  Cells often perform computations in response to environmental cues. A simple
example is the classic problem, first considered by Berg and Purcell, of
determining the concentration of a chemical ligand in the surrounding media. On
general theoretical grounds (Landuer's Principle), it is expected that such
computations require cells to consume energy. Here, we explicitly calculate the
energetic costs of computing ligand concentration for a simple two-component
cellular network that implements a noisy version of the Berg-Purcell strategy.
We show that learning about external concentrations necessitates the breaking
of detailed balance and consumption of energy, with greater learning requiring
more energy. Our calculations suggest that the energetic costs of cellular
computation may be an important constraint on networks designed to function in
resource poor environments such as the spore germination networks of bacteria.
