Principal cells in the olfactory bulb (OB), mitral and tufted cells, receive direct sensory input and generate output signals that are transmitted to downstream cortical targets. Excitatory input from glutamatergic receptor neurons are the primary known sources of rapid excitation to OB principal cells. Principal cells also receive inhibitory input from local GABAergic interneurons in both the glomerular and plexiform layers. Previous work suggests that the functional effect of these inhibitory inputs, including numerous dendrodendritic synapses with GABAergic granule cells, is to reduce firing probability. In this study, we use in vitro patch clamp recordings to demonstrate that rat (of both sexes) OB mitral cells also can be excited by GABAergic synapses formed outside the glomerular layer. Depolarizing GABAergic responses to focal extracellular stimulation were revealed when fast ionotropic glutamate receptors were blocked, and occurred with short, monosynaptic latencies. These novel synaptic responses ...