V1 activity during feedforward and early feedback processing is necessary for both conscious and unconscious motion perception.

The study of blindsight has revealed a seminal dissociation between conscious vision and visually guided behavior: some patients who are blind due to V1 lesions seem to be able to employ unconscious visual information in their behavior. The standard assumption is that these findings generalize to the neurologically healthy. We tested whether unconscious processing of motion is possible without the contribution of V1 in neurologically healthy participants by disturbing activity in V1 using transcranial magnetic stimulation (TMS). Unconscious processing was measured with redundant target effect (RTE), a phenomenon where participants respond faster to two stimuli than to one stimulus, when the task is just to respond as fast as possible when one stimulus or two simultaneous stimuli are presented. We measured the RTE caused by a motion stimulus. V1 activity was interfered with different stimulus onset asynchronies (SOA) to test whether TMS delivered in a specific time window suppresses conscious perception (participant reports seeing only one of the two stimuli) but does not affect unconscious processing (RTE). We observed that at each SOA, when TMS suppressed conscious perception of the stimulus, the RTE was also eliminated. However, when visibility of the redundant target was suppressed with a visual mask, we found unconscious processing of motion. This suggests that unconscious processing of motion depends on V1 in neurologically healthy humans. We conclude that the neural mechanisms that enable motion processing in blindsight are modulated by neuroplastic changes in connectivity between subcortical areas and the visual cortex after the V1 lesion. Neurologically healthy observers cannot process motion unconsciously without functioning of V1.

Early processing in primary visual cortex is necessary for conscious and unconscious vision while late processing is necessary only for conscious vision in neurologically healthy humans.

The neural mechanisms underlying conscious and unconscious visual processes remain controversial. Blindsight patients may process visual stimuli unconsciously despite their V1 lesion, promoting anatomical models, which suggest that pathways bypassing the V1 support unconscious vision. On the other hand, physiological models argue that the major geniculostriate pathway via V1 is involved in both unconscious and conscious vision, but in different time windows and in different types of neural activity. According to physiological models, feedforward activity via V1 to higher areas mediates unconscious processes whereas feedback loops of recurrent activity from higher areas back to V1 support conscious vision. With transcranial magnetic stimulation (TMS) it is possible to study the causal role of a brain region during specific time points in neurologically healthy participants. In the present study, we measured unconscious processing with redundant target effect, a phenomenon where participants respond faster to two stimuli than one even when one of the stimuli is not consciously perceived. We tested the physiological feedforward-feedback model of vision by suppressing conscious vision by interfering selectively either with early or later V1 activity with TMS. Our results show that early V1 activity (60ms) is necessary for both unconscious and conscious vision. During later processing stages (90ms), V1 contributes selectively to conscious vision. These findings support the feedforward-feedback-model of consciousness.

Psychophysical "blinding" methods reveal a functional hierarchy of unconscious visual processing.

Numerous non-invasive experimental "blinding" methods exist for suppressing the phenomenal awareness of visual stimuli. Not all of these suppressive methods occur at, and thus index, the same level of unconscious visual processing. This suggests that a functional hierarchy of unconscious visual processing can in principle be established. The empirical results of extant studies that have used a number of different methods and additional reasonable theoretical considerations suggest the following tentative hierarchy. At the highest levels in this hierarchy is unconscious processing indexed by object-substitution masking. The functional levels indexed by crowding, the attentional blink (and other attentional blinding methods), backward pattern masking, metacontrast masking, continuous flash suppression, sandwich masking, and single-flash interocular suppression, fall at progressively lower levels, while unconscious processing at the lowest levels is indexed by eye-based binocular-rivalry suppression. Although unconscious processing levels indexed by additional blinding methods is yet to be determined, a tentative placement at lower levels in the hierarchy is also given for unconscious processing indexed by Troxler fading and adaptation-induced blindness, and at higher levels in the hierarchy indexed by attentional blinding effects in addition to the level indexed by the attentional blink. The full mapping of levels in the functional hierarchy onto cortical activation sites and levels is yet to be determined. The existence of such a hierarchy bears importantly on the search for, and the distinctions between, neural correlates of conscious and unconscious vision.

Neuronavigated TMS of early visual cortex eliminates unconscious processing of chromatic stimuli.

Some neurological patients with primary visual cortex (V1) lesions can guide their behavior based on stimuli presented to their blind visual field. One example of this phenomenon is the ability to discriminate colors in the absence of awareness. These so-called patients with blindsight must have a neural pathway that bypasses V1, explaining their ability to unconsciously process stimuli. The pathways that have been most often hypothesized to be the cause of blindsight connect lateral geniculate nucleus (LGN) or superior colliculus (SC) to extrastriate cortex, most likely V5, and parietal areas. To test if similar pathways function in neurologically healthy individuals or if unconscious processing depends on early visual cortex, we disturbed the visibility of a chromatic stimulus with metacontrast masking (Experiment 1) or neuronavigated transcranial magnetic stimulation (TMS) of early visual cortex, exact target being retinotopically mapped V1 (Experiment 2). We measured unconscious processing using the redundant target effect (RTE), which is the speeding up of reaction times in response to dual stimuli compared with one stimulus, when the task is to respond to any number of stimuli. An unconscious chromatic RTE was found when the visibility of the redundant chromatic stimulus was suppressed with a visual mask. When TMS was targeted to the correct retinotopic location of V1, and conscious perception of the redundant chromatic stimulus suppressed, the RTE was eliminated. Whether the elimination of unconscious RTE during TMS was exclusively due to disruption of V1 activity, or whether it was due to the possible interference with processing in V2 or even V3, is discussed. Based on our results and converging evidence from previous studies, we conclude that unconscious processing of chromatic information depends on the early visual cortex, in neurologically healthy participants.

The chronometry of visual perception: review of occipital TMS masking studies.

Transcranial magnetic stimulation (TMS) continues to deliver on its promise as a research tool. In this review article we focus on the application of TMS to early visual cortex (V1, V2, V3) in studies of visual perception and visual awareness. Depending on the asynchrony between visual stimulus onset and TMS pulse (SOA), TMS can suppress visual perception, allowing one to track the time course of functional relevance (chronometry) of early visual cortex for vision. This procedure has revealed multiple masking effects ('dips'), some consistently (∼+100ms SOA) but others less so (∼-50ms, ∼-20ms, ∼+30ms, ∼+200ms SOA). We review the state of TMS masking research, focusing on the evidence for these multiple dips, the relevance of several experimental parameters to the obtained 'masking curve', and the use of multiple measures of visual processing (subjective measures of awareness, objective discrimination tasks, priming effects). Lastly, we consider possible future directions for this field. We conclude that while TMS masking has yielded many fundamental insights into the chronometry of visual perception already, much remains unknown. Not only are there several temporal windows when TMS pulses can induce visual suppression, even the well-established 'classical' masking effect (∼+100ms) may reflect more than one functional visual process.