Every cognitive act entails the participation of multiple brain regions. In visual short-term memory, for example, visual information is initially encoded in sensory brain areas and then communicated to regions that mediate the retention, manipulation and retrieval of information. One prominent hypothesis that addresses the question of how communication between neural ensembles is achieved claims that neuronal oscillations support the timely coordination of neural activity between different brain regions 1,2 . Specifically, neuronal oscillations in the theta frequency band (3-9 Hz) have been suggested to underlie the interaction between neural ensembles during mnemonic processing 3,4 .A line of evidence supporting this hypothesis stems from studies investigating the role of hippocampal theta in memory formation in rodents [5][6][7] . Additional evidence comes from studies measuring surface-based or intracortical electroencephalography (EEG) in human subjects. In these cases, memory performance correlates with an increase in theta power [8][9][10] . Moreover, enhanced theta synchrony between electric potentials recorded from memory-related areas has been observed 11,12 .These findings raise the question of whether theta synchrony measured at the mesoscopic level of LFPs provides a basis for the timely coordination of spiking output between distant cortical areas that have been traditionally associated with the sensory encoding of visual information on the one hand and mnemonic processing on the other. Moreover, is the precision of coordination between these regions associated with changes in memory performance? To answer these questions, we studied neuronal interactions between the extrastriate visual area V4 and the lateral prefrontal cortex (lPF) while monkeys performed a visual memory task.Although neural activity in V4 has been related to color and shape processing of visual objects [13][14][15][16] and visual attention 17 , neural responses in lPF have been traditionally associated with working memory, that is, the short-term maintenance and manipulation of sensory information in memory tasks 18,19 . More recently, however, an increasing number of studies have found that the neural circuitry underlying short-term retention of sensory information likely entails earlier sensory cortical areas as well 20 . In these cases, both prefrontal regions 21 and V4 have been linked to memory-related oscillatory synchrony. Specifically, theta oscillations are enhanced during the delay period of memory tasks in both lPF and V4 (ref. 22), and increased oscillatory theta synchrony is accompanied by a phase-dependent coding of visual stimuli retained in short-term memory 23,24 .We observed enhanced phase locking between local field potentials recorded in V4 and lPF (inter-area LFP phase locking) that occurred in the theta range (~3-9 Hz) during the memory period of a visual short-term memory task. Increased LFP-phase locking was associated with greater locking of spikes to the phase of the theta oscillations in the respective...