Single-cell analysis of habituation in Stentor coeruleus

Although learning is often viewed as a unique feature of organisms with complex nervous systems, single -celled organisms also demonstrate basic forms of learning. The giant ciliate Stentor coeruleus responds to me-chanical stimuli by contracting into a compact shape, presumably as a defense mechanism. When a Stentor cell is repeatedly stimulated at a constant level of force, it will learn to ignore that stimulus but will still respond to stronger stimuli. Prior studies of habituation in Stentor reported a graded response, suggesting that cells transition through a continuous range of response probabilities. By analyzing single cells using an automated apparatus to deliver calibrated stimuli, we find that habituation occurs via a single step-like switch in contrac-tion probability within each cell, with the graded response in a population arising from the random distribution of switching times in individual cells. This step-like response allows Stentor behavior to be represented by a simple two-state model whose parameters can be estimated from experimental measurements. We find that transition rates depend on stimulus force and also on the time between stimuli. The ability to measure the behavior of the same cell to the same stimulus allowed us to quantify the functional heterogeneity among sin-gle cells. Together, our results suggest that the behavior of Stentor is governed by a two-state stochastic ma-chine whose transition rates are sensitive to the time series properties of the input stimuli.