A model of chemotaxis and associative learning in C. elegans.

The nematode C. elegans has attracted a great deal of interest from the neuroscience community due to the simplicity of its nervous system, which in the hermaphrodite is composed of just 302 neurons. C. elegans is known to engage in a number of sophisticated behaviours such as chemo- and thermotaxis. Experimental work has shown that these behaviours can be modified by experience and that C. elegans is capable of associative learning. In this paper, we focus on the chemotactic response of C. elegans to sodium chloride mediated by the ASE sensory neurons. We construct a biophysical model of the ASEL and ASER neurons that captures the time course of the ASE responses in response to up- and down-steps in NaCl concentration. We use this model to show that the time course of the ASE responses provide sufficient temporal resolution to successfully drive chemotaxis in C. elegans via steering, pirouettes and control of final turn angle. We show that these different locomotion strategies are individually capable of driving chemotaxis and that by working together they produce the best chemotactic response. We find that there is a separation into upward and downward drives mediated by the left and right ASE neurons. We show that the connectivity from ASEL and ASER must be of opposite polarity and that ASER, and the concomitant ability to sense when the worm is moving down the gradient, is more important for chemotaxis than ASEL, findings that are consistent with existing modelling studies in the literature. Finally, we examine associative learning in the network and show that experimental data can be explained by changes that occur at either the synaptic or sensory neuron level, the choice of which has distinct consequences for network function.