There are clear examples where spike timing matters a great deal and the Jeffres model of sound localization, as you point out, is a great example of this. Howe

There are clear examples where spike timing matters a great deal and the Jeffres model of sound localization, as you point out, is a great example of this. However, spike timing codes seem to be more prevalent in the periphery; rate codes dominate in central structures, and in particular the cortex. The receptors of sensory afferents faithfully report the timing of individual events (of photons, pinpricks of the skin, etc) with single spikes. But after a few synapses the pattern of spikes looks much more like a (noisy) rate code. The origin of the noise, whether it even is noise, how the temporal code is transformed into a rate code, and many other questions along these lines are active research topics. All that aside, Koch is well aware that there is evidence for both rate and temporal codes. As he wrote, "spikes mean different things in different contexts."