Cortical quantity representations of visual numerosity and timing overlap increasingly into superior cortices but remain distinct.

Many sensory brain areas are organized as topographic maps where neural response preferences change gradually across the cortical surface. Within association cortices, 7-Tesla fMRI and neural model-based analyses have also revealed many topographic maps for quantities like numerosity and event timing, often in similar locations. Numerical and temporal quantity estimations also show behavioral similarities and even interactions. For example, the duration of high-numerosity displays is perceived as longer than that of low-numerosity displays. Such interactions are often ascribed to a generalized magnitude system with shared neural responses across quantities. Anterior quantity responses are more closely linked to behavior. Here, we investigate whether common quantity representations hierarchically emerge by asking whether numerosity and timing maps become increasingly closely related in their overlap, response preferences, and topography. While the earliest quantity maps do not overlap, more superior maps overlap increasingly. In these overlapping areas, some intraparietal maps have consistently correlated numerosity and timing preferences, and some maps have consistent angles between the topographic progressions of numerosity and timing preferences. However, neither of these relationships increases hierarchically like the amount of overlap does. Therefore, responses to different quantities are initially derived separately, then progressively brought together, without generally becoming a common representation. Bringing together distinct responses to different quantities may underlie behavioral interactions and allow shared access to comparison and action planning systems.

Visual timing-tuned responses in human association cortices and response dynamics in early visual cortex.

Quantifying the timing (duration and frequency) of brief visual events is vital to human perception, multisensory integration and action planning. Tuned neural responses to visual event timing have been found in association cortices, in areas implicated in these processes. Here we ask how these timing-tuned responses are related to the responses of early visual cortex, which monotonically increase with event duration and frequency. Using 7-Tesla functional magnetic resonance imaging and neural model-based analyses, we find a gradual transition from monotonically increasing to timing-tuned neural responses beginning in the medial temporal area (MT/V5). Therefore, across successive stages of visual processing, timing-tuned response components gradually become dominant over inherent sensory response modulation by event timing. This additional timing-tuned response component is independent of retinotopic location. We propose that this hierarchical emergence of timing-tuned responses from sensory processing areas quantifies sensory event timing while abstracting temporal representations from spatial properties of their inputs.

Auditory timing-tuned neural responses in the human auditory cortices.

Perception of sub-second auditory event timing supports multisensory integration, and speech and music perception and production. Neural populations tuned for the timing (duration and rate) of visual events were recently described in several human extrastriate visual areas. Here we ask whether the brain also contains neural populations tuned for auditory event timing, and whether these are shared with visual timing. Using 7T fMRI, we measured responses to white noise bursts of changing duration and rate. We analyzed these responses using neural response models describing different parametric relationships between event timing and neural response amplitude. This revealed auditory timing-tuned responses in the primary auditory cortex, and auditory association areas of the belt, parabelt and premotor cortex. While these areas also showed tonotopic tuning for auditory pitch, pitch and timing preferences were not consistently correlated. Auditory timing-tuned response functions differed between these areas, though without clear hierarchical integration of responses. The similarity of auditory and visual timing tuned responses, together with the lack of overlap between the areas showing these responses for each modality, suggests modality-specific responses to event timing are computed similarly but from different sensory inputs, and then transformed differently to suit the needs of each modality.

Numerosity tuning in human association cortices and local image contrast representations in early visual cortex.

Human early visual cortex response amplitudes monotonically increase with numerosity (object number), regardless of object size and spacing. However, numerosity is typically considered a high-level visual or cognitive feature, while early visual responses follow image contrast in the spatial frequency domain. We find that, at fixed contrast, aggregate Fourier power (at all orientations and spatial frequencies) follows numerosity closely but nonlinearly with little effect of object size, spacing or shape. This would allow straightforward numerosity estimation from spatial frequency domain image representations. Using 7T fMRI, we show monotonic responses originate in primary visual cortex (V1) at the stimulus's retinotopic location. Responses here and in neural network models follow aggregate Fourier power more closely than numerosity. Truly numerosity tuned responses emerge after lateral occipital cortex and are independent of retinotopic location. We propose numerosity's straightforward perception and neural responses may result from the pervasive spatial frequency analyses of early visual processing.

Adaptation to visual numerosity changes neural numerosity selectivity.

Perceiving numerosity, i.e. the set size of a group of items, is an evolutionarily preserved ability found in humans and animals. A useful method to infer the neural underpinnings of a given perceptual property is sensory adaptation. Like other primary perceptual attributes, numerosity is susceptible to adaptation. Recently, we have shown numerosity-selective neural populations with a topographic organization in the human brain. Here, we investigated whether numerosity adaptation can affect the numerosity selectivity of these populations using ultra-high field (7 Tesla) functional magnetic resonance imaging (fMRI). Participants viewed stimuli of changing numerosity (1 to 7 dots), which allowed the mapping of numerosity selectivity. We interleaved a low or high numerosity adapter stimulus with these mapping stimuli, repeatedly presenting 1 or 20 dots respectively to adapt the numerosity-selective neural populations. We analyzed the responses using custom-build population receptive field neural models of numerosity encoding and compared estimated numerosity preferences between adaptation conditions. We replicated our previous studies where we found several topographic maps of numerosity-selective responses. We found that overall, numerosity adaptation altered the preferred numerosities within the numerosity maps, resulting in predominantly attractive biases towards the numerosity of the adapter. The differential biases could be explained by the difference between the unadapted preferred numerosity and the numerosity of the adapter, with attractive biases being observed with higher difference. The results could link perceptual numerosity adaptation effects to changes in neural numerosity selectivity.