Here we propose that earlier-demonstrated details in the primate visual cortical map may account for an otherwise puzzling (and problematic) finding in the current human fMRI literature. Specifically, the well-known regions LO and MT(+) reportedly overlap in the human cortical visual map, when those two regions are localized using standard stimulus comparisons in conventional fMRI experiments. Here we describe evidence supporting the idea that the apparent functional overlap between LO and MT arises from a third area (the MT crescent: “MTc”), which is well known to surround posterior MT based on earlier histological, neuroanatomical, and electrophysiological studies in nonhuman primates. If we assume that MTc also exists in human visual cortex, and that it has a location and functional properties intermediate to those in LO and MT, simplistic modeling confirmed that this arrangement could produce apparent overlap between localizers for LO and MT in conventional fMRI maps in human visual cortex.

The nucleoredoxin-like gene Nxnl1 (Txnl6) and its paralogue Nxnl2 encode the rod-derived cone viability factors (RdCVF and RdCVF2), which increase the resistance to photooxidative damage and have therapeutic potential for the survival of cones in retinitis pigmentosa. In this study, the transcription of Nxnl genes was investigated as a function of the day/night cycle in rats. The transcript levels of Nxnl1 and Nxnl2 were seen to display daily rhythms with steadily increasing values during the light phase and peak expression around dark onset in preparations of whole retina, photoreceptor cells and—but only in regard to Nxnl1—in photoreceptor-related pinealocytes. The cycling of Nxnl1 but not that of Nxnl2 persisted in constant darkness in the retina. This suggests that daily regulation of Nxnl1 is driven by a circadian clock, whereas that of Nxnl2 is promoted by environmental light. The present data indicate clock- and light-dependent regulations of nucleoredoxin-like genes that may be part of a protective shield against photooxidative damage.

Early in development, before the onset of vision, the retina establishes direction-selective responses. During this time period, the retina spontaneously generates bursts of action potentials that propagate across its extent. The precise spatial and temporal properties of these “retinal waves” have been implicated in the formation of retinal projections to the brain. However, their role in the development of direction selective circuits within the retina has not yet been determined. We addressed this issue by combining multielectrode array and cell-attached recordings to examine mice that lack the CaV3.2 subunit of T-type Ca2+ channels (CaV3.2 KO) because these mice exhibit disrupted waves during the period that direction selective circuits are established. We found that the spontaneous activity of these mice displays wave-associated bursts of action potentials that are altered from that of control mice: the frequency of these bursts is significantly decreased and the firing rate within each burst is reduced. Moreover, the projection patterns of the retina demonstrate decreased eye-specific segregation in the dorsal lateral geniculate nucleus (dLGN). However, after eye-opening, the direction selective responses of CaV3.2 KO direction selective ganglion cells (DSGCs) are indistinguishable from those of wild-type DSGCs. Our data indicate that although the temporal properties of the action potential bursts associated with retinal waves are important for activity-dependent refining of retinal projections to central targets, they are not critical for establishing direction selectivity in the retina.

Electroretinogram (ERG) studies identified a new mouse line with a normal a-wave but lacking the b-wave component. The ERG phenotype of this new allele, nob7, matched closely that of mouse mutants for Grm6, Lrit3, Trpm1, and Nyx, which encode for proteins expressed in depolarizing bipolar cells (DBCs). To identify the underlying mutation, we first crossed nob7 mice with Grm6nob3 mutants and measured the ERGs in offspring. All the offspring lacked the b-wave, indicating that nob7 is a new allele for Grm6: Grm6nob7. Sequence analyses of Grm6nob7 cDNAs identified a 28 base pair insertion between exons 8 and 9, which would result in a frameshift mutation in the open reading frame that encodes the metabotropic glutamate receptor 6 (Grm6). Sequencing both the cDNA and genomic DNA from exon 8 and intron 8, respectively, from the Grm6nob7 mouse revealed a G to A transition at the last position in exon 8. This mutation disrupts splicing and the normal exon 8 is extended by 28 base pairs, because splicing occurs 28 base pairs downstream at a cryptic splice donor. Consistent with the impact of the resulting frameshift mutation, there is a loss of mGluR6 protein (encoded by Grm6) from the dendritic tips of DBCs in the Grm6nob7 retina. These results indicate that Grm6nob7 is a new model of the complete form of congenital stationary night blindness, a human condition that has been linked to mutations of GRM6.

Unlike the mammalian retina, the teleost fish retina undergoes persistent neurogenesis from intrinsic stem cells. In marine teleosts, most cone photoreceptor genesis occurs early in the embryonic and larval stages, and rods are added primarily during and after metamorphosis. This study demonstrates a developmental paradigm in elopomorph fishes in which retinas are rod-dominated in larvae, but undergo periods of later cone genesis. Retinal characteristics were compared at different developmental stages among three ecologically distinct elopomorph fishes—ladyfish (Elops saurus), bonefish (Albula vulpes), and speckled worm eel (Myrophis punctatus). The objectives were to improve our understanding of (1) the developmental strategy in the elopomorph retina, (2) the functional architecture of the retina as it relates to ecology, and (3) how the light environment influences photoreceptor genesis. Photoreceptor morphologies, distributions, and spectral absorption were studied at larval, juvenile, and adult stages. Premetamorphic retinas in all three species are rod-dominated, but the retinas of these species undergo dramatic change over the course of development, resulting in juvenile and adult retinal characteristics that correlate closely with ecology. Adult E. saurus has high rod densities, grouped photoreceptors, a reflective tapetum, and longer-wavelength photopigments, supporting vision in turbid, low-light conditions. Adult A. vulpes has high cone densities, low rod densities, and shorter-wavelength photopigments, supporting diurnal vision in shallow, clear water. M. punctatus loses cones during metamorphosis, develops new cones after settlement, and maintains high rod but low cone densities, supporting primarily nocturnal vision. M. punctatus secondary cone genesis occurs rapidly throughout the retina, suggesting a novel mechanism of vertebrate photoreceptor genesis. Finally, in postsettlement M. punctatus, the continuous presence or absence of visible light modulates rod distribution but does not affect secondary cone genesis, suggesting some degree of developmental plasticity influenced by the light environment.

The goals of these experiments were to describe the morphology and synaptic connections of amacrine cells in the baboon retina that contain immunoreactive vesicular glutamate transporter 3 (vGluT3). These amacrine cells had the morphology characteristic of knotty bistratified type 1 cells, and their dendrites formed two plexuses on either side of the center of the inner plexiform layer. The primary dendrites received large synapses from amacrine cells, and the higher-order dendrites were both pre- and postsynaptic to other amacrine cells. Based on light microscopic immunolabeling results, these include AII cells and starburst cells, but not the polyaxonal amacrine cells tracer-coupled to ON parasol ganglion cells. The vGluT3 cells received input from ON bipolar cells at ribbon synapses and made synapses onto OFF bipolar cells, including the diffuse DB3a type. Many synapses from vGluT3 cells onto retinal ganglion cells were observed in both plexuses. At synapses where vGluT3 cells were presynaptic, two types of postsynaptic densities were observed; there were relatively thin ones characteristic of inhibitory synapses and relatively thick ones characteristic of excitatory synapses. In the light microscopic experiments with Neurobiotin-injected ganglion cells, vGluT3 cells made contacts with midget and parasol ganglion cells, including both ON and OFF types. Puncta containing immunoreactive gephyrin, an inhibitory synapse marker, were found at appositions between vGluT3 cells and each of the four types of labeled ganglion cells. The vGluT3 cells did not have detectable levels of immunoreactive γ-aminobutyric acid (GABA) or immunoreactive glycine transporter 1. Thus, the vGluT3 cells would be expected to have ON responses to light and make synapses onto neurons in both the ON and the OFF pathways. Taken with previous results, these findings suggest that vGluT3 cells release glycine at some of their output synapses and glutamate at others.

In macaque, it has long been known since the late nineties that the medial parieto-occipital sulcus (POS) contains two regions, V6 and V6A, important for visual motion and action. While V6 is a retinotopically organized extrastriate area, V6A is a broadly retinotopically organized visuomotor area constituted by a ventral and dorsal subdivision (V6Av and V6Ad), both containing arm movement-related cells active during spatially directed reaching movements. In humans, these areas have been mapped only in recent years thanks to neuroimaging methods. In a series of brain mapping studies, by using a combination of functional magnetic resonance imaging methods such as wide-field retinotopy and task-evoked activity, we mapped human areas V6 (Pitzalis et al., 2006) and V6Av (Pitzalis et al., 2013d) retinotopically and defined human V6Ad functionally as a pointing-selective region situated anteriorly in the close proximity of V6Av (Tosoni et al., 2014). Like in macaque, human V6 is a motion area (e.g., Pitzalis et al., 2010, 2012, 2013a,b,c), while V6Av and V6Ad respond to pointing movements (Tosoni et al., 2014). The retinotopic organization (when present), anatomical position, neighbor relations, and functional properties of these three areas closely resemble those reported for macaque V6 (Galletti et al., 1996, 1999a), V6Av, and V6Ad (Galletti et al., 1999b; Gamberini et al., 2011). We suggest that information on objects in depth which are translating in space, because of the self-motion, is processed in V6 and conveyed to V6A for evaluating object distance in a dynamic condition such as that created by self-motion, so to orchestrate the eye and arm movements necessary to reach or avoid static and moving objects in the environment.

Visual alterations, peripheral light loss (PLL) and blackout (BO), are components of acceleration (+Gz) induced loss of consciousness (LOC) and recovery of consciousness (ROC). The kinetics of loss of vision (LOV) and recovery of vision (ROV) were determined utilizing ocular pressure induced retinal ischemia and compared to the kinetics of LOC and ROC resulting from +Gz-induced cephalic nervous system (CPNS) ischemia. The time from self-induced retinal ischemia in completely healthy subjects (N = 104) to the onset of PLL and complete BO was measured. The time from release of ocular pressure, with return of normal retinal circulation, to the time for complete recovery of visual fields was also measured. The kinetics of pressure induced LOV and ROV was compared with previously developed kinetics of +Gz-induced LOC and ROC focusing on the rapid onset, vertical arm, of the +Gz-induced LOC and ROC curves. The time from onset of increased ocular pressure, immediately inducing retinal ischemia, to PLL was 5.04 s with the time to BO being 8.73 s. Complete recovery of the visual field from BO following release of ocular pressure, immediately abolishing retinal ischemia, was 2.74 s. These results confirm experimental findings that visual loss is frequently not experienced prior to LOC during exposure to rapid onset, high levels of +Gz-stress above tolerance. Offset of pressure induced retinal ischemia to ROV was 2.74 s, while the time from offset of +Gz-induced CPNS ischemia to ROC was 5.29 s. Recovery of retinal function would be predicted to be complete before consciousness is regained following +Gz-induced LOC. Ischemia onset time normalization in neurologic tissues permits comparison between different stress-induced times to altered function. The +Gz-time tolerance curves for LOV and LOC provide comparison and integration of neurologic state transition kinetics in the retina and CPNS.

The gap-junction-forming protein connexin36 (Cx36) represents the anatomical substrate of photoreceptor electrical coupling in mammals. The strength of coupling is directly correlated to the phosphorylation of Cx36 at two regulatory sites: Ser110 and Ser293. Our previous work demonstrated that the extent of biotinylated tracer coupling between photoreceptor cells, which provides an index of the extent of electrical coupling, depends on the mouse strain. In the C57Bl/6J strain, light or dopamine reduces tracer coupling and Cx36 phosphorylation in photoreceptors. Conversely, darkness or a dopaminergic antagonist increases tracer coupling and Cx36 phosphorylation, regardless of the daytime. In the CBA/CaJ strain, photoreceptor tracer coupling is not only regulated by light and dopamine, but also by a circadian clock, a type of oscillator with a period close to 24 h and intrinsic to the retina, so that under prolonged dark-adapted conditions tracer coupling is broader at night compared to daytime. In the current study, we examined whether the modulation of photoreceptor coupling by a circadian clock in the CBA/CaJ mouse photoreceptors reflected a change in Cx36 protein expression and/or phosphorylation. We found no significant change in Cx36 expression or in the number of Cx36 gap junction among the conditions examined. However, we found that Cx36 phosphorylation is higher under dark-adapted conditions at night than in the daytime, and is the lowest under prolonged illumination at any time of the day/night cycle. Our observations are consistent with the view that the circadian clock regulation of photoreceptor electrical coupling is mouse strain-dependent and highlights the critical position of Cx36 phosphorylation in the control of photoreceptor coupling.

As highlighted by several contributions to this special issue, there is still ongoing debate about the number, exact location, and boundaries of the visual areas located in cortex immediately rostral to the second visual area (V2), i.e., the “third tier” visual cortex, in primates. In this review, we provide a historical overview of the main ideas that have led to four models of third tier cortex organization, which are at the center of today's debate. We formulate specific predictions of these models, and compare these predictions with experimental evidence obtained primarily in New World primates. From this analysis, we conclude that only one of these models (the “multiple-areas” model) can accommodate the breadth of available experimental evidence. According to this model, most of the third tier cortex in New World primates is occupied by two distinct areas, both representing the full contralateral visual quadrant: the dorsomedial area (DM), restricted to the dorsal half of the third visual complex, and the ventrolateral posterior area (VLP), occupying its ventral half and a substantial fraction of its dorsal half. DM belongs to the dorsal stream of visual processing, and overlaps with macaque parietooccipital (PO) area (or V6), whereas VLP belongs to the ventral stream and overlaps considerably with area V3 proposed by others. In contrast, there is substantial evidence that is inconsistent with the concept of a single elongated area V3 lining much of V2. We also review the experimental evidence from macaque monkey and humans, and propose that, once the data are interpreted within an evolutionary-developmental context, these species share a homologous (but not necessarily identical) organization of the third tier cortex as that observed in New World monkeys. Finally, we identify outstanding issues, and propose experiments to resolve them, highlighting in particular the need for more extensive, hypothesis-driven investigations in macaque and humans.

The second-order visual mechanisms perform the operation of integrating the spatially distributed local visual information. Their organization is traditionally considered within the framework of the filter-rectify-filter model. These are the second-order filters that provide the ability to detect texture gradients. However, the question of the mechanisms' selectivity to the modulation dimension remains open. The aim of this investigation is to answer the above question by using visual evoked potentials (VEPs). Stimuli were textures consisting of staggered Gabor patches. The base texture was nonmodulated (NM). Three other textures represented the base texture which was sinusoidally modulated in different dimensions: contrast, orientation, or spatial frequency. EEG was recorded with 20 electrodes. VEPs of 500 ms duration were obtained for each of the four textures. After that, VEP to the NM texture was subtracted from VEP to each modulated texture. As a result, three different waves (d-waves) were obtained for each electrode site. Each d-wave was then averaged across all the 48 observers. The revealed d-waves have a latency of about 200 ms and, in our opinion, reflect the second-order filters reactivation through the feedback connection. The d-waves for different modulation dimensions were compared with each other in time, amplitude, topography, and localization of the sources of activity that causes the d-wave (with sLORETA). We proceeded from the assumption that the d-wave (its first component) represents functioning of the second-order visual mechanisms and activity changes at the following processing stages. It was found that the d-waves for different modulation dimensions significantly differ in all parameters. The obtained results indicate that the spatial modulations of different texture parameters caused specific changes in the brain activity, which could be evidence supporting the specificity of the second-order visual mechanisms to modulation dimension.

The organization of the cortex located immediately anterior to the second visual area (V2), i.e., the third tier visual cortex, remains controversial, especially in New World primates. In particular, there is lack of consensus regarding the exact location and extent of the lower visual quadrant representation of the third visual area V3 (or ventrolateral posterior –VLP – of a different nomenclature). Microelectrode and connectional mapping studies have revealed the existence of an upper visual quadrant representation abutting dorsal V2 anteriorly, and bordered medially and laterally by representations of the lower visual quadrant. It remains unclear whether these lower field regions are both part of a single area V3, which is split into two patches by an interposed region of upper field representation, or whether they are the lower field representations of two different areas, the dorsomedial area (DM) and area V3/VLP, respectively. To address this question, we quantitatively analyzed the patterns of corticocortical afferent connections labeled by tracer injections targeted to these two lower field regions in the dorsal aspect of the third tier cortex. We found different inter-areal connectivity patterns arising from these two regions, strongly suggesting that they belong to two different visual areas. In particular, our results indicate that the dorsal aspect of the third tier cortex consists of two distinct areas: a full area DM, representing the lower quadrant medially, and the upper quadrant laterally, and the lower quadrant representation of V3/VLP, located laterally to upper field DM. DM is predominantly connected with areas of the dorsal visual stream, and V3/VLP with areas of the ventral stream. These results prompt further functional investigations of the third tier cortex, as previous studies of this cortical territory may have pooled response properties of two very different areas into a single area V3.

The number, location, extent, and functional properties of the cortical areas that occupy the medial parieto-occipital cortex (mPOC) have been, and still is, a matter of scientific debate. The mPOC is a convoluted region of the brain that presents a high level of individual variability, and the fact that many areas of mPOC are located within very deep sulci further limits the possibility to investigate their anatomo-functional properties. In the present review, we summarize the location and extent of mPOC areas in the macaque brain as obtained by architectural, connectional, and functional data. The different approaches lead to a subdivision of mPOC that includes areas V2, V3, V6, V6Av, and V6Ad. Extrastriate areas V2 and V3 occupy the posterior wall of the parieto-occipital sulcus (POs). The fundus of POs and the ventralmost part of the anterior wall of the sulcus are occupied by a retinotopically organized visual area, called V6, which represents the contralateral part of the visual field and emphasizes its periphery. The remaining part of the anterior wall of POs is occupied by two areas, V6Av and V6Ad, which contain visual as well as arm reaching neurons. Our analyses suggest that areas V6 and V6Av, together, occupy the cortical territory previously described as area PO. Functionally, area V6 is a motion area particularly sensitive to the real motion of objects in the animal's field of view, while V6Av and V6Ad are visuomotor areas likely involved in the visual guidance of arm movement and object prehension.

Areas V3 and V4 are commonly thought of as individual entities in the primate visual system, based on definition criteria such as their representation of visual space, connectivity, functional response properties, and relative anatomical location in cortex. Yet, large-scale functional and anatomical organization patterns not only emphasize distinctions within each area, but also links across visual cortex. Specifically, the visuotopic organization of V3 and V4 appears to be part of a larger, supra-areal organization, clustering these areas with early visual areas V1 and V2. In addition, connectivity patterns across visual cortex appear to vary within these areas as a function of their supra-areal eccentricity organization. This complicates the traditional view of these regions as individual functional “areas.” Here, we will review the criteria for defining areas V3 and V4 and will discuss functional and anatomical studies in humans and monkeys that emphasize the integration of individual visual areas into broad, supra-areal clusters that work in concert for a common computational goal. Specifically, we propose that the visuotopic organization of V3 and V4, which provides the criteria for differentiating these areas, also unifies these areas into the supra-areal organization of early visual cortex. We propose that V3 and V4 play a critical role in this supra-areal organization by filtering information about the visual environment along parallel pathways across higher-order cortex.

Inferring neural responses from functional magnetic resonance imaging (fMRI) data is challenging. Even if we take advantage of high-field systems to acquire data with submillimeter resolution, we are still acquiring data in which a single datum summarizes the responses of tens of thousands of neurons. Excitation and inhibition, spikes and subthreshold membrane potential modulations, local and long-range computations, and tuned and nonselective responses are mixed together in one signal. With a priori knowledge of the underlying neural population responses, careful experiment design allows us to manipulate the experiment or task design so that subpopulations are selectively modulated, and our experiments can reveal those tuning functions. However, because we want to be able to use fMRI to discover new kinds of tuning functions and selectivity, we cannot limit ourselves to experiments in which we already know what we are looking for. Broadly speaking, analyses that rely on classification of responses that are distributed across the local neural population [multi-voxel pattern analyses (MVPA)] offer the ability to discover new kinds of information representation and selectivities in neural subpopulations. There is, however, no way to determine how the information discovered with MVPA or other analyses is related to the underlying neuronal tuning functions. Therefore, we must continue to rely on behavioral, computational, and animal models to develop theories of information representation in mid-tier visual cortical areas. Once encoding models exist, fMRI can be powerful for testing these a priori models of information representation. As an aide in developing these models, an important contribution that fMRI can make to our understanding of mid-tier visual areas is derived from connectivity analyses and experiments that study information sharing between visual areas. This ability to quantify localized population average responses throughout the brain is the strength we can best leverage to discover new properties of local and long-range neural networks.

In primates, the cortex adjoining the rostral border of V2 has been variously interpreted as belonging to a single visual area, V3, with dorsal V3 (V3d) representing the lower visual quadrant and ventral V3 (V3v) representing the upper visual quadrant, V3d and V3v constituting separate, incomplete visual areas, V3d and ventral posterior (VP), or V3d being divided into several visual areas, including a dorsomedial (DM) visual area, a medial visual area (M), and dorsal extension of VP (or VLP). In our view, the evidence from V1 connections strongly supports the contention that V3v and V3d are parts of a single visual area, V3, and that DM is a separate visual area along the rostral border of V3d. In addition, the retinotopy revealed by V1 connection patterns, microelectrode mapping, optical imaging mapping, and functional magnetic resonance imaging (fmri) mapping indicates that much of the proposed territory of V3d corresponds to V3. Yet, other evidence from microelectrode mapping and anatomical connection patterns supports the possibility of an upper quadrant representation along the rostral border of the middle of dorsal V2 (V2d), interpreted as part of DM or DM plus DI, and along the midline end of V2d, interpreted as the visual area M. While the data supporting these different interpretations appear contradictory, they also seem, to some extent, valid. We suggest that V3d may have a gap in its middle, possibly representing part of the upper visual quadrant that is not part of DM. In addition, another visual area, M, is likely located at the DM tip of V3d. There is no evidence for a similar disruption of V3v. For the present, we favor continuing the traditional concept of V3 with the possible modification of a gap in V3d in at least some primates.

A basic principle in visual neuroscience is the retinotopic organization of neural receptive fields. Here, we review behavioral, neurophysiological, and neuroimaging evidence for nonretinotopic processing of visual stimuli. A number of behavioral studies have shown perception depending on object or external-space coordinate systems, in addition to retinal coordinates. Both single-cell neurophysiology and neuroimaging have provided evidence for the modulation of neural firing by gaze position and processing of visual information based on craniotopic or spatiotopic coordinates. Transient remapping of the spatial and temporal properties of neurons contingent on saccadic eye movements has been demonstrated in visual cortex, as well as frontal and parietal areas involved in saliency/priority maps, and is a good candidate to mediate some of the spatial invariance demonstrated by perception. Recent studies suggest that spatiotopic selectivity depends on a low spatial resolution system of maps that operates over a longer time frame than retinotopic processing and is strongly modulated by high-level cognitive factors such as attention. The interaction of an initial and rapid retinotopic processing stage, tied to new fixations, and a longer lasting but less precise nonretinotopic level of visual representation could underlie the perception of both a detailed and a stable visual world across saccadic eye movements.

This study defined the amplitude and phase characteristics of rod- and cone-isolated flicker electroretinograms (ERGs) and determined how these responses summate to generate the nonreceptor-specific ERG. Full-field ERGs were obtained from six normally sighted subjects (age 26 to 44 years) using a four-primary LED-based photostimulator and standard recording techniques. The four primaries were either modulated sinusoidally in phase to achieve simultaneous rod and cone activation (ERGR+C; nonreceptor-specific) or in different phases to achieve rod-isolated (ERGR) and cone-isolated (ERGC) responses by means of triple silent substitution. ERGs were measured at two mean luminance levels (2.4 and 24 cd/m2), two contrasts (20 and 40%), and four temporal frequencies (2–15 Hz). Fundamental amplitude and phase for each condition were derived by Fourier analysis. Response amplitude and phase depended on the stimulus conditions (frequency, mean luminance, and contrast), however, for all conditions: 1) response phase decreased monotonically as stimulus frequency increased; 2) response amplitude tended to decrease monotonically as stimulus frequency increased, with the exception of the 24 cd/m2, 40% contrast ERGR+C that was sharply V-shaped; 3) ERGR phase was delayed (32 to 210 deg) relative to the ERGC phase; 4) ERGR amplitude was typically equal to or lower than the ERGC amplitude, with the exception of the 2.4 cd/m2, 40% contrast condition; and 5) the pattern of ERGR+C responses could be accounted for by a vector summation model of the rod and cone pathway signals. The results show that the ERGR+C amplitude and phase can be predicted from ERGR and ERGC amplitude and phase. For conditions that elicit ERGR and ERGC responses that have approximately equal amplitude and opposite phase, there is strong destructive interference between the rod and cone responses that attenuates the ERGR+C. Conditions that elicit equal amplitude and opposite phase rod and cone responses may be particularly useful for evaluating rod–cone interactions.

Anatomical and electrophysiological studies have provided us with detailed information regarding the extent and topography of the primary (V1) and secondary (V2) visual areas in primates. The consensus about the V1 and V2 maps, however, is in sharp contrast with controversies regarding the organization of the cortical areas lying immediately rostral to V2. In this review, we address the contentious issue of the extent of the third visual area (V3). Specifically, we will argue for the existence of both ventral (V3v) and dorsal (V3d) segments of V3, which are located, respectively, adjacent to the anterior border of ventral and dorsal V2. V3v and V3d would together constitute a single functional area with a complete representation of both upper and lower visual hemifields. Another contentious issue is the organization of the parietal-occipital (PO) area, which also borders the rostral edge of the medial portion of dorsal V2. Different from V1, V2, and V3, which exhibit a topography based on the defined lines of isoeccentricity and isopolar representation, area PO only has a systematic representation of polar angles, with an emphasis on the peripheral visual field (isoeccentricity lines are not well defined). Based on the connectivity patterns of area PO with distinct cytochrome oxidase modules in V2, we propose a subdivision of the dorsal stream of visual information processing into lateral and medial domains. In this model, area PO constitutes the first processing instance of the dorsal-medial stream, coding for the full-field flow of visual cues during navigation. Finally, we compare our findings with those in other species of Old and New World monkeys and argue that larger animals, such as macaque and capuchin monkeys, have similar organizations of the areas rostral to V2, which is different from that in smaller New World monkeys.

The ventral surface of the human occipital lobe contains multiple retinotopic maps. The most posterior of these maps is considered a potential homolog of macaque V4, and referred to as human V4 (“hV4”). The location of the hV4 map, its retinotopic organization, its role in visual encoding, and the cortical areas it borders have been the subject of considerable investigation and debate over the last 25 years. We review the history of this map and adjacent maps in ventral occipital cortex, and consider the different hypotheses for how these ventral occipital maps are organized. Advances in neuroimaging, computational modeling, and characterization of the nearby anatomical landmarks and functional brain areas have improved our understanding of where human V4 is and what kind of visual representations it contains.