The brain site of perceptual learning has been frequently debated. Recent psychophysical evidence for complete learning transfer to new retinal locations and orientations/directions suggests that perceptual learning may mainly occur in high-level brain areas. Contradictorily, ERP C1 changes associated with perceptual learning are cited as evidence for training-induced plasticity in the early visual cortex. However, C1 can be top-down modulated, which suggests the possibility that C1 changes may result from top-down modulation of the early visual cortex by high-level perceptual learning. To single out the potential top-down impact, we trained observers with a peripheral orientation discrimination task and measured C1 changes at an untrained diagonal quadrant location where learning transfer was previously known to be significant. Our assumption was that any C1 changes at this untrained location would indicate top-down modulation of the early visual cortex, rather than plasticity in the early visual cortex. The expected learning transfer was indeed accompanied with significant C1 changes. Moreover, C1 changes were absent in an untrained shape discrimination task with the same stimuli. We conclude that ERP C1 can be top-down modulated in a task-specific manner by high-level perceptual learning, so that C1 changes may not necessarily indicate plasticity in the early visual cortex. Moreover, learning transfer and associated C1 changes may indicate that learning-based top-down modulation can be remapped to early visual cortical neurons at untrained locations to enable learning transfer.

When an observer reports a letter flanked by additional letters in the visual periphery, the response errors (the crowding effect) may result from failure to recognize the target letter (recognition errors), from mislocating a correctly recognized target letter at a flanker location (target misplacement errors), or from reporting a flanker as the target letter (flanker substitution errors). Crowding can be reduced through perceptual learning. However, it is not known how perceptual learning operates to reduce crowding. In this study we trained observers with a partial-report task (Experiment 1), in which they reported the central target letter of a three-letter string presented in the visual periphery, or a whole-report task (Experiment 2), in which they reported all three letters in order. We then assessed the impact of training on recognition of both unflanked and flanked targets, with particular attention to how perceptual learning affected the types of errors. Our results show that training improved target recognition but not single-letter recognition, indicating that training indeed affected crowding. However, training did not reduce target misplacement errors or flanker substitution errors. This dissociation between target recognition and flanker substitution errors supports the view that flanker substitution may be more likely a by-product (due to response bias), rather than a cause, of crowding. Moreover, the dissociation is not consistent with hypothesized mechanisms of crowding that would predict reduced positional errors.