A test of balanced fitness limitations theory: Pollen limitation in plants.

When reproductive success is determined by the relative availabilities of a series of essential, non-substitutable resources, the theory of balanced fitness limitations predicts that the cost of harvesting a particular resource shapes the likelihood that a shortfall of that resource will constrain realized fitness. Plant reproduction through female function offers a special opportunity to test this theory; essential resources in this context include, first, the pollen received from pollinators or abiotic vectors that is used to fertilize ovules, and, second, the resources needed to provision the developing seeds and fruit. For many plants realized reproductive success through female function can be readily quantified in the field, and one key potential constraint on fitness, pollen limitation, can be assessed experimentally by manually supplementing pollen receipt. We assembled a comparative dataset of pollen limitation using only studies that supplement pollen to all flowers produced over the plant's reproductive lifespan. Pre- and post-pollination costs were estimated using the weight of flowers and fruits and estimates of fruit set. Consistent with expectations, we find self-incompatible plants make greater pre-pollination investments and experience greater pollen limitation. However, contrary to theoretical expectations, when variation due to self-compatibility is accounted for by including self-compatibility in the statistical model as a covariate, we find no support for the prediction that plants that invest more heavily in pre-pollination costs are subject to greater pollen limitation. Strong within-species, between-population variation in the expression of pollen limitation makes the quantification of mean pollen limitation difficult. We urge plant ecologists to conduct more studies of pollen limitation using whole-plant pollen supplementation to produce a richer comparative dataset that would support a more robust test of the balanced limitations hypothesis.