The group I metabotropic glutamate receptor 5 (mGluR5) has been implicated in the development of cortical sensory maps. However its precise roles in the synaptic function and plasticity of thalamocortical connections remain unknown. Here we first show that in mGluR5 knockout mice bred onto a C57BL6 background cyto-architectonic differentiation into barrels is missing, but the representations for large whiskers are identifiable as clusters of thalamocortical afferents. The altered dendritic morphology of cortical layer IV spiny stellate neurons in mGluR5 knockout mice implicates a role for mGluR5 in the dendritic morphogenesis of excitatory neurons. Next, in vivo single unit recordings of whisker evoked activity in mGluR5 knockout adults demonstrated a preserved topographical organization of the whisker representation, but a significantly diminished temporal discrimination of center to surround whiskers in the responses of individual neurons. To evaluate synaptic function at thalamocortical synapses in mGluR5 knockout mice, whole-cell voltage clamp recording was conducted in acute thalamocortical brain slices prepared from postnatal day 4-11 mice. At mGluR5 knockout thalamocortical synapses, NMDA currents decayed faster and synaptic strength was more easily reduced, but more difficult to strengthen by Hebbian-type pairing protocols, despite a normal developmental increase in AMPAR-mediated currents and presynaptic function. We have therefore demonstrated that mGluR5 is required for synaptic function/plasticity at thalamocortical synapses as barrels are forming and we propose that these functional alterations at the thalamocortical synapse are the basis of the abnormal anatomical and functional development of the somatosensory cortex in the mGluR5 knockout mouse.