Perceptual grouping of line segments into object contours has been thought to be mediated, in part, by long-range horizontal connectivity intrinsic to the primary visual cortex (V1), with a contribution by top-down feedback projections. To dissect the contributions of intraareal and interareal connections during contour integration, we applied conditional Granger causality analysis to assess directional influences among neural signals simultaneously recorded from visual cortical areas V1 and V4 of monkeys performing a contour detection task. Our results showed that discounting the influences from V4 markedly reduced V1 lateral interactions, indicating dependence on feedback signals of the effective connectivity within V1. On the other hand, the feedback influences were reciprocally dependent on V1 lateral interactions because the modulation strengths from V4 to V1 were greatly reduced after discounting the influences from other V1 neurons. Our findings suggest that feedback and lateral connections closely interact to mediate image grouping and segmentation.perceptual grouping | contour integration | Granger causality | horizontal connection | feedback connection A key step in the visual system's analysis of object shape is to group line segments into global contours and segregate these contours from background features. This process is critical to identifying object boundaries in complex visual scenes, and thus particularly important for performing shape discrimination; image segmentation; and, ultimately, object recognition.Contour integration follows the Gestalt principle of good continuation (1). The underlying neural underpinnings have been characterized as an association field (2), which links contour elements that are part of smooth contours. Neurophysiological studies in monkeys have identified that the primary visual cortex (V1) makes a fundamental contribution to contour integration (3-6), and anatomical studies have shown that the topology of horizontal connections in V1 is well suited for mediating interactions between neurons with a similar orientation preference (7-10). Such intracortical circuitry in V1 has been implemented in many computational models to account for the successful process of contour integration (11-13). Although many lines of converging evidence suggest that V1 is intimately involved in contour integration, circuitbased models have to take into account the findings that contour grouping is more than a bottom-up or hard-wired process, but that it is strongly dependent on top-down feedback influences (5, 14-17). Surface segmentation, another important intermediate stage in processing of visual images, is also mediated by interactions between feedforward and feedback connections (18).We have proposed a model whereby cortical feedback contributes to the effective connectivity of horizontal connections within V1 (13,19). A possible role of higher cortical areas in this process is to disambiguate local image components by creating a template that is fed back to V1, which then can select...