“…At least for simple cells in layer 4B, this RF structure determines the response to moving visual features (McLean & Palmer, 1989;McLean et al, 1994;Reid et al, 1991;Albrecht & Geisler, 1991;DeAngelis et al, 1993;DeAngelis et al, 1995;Jagadeesh et al, 1993;Jagadeesh et al, 1997;Murthy et al, 1998), and hence the cell's tuning for direction and speed 1 . Lagged and nonlagged inputs that converge, either directly or via other cortical neurons, on simple cells and segregate in separate subregions of the simple cells' RF are thus likely to contribute to the earliest level of cortical velocity selectivity (Saul & Humphrey, 1990;Saul & Humphrey, 1992a;Saul & Humphrey, 1992b;Ferster et al, 1996; 1 To avoid confusion, we point out that the term 'speed tuning' is sometimes used in a more restricted sense. Simple cells exhibit tuning for spatial and temporal frequencies that results in preference for speeds of moving gratings depending on their spatial frequency.…”