How depressed? Estimates of inbreeding effects during seed development depend on reproductive conditions

Inbreeding depression can reduce the performance of offspring produced by mating between relatives, with consequences for population dynamics and sexual-system evolution. In flowering plants, inbreeding depression commonly acts most intensely during seed development. This predispersal component is typically estimated by comparing seed production following exclusive self- and cross-pollination, but such estimates are unbiased only if seed production is limited by ovule availability, rather than by pollen receipt or seed-development resources. To overcome this problem, we propose experimental and statistical methods based on a model of ovule fertilization and seed development that accounts for differential fertilization by self- and cross-pollen, limited ovule viability or receptivity, differential survival of self- and cross-zygotes and limited resource availability. Simulations illustrate that the proposed methods eliminate bias in estimated predispersal inbreeding depression caused by pollen limitation and can improve estimates under resource limitation. Application of these methods to two orchid species further demonstrates their utility in identifying and estimating diverse influences on reproductive performance under typical conditions. Although our theoretical results raise questions about the reported intensity of predispersal inbreeding depression, our proposed methods guard against bias while also providing insight into plant reproduction.