Signal-to-noise ratios in physical systems can be significantly degraded if the outputs of the systems are highly variable. Biological processes for which highly stereotyped signal generations are necessary features appear to have reduced their signal variabilities by employing multiple processing steps. To better understand why this multistep cascade structure might be desirable, we prove that the reliability of a signal generated by a multistate system with no memory (i.e., a Markov chain) is maximal if and only if the system topology is such that the process steps irreversibly through each state, with transition rates chosen such that an equal fraction of the total signal is generated in each state. Furthermore, our result indicates that by increasing the number of states, it is possible to arbitrarily increase the reliability of the system. In a physical system, however, an energy cost is associated with maintaining irreversible transitions, and this cost increases with the number of such transitions (i.e., the number of states). Thus, an infinite-length chain, which would be perfectly reliable, is infeasible. To model the effects of energy demands on the maximally reliable solution, we numerically optimize the topology under two distinct energy functions that penalize either irreversible transitions or incommunicability between states, respectively. In both cases, the solutions are essentially irreversible linear chains, but with upper bounds on the number of states set by the amount of available energy. We therefore conclude that a physical system for which signal reliability is important should employ a linear architecture, with the number of states (and thus the reliability) determined by the intrinsic energy constraints of the system.

Cortical synaptic plasticity depends on the relative timing of pre- and postsynaptic spikes and also on the temporal pattern of presynaptic spikes and of postsynaptic spikes. We study the hypothesis that cortical synap- tic plasticity does not associate individual spikes, but rather whole fir- ing episodes, and depends only on when these episodes start and how long they last, but as little as possible on the timing of individual spikes. Here we present the mathematical background for such a study. Stan- dard methods from hidden Markov models are used to define what “fir- ing episodes” are. Estimating the probability of being in such an episode requires not only the knowledge of past spikes, but also of future spikes. We show how to construct a causal learning rule, which depends only on past spikes, but associates pre- and postsynaptic firing episodes as if it also knew future spikes. We also show that this learning rule agrees with some features of synaptic plasticity in superficial layers of rat visual cortex (Froemke and Dan, Nature 416:433, 2002).