The computational principles underlying the processing of sensoryevoked synaptic inputs are understood only rudimentarily. A critical missing factor is knowledge of the activation patterns of the synaptic inputs to the processing neurons. Here we use welldefined, reproducible skin stimulation to describe the specific signal transformations that occur in different parallel mossy fiber pathways and analyze their representation in the synaptic inputs to cerebellar granule cells. We find that mossy fiber input codes are preserved in the synaptic responses of granule cells, suggesting a coding-specific innervation. The computational consequences of this are that it becomes possible for granule cells to also transmit weak sensory inputs in a graded fashion and to preserve the specific activity patterns of the mossy fibers.cerebellum ͉ cuneate ͉ lateral reticular nucleus F irst-order processing of sensory inputs within the mammalian brain occurs within the spinal cord, brainstem nuclei, and thalamus, as well as in the input layers of specific parts of the cerebral and cerebellar cortices. For example, skin sensory input destined for the cerebellar granule layer can be preprocessed through either the cuneate nucleus pathway (1) or the spinal cord-lateral reticular nucleus (LRN) pathways (2) before being processed further by the granule cells. In the cuneate nucleus, neurons process raw primary afferent input mediated directly from skin receptors, and a set of rather intricate connectivity patterns ensures that the information conveyed through the cuneate neurons represent a synthesized receptive field that have sharp borders (3, 4). Thus, it may be speculated that the purpose of the preprocessing is to present the input layers with sensory information in a form that can be more readily used by downstream neurons, eliminating properties of the skin sensory sheet that are not relevant to the central processing. In contrast, the LRN, another important precerebellar source, receives skin sensory input through neurons of the spinal cord that are involved in the mediation of descending motor commands (5, 6). Although both the cuneate and the LRN are strongly influenced by skin input, they are likely to code the same skin input in different ways because of their differences in convergent synaptic inputs and intrinsic cellular properties.The understanding of the function of the input layers is more limited, but at least for the cerebellar granule cells, various contrasting theories have been proposed (7-9). Cerebellar granule cells have attracted considerable interest because of their relative simplicity; they receive only about 4 mossy fiber synaptic inputs, each of which evokes strong postsynaptic responses (7, 10). Therefore, actually determining the precise role of each individual synapse in neuronal information processing is feasible, if intracellular recordings and a description of the natural activation patterns in vivo of each synaptic input can be obtained. Here we approach this aim by identifying the specific coding o...