It has been known for >30 years that neuronal spike trains exhibit correlations, that is, the occurrence of a spike at one time is not independent of the occurrence of spikes at other times, both within spike trains from single neurons and across spike trains from multiple neurons. The presence of these correlations has led to the proposal that they might form a key element of the neural code. Specifically, they might act as an extra channel for information, carrying messages about events in the outside world that are not carried by other aspects of the spike trains, such as firing rate. Currently, there is no general consensus about whether this proposal applies to real spike trains in the nervous system. This is largely because it has been hard to separate information carried in correlations from that not carried in correlations. Here we propose a framework for performing this separation. Specifically, we derive an information-theoretic cost function that measures how much harder it is to decode neuronal responses when correlations are ignored than when they are taken into account. This cost function can be readily applied to real neuronal data. E ver since Adrian and Zotterman observed that the firing rate of peripheral touch receptors coded for the pressure applied to a patch of skin (1), neuroscientists have been trying to crack the neural code, that is, to understand the relationship between neuronal activity and events in the outside world. For much of that time, the working hypothesis was that information is carried by firing rate. More recently it has been proposed that firing rate is not the whole story: Information might also be carried in spike patterns, both within spike trains from single neurons (2-6) and across spike trains from multiple neurons (7-9).One aspect of this proposal, an aspect that has led to a great deal of debate, is that correlations in spike patterns may be of particular importance (7)(8)(9)(10)12). It has been known for many years that spike trains contain correlations; that is, the presence of a spike at one time is not independent of the presence of spikes at other times. These correlations exist not just within spike trains but across them as well, with the most common example being synchronous spikes across pairs of cells (13)(14)(15)(16). What has led to the debate is the suggestion that these correlations might form a key aspect of the code. The idea is that they might serve as an extra information channel, conveying messages not carried elsewhere in the spike trains.How might the correlations do this? An example, using synchronous spikes, is shown in Fig. 1a. In this example there are two stimuli, A and B, and two neurons. When stimulus A is presented, the two neurons produce five spikes on average. When stimulus B is presented, they also produce five spikes on average. What is different, though, is the correlational structure of the responses: When stimulus A is presented, the two neurons tend to produce few synchronous spikes, whereas when stimulus B is presented, they ...