Pollination efficiency and the pollen-ovule ratio.

Pollination presents a risky journey for pollen grains. Pollen loss is sometimes thought to favour greater pollen investment to compensate for the inefficiency of transport. Sex allocation theory, to the contrary, has consistently concluded that postdispersal loss should have no selective effect on investment in either sex function. But the intuitively appealing compensation idea continues to be raised despite the lack of theoretical endorsement. We address the theoretical issue with a model that directly represents pollen loss (and ovule loss through floral demise or loss of receptivity) as rate-dependent dynamical processes. These loss rates can be varied to examine the effect of pollination efficiency on optimal sex allocation. Pollen-ovule ratios follow from the sex allocation based on the resource costs of pollen and ovule production. This model confirms conventional findings that pollen loss should have essentially no effect on sexual resource allocation in large, panmictic populations. Pollen limitation of seed set does not alter this conclusion. These results force us to rethink the empirical association of pollination efficiency with low pollen-ovule ratios. This pattern could arise if efficient pollen transport commonly results in stigmatic deposition of cohorts of related pollen. Empirical evidence of correlated paternity supports this explanation.

Infrastructure construction without information exchange: the trail clearing mechanism in Atta leafcutter ants.

A wide range of group-living animals construct tangible infrastructure networks, often of remarkable size and complexity. In ant colonies, infrastructure construction may require tens of thousands of work hours distributed among many thousand individuals. What are the individual behaviours involved in the construction and what level of complexity in inter-individual interaction is required to organize this effort? We investigate this question in one of the most sophisticated trail builders in the animal world: the leafcutter ants, which remove leaf litter, cut through overhangs and shift soil to level the path of trail networks that may cumulatively extend for kilometres. Based on obstruction experiments in the field and the laboratory, we identify and quantify different individual trail clearing behaviours. Via a computational model, we further investigate the presence of recruitment, which-through direct or indirect information transfer between individuals-is one of the main organizing mechanisms of many collective behaviours in ants. We show that large-scale transport networks can emerge purely from the stochastic process of workers encountering obstructions and subsequently engaging in removal behaviour with a fixed probability. In addition to such incidental removal, we describe a dedicated clearing behaviour in which workers remove additional obstructions independent of chance encounters. We show that to explain the dynamics observed in the experiments, no information exchange (e.g. via recruitment) is required, and propose that large-scale infrastructure construction of this type can be achieved without coordination between individuals.

Physical and biological determinants of collective behavioural dynamics in complex systems: pulling chain formation in the nest-weaving ant Oecophylla smaragdina.

The evolution of nest weaving, the inclusion of larval silk in the nest walls, is considered one of the pinnacles of cooperative behaviour in social insects. Within the four ant genera in which this has evolved, Oecophylla are unique in being the only group that precedes the deposition of larval silk by actively manipulating the leaf substrate to form a nest chamber. Here we provide the first descriptions of the manipulation process within a complex-systems framework. Substrate manipulation involves individual ants selecting, grasping and attempting to pull the edge of the substrate. These individuals are then joined by nest mates at the work site, who either select a site beside the first individual or grasp the body of the first or preceding worker to form a chain of pulling ants that together drag and bend the substrate. Site selection by individual workers is not random when confronted with an artificial leaf, with individuals more likely to grasp a substrate at its tip rather than along a more broad edge. The activity of additional individuals is also not random, with their activity being grouped in both space and time. Additional individuals are more likely to join an existing biting individual or pulling group. The positive feedback associated with the early stages of pulling behaviour appears typical for many of the collective actions observed in social insects.