The amount of PI supplied at resolution depends not on who controls food allocation, however, but on the nature of the supply and demand mechanisms. These have yet to be established experimentally, but a recent regression model illustrates how this could be achieved in the field. Determination of the effect of supply on demand (ESD) and the effect of demand on supply (EDS) mechanisms is critical to parent-offspring conflict theory, which has not been adequately tested empirically. There is an underlying, and until recently untested, assumption of models of intrafamilial conflict that there is genetic variation for both offspring demand and parental supply behaviours, so that the behaviours can coevolve. Recent studies on great tits, burrower bugs and mice all found evidence for genetic variation in supply and demand behaviours, but the predicted negative correlation between genes expressed in mothers and their offspring (i.e. parent-offspring coevolution), was found only for burrower bugs. The lack of a negative relationship for great tits and mice may have been a consequence of antagonistic coevolution between the sexes (sexual conflict). These studies illustrate the importance of the underlying genetics and mating system in determining conflict resolution, and point to the need for new models (especially of interbrood competition) taking differences in the genetics and the co-evolution of the ESD and EDS mechanisms into account. We also discuss the importance of the comparative approach in determining evolutionary consequences of conflicts, and use the recent work on growth costs of begging to illustrate the difficulties of measuring costs of conflict in an evolutionary currency. The recent growth in empirical work on conflicts in families illustrates an increasing, and increasingly productive, integration between theoreticians and empiricists.