Biphasic responses, quantal signals and cellular behaviour.

It is commonplace to think of thresholds in biological systems. Biphasic responses, with both thresholds and upper limits, or lintels, are also surprisingly common. In this paper we show that they are found in many systems in which an aspect of cellular behaviour is controlled by chemical signals. In some cases the biphasic response can lead to the partitioning of a tissue into regions expressing different behaviours and, therefore, in principle able to take different developmental and evolutionary paths within the same organism. Several other features are common; these include brief, all-or-nothing responses and the expression of different behaviours evoked by a signal of a single chemical species in one or more cell types, but over different concentration ranges. Such behaviour is illustrated very clearly by the differentiation of cells in the mammalian immune system as well as by developing slime mould cells, so the underlying principle is widespread. We suggest that the interaction of unitary behaviours with chemical signals showing such non-linear concentration dependences will account for the complexity of development.