Presaccadic Attention Depends on Eye Movement Direction and Is Related to V1 Cortical Magnification.

With every saccadic eye movement, humans bring new information into their fovea to be processed with high visual acuity. Notably, perception is enhanced already before a relevant item is foveated: During saccade preparation, presaccadic attention shifts to the upcoming fixation location, which can be measured via behavioral correlates such as enhanced visual performance or modulations of sensory feature tuning. The coupling between saccadic eye movements and attention is assumed to be robust and mandatory and considered a mechanism facilitating the integration of pre- and postsaccadic information. However, until recently it had not been investigated as a function of saccade direction. Here, we measured contrast response functions during fixation and saccade preparation in male and female observers and found that the pronounced response gain benefit typically elicited by presaccadic attention is selectively lacking before upward saccades at the group level-some observers even showed a cost. Individual observer's sensitivity before upward saccades was negatively related to their amount of surface area in primary visual cortex representing the saccade target, suggesting a potential compensatory mechanism that optimizes the use of the limited neural resources processing the upper vertical meridian. Our results raise the question of how perceptual continuity is achieved and how upward saccades can be accurately targeted despite the lack of-theoretically required-presaccadic attention.

Variability of the Surface Area of the V1, V2, and V3 Maps in a Large Sample of Human Observers.

How variable is the functionally defined structure of early visual areas in human cortex and how much variability is shared between twins? Here we quantify individual differences in the best understood functionally defined regions of cortex: V1, V2, V3. The Human Connectome Project 7T Retinotopy Dataset includes retinotopic measurements from 181 subjects (109 female, 72 male), including many twins. We trained four "anatomists" to manually define V1-V3 using retinotopic features. These definitions were more accurate than automated anatomical templates and showed that surface areas for these maps varied more than threefold across individuals. This threefold variation was little changed when normalizing visual area size by the surface area of the entire cerebral cortex. In addition to varying in size, we find that visual areas vary in how they sample the visual field. Specifically, the cortical magnification function differed substantially among individuals, with the relative amount of cortex devoted to central vision varying by more than a factor of 2. To complement the variability analysis, we examined the similarity of visual area size and structure across twins. Whereas the twin sample sizes are too small to make precise heritability estimates (50 monozygotic pairs, 34 dizygotic pairs), they nonetheless reveal high correlations, consistent with strong effects of the combination of shared genes and environment on visual area size. Collectively, these results provide the most comprehensive account of individual variability in visual area structure to date, and provide a robust population benchmark against which new individuals and developmental and clinical populations can be compared.SIGNIFICANCE STATEMENT Areas V1, V2, and V3 are among the best studied functionally defined regions in human cortex. Using the largest retinotopy dataset to date, we characterized the variability of these regions across individuals and the similarity between twin pairs. We find that the size of visual areas varies dramatically (up to 3.5×) across healthy young adults, far more than the variability of the cerebral cortex size as a whole. Much of this variability appears to arise from inherited factors, as we find very high correlations in visual area size between monozygotic twin pairs, and lower but still substantial correlations between dizygotic twin pairs. These results provide the most comprehensive assessment of how functionally defined visual cortex varies across the population to date.

Cross-dataset reproducibility of human retinotopic maps.

Population receptive field (pRF) models fit to fMRI data are used to non-invasively measure retinotopic maps in human visual cortex, and these maps are a fundamental component of visual neuroscience experiments. Here, we examined the reproducibility of retinotopic maps across two datasets: a newly acquired retinotopy dataset from New York University (NYU) (n = 44) and a public dataset from the Human Connectome Project (HCP) (n = 181). Our goal was to assess the degree to which pRF properties are similar across datasets, despite substantial differences in their experimental protocols. The two datasets simultaneously differ in their stimulus apertures, participant pool, fMRI protocol, MRI field strength, and preprocessing pipeline. We assessed the cross-dataset reproducibility of the two datasets in terms of the similarity of vertex-wise pRF estimates and in terms of large-scale polar angle asymmetries in cortical magnification. Within V1, V2, V3, and hV4, the group-median NYU and HCP vertex-wise polar angle estimates were nearly identical. Both eccentricity and pRF size estimates were also strongly correlated between the two datasets, but with a slope different from 1; the eccentricity and pRF size estimates were systematically greater in the NYU data. Next, to compare large-scale map properties, we quantified two polar angle asymmetries in V1 cortical magnification previously identified in the HCP data. The NYU dataset confirms earlier reports that more cortical surface area represents horizontal than vertical visual field meridian, and lower than upper vertical visual field meridian. Together, our findings show that the retinotopic properties of V1, V2, V3, and hV4 can be reliably measured across two datasets, despite numerous differences in their experimental design. fMRI-derived retinotopic maps are reproducible because they rely on an explicit computational model of the fMRI response. In the case of pRF mapping, the model is grounded in physiological evidence of how visual receptive fields are organized, allowing one to quantitatively characterize the BOLD signal in terms of stimulus properties (i.e., location and size). The new NYU Retinotopy Dataset will serve as a useful benchmark for testing hypotheses about the organization of visual areas and for comparison to the HCP 7T Retinotopy Dataset.

Radial asymmetries in population receptive field size and cortical magnification factor in early visual cortex.

Human visual cortex does not represent the whole visual field with the same detail. Changes in receptive field size, population receptive field (pRF) size and cortical magnification factor (CMF) with eccentricity are well established, and associated with changes in visual acuity with eccentricity. Visual acuity also changes across polar angle. However, it remains unclear how RF size, pRF size and CMF change across polar angle. Here, we examine differences in pRF size and CMF across polar angle in V1, V2 and V3 using pRF modeling of human fMRI data. In these visual field maps, we find smaller pRFs and larger CMFs in horizontal (left and right) than vertical (upper and lower) visual field quadrants. Differences increase with eccentricity, approximately in proportion to average pRF size and CMF. Similarly, we find larger CMFs in the lower than upper quadrant, and again differences increase with eccentricity. However, pRF size differences between lower and upper quadrants change direction with eccentricity. Finally, we find slightly smaller pRFs in the left than right quadrants of V2 and V3, though this difference is very small, and we find no differences in V1 and no differences in CMF. Moreover, differences in pRF size and CMF vary gradually with polar angle and are not limited to the meridians or visual field map discontinuities. PRF size and CMF differences do not consistently follow patterns of cortical curvature, despite the link between cortical curvature and polar angle in V1. Thus, the early human visual cortex has a radially asymmetric representation of the visual field. These asymmetries may underlie consistent reports of asymmetries in perceptual abilities.

Improving the Reliability and Accuracy of Population Receptive Field Measures Using a Logarithmically Warped Stimulus.

The population receptive field method, which measures the region in visual space that elicits a BOLD signal in a voxel in retinotopic cortex, is a powerful tool for investigating the functional organization of human visual cortex with fMRI (Dumoulin & Wandell, 2008). However, recent work has shown that population receptive field (pRF) estimates for early retinotopic visual areas can be biased and unreliable, especially for voxels representing the fovea. Here, we show that a 'log-bar' stimulus that is logarithmically warped along the eccentricity dimension produces more reliable estimates of pRF size and location than the traditional moving bar stimulus. The log-bar stimulus was better able to identify pRFs near the foveal representation, and pRFs were smaller in size, consistent with simulation estimates of receptive field sizes in the fovea.

Visual field asymmetries in visual word form identification.

Visual performance across the visual fields interacts with visual tasks and visual stimuli, and visual resolution decreases as a function of eccentricity, varying at isoeccentric locations. In this study, we investigated the extent of asymmetry and the rate of change in visual acuity threshold for visual word form (VWF) identification at horizontal and vertical azimuths across the fovea, and at eccentricities of 1°, 2°, 4°, 6° and 8° for 10%, 20%, 40%, and 80% contrast levels, to determine whether and how the eccentricities, meridians, and contrasts modulated the VWF identification acuity threshold. The stimuli were 16 traditional Chinese characters of similar legibility. Participants pressed a key to indicate the character presented, either monocularly or binocularly, at one of 21 randomly selected locations. A staircase procedure was used to determine the threshold, and a multiple linear regression model was used to fit the linear cortical magnification factor (CMF). We found that (1) the asymmetry was most pronounced on the vertical and superior azimuths, (2) the asymmetry between the right and left azimuths was not significant, (3) the CMF was significantly smaller on the vertical azimuth than on the horizontal azimuth, (4) the CMF was smaller on the superior vertical azimuth than on the inferior azimuth, and (5) monocular viewing and low contrast enhanced the CMF difference between azimuths. In conclusion, vertical and horizontal azimuths, location of eccentricity, contrast levels of word symbols, and monocular/binocular viewing have different effects on visual field asymmetry and cortical magnification factors.

The role of extrinsic and intrinsic factors in perceptual filling-in of the blind-spot with variegated color and texture stimuli.

Vision scientists dedicated their efforts to unraveling the mechanism of filling-in at the blind-spot (BS) through numerous psychophysical experiments. The prevalent interpretation, emphasizing active filling-in, has spurred extensive research endeavors. In a parallel vein, a pertinent study highlighted the predominance of the nasal Visual Field (VF) over the temporal one and postulated the role of the Cortical Magnification Factor (CMF) in explaining the asymmetry of filling-in. In this study, we first replicated this experiment and then conducted BS-specific psychophysical experiments employing various bi-colored and bi-textured (patterned) stimuli. We observed that nasal dominance is not persistent in the context of the spread of perception for BS filling-in. We posit that the visual information processing priority index (VIPPI), comprising the CMF (an intrinsic factor unaffected by stimulus characteristics) and relative luminance (an extrinsic factor dependent on stimulus characteristics), governs the spread of perception for filling-in in case of diverse neighborhoods of the BS.

Modeling a space-variant cortical representation for apparent motion.

Receptive field sizes of neurons in early primate visual areas increase with eccentricity, as does temporal processing speed. The fovea is evidently specialized for slow, fine movements while the periphery is suited for fast, coarse movements. In either the fovea or periphery discrete flashes can produce motion percepts. Grossberg and Rudd (1989) used traveling Gaussian activity profiles to model long-range apparent motion percepts. We propose a neural model constrained by physiological data to explain how signals from retinal ganglion cells to V1 affect the perception of motion as a function of eccentricity. Our model incorporates cortical magnification, receptive field overlap and scatter, and spatial and temporal response characteristics of retinal ganglion cells for cortical processing of motion. Consistent with the finding of Baker and Braddick (1985), in our model the maximum flash distance that is perceived as an apparent motion (Dmax) increases linearly as a function of eccentricity. Baker and Braddick (1985) made qualitative predictions about the functional significance of both stimulus and visual system parameters that constrain motion perception, such as an increase in the range of detectable motions as a function of eccentricity and the likely role of higher visual processes in determining Dmax. We generate corresponding quantitative predictions for those functional dependencies for individual aspects of motion processing. Simulation results indicate that the early visual pathway can explain the qualitative linear increase of Dmax data without reliance on extrastriate areas, but that those higher visual areas may serve as a modulatory influence on the exact Dmax increase.

Figure-Ground Segmentation and Biological Motion Perception in Peripheral Visual Field.

Biological motion perception is a specific type of perceptual organization, during which a clear image of a moving human body is perceptually generated in virtue of certain core light dots representing the major joint movements. While the processes of biological motion perception have been studied extensively for almost a century, there is still a debate on whether biological motion task performance can be equally precise across all visual field or is central visual field specified for biological motion perception. The current study explores the processes of biological motion perception and figure-ground segmentation in the central and peripheral visual field, expanding the understanding of perceptual organization across different eccentricities. The method involved three different tasks of visual grouping: (1) a static visual grouping task, (2) a dynamic visual grouping task, and (3) a biological motion detection task. The stimuli in (1) and (2) were generated from 12-13 dots grouped by proximity and common fate, and, in (3), light dots representing human motion. All stimuli were embedded in static or dynamics visual noise and the threshold value for the number of noise dots in which the elements could still be grouped by proximity and/or common fate was determined. The results demonstrate that biological motion can be differentiated from the scrambled set of moving dots in a more intensive visual noise than static and dynamic visual grouping tasks. Furthermore, in all three visual tasks (static and dynamic grouping, and biological motion detection) the performance was significantly worse in the periphery than in the central visual field, and object magnification could not compensate for the reduced performance in any of the three grouping tasks. The preliminary results of nine participants indicate that (a) human motion perception involves specific perceptual processes, providing the high-accuracy perception of the human body and (b) the processes of figure-ground segmentation are governed by the bottom-up processes and the best performance can be achieved only when the object is demonstrated in the central visual field.

Visually evoked potentials (VEPs) across the visual field in hearing and deaf cats.

Introduction: Congenitally deaf cats perform better on visual localization tasks than hearing cats, and this advantage has been attributed to the posterior auditory field. Successful visual localization requires both visual processing of the target and timely generation of an action to approach the target. Activation of auditory cortex in deaf subjects during visual localization in the peripheral visual field can occur either via bottom-up stimulus-driven and/or top-down goal-directed pathways. Methods: In this study, we recorded visually evoked potentials (VEPs) in response to a reversing checkerboard stimulus presented in the hemifield contralateral to the recorded hemisphere in both hearing and deaf cats under light anesthesia. Results: Although VEP amplitudes and latencies were systematically modulated by stimulus eccentricity, we found little evidence of changes in VEP in deaf cats that can explain their behavioral advantage. A statistical trend was observed, showing larger peak amplitudes and shorter peak latencies in deaf subjects for stimuli in the near- and mid-peripheral field. Additionally, latency of the P1 wave component had a larger inter-sweep variation in deaf subjects. Discussion: Our results suggested that cross-modal plasticity following deafness does not play a major part in cortical processing of the peripheral visual field when the "vision for action" system is not recruited.

Polar angle asymmetries in visual perception and neural architecture.

Human visual performance changes with visual field location. It is best at the center of gaze and declines with eccentricity, and also varies markedly with polar angle. These perceptual polar angle asymmetries are linked to asymmetries in the organization of the visual system. We review and integrate research quantifying how performance changes with visual field location and how this relates to neural organization at multiple stages of the visual system. We first briefly review how performance varies with eccentricity and the neural foundations of this effect. We then focus on perceptual polar angle asymmetries and their neural foundations. Characterizing perceptual and neural variations across and around the visual field contributes to our understanding of how the brain translates visual signals into neural representations which form the basis of visual perception.

Cortical magnification eliminates differences in contrast sensitivity across but not around the visual field.

Human visual performance changes dramatically both across (eccentricity) and around (polar angle) the visual field. Performance is better at the fovea, decreases with eccentricity, and is better along the horizontal than vertical meridian and along the lower than the upper vertical meridian. However, all neurophysiological and virtually all behavioral studies of cortical magnification have investigated eccentricity effects without considering polar angle. Most performance differences due to eccentricity are eliminated when stimulus size is cortically magnified (M-scaled) to equate the size of its cortical representation in primary visual cortex (V1). But does cortical magnification underlie performance differences around the visual field? Here, to assess contrast sensitivity, human adult observers performed an orientation discrimination task with constant stimulus size at different locations as well as when stimulus size was M-scaled according to stimulus eccentricity and polar angle location. We found that although M-scaling stimulus size eliminates differences across eccentricity, it does not eliminate differences around the polar angle. This finding indicates that limits in contrast sensitivity across eccentricity and around polar angle of the visual field are mediated by different anatomical and computational constraints.

Somatotopic Mapping of the Fingers in the Somatosensory Cortex Using Functional Magnetic Resonance Imaging: A Review of Literature.

Multiple studies have demonstrated finger somatotopy in humans and other primates using a variety of brain mapping techniques including functional magnetic resonance imaging (fMRI). Here, we review the literature to better understand the reliability of fMRI for mapping the somatosensory cortex. We have chosen to focus on the hand and fingers as these areas have the largest representation and have been the subject of the largest number of somatotopic mapping experiments. Regardless of the methods used, individual finger somatosensory maps were found to be organized across Brodmann areas (BAs) 3b, 1, and 2 in lateral-to-medial and inferior-to-superior fashion moving from the thumb to the pinky. However, some consistent discrepancies are found that depend principally on the method used to stimulate the hand and fingers. Therefore, we suggest that a comparative analysis of different types of stimulation be performed to address the differences described in this review.

On the cortical mapping function - Visual space, cortical space, and crowding.

The retino-cortical visual pathway is retinotopically organized: Neighbourhood relationships on the retina are preserved in the mapping. Size relationships in that mapping are also highly regular: The size of a patch in the visual field that maps onto a cortical patch of fixed size follows, along any radius and over a wide range, simply a linear function with retinal eccentricity. As a consequence, the mapping of retinal to cortical locations follows a logarithmic function along that radius. While this has already been shown by Fischer (1973, Vision Research, 13, 2113-2120), the link between the linear function - which describes the local behaviour by the cortical magnification factor M - and the logarithmic location function for the global behaviour, has never been made explicit. The present paper provides such a link as a set of ready-to-use equations using Levi and Klein's E2 nomenclature, and examples for their validity and applicability in the mapping literature are discussed. The equations allow estimating M in the retinotopic centre; values thus derived from the literature show enormous, hitherto unnoticed, variability. A new structural parameter, d2, is proposed to characterize the cortical map, as a counterpart to E2; it shows much more stability. One pitfall is discussed and spelt out, namely the common myth that a pure logarithmic function, without constant term, will give an adequate map. The correct equations are finally extended to describe the cortical map of Bouma's law on visual crowding. The result contradicts recent suggestions that critical crowding distance corresponds to constant cortical distance.

Role of the Dorsal Posterior Parietal Cortex in the Accurate Perception of Object Magnitude in Peripheral Vision.

Following superior parietal lobule and intraparietal sulcus (SPL-IPS) damage, optic ataxia patients underestimate the distance of objects in the ataxic visual field such that they produce hypometric pointing errors. The metrics of these pointing errors relative to visual target eccentricity fit the cortical magnification of central vision. The SPL-IPS would therefore implement an active "peripheral magnification" to match the real metrics of the environment for accurate action. We further hypothesized that this active compensation of the central magnification by the SPL-IPS contributes to actual object' size perception in peripheral vision. Three optic ataxia patients and 10 age-matched controls were assessed in comparing the thickness of two rectangles flashed simultaneously, one in central and another in peripheral vision. The bilateral optic ataxia patient exhibited exaggerated underestimation bias and uncertainty compared to the control group in both visual fields. The two unilateral optic ataxia patients exhibited a pathological asymmetry between visual fields: size perception performance was affected in their contralesional peripheral visual field compared to their healthy side. These results demonstrate that the SPL-IPS contributes to accurate size perception in peripheral vision.

Cortical magnification in human visual cortex parallels task performance around the visual field.

Human vision has striking radial asymmetries, with performance on many tasks varying sharply with stimulus polar angle. Performance is generally better on the horizontal than vertical meridian, and on the lower than upper vertical meridian, and these asymmetries decrease gradually with deviation from the vertical meridian. Here, we report cortical magnification at a fine angular resolution around the visual field. This precision enables comparisons between cortical magnification and behavior, between cortical magnification and retinal cell densities, and between cortical magnification in twin pairs. We show that cortical magnification in the human primary visual cortex, measured in 163 subjects, varies substantially around the visual field, with a pattern similar to behavior. These radial asymmetries in the cortex are larger than those found in the retina, and they are correlated between monozygotic twin pairs. These findings indicate a tight link between cortical topography and behavior, and suggest that visual field asymmetries are partly heritable.

Shape discrimination in peripheral vision: Addressing pragmatic limitations of M-scaling radial frequency patterns.

Peripheral worsening in shape discrimination (SD) can be compensated by size-scaling of peripheral stimuli. However, such scaling results in production of large stimuli that occupy a vast range of eccentricities. We used six proportionally decreasing spatial scales to address this pragmatic limitation and to explore how shape discrimination varies with radius in the nasal visual field. Five participants with normal vision discriminated circles and radial frequency (RF) patterns presented nasally to the fixation point at 5°, 10°, 15° and 20°. Stimuli were scaled with the nasal cortical magnification factor (nCMF) from a central stimulus in six spatial scales, which varied from 0.125 to 1, where 1 corresponded to 1.2° radius. Thresholds expressed in Weber fractions remained constant at eccentricities up to 20° regardless of the spatial scale. Weber fractions for the smaller spatial scales (0.125-0.5) were higher and more variable than for the larger spatial scales (0.75-1), yet still constant across periphery. The results provide evidence that peripheral shape discrimination is constrained by low-level properties, such as eccentricity, and can be predicted by the cortical magnification theory. However, above the peripheral modulation resolution limits, RF shape discrimination is based on the proportion between the modulation amplitude and the radius for larger scales (0.75-1), and demonstrates peripheral scale invariance for these stimuli. For eccentric shape discrimination tests, stimuli with low spatial frequency, high contrast, and radii corresponding to SS 0.75-0.875 should be used to ensure constant Weber fractions, small variability, and peripheral stimuli that are not excessively magnified.

The Mechanism of Macular Sparing.

Patients with homonymous hemianopia sometimes show preservation of the central visual fields, ranging up to 10°. This phenomenon, known as macular sparing, has sparked perpetual controversy. Two main theories have been offered to explain it. The first theory proposes a dual representation of the macula in each hemisphere. After loss of one occipital lobe, the back-up representation in the remaining occipital lobe is postulated to sustain ipsilateral central vision in the blind hemifield. This theory is supported by studies showing that some midline retinal ganglion cells project to the wrong hemisphere, presumably driving neurons in striate cortex that have ipsilateral receptive fields. However, more recent electrophysiological recordings and neuroimaging studies have cast doubt on this theory by showing only a minuscule ipsilateral field representation in early visual cortical areas. The second theory holds that macular sparing arises because the occipital pole, where the macula is represented, remains perfused after occlusion of the posterior cerebral artery because it receives collateral flow from the middle cerebral artery. An objection to this theory is that it cannot account for reports of macular sparing in patients after loss of an entire occipital lobe. On close scrutiny, such reports turn out to be erroneous, arising from inadequate control of fixation during visual field testing. Patients seem able to detect test stimuli on their blind side within the macula or along the vertical meridian because they make surveillance saccades. A purported treatment for hemianopia, called vision restoration therapy, is based on this error. The dual perfusion theory is supported by anatomical studies showing that the middle cerebral artery perfuses the occipital pole in many individuals.In patients with hemianopia from stroke, neuroimaging shows preservation of the occipital pole when macular sparing is present. The frontier dividing the infarcted territory of the posterior cerebral artery and the preserved territory of the middle cerebral artery is variable, but always falls within the representation of the macula, because the macula is so highly magnified. For physicians, macular sparing was an important neurological sign in acute hemianopia because it signified a posterior cerebral artery occlusion. Modern neuroimaging has supplanted the importance of that clinical sign but at the same time confirmed its validity. For patients, macular sparing remains important because it mitigates the impact of hemianopia and preserves the ability to read fluently.

The striate cortex and hemianopia.

We begin with the functions of the striate cortex (area V1 of the visual cortex) and end with a review of the effects of damage to striate cortex or its inputs; namely, homonymous hemifield defects. Clinical and anatomical studies accrued over the past 25 years have modified our understanding of the role of V1 in vision. We discuss the evidence that V1 is not the sole recipient of visual signals; is not the earliest recipient of visual signals; and is not essential for conscious vision. In the second section, we give a brief history of how the visual field was found to be represented in striate cortex, then cover the work that has demonstrated the overrepresentation of the central region of vision in humans. The common patterns of visual field disturbance caused by damage to the retrochiasmal visual system are discussed, with some less common examples shown as brief case studies.

Postnatal development of subfields in the core region of the mouse auditory cortex.

The core region of the rodent auditory cortex has two subfields: the primary auditory area (A1) and the anterior auditory field (AAF). Although the postnatal development of A1 has been studied in several mammalian species, few studies have been conducted on the postnatal development of AAF. Using a voltage-sensitive-dye-based imaging method, we examined and compared the postnatal development of AAF and A1 in mice from postnatal day 11 (P11) to P40. We focused on the postnatal development of tonotopy, the relative position between A1 and AAF, and the properties of tone-evoked responses in the subfields. Tone-evoked responses in the mouse auditory cortex were first observed at P12, and tonotopy was found in both A1 and AAF at this age. Quantification of tonotopy using the cortical magnification factor (CMF; octave difference per unit cortical distance) revealed a rapid change from P12 to P14 in both A1 and AAF, and a stable level from P14. A similar time course of postnatal development was found for the distance between the 4 kHz site in A1 and AAF, the distance between the 16 kHz site in A1 and AAF, and the angle between the frequency axis of A1 and AAF. The maximum amplitude and rise time of tone-evoked signals in both A1 and AAF showed no significant change from P12 to P40, but the latency of the responses to both the 4 kHz and 16 kHz tones decreased during this period, with a more rapid decrease in the latency to 16 kHz tones in both subfields. The duration of responses evoked by 4 kHz tones in both A1 and AAF showed no significant postnatal change, but the duration of responses to 16 kHz tones decreased exponentially in both subfields. The cortical area activated by 4 kHz tones in AAF was always larger than that in A1 at all ages (P12-P40). Our results demonstrated that A1 and AAF developed in parallel postnatally, showing a rapid maturation of tonotopy, slow maturation of response latency and response duration, and a dorsal-to-ventral order (high-frequency site to low-frequency site) of functional maturation.