The corticogeniculate circuit is an evolutionarily conserved pathway linking the primary visual cortex with the visual thalamus in the feedback direction. While the corticogeniculate circuit is anatomically robust, the impact of corticogeniculate feedback on the visual response properties of visual thalamic neurons is subtle. Accordingly, discovering the function of corticogeniculate feedback in vision has been a particularly challenging task. In this review, the morphology, organization, physiology, and function of corticogeniculate feedback is compared across mammals commonly studied in visual neuroscience: primates, carnivores, rabbits, and rodents. Common structural and organizational motifs are present across species, including the organization of corticogeniculate feedback into parallel processing streams in highly visual mammals.

The patches in primary visual cortex constitute hot spots of metabolic activity, manifested by enhanced levels of cytochrome oxidase (CO) activity. They are also labeled preferentially by immunostaining for glutamic acid decarboxylase (GAD), γ-aminobutyric acid (GABA), and parvalbumin. However, calbindin shows stronger immunoreactivity outside patches. In light of this discrepancy, the distribution of the vesicular GABA transporter (VGAT) was examined in striate cortex of two normal macaques. VGAT immunoreactivity was strongest in layers 4B, 4Cα, and 5. In tangential sections, the distribution of CO, GAD, and VGAT was compared in layer 2/3. There was a close match between all three labels. This finding indicates that GABA synthesis is enriched in patches, and that inhibitory synapses are more active in patches than interpatches.

Although the primary visual cortex (V1) is one of the most extensively studied areas of the primate brain, very little is known about how the far periphery of visual space is represented in this area. We characterized the physiological response properties of V1 neurons in anaesthetized marmoset monkeys, using high-contrast drifting gratings. Comparisons were made between cells with receptive fields located in three regions of V1, defined by eccentricity: central (3–5°), near peripheral (5–15°), and far peripheral (>50°). We found that orientation selectivity of individual cells was similar from the center to the far periphery. Nonetheless, the proportion of orientation-selective neurons was higher in central visual field representation than in the peripheral representations. In addition, there were similar proportions of cells representing all orientations, with the exception of the representation of the far periphery, where we detected a bias favoring near-horizontal orientations. The proportions of direction-selective cells were similar throughout V1. When the center/surround organization of the receptive fields was tested with gratings with varying diameters, we found that the population of neurons that was suppressed by large gratings was smaller in the far periphery, although the strength of suppression in these cells tended to be stronger. In addition, the ratio between the diameters of the excitatory centers and suppressive surrounds was similar across the entire visual field. These results suggest that, superimposed on the broad uniformity of V1, there are subtle physiological differences, which indicate that spatial information is processed differently in the central versus far peripheral visual fields.

Monocular deprivation (MD) by lid suture is one of the classic paradigms for the study of developmental plasticity in the cerebral cortex, and we have detailed knowledge of its anatomical and physiological consequences as well as underlying molecular and cellular mechanisms. However, the effects of other forms of manipulating visual input through one eye on the functional architecture of the primary visual cortex (V1) have not yet been examined directly. We compared MD by lid suture with the effects of daily monocular lens wear using either a frosted lens or a neutral density (ND) filter. We used optical imaging of intrinsic signals and visually evoked potentials (VEPs) to assess responses in V1 to monocular stimulation. We found that loss of stimulus contrast through monocular frosted lens wear resulted in marked takeover of cortical territory by the nondeprived eye (NDE) similar to that caused by classic MD, and in virtual absence of orientation-selective responses following stimulation of the deprived eye (DE). Furthermore, amplitudes of VEPs in response to gratings of a range of spatial frequencies were significantly reduced in the DE compared to the NDE. In contrast, differences in luminance between two eyes caused by an ND filter in front of one eye did not affect ocular dominance and orientation maps, and there was no significant difference in the amplitude of VEPs elicited through the two eyes. Our results are consistent with previous electrophysiological studies in demonstrating that binocular pattern information is necessary to maintain normal functional maps in both eyes, while reduced luminance in one eye has little effect on the overall functional architecture and visual responses in V1.

Recent results indicate that astrocytic β-adrenergic receptors (βAR) participate in noradrenergic modulation of synaptic activity. In this study, we sought to examine whether neural activity can, in turn, regulate astrocytic βAR. To address this question, an antiserum that recognizes β-adrenergic receptors (βAR) specifically in astrocytes was used to assess the distribution of the receptors across ocular dominance columns in VI of two monocular and four visually intact adult monkeys. Cytochrome oxidase histochemistry (CO) was used to identify the position of the cortical laminae and of the ocular dominance columns receiving visual inputs from the intact and enucleated eyes. This stain revealed the expected pattern within V1 of monocular monkeys–i.e. darker and lighter bands of equal widths (ca. 500μm) spanning laminae 4–6, each associated with larger and smaller blobs, respectively, in lamina 2/3;. Alignment of CO sections with adjacent sections stained for astrocytic βAR by the immunoperoxidase method revealed intense βAR-like immunoreactivity (βAR-li) in the superficial laminae, a slightly weaker staining in the infragranular laminae and weakest staining in lamina 4C. Within lamina 4C., a prominent striped pattern was evident. The darker bands of the stripe closely matched widths and positions of the lighter CO columns associated with the enucleated eye. On the other hand, immunocytochemical staining for the astrocytic intermediate filament protein, GFAP, within V1 of monocular monkeys revealed no inter-columnar difference in the density of astrocytic cell bodies or processes. Nissl stain also revealed no overt inter-columnar differences in cell density. V1 of visually intact monkeys exhibited a similar laminar distribution pattern of βAR-li and of CO. Within lamina 4C., βAR-li was uniformly faint and CO staining was uniformly intense. This suggests that the striped pattern of βAR-li seen in lamina 4C of monocular monkeys results from elevation of the βAR-antigen within the inactive columns. The results indicate that astrocytic βAR density is regulated by local neural activity. The mechanisms regulating βAR density are likely to be independent of those regulating glial cell proliferation or GFAP synthesis. In vitro experimental results of others suggest that elevation of astrocytic βAR may be a mechanism compensating for chronic neural inactivity, since the coincident release of noradrenaline with visual stimulation would elevate neuropil excitability via the astrocytic mechanism of (1) decreasing the uptake of neuronally released L-glutamate; (2) increasing GABA uptake; and (3) stimulating glycogenolysis. Alternatively, the changes in βAR-li may reflect an up-regulation of the receptors within inactive columns due to reduced levels of noradrenaline release.

Extrastriate areas TEO and V4 have been associated with form and color vision. Area V4 has also been suggested to participate in processes concerned with attention, stimulus salience, and perceptual learning. In a continuing effort to elucidate the connectional interactions and microcircuitry of these areas, we describe in this report the pattern of feedback connections from TEO and V4. Connections were demonstrated by injections of the high-resolution anterograde tracers PHA-L or biocytin and further analyzed by reconstruction of 25 individual axons through serial sections. This analysis yielded several new results: (1) Both areas TEO and V4 have widespread feedback connections (defined by their preferential termination in layer 1 and avoidance of layer 4). From TEO, there are dense projections to area V4 and moderate ones to V2 and V1. From V4, there are dense projections to V2 and moderate ones to V3 and V1. (2) Terminal fields span large territories in area V1, up to 6.0 mm in the case of axons originating from TEO; up to 5.0 mm in the case of axons originating from V4. In V2, fields tend to be smaller, between 3.0–5.0 mm. (3) Many axons from TEO and some from V4 have terminations in both areas V1 and V2. (4) Because individual terminal clusters and segments are often larger than cytochrome oxidase compartments, especially in V1, we suggest they may not be correlated with this compartmental organization. These results are consistent with the hypothesis that feedback connections may contribute to processes other than perceptual discrimination.

To study the relationship between ocular dominance columns (ODCs) and axonal projections of individual layer 6 pyramidal neurons in the primary visual cortex, neurons were intracellularly labeled with biocytin in live slices prepared from macaque monkeys that had received an intravitreal injection of tetrodotoxin (TTX). The TTX injection indirectly causes a decrease in cytochrome oxidase (CO) expression in the cortical ODCs corresponding to the treated eye (Wong-Riley & Carroll, 1984). Sections from slices with labeled layer 6 neurons were double stained for biocytin and CO, to allow visualization of neuronal processes as well as ODCs. Twenty-seven layer 6 pyramidal neurons in ODC-labeled slices were analyzed. These neurons were classified according to the criteria of Wiser and Callaway (1996). Eight of these are class I neurons, which have dense axonal projections to the monocular layer 4C. The remaining 19 are class II neurons which project primarily to the binocular layers outside 4C. Among class I neurons, two have dense axonal arbors in layer 4Cα (type Iα), one in layer 4Cβ (type Iβ), and two throughout the depth of layer 4C (type IC). None of these neurons have ODC-specific axonal arbors. The remaining three class I neurons have focused axonal projections in layers 4Cβ and 4A (type IβA). All three appear to have axonal arbors predominantly within their home ODC in layer 4C. The axonal arbors of class II neurons do not appear to relate to ODCs in any specific fashion.

We studied excitatory local circuits in the macaque primary visual cortex (V1) to investigate their relationships to the magnocellular (M) and parvocellular (P) streams. Sixty-two intracellularly labeled spiny neurons in layers 2–5 were analyzed. We made detailed observations of the laminar and columnar specificity of axonal arbors and noted correlations with dendritic arbors. We find evidence for considerable mixing of M and P streams by the local circuitry in V1. Such mixing is provided by neurons in the primary geniculate recipient layer 4C, as well as by neurons in both the supragranular and infragranular layers. We were also interested in possible differences in the axonal projections of neurons with different dendritic morphologies. We found that layer 4B spiny stellate and pyramidal neurons have similar axonal arbors. However, we identified two types of layer 5 pyramidal neuron. The majority have a conventional pyramidal dendritic morphology, a dense axonal arbor in layers 2–4B, and do not project to the white matter. Layer 5 projection neurons have an unusual “backbranching” dendritic morphology (apical dendritic branches arc downward rather than upward) and weak or no axonal arborization in layers 2–4B, but have long horizontal axonal projections in layer 5B. We find no strong projection from layer 5 pyramidal neurons to layer 6. In macaque V1 there appears to be no single source of strong local input to layer 6; only a minority of cells in layers 2–5 have axonal branches in layer 6 and these are sparse. Our results suggest that local circuits in V1 mediate interactions between M and P input that are complex and not easily incorporated into a simple framework.

We measured the latency of 35 cells from V1 in two rhesus monkeys, to dynamic random dot stimuli monocular and binocularly presented. Mean latencies after non-dominant eye stimulation (97.9 ms) were longer than those for dominant eye (78.2 ms) and binocular (70.7 ms) stimulation. Differences between latencies for dominant eye and binocular stimulation were not statistically significant. For dominant eye, there was a significant statistical correlation between dominance strength and latency (R = −0.36; p = 0.03). We failed to find significant statistical differences between latencies for cells with temporal and nasal dominant receptive-field. We conclude that, in V1, the response latency is largely determined by the dominant eye, whereas interocular interactions do not seem to play a relevant role regarding response latency.

We measured the responses of single neurons in marmoset visual cortex (V1, V2, and the third visual complex) to moving first-order stimuli and to combined first- and second-order stimuli in order to determine whether first-order motion processing was influenced by second-order motion. Beat stimuli were made by summing two gratings of similar spatial frequency, one of which was static and the other was moving. The beat is the product of a moving sinusoidal carrier (first-order motion) and a moving low-frequency contrast envelope (second-order motion). We compared responses to moving first-order gratings alone with responses to beat patterns with first-order and second-order motion in the same direction as each other, or in opposite directions to each other in order to distinguish first-order and second-order direction-selective responses. In the majority (72%, 67/93) of cells (V1 73%, 45/62; V2 70%, 16/23; third visual complex 75%, 6/8), responses to first-order motion were significantly influenced by the addition of a second-order signal. The second-order envelope was more influential when moving in the opposite direction to the first-order stimulus, reducing first-order direction sensitivity in V1, V2, and the third visual complex. We interpret these results as showing that first-order motion processing through early visual cortex is not separate from second-order motion processing; suggesting that both motion signals are processed by the same system.

During development, the projection from the lateral geniculate nucleus to striate cortex becomes segregated into monocular regions called ocular dominance columns. Prior studies in cats have suggested that experimental strabismus or alternating monocular occlusion increases the width and segregation of columns. In the squirrel monkey, strabismus has been reported to induce the formation of ocular dominance columns. However, these studies are difficult to interpret because no animal can serve as its own control and the degree of inter-individual variability among normal subjects is considerable. We have re-examined the effect of strabismus on ocular dominance columns in a large group of strabismic and normal squirrel monkeys. Five animals rendered strabismic at age one week had well-developed, widely spaced columns. Among 16 control animals, a wide spectrum of column morphology was encountered. Some control animals lacked ocular dominance columns, whereas others had columns similar to those observed in strabismic animals. Natural variation in column expression in normal squirrel monkeys, and potential uncontrolled genetic influences, made it impossible to determine if strabismus affects ocular dominance columns. It was evident however, that strabismus does not affect the binocular projection from the lateral geniculate nucleus to each CO patch in the upper layers. In strabismic monkeys, just as in normal animals, each patch received input from geniculate afferents serving both the left eye and the right eye. In addition, in strabismic monkeys, as in normal animals, patches were not aligned with ocular dominance columns.

This paper explores different syntactic variations and functional properties of clausal units that are initiated by the finite verb in Swedish. We focus on V1 constructions that are basically declarative in function, thus excluding interrogative, conditional and directive uses. Because V1 constructions, and particularly certain variants of them, are typical of spoken Swedish, our examples and analyses focus primarily on the usage in speaking. The V1 constructions studied include whole-utterance constructions in conversational sequences, utterance-internal extensions, and utterance-internal reshapings, such as syntactic blends. We will offer an analysis of (declarative) V1 constructions, which in some respects differs from analyses proposed by generative syntacticians or traditional grammarians. Our analysis of the constructional resource is discourse oriented. In this perspective, V1 constructions are analyzable as sequentially dependent, second, or ‘subsequent’ moves, viz. units that can scarcely initiate a communicative project but which instead continue, extend or comment on an already initiated project. Thus, rather than analyzing V1 constructions as one type of ellipsis they could be seen as full-fledged clausal units whose existence is conditioned by their position in a discourse or utterance context. From a theoretical perspective, this paper is a contribution to an emerging dialogical or interactional model of (Swedish) grammar.

This study examines the temporal properties of geniculocortical and corticogeniculate (CG) pathways that link the lateral geniculate nucleus (LGN) and primary visual cortex in the ferret. Using electrical stimulation in the LGN to evoke action potentials in geniculocortical and CG axons, results show that conduction latencies are significantly faster in geniculocortical neurons than in CG neurons. Within each pathway, axonal latency and visual physiology support the view of sub-classes of neurons. By examining the timing of visual responses and the latency of CG feedback, estimates indicate that visual information can reach the cortex and return to the LGN as early as 60 msec following the onset of a visual stimulus. These findings place constraints on the functional role of corticogeniculate feedback for visual processing.

Video cathode ray tube (CRT) technology has proven to be extremely valuable for performing research in the visual system. However, the image on a CRT monitor is not constant, but consists of a series of brief pulses. This has implications for any study that explores the responses of neurons in the visual system on short time scales. In particular, there is no unambiguous time point at which a visual stimulus presented via CRT may be said to have ended. Recordings from single units in visual cortical area V1 of an awake primate demonstrate that, when studying changes in response timing on the order of 10 ms or less, stimuli delivered at video frame rates do not duplicate the effects seen with stimuli that have continuous functions of luminance versus time. Additionally, there does not seem to be any clear method of comparing the results obtained with video-rate stimuli with results obtained with continuous-time stimuli that holds for all conditions. These effects are especially critical when exploring the time course of the neuronal responses to the ending of a visual stimulus (off-response). Our findings cast doubt upon the recently reported result that off-responses have consistently shorter latencies than on-responses.

Mature visual cortex shows a single, binocularly matched orientation map. This matching develops without visual experience. It persists despite early monocular deprivation that largely eliminates one eye's map, followed by reverse suture (deprivation of the previously open eye and opening of the previously deprived eye), even though the two eyes lack common visual experience in this case. These results have been interpreted to suggest that the structure of orientation maps either is innately predetermined or, if it arises through self-organization, is determined by external cues such as boundary conditions or a “scaffolding” of horizontal connections. We show, to the contrary, that these results are the expected outcomes if orientation maps develop through activity-instructed, correlation-based development of the geniculocortical connections without additional cues. A weak, binocularly correlated orientation map is known to exist before deprivation onset; we previously showed how this can arise through activity-instructed development. Now we show that this initial correlation between the two eyes' maps can persist or increase despite deprivation sufficient to cause massive loss of the deprived eye's geniculocortical synaptic strength, followed by reverse suture. Given sufficient early correlated map development, each map's fate is “dynamically committed”: the two eyes' maps will converge upon a common outcome, even if developing independently. This dynamic fate commitment is retained even after severe deprivation.

During normal vision, when subjects attempt to fix their gaze on a small stimulus feature, small fixational eye movements persist. We have recorded the impulse activity of single neurons in primary visual cortex (V1) of macaque monkeys while their fixational eye movements moved the receptive-field activating region (AR) over and around a stationary stimulus. Three types of eye movement activation were found. (1) Saccade cells discharged when a fixational saccade moved the AR onto the stimulus, off the stimulus, or across the stimulus. (2) Position/drift cells discharged during the intersaccadic (drift) intervals and were not activated by saccades that swept the AR across the stimulus without remaining on it. To activate these neurons, it was essential that the AR be placed on the stimulus and many of these cells were selective for the sign of contrast. They had smaller ARs than the other cell types. (3) Mixed cells fired bursts of activity immediately following saccades and continued to fire at a lower rate during intersaccadic intervals. The tendency of each neuron to fire transient bursts or sustained trains of impulses following saccades was strongly correlated with the transiency of its response to stationary flashed stimuli. For one monkey, an extraretinal influence accompanying fixational saccades was identified. During natural viewing, the different eye movement classes probably make different contributions to visual processing. Position/drift neurons are well suited for coding spatial details of the visual scene because of their small AR size and their selectivity for sign of contrast and retinal position. However, saccade neurons transmit information that is ambiguous with respect to the spatial details of the retinal image because they are activated whether the AR lands on a stimulus contour, or the AR leaves or crosses the contour and lands in another location. Saccade neurons may be involved in constructing a stable world in spite of incessant retinal image motion, as well as in suppressing potentially confusing input associated with saccades.