Circuits and Mechanisms for Surround Modulation in Visual Cortex.

Surround modulation (SM) is a fundamental property of sensory neurons in many species and sensory modalities. SM is the ability of stimuli in the surround of a neuron's receptive field (RF) to modulate (typically suppress) the neuron's response to stimuli simultaneously presented inside the RF, a property thought to underlie optimal coding of sensory information and important perceptual functions. Understanding the circuit and mechanisms for SM can reveal fundamental principles of computations in sensory cortices, from mouse to human. Current debate is centered over whether feedforward or intracortical circuits generate SM, and whether this results from increased inhibition or reduced excitation. Here we present a working hypothesis, based on theoretical and experimental evidence, that SM results from feedforward, horizontal, and feedback interactions with local recurrent connections, via synaptic mechanisms involving both increased inhibition and reduced recurrent excitation. In particular, strong and balanced recurrent excitatory and inhibitory circuits play a crucial role in the computation of SM.

Patient characteristics associated with the acceptability of teleconsultation: a retrospective study of osteoporotic patients post-COVID-19.

BACKGROUND: Due to the COVID-19 pandemic, teleconsultations (TCs) have become common practice for many chronic conditions, including osteoporosis. While satisfaction with TCs among patients increases in times of emergency, we have little knowledge of whether the acceptability of TCs persists once in-person visits return to being a feasible and safe option. In this study, we assess the acceptability of TCs across five dimensions for osteoporosis care among patients who started or continued with TCs after the COVID-19 pandemic had waned. We then explore the patient characteristics associated with these perceptions. METHODS: Between January and April 2022, 80 osteoporotic patients treated at the Humanitas Hospital in Milan, Italy, were recruited to answer an online questionnaire about the acceptability of TCs for their care. The acceptability of TCs was measured using a modified version of the Service User Technology Acceptability Questionnaire (SUTAQ), which identifies five domains of acceptability: perceived benefits, satisfaction, substitution, privacy and discomfort, and care personnel concerns. Multivariable ordinary least squares (OLS) linear regression analysis was performed to assess which patient characteristics in terms of demographics, socio-economic conditions, digital skills, social support, clinical characteristics and pattern of TC use were correlated with the five domains of acceptability measured through the SUTAQ. RESULTS: The degree of acceptability of TCs was overall good across the 80 respondents and the five domains. Some heterogeneity in perceptions emerged with respect to TCs substituting for in-person visits, negatively impacting continuity of care and reducing the length of consultations. For the most part, acceptability was not affected by patient characteristics with a few exceptions related to treatment time and familiarity with the TC service modality (i.e., length of osteoporosis treatment and number of TCs experienced by the patient). CONCLUSIONS: TCs appear to be an acceptable option for osteoporosis care in the aftermath of the COVID-19 pandemic. This study suggests that other characteristics besides age, digital skills and social support, which are traditionally relevant to TC acceptability, should be taken into account in order to better target this care delivery modality.

Fitbit Data to Assess Functional Capacity in Patients Before Elective Surgery: Pilot Prospective Observational Study.

BACKGROUND: Preoperative assessment is crucial to prevent the risk of complications of surgical operations and is usually focused on functional capacity. The increasing availability of wearable devices (smartwatches, trackers, rings, etc) can provide less intrusive assessment methods, reduce costs, and improve accuracy. OBJECTIVE: The aim of this study was to present and evaluate the possibility of using commercial smartwatch data, such as those retrieved from the Fitbit Inspire 2 device, to assess functional capacity before elective surgery and correlate such data with the current gold standard measure, the 6-Minute Walk Test (6MWT) distance. METHODS: During the hospital visit, patients were evaluated in terms of functional capacity using the 6MWT. Patients were asked to wear the Fitbit Inspire 2 for 7 days (with flexibility of -2 to +2 days) after the hospital visit, before their surgical operation. Resting heart rate and daily steps data were retrieved directly from the smartwatch. Feature engineering techniques allowed the extraction of heart rate over steps (HROS) and a modified version of Non-Exercise Testing Cardiorespiratory Fitness. All measures were correlated with 6MWT. RESULTS: In total, 31 patients were enrolled in the study (n=22, 71% men; n=9, 29% women; mean age 76.06, SD 4.75 years). Data were collected between June 2021 and May 2022. The parameter that correlated best with the 6MWT was the Non-Exercise Testing Cardiorespiratory Fitness index (r=0.68; P<.001). The average resting heart rate over the whole acquisition period for each participant had r=-0.39 (P=.03), even if some patients did not wear the device at night. The correlation of the 6MWT distance with the HROS evaluated at 1% quantile was significant, with Pearson coefficient of -0.39 (P=.04). Fitbit step count had a fair correlation of 0.59 with 6MWT (P<.001). CONCLUSIONS: Our study is a promising starting point for the adoption of wearable technology in the evaluation of functional capacity of patients, which was strongly correlated with the gold standard. The study also identified limitations in the availability of metrics, variability of devices, accuracy and quality of data, and accessibility as crucial areas of focus for future studies.

Electrical Impedance Tomography: From the Traditional Design to the Novel Frontier of Wearables.

Electrical impedance tomography (EIT) is a medical imaging technique based on the injection of a current or voltage pattern through electrodes on the skin of the patient, and on the reconstruction of the internal conductivity distribution from the voltages collected by the electrodes. Compared to other imaging techniques, EIT shows significant advantages: it does not use ionizing radiation, is non-invasive and is characterized by high temporal resolution. Moreover, its low cost and high portability make it suitable for real-time, bedside monitoring. However, EIT is also characterized by some technical limitations that cause poor spatial resolution. The possibility to design wearable devices based on EIT has recently given a boost to this technology. In this paper we reviewed EIT physical principles, hardware design and major clinical applications, from the classical to a wearable setup. A wireless and wearable EIT system seems a promising frontier of this technology, as it can both facilitate making clinical measurements and open novel scenarios to EIT systems, such as home monitoring.

A Scoping Review on Wearable Devices for Environmental Monitoring and Their Application for Health and Wellness.

This scoping review is focused on wearable devices for environmental monitoring. First, the main pollutants are presented, followed by sensing technologies that are used for the parameters of interest. Selected examples of wearables and portables are divided into commercially available and research-level projects. While many commercial products are in fact portable, there is an increasing interest in using a completely wearable technology. This allows us to correlate the pollution level to other personal information (performed activity, position, and respiratory parameters) and thus to estimate personal exposure to given pollutants. The fact that there are no univocal indices to estimate outdoor or indoor air quality is also an open problem. Finally, applications of wearables for environmental monitoring are discussed. Combining environmental monitoring with other devices would permit better choices of where to perform sports activities, especially in highly polluted areas, and provide detailed information on the living conditions of individuals.

Anatomy and Physiology of Macaque Visual Cortical Areas V1, V2, and V5/MT: Bases for Biologically Realistic Models.

The cerebral cortex of primates encompasses multiple anatomically and physiologically distinct areas processing visual information. Areas V1, V2, and V5/MT are conserved across mammals and are central for visual behavior. To facilitate the generation of biologically accurate computational models of primate early visual processing, here we provide an overview of over 350 published studies of these three areas in the genus Macaca, whose visual system provides the closest model for human vision. The literature reports 14 anatomical connection types from the lateral geniculate nucleus of the thalamus to V1 having distinct layers of origin or termination, and 194 connection types between V1, V2, and V5, forming multiple parallel and interacting visual processing streams. Moreover, within V1, there are reports of 286 and 120 types of intrinsic excitatory and inhibitory connections, respectively. Physiologically, tuning of neuronal responses to 11 types of visual stimulus parameters has been consistently reported. Overall, the optimal spatial frequency (SF) of constituent neurons decreases with cortical hierarchy. Moreover, V5 neurons are distinct from neurons in other areas for their higher direction selectivity, higher contrast sensitivity, higher temporal frequency tuning, and wider SF bandwidth. We also discuss currently unavailable data that could be useful for biologically accurate models.

Smart Textiles and Sensorized Garments for Physiological Monitoring: A Review of Available Solutions and Techniques.

Several wearable devices for physiological and activity monitoring are found on the market, but most of them only allow spot measurements. However, the continuous detection of physiological parameters without any constriction in time or space would be useful in several fields such as healthcare, fitness, and work. This can be achieved with the application of textile technologies for sensorized garments, where the sensors are completely embedded in the fabric. The complete integration of sensors in the fabric leads to several manufacturing techniques that allow dealing with both the technological challenges entailed by the physiological parameters under investigation, and the basic requirements of a garment such as perspiration, washability, and comfort. This review is intended to provide a detailed description of the textile technologies in terms of materials and manufacturing processes employed in the production of sensorized fabrics. The focus is pointed at the technical challenges and the advanced solutions introduced with respect to conventional sensors for recording different physiological parameters, and some interesting textile implementations for the acquisition of biopotentials, respiratory parameters, temperature and sweat are proposed. In the last section, an overview of the main garments on the market is depicted, also exploring some relevant projects under development.

Comparison between PtCO2 and PaCO2 and Derived Parameters in Heart Failure Patients during Exercise: A Preliminary Study.

Evaluation of arterial carbon dioxide pressure (PaCO2) and dead space to tidal volume ratio (VD/VT) during exercise is important for the identification of exercise limitation causes in heart failure (HF). However, repeated sampling of arterial or arterialized ear lobe capillary blood may be clumsy. The aim of our study was to estimate PaCO2 by means of a non-invasive technique, transcutaneous PCO2 (PtCO2), and to verify the correlation between PtCO2 and PaCO2 and between their derived parameters, such as VD/VT, during exercise in HF patients. 29 cardiopulmonary exercise tests (CPET) performed on a bike with a ramp protocol aimed at achieving maximal effort in ≈10 min were analyzed. PaCO2 and PtCO2 values were collected at rest and every 2 min during active pedaling. The uncertainty of PCO2 and VD/VT measurements were determined by analyzing the error between the two methods. The accuracy of PtCO2 measurements vs. PaCO2 decreases towards the end of exercise. Therefore, a correction to PtCO2 that keeps into account the time of the measurement was implemented with a multiple regression model. PtCO2 and VD/VT changes at 6, 8 and 10 min vs. 2 min data were evaluated before and after PtCO2 correction. PtCO2 overestimates PaCO2 for high timestamps (median error 2.45, IQR -0.635-5.405, at 10 min vs. 2 min, p-value = 0.011), while the error is negligible after correction (median error 0.50, IQR = -2.21-3.19, p-value > 0.05). The correction allows removing differences also in PCO2 and VD/VT changes. In HF patients PtCO2 is a reliable PaCO2 estimation at rest and at low exercise intensity. At high exercise intensity the overall response appears delayed but reproducible and the error can be overcome by mathematical modeling allowing an accurate estimation by PtCO2 of PaCO2 and VD/VT.

Morphological Cell Types Projecting from V1 Layer 4B to V2 Thick and Thin Stripes.

In macaque visual cortex, different cytochrome oxidase stripes of area V2 receive segregated projections from layers (L)2/3 and 4B of the primary visual cortex (V1), and project to dorsal or ventral stream extrastriate areas. Parallel V1-to-V2 pathways suggest functionally specialized circuits, but it is unknown whether these circuits arise from distinct cell types. V1 L4B includes two morphological types of excitatory projection neurons: pyramids, which carry mixed magnocellular (M) and parvocellular (P) information to downstream areas, and spiny stellates, which carry only M information. Previous studies have shown that, overall, V2 receives ∼80% of its L4B inputs from pyramids, thus receiving mixed M and P signals. However, it is unknown how pyramids and stellates distribute their outputs to the different V2 stripes, and whether different stripes receive inputs from morphologically distinct neuron types. Using viral-mediated labeling of V2-projecting L4B neurons in male macaques, we show that thick stripes receive a greater contribution of L4B inputs from M-dominated spiny stellates compared with thin stripes. Both stripe types, however, receive a much larger contribution from spiny stellates than previously shown for V2 overall, indicating that a larger amount of M information than previously thought flows into both the dorsal and ventral streams via the V2 thick and thin stripes, respectively. Moreover, we identify four types of V2-projecting L4B cells differing in size and complexity. Three such cell types project to both thin and thick stripes, but one type, the giant spiny-stellate neuron, resembling L4B neurons projecting to motion-sensitive area MT, was only found to project to thick stripes.SIGNIFICANCE STATEMENT Area V1 partitions visual information into functionally specialized parallel pathways which terminate into distinct stripes of area V2. We asked whether V1 inputs to different V2 stripes arise from morphologically different cell types. V1 layer (L)4B has two cell types: pyramids, which carry both magnocellular (M) and parvocellular (P) visual signals, and spiny stellates, which carry only M signals. We find that V2 thick stripes, which project to areas processing object motion, receive a larger fraction of L4B input from M-dominated stellates compared with thin stripes, which project to areas processing object attributes. We also identify four morphological types of V2-projecting L4B neurons, suggestive of four functionally specialized cell types.

Bulk tank milk quality data is unlikely to give useful information about dairy cow welfare at herd level.

This research communication explores the value of routinely collected bulk tank milk quality data for estimating dairy cattle welfare at herd level. Selected bulk tank milk quality parameters (somatic cell count, total bacterial count, urea, protein and fat contents) recorded during the years 2014-2016 in 287 Italian dairy farms were compared with the animal welfare data of each farm. The welfare assessment data were extracted from the database of the Italian Reference Centre for Animal Welfare (CReNBA), which includes the outputs of the application of the CReNBA welfare assessment protocol for dairy cows, used at national level for on-farm controls. The statistical analysis was carried out using the correlation coefficient for Kendall's Tau ranks, in order to investigate the presence of a categoric relationship between the selected bulk tank milk quality parameters and the overall animal welfare score or the scores of the single areas A (farm management and staff training), B (housing) and C (animal-based measures). Somatic cell count, total bacterial count, urea and proteins demonstrated only a few statistically significant and very weak correlations with farm animal welfare data, while no significant correlations were obtained for milk fat content. Given the weak correlations found, the selected bulk tank milk parameters seems to be able to provide only limited information about the welfare level of the herd, thus it could be difficult to use them for drawing up a pre-screening model for identifying herds at risk of poor welfare.

Comparison between the Airgo™ Device and a Metabolic Cart during Rest and Exercise.

The aim of this study is to compare the accuracy of Airgo™, a non-invasive wearable device that records breath, with respect to a gold standard. In 21 healthy subjects (10 males, 11 females), four parameters were recorded for four min at rest and in different positions simultaneously by Airgo™ and SensorMedics 2900 metabolic cart. Then, a cardio-pulmonary exercise test was performed using the Erg 800S cycle ergometer in order to test Airgo™'s accuracy during physical effort. The results reveal that the relative error median percentage of respiratory rate was of 0% for all positions at rest and for different exercise intensities, with interquartile ranges between 3.5 (standing position) and 22.4 (low-intensity exercise) breaths per minute. During exercise, normalized amplitude and ventilation relative error medians highlighted the presence of an error proportional to the volume to be estimated. For increasing intensity levels of exercise, Airgo™'s estimate tended to underestimate the values of the gold standard instrument. In conclusion, the Airgo™ device provides good accuracy and precision in the estimate of respiratory rate (especially at rest), an acceptable estimate of tidal volume and minute ventilation at rest and an underestimation for increasing volumes.

Local Circuits of V1 Layer 4B Neurons Projecting to V2 Thick Stripes Define Distinct Cell Classes and Avoid Cytochrome Oxidase Blobs.

Decades of anatomical studies on the primate primary visual cortex (V1) have led to a detailed diagram of V1 intrinsic circuitry, but this diagram lacks information about the output targets of V1 cells. Understanding how V1 local processing relates to downstream processing requires identification of neuronal populations defined by their output targets. In primates, V1 layers (L)2/3 and 4B send segregated projections to distinct cytochrome oxidase (CO) stripes in area V2: neurons in CO blob columns project to thin stripes while neurons outside blob columns project to thick and pale stripes, suggesting functional specialization of V1-to-V2 CO streams. However, the conventional diagram of V1 shows all L4B neurons, regardless of their soma location in blob or interblob columns, as projecting selectively to CO blobs in L2/3, suggesting convergence of blob/interblob information in L2/3 blobs and, possibly, some V2 stripes. However, it is unclear whether all L4B projection neurons show similar local circuitries. Using viral-mediated circuit tracing, we have identified the local circuits of L4B neurons projecting to V2 thick stripes in macaque. Consistent with previous studies, we found the somata of this L4B subpopulation to reside predominantly outside blob columns; however, unlike previous descriptions of local L4B circuits, these cells consistently projected outside CO blob columns in all layers. Thus, the local circuits of these L4B output neurons, just like their extrinsic projections to V2, preserve CO streams. Moreover, the intra-V1 laminar patterns of axonal projections identify two distinct neuron classes within this L4B subpopulation, including a rare novel neuron type, suggestive of two functionally specialized output channels. SIGNIFICANCE STATEMENT: Conventional diagrams of primate primary visual cortex (V1) depict neuronal connections within and between different V1 layers, but lack information about the cells' downstream targets. This information is critical to understanding how local processing in V1 relates to downstream processing. We have identified the local circuits of a population of cells in V1 layer (L)4B that project to area V2. These cells' local circuits differ from classical descriptions of L4B circuits in both the laminar and functional compartments targeted by their axons, and identify two neuron classes. Our results demonstrate that both local intra-V1 and extrinsic V1-to-V2 connections of L4B neurons preserve CO-stream segregation, suggesting that across-stream integration occurs downstream of V1, and that output targets dictate local V1 circuitry.

Informative features of local field potential signals in primary visual cortex during natural image stimulation.

The local field potential (LFP) is of growing importance in neurophysiology as a metric of network activity and as a readout signal for use in brain-machine interfaces. However, there are uncertainties regarding the kind and visual field extent of information carried by LFP signals, as well as the specific features of the LFP signal conveying such information, especially under naturalistic conditions. To address these questions, we recorded LFP responses to natural images in V1 of awake and anesthetized macaques using Utah multielectrode arrays. First, we have shown that it is possible to identify presented natural images from the LFP responses they evoke using trained Gabor wavelet (GW) models. Because GW models were devised to explain the spiking responses of V1 cells, this finding suggests that local spiking activity and LFPs (thought to reflect primarily local synaptic activity) carry similar visual information. Second, models trained on scalar metrics, such as the evoked LFP response range, provide robust image identification, supporting the informative nature of even simple LFP features. Third, image identification is robust only for the first 300 ms following image presentation, and image information is not restricted to any of the spectral bands. This suggests that the short-latency broadband LFP response carries most information during natural scene viewing. Finally, best image identification was achieved by GW models incorporating information at the scale of ∼ 0.5° in size and trained using four different orientations. This suggests that during natural image viewing, LFPs carry stimulus-specific information at spatial scales corresponding to few orientation columns in macaque V1.

Resolving the organization of the third tier visual cortex in primates: a hypothesis-based approach.

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.

Corticocortical connection patterns reveal two distinct visual cortical areas bordering dorsal V2 in marmoset monkey.

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.

Different orientation tuning of near- and far-surround suppression in macaque primary visual cortex mirrors their tuning in human perception.

In primary visual cortex (V1), neuronal responses to stimuli inside the receptive field (RF) are usually suppressed by stimuli in the RF surround. This suppression is orientation specific. Similarly, in human vision surround stimuli can suppress perceived contrast of a central stimulus in an orientation-dependent manner. The surround consists of two regions likely generated by different circuits: a near-surround generated predominantly by geniculocortical and intra-V1 horizontal connections, and a far-surround generated exclusively by interareal feedback. Using stimuli confined to the near- or far-surround of V1 neurons, and similar stimuli in human psychophysics, we find that near-surround suppression is more sharply orientation tuned than far-surround suppression in both macaque V1 and human perception. These results point to a similarity between surround suppression in macaque V1 and human vision, and suggest that feedback circuits are less orientation biased than horizontal circuits. We find the sharpest tuning of near-surround suppression in V1 layers (3, 4B, 4Cα) with patterned and orientation-specific horizontal connections. Sharpest tuning of far-surround suppression occurs in layer 4B, suggesting greater orientation specificity of feedback to this layer. Different orientation tuning of near- and far-surround suppression may reflect a statistical bias in natural images, whereby nearby edges have higher probability than distant edges of being co-oriented and belonging to the same contour. Surround suppression would, thus, increase the coding efficiency of frequently co-occurring contours and the saliency of less frequent ones. Such saliency increase can help detect small orientation differences in nearby edges (for contour completion), but large orientation differences in distant edges (for directing saccades/attention).

Two projection streams from macaque V1 to the pale cytochrome oxidase stripes of V2.

In the primate visual cortex, areas V1 and V2 distribute information they receive from the retina to virtually all extrastriate cortex, parsing this information into dorsal and ventral streams. Therefore, understanding the connectivity between V1 and V2 is crucial to understand visual cortical processing. Cytochrome oxidase staining in V2 reveals a repeating pattern of pale-thick-pale-thin stripes. V1 sends parallel output pathways to distinct V2 stripes. Previous models proposed either three or two parallel V1-to-V2 pathways in macaque, but both models viewed the two pale stripes within a single stripe cycle as a single compartment. However, recent studies have suggested that the two pale stripes may be functionally distinct, and in marmosets they also differ anatomically in the laminar origin of projections they receive from V1. Here we have asked whether the two pale stripes are also anatomically distinct in macaque. We made small retrograde tracer injections in different pale stripe types. We found that while both pale stripes receive a predominant V1 input from layers 2/3, only one set of pale stripes (pale lateral) receives significant projections from layer 4B, while the other set (pale medial) receives few or no layer 4B projections. Moreover, different tracer injections in nearby pale stripe types revealed that 97-99% of layer 2/3 cells only project to a single pale stripe type. These results demonstrate that in macaque, the two pale stripes are anatomically distinct compartments, and support the notion of two distinct projection streams from V1 to the two pale stripes of V2.

High-resolution mapping of anatomical connections in marmoset extrastriate cortex reveals a complete representation of the visual field bordering dorsal V2.

The primate visual cortex consists of many areas. The posterior areas (V1, V2, V3, and middle temporal) are thought to be common to all primate species. However, the organization of cortex immediately anterior to area V2 (the "third tier" cortex) remains controversial, particularly in New World primates. The main point of contention has been whether the third tier cortex consists of a single area V3, representing lower and upper visual quadrants in dorsal and ventral cortex, respectively, or of 2 distinct areas (the dorsomedial [DM] area and a V3-like area). Resolving this controversy is crucial to understand the function and evolution of the third tier cortex. We have addressed this issue in marmosets, by performing high-precision mapping of corticocortical connections in cortex bordering dorsal V2. Multiple closely spaced neuroanatomical tracer injections were placed across the full width of dorsal V2 or adjacent anterior cortex, and the location of resulting labeled cells mapped throughout whole flattened visual cortex. The resulting topographic patterns of labeled connections allowed us to define areas and their boundaries. We found that a complete representation of the visual field borders dorsal V2 and that the third tier cortex consists of 2 distinct areas. These results unequivocally support the DM model.

Primate V2 Receptive Fields Derived from Anatomically Identified Large-Scale V1 Inputs.

In the primate visual system, visual object recognition involves a series of cortical areas arranged hierarchically along the ventral visual pathway. As information flows through this hierarchy, neurons become progressively tuned to more complex image features. The circuit mechanisms and computations underlying the increasing complexity of these receptive fields (RFs) remain unidentified. To understand how this complexity emerges in the secondary visual area (V2), we investigated the functional organization of inputs from the primary visual cortex (V1) to V2 by combining retrograde anatomical tracing of these inputs with functional imaging of feature maps in macaque monkey V1 and V2. We found that V1 neurons sending inputs to single V2 orientation columns have a broad range of preferred orientations, but are strongly biased towards the orientation represented at the injected V2 site. For each V2 site, we then constructed a feedforward model based on the linear combination of its anatomically-identified large-scale V1 inputs, and studied the response proprieties of the generated V2 RFs. We found that V2 RFs derived from the linear feedforward model were either elongated versions of V1 filters or had spatially complex structures. These modeled RFs predicted V2 neuron responses to oriented grating stimuli with high accuracy. Remarkably, this simple model also explained the greater selectivity to naturalistic textures of V2 cells compared to their V1 input cells. Our results demonstrate that simple linear combinations of feedforward inputs can account for the orientation selectivity and texture sensitivity of V2 RFs.

Primate V2 Receptive Fields Derived from Anatomically Identified Large-Scale V1 Inputs.

In the primate visual system, visual object recognition involves a series of cortical areas arranged hierarchically along the ventral visual pathway. As information flows through this hierarchy, neurons become progressively tuned to more complex image features. The circuit mechanisms and computations underlying the increasing complexity of these receptive fields (RFs) remain unidentified. To understand how this complexity emerges in the secondary visual area (V2), we investigated the functional organization of inputs from the primary visual cortex (V1) to V2 by combining retrograde anatomical tracing of these inputs with functional imaging of feature maps in macaque monkey V1 and V2. We found that V1 neurons sending inputs to single V2 orientation columns have a broad range of preferred orientations, but are strongly biased towards the orientation represented at the injected V2 site. For each V2 site, we then constructed a feedforward model based on the linear combination of its anatomically-identified large-scale V1 inputs, and studied the response proprieties of the generated V2 RFs. We found that V2 RFs derived from the linear feedforward model were either elongated versions of V1 filters or had spatially complex structures. These modeled RFs predicted V2 neuron responses to oriented grating stimuli with high accuracy. Remarkably, this simple model also explained the greater selectivity to naturalistic textures of V2 cells compared to their V1 input cells. Our results demonstrate that simple linear combinations of feedforward inputs can account for the orientation selectivity and texture sensitivity of V2 RFs.