The role of the ventral and dorsal pathways in reading Chinese characters and English words

Previous literature in alphabetic languages suggests that the occipital-temporal region (the ventral pathway) is specialized for automatic parallel word recognition, whereas the parietal region (the dorsal pathway) is specialized for serial letter-by-letter reading ( [Cohen et al., 2008] and [Ho et al., 2002] ). However, few studies have directly examined the role of the ventral and dorsal pathways in Chinese reading compared to English reading. To investigate this issue, we adopted the degraded word processing paradigm used by Cohen et al. (2008) and compared brain regions involved in the processing of degraded Chinese characters and English words during lexical decision, using functional magnetic resonance imaging (fMRI). The degraded characters/words were created by inserting blank spaces between radicals of Chinese characters or syllables of English polysyllabic words. Generally, the current study replicated the effects of Cohen et al. (2008), showing that in Chinese – like in alphabetic languages – character spacing modulates both ventral (bilateral cuneus, left middle occipital gyrus) and dorsal (left superior parietal lobule and middle frontal gyrus) pathways. In addition, the current study showed greater activation in bilateral cuneus and right lingual gyrus for Chinese versus English when comparing spaced to normal stimuli, suggesting that Chinese character recognition relies more on ventral visual-spatial processing than English word recognition. Interestingly, bilateral cuneus showed monotonic patterns in response to increasing spacing, while the rest of the regions of interest showed non-monotonic patterns, indicating different profiles for these regions in visual-spatial processing.     

Interactions between the dorsal and the ventral pathways in mental rotation: An fMRI study

In this fMRI study, we examined the relationship between activations in the inferotemporal region (ventral pathway) and the parietal region (dorsal pathway), as well as in the prefrontal cortex (associated with working memory), in a modified mental rotation task. We manipulated figural complexity (simple vs. complex) to affect the figure recognition process (associated with the ventral pathway) and the amount of rotation (0° vs. 90°), typically associated with the dorsal pathway. The pattern of activation not only showed that both streams are affected by both manipulations, but also showed an overadditive interaction. The effect of figural complexity was greater for 90° rotation than for 0° in multiple regions, including the ventral, dorsal, and prefrontal regions. In addition, functional connectivity analyses on the correlations across the time courses of activation between regions of interest showed increased synchronization among multiple brain areas as task demand increased. The results indicate that both the dorsal and the ventral pathways show interactive effects of object and spatial processing, and they suggest that multiple regions interact to perform mental rotation.     

Integrating faces, houses, motion, and action: spontaneous binding across ventral and dorsal processing streams

Perceiving an event requires the integration of its features across numerous brain maps and modules. Visual object perception is thought to be mediated by a ventral processing stream running from occipital to inferotemporal cortex, whereas most spatial processing and action control is attributed to the dorsal stream connecting occipital, parietal, and frontal cortex. Here we show that integration operates not only on ventral features and objects, such as faces and houses, but also across ventral and dorsal pathways, binding faces and houses to motion and manual action. Furthermore, these bindings seem to persist over time, as they influenced performance on future task-relevant visual stimuli. This is reflected by longer reaction times for repeating one, but alternating other features in a sequence, compared to complete repetition or alternation of features. Our findings are inconsistent with the notion that the dorsal stream is operating exclusively online and has no access to memory.     

The role of the dorsal and ventral hippocampus in olfactory working memory

Olfactory working memory and pattern separation for odor information was assessed in male rats using a matching-to-sample for odors paradigm. The odor set consisted of a five aliphatic acids with unbranched carbon chains that varied from two- to six-carbons in length. Each trial consisted of a sample phase followed by a choice phase. During the sample phase, rats would receive one of five different odors. Fifteen seconds later during the choice phase one of the previous odors was presented simultaneously side by side with a different odor that was based on the number of aliphatic acids that varied in the carbon chains from two- to six-carbons in length and rats were allowed to choose between the two odors. The rule to be learned in order to receive a food reward was to always choose the odor that occurred during the study phase. Odor separations of 1, 2, 3 or 4 were selected for each choice phase and represented the carbon chain difference between the study phase odor and the test phase odor. Once an animal reached a criterion of 80–90% correct across all temporal separations based on 40 trials, rats received a control, dorsal hippocampal, or ventral hippocampal lesion and were retested on the task. On postoperative trials, only the ventral hippocampal lesion group was impaired relative to both control and dorsal hippocampal groups groups. There were no effects on odor pattern separation. All groups of rats could discriminate between the odors. The data suggest that the ventral hippocampus, but not dorsal hippocampus, supports working memory for odor information.     

Visual awareness of objects and their colour

At any given moment, our awareness of what we 'see' before us seems to be rather limited. If, for instance, a display containing multiple objects is shown (red or green disks), when one object is suddenly covered at random, observers are often little better than chance in reporting about its colour (Wolfe, Reinecke, & Brawn, Visual Cognition, 14, 749-780, 2006). We tested whether, when object attributes (such as colour) are unknown, observers still retain any knowledge of the presence of that object at a display location. Experiments 1-3 involved a task requiring two-alternative (yes/no) responses about the presence or absence of a colour-defined object at a probed location. On this task, if participants knew about the presence of an object at a location, responses indicated that they also knew about its colour. A fourth experiment presented the same displays but required a three-alternative response. This task did result in a data pattern consistent with participants' knowing more about the locations of objects within a display than about their individual colours. However, this location advantage, while highly significant, was rather small in magnitude. Results are compared with those of Huang (Journal of Vision, 10(10, Art. 24), 1-17, 2010), who also reported an advantage for object locations, but under quite different task conditions.     

Perceptual awareness and its loss in unilateral neglect and extinction

We review recent evidence from studies of patients with unilateral neglect and/or extinction, who suffer from a loss of awareness for stimuli towards the affected side of space. We contrast their deficit with the effects of damage to primary sensory areas, noting that such areas can remain structurally intact in neglect, with lesions typically centred on the right inferior parietal lobe. In keeping with preservation of initial sensory pathways, many recent studies have shown that considerable residual processing can still take place for neglected or extinguished stimuli, yet without reaching the patient's awareness. This ranges from preserved visual grouping processes through to activation of identity, semantics and emotional significance. Similarly to 'preattentive' processing in normals, such residual processing can modulate what will enter the patient's awareness. Recent studies have used measures such as ERPs and fMRI to determine the neural correlates of conscious versus unconscious perception in the patients, which in turn can be related to the anatomy of their lesions. We relate the patient findings to neurophysiological data from areas in the monkey parietal lobe, which indicate that these serve as cross-modal and sensorimotor interfaces highlighting currently relevant locations as targets for intentional action. We speculate on the special role such brain regions may play in perceptual awareness, seeking to explain how damage to a system which appears primarily to code space could eliminate awareness even for non-spatial stimulus properties at affected locations. This may relate to the extreme nature of 'winner-takes-all' functions within the parietal lobe, and their correspondingly strong influence on other brain areas.     

Individual differences in verbal conditioning, extinction, and awareness

The present study examined the relationship of three individual difference variables—need for approval, a belief in internal versus external control of reinforcement, and verbal intelligence— to acquisition, extinction, and awareness in a verbal conditioning task Based on a postexperiment interview and recognition task, subjects were grouped according to their awareness of the response-reinforcement contingencies and the degree to which they admitted influence by the experimenter Approval motivated subjects were more likely to acquire the reinforced response regardless of awareness levels than were low need for approval subjects. Subjects assessed as internal were more likely to deny influence by the experimenter and in some instances were more resistant to extinction than subjects called external Subjects higher in verbal intelligence evidenced more awareness than subjects with lower intelligence scores Results suggest that the inclusion of individual difference variable is of crucial importance to a clear understanding of the experimenter-subject interaction in the verbal conditioning situation Implications for the other dyadic interactions, particularly psychotherapy, were discussed     

Source monitoring: The importance of feature binding at encoding

This paper is based on the source monitoring framework (Johnson, Hashtroudi, & Lindsay, 1993) and provides a review of data that suggests the involvement of working memory encoding processes in source monitoring tasks. Source monitoring refers to diverse mental processes used to distinguish the origin of our memories. A key assumption is that, when people encode an event, component processes of working memory play a crucial role in maintaining and binding the various features (e.g., semantic, perceptual) of an event together. It follows that success on a working memory binding test should be a good predictor of proficiency on a long-term source monitoring task, and that individual differences in working memory binding ability should account for the different pattern of source monitoring errors. Interestingly, these predictions have not been directly tested.     

Visual awareness and the detection of fearful faces

A commonly held view is that emotional stimuli are processed independently of awareness. Here, the authors parametrically varied the duration of a fearful face target stimulus that was backward masked by a neutral face. The authors evaluated awareness by characterizing behavioral performance using receiver operating characteristic curves from signal detection theory. Their main finding was that no universal objective awareness threshold exists for fear perception. Although several subjects displayed a behavioral pattern consistent with previous reports (i.e., targets masked at 33 ms), a considerable percentage of their subjects (64%) were capable of reliably detecting 33-ms targets. Their findings suggest that considerable information is available even in briefly presented stimuli (possibly as short as 17 ms) to support masked fear detection.     

Understanding the dorsal and ventral systems of the human cerebral cortex: beyond dichotomies

Traditionally, characterizations of the macrolevel functional organization of the human cerebral cortex have focused on the left and right cerebral hemispheres. However, the idea of left brain versus right brain functions has been shown to be an oversimplification. We argue here that a top-bottom divide, rather than a left-right divide, is a more fruitful way to organize human cortical brain functions. However, current characterizations of the functions of the dorsal (top) and ventral (bottom) systems have rested on dichotomies, namely where versus what and how versus what. We propose that characterizing information-processing systems leads to a better macrolevel organization of cortical function; specifically, we hypothesize that the dorsal system is driven by expectations and processes sequences, relations, and movement, whereas the ventral system categorizes stimuli in parallel, focuses on individual events, and processes object properties (such as shape in vision and pitch in audition). To test this hypothesis, we reviewed over 100 relevant studies in the human neuroimaging and neuropsychological literatures and coded them relative to 11 variables, some of which characterized our hypothesis and some of which characterized the previous dichotomies. The results of forward stepwise logistic regressions supported our characterization of the 2 systems and showed that this model predicted the empirical findings better than either the traditional dichotomies or a left-right difference.     

Visual awareness and the levels of processing hypothesis: A critical review

Does a visual percept emerge to consciousness in a graded manner (i.e. evolving through increasing degrees of clarity), or according to a dichotomous, “all-or-none” pattern (i.e. abruptly transitioning from unawareness to awareness)? The level of processing hypothesis (LoP; B. Windey and A. Cleeremans, 2015) recently proposed a theoretical framework where the transition from unaware to aware visual experience is graded for low-level stimulus representations (i.e. stimulus “energy” or “feature” levels) whereas it is dichotomous for high-level (i.e. the perception of “letters”, “words” or “meaning”) stimulus perception. Here, we will critically review current behavioral and brain-based evidence on the LoP hypothesis and discuss potential challenges (such as differences in LoP conceptualizations, awareness scale related issues, attentional confounds and divergences on experimental factors or statistical analyses) which might be of use for future research within the field. Overall, the LoP hypothesis is a recent and promising proposal that attempts to integrate divergent evidence on the graded vs. dichotomous emergence of awareness debate. Whereas current evidence validates some of the assumptions proposed by the LoP account, there is still much work to do on both methodological and experimental levels. Future neuroimaging studies might help to disentangle the current complex pattern of results found in LoP studies and, importantly, shed some light on the ongoing debate about the search for the Neural Correlates of Consciousness (NCC).     

Visual awareness and the on-line modification of action.

An influential theory of visually guided action proposes that (a) conscious perception of target displacement disrupts on-line action and (b) small target perturbations are inconsequential, provided the participant is unaware of them. This study examined these claims in a study of rapid aiming movements to targets. Novel features included on-line verbal reports of target displacement, and the factorial combination of small versus large displacements occurring near peak saccadic velocity or 100 ms later. Although awareness of target displacement had no effect on movement kinematics, even small target displacements near peak saccadic velocity affected kinematic measures. These results support both a strong view of visual stream separation in the on-line control of action and richer spatial coding by unconscious processes than has previously been acknowledged.     

Visual search while driving: skill and awareness during inspection of the scene

Novice drivers tend to restrict their search of the road on dual-carriageways, relative to the scanning observed in experienced drivers. The present study determined whether the difference was the result of novices having limited mental capacity remaining after vehicle control had been maintained, or whether it resulted from an impoverished mental model of the events likely to occur on a dual-carriageway. Novice and experienced drivers watched video-recordings taken from a car travelling along a variety of roads, including dual-carriageways, and their eye movements were recorded to determine the scanning patterns as they followed instructions to indicate hazardous events. The experienced drivers showed more extensive scanning on the demanding sections of dual-carriageway in this task. This supports the hypothesis that the inspection of the roadway by novices is limited not because they have limited mental resources residual from the task of vehicle control, but that they have an impoverished mental model of what is likely to happen on dual-carriageways.     

Visual awareness negativity is an early neural correlate of awareness: A preregistered study with two Gabor sizes

Electrophysiological recordings are commonly used to study the neural correlates of consciousness in humans. Previous research is inconsistent as to whether awareness can be indexed with visual awareness negativity (VAN) at about 200 ms or if it occurs later. The present study was preregistered with two main aims: First, to provide independent evidence for or against the presence of VAN, and second, to study whether stimulus size may account for the inconsistent findings. Subjects were shown low-contrast Gaussian filtered gratings (Gabor patches) in the four visual quadrants. Gabor size (large and small) was varied in different sessions and calibrated to each subject’s threshold of visual awareness. Event-related potentials were derived from trials in which subjects localized the Gabors correctly to capture the difference between trials in which they reported awareness versus no awareness. Bayesian analyses revealed very strong evidence for the presence of VAN for both Gabor sizes. However, there was no evidence for or against an effect of stimulus size. The present findings provide evidence for VAN as an early neural correlate of awareness.     

Tuning perception: Visual working memory biases the quality of visual awareness

Studies of consciousness reveal that it is possible to manipulate subjective awareness of a visual stimulus. For example, items held in visual working memory (VWM) that match target features increase the speed with which the target reaches visual awareness. To examine the effect of VWM on perception, previous studies have mainly used coarse measures of awareness, such as present/absent or forced-choice judgments. These methods can reveal whether or not an individual has seen an item, but they do not provide information about the quality with which the item was seen. Using continuous report methods it has been shown that the fidelity of a perceived item can be affected by whether or not that item is masked. In the present study, we used an object-substitution masking task to examine whether items held in VWM would influence the quality with which a masked target reached awareness, or whether the threshold for awareness was instead affected by stimuli held in memory. We observed that targets matching the contents of VWM were recalled with greater precision compared to items that did not match the contents of VWM. Importantly, this effect occurred without affecting the likelihood of the target being perceived. These results suggest that VWM plays a greater role in modulating the fidelity of perceived representations than in lowering the overall threshold of awareness.     

Testing the dorsal stream attention hypothesis: Electrophysiological correlates and the effects of ventral stream damage

The roles of dorsal and ventral processing streams in visual orienting and conscious perception were examined in two experiments. The first employed high density EEG with source localization. The second comprised a neuropsychological case study. Visual orienting was assessed with an attention procedure, where peripheral letters cued participants towards a target location. In the perception procedure participants responded to the same letters by performing an explicit conscious discrimination. In Experiment 1, the peripheral letters elicited rapid dorsal stream activation in the attention procedure, and this activation preceded top-down enhancement of target processing in occipital cortex. In the perception procedure early ventral stream activation was seen. In addition, peripheral letters elicited an “early directing attention negativity” (EDAN) over parietal recording sites in the attention procedure, but not in the perception procedure. In Experiment 2, a patient with a bilateral ventral stream lesion but preserved dorsal stream function showed clear disruption to performance in the perception procedure, whilst exhibiting a normal visual orienting effect in the attention procedure. Taken together these findings (1) highlight the distinct roles of the dorsal and ventral streams in attention and perception, and (2) suggest how these streams might interact, via reentrant effects of attention on perceptual processing.     

Relationship between visual binding, reentry and awareness

Visual feature binding has been suggested to depend on reentrant processing. We addressed the relationship between binding, reentry, and visual awareness by asking the participants to discriminate the color and orientation of a colored bar (presented either alone or simultaneously with a white distractor bar) and to report their phenomenal awareness of the target features. The success of reentry was manipulated with object substitution masking and backward masking. The results showed that late reentrant processes are necessary for successful binding but not for phenomenal awareness of the bound features. Binding errors were accompanied by phenomenal awareness of the misbound feature conjunctions, demonstrating that they were experienced as real properties of the stimuli (i.e., illusory conjunctions). Our results suggest that early preattentive binding and local recurrent processing enable features to reach phenomenal awareness, while later attention-related reentrant iterations modulate the way in which the features are bound and experienced in awareness.     

Visual awareness due to neuronal activities in subcortical structures: a proposal

It has been shown that visual awareness in the blind hemifield of hemianopic cats that have undergone unilateral ablations of visual cortex can be restored by sectioning the commissure of the superior colliculus or by destroying a portion of the substantia nigra contralateral to the cortical lesion (the Sprague effect). We propose that the visual awareness that is recovered is due to synchronized oscillatory activities in the superior colliculus ipsilateral to the cortical lesion. These oscillatory activities are normally partially suppressed by the inhibitory, GABAergic contralateral nigrotectal projection, and the destruction of the substantia nigra, or the sectioning of the collicular commissure, disinhibits the collicular neurons, causing an increase in the extent of oscillatory activity and/or synchronization between activities at different sites. This increase in the oscillatory and synchronized character is sufficient for the activities to give rise to visual awareness. We argue that in rodents and lower vertebrates, normal visual awareness is partly due to synchronized oscillatory activities in the optic tectum and partly due to similar activities in visual cortex. It is only in carnivores and primates that visual awareness is wholly due to cortical activities. Based on von Baerian recapitulation theory, we propose that, even in humans, there is a period in early infancy when visual awareness is partially due to activities in the superior colliculus, but that this awareness gradually disappears as the nigrotectal projection matures.     

Effects of ventral and dorsal CA1 subregional lesions on trace fear conditioning

Recent lines of research have focused on dissociating function between the dorsal and ventral hippocampus along space and anxiety dimensions. In the dorsal hippocampus, the CA1 subregion has been implicated in the acquisition of contextual fear as well as in the trace interval in trace fear conditioning. The present study was designed to test the relative contributions of dorsal (dCA1) and ventral CA1 (vCA1) in trace fear conditioning. Long-Evans rats received ibotenate lesions of the ventral CA1 (n=7), dorsal CA1 (n=9), or vehicle control lesions (n=8) prior to trace fear conditioning acquisition. Results suggest dCA1 and vCA1 groups show no significant deficits during acquisition when compared to control groups. dCA1 and vCA1 both show deficits in the retention of contextual fear when tested 24 h post-acquisition (P<.05 and P<.01, respectively), and vCA1 was impaired relative to dCA1 (P<.05). This is suggestive of a graded involvement in contextual retention between the dorsal and ventral aspects of CA1. dCA1 showed no deficit for retention of conditioned fear to the tone or the trace when tested 48 h post-acquisition, whereas vCA1 did show a significant deficit for the trace interval and a slight, non-significant reduction in freezing to the tone, when compared to the control group (p<.05). Overall the data are suggestive of a graded involvement in retention of fear conditioning between the dorsal and ventral aspects of CA1, but it is likely that vCA1 may be critically involved in retention of trace fear conditioning.