Understanding interactions between populations: Individual based modelling and quantification using pair correlation functions.

Understanding the underlying mechanisms that produce the huge variety of swarming and aggregation patterns in animals and cells is fundamental in ecology, developmental biology, and regenerative medicine, to name but a few examples. Depending upon the nature of the interactions between individuals (cells or animals), a variety of different large-scale spatial patterns can be observed in their distribution; examples include cell aggregates, stripes of different coloured skin cells, etc. For the case where all individuals are of the same type (i.e., all interactions are alike), a considerable literature already exists on how the collective organisation depends on the inter-individual interactions. Here, we focus on the less studied case where there are two different types of individuals present. Whilst a number of continuum models of this scenario exist, it can be difficult to compare these models to experimental data, since real cells and animals are discrete. In order to overcome this problem, we develop an agent-based model to simulate some archetypal mechanisms involving attraction and repulsion. However, with this approach (as with experiments), each realisation of the model is different, due to stochastic effects. In order to make useful comparisons between simulations and experimental data, we need to identify the robust features of the spatial distributions of the two species which persist over many realisations of the model (for example, the size of aggregates, degree of segregation or intermixing of the two species). In some cases, it is possible to do this by simple visual inspection. In others, the features of the pattern are not so clear to the unaided eye. In this paper, we introduce a pair correlation function (PCF), which allows us to analyse multi-species spatial distributions quantitatively. We show how the differing strengths of inter-individual attraction and repulsion between species give rise to different spatial patterns, and how the PCF can be used to quantify these differences, even when it might be impossible to recognise them visually.