Attention without awareness in blindsight.

The act of attending has frequently been equated with visual awareness. We examined this relationship in 'blindsight'--a condition in which the latter is absent or diminished as a result of damage to the primary visual cortex. Spatially selective visual attention is demonstrated when information that stimuli are likely to appear at a specific location enhances the speed or accuracy of detection of stimuli subsequently presented at that location. In a blindsight subject, we showed that attention can confer an advantage in processing stimuli presented at an attended location, without those stimuli entering consciousness. Attention could be directed both by symbolic cues in the subject's spared field of vision or cues presented in his blind field. Cues in his blind field were even effective in directing his attention to a second location remote from that at which the cue was presented. These indirect cues were effective whether or not they themselves elicited non-visual awareness. We concluded that the spatial selection of information by an attentional mechanism and its entry into conscious experience cannot be one and the same process.     

Differential roles of distracters in reflexive and memory-based localization

We investigated the effects of spatial and temporal factors on manual localization of a visual target by measuring accuracy, precision, and bias. Spatial factors included manipulation of display as with or without distracters, with invariant or variant distracters, and with near or far distracters, respectively, in Experiments 1, 2, and 3. The target and distracters were of 1degrees dots differing only by luminance parameter; they were presented concurrently for 150 or 1000 ms while observers had to memorize the target location maintaining a fixed gaze. The observers' task was to reproduce the location of the target with a mouse cursor available 150 ms following stimuli offset. Results from all experiments showed that localization performance for a briefly exposed target was as accurate and precise as that for a long exposed target. Moreover, manipulation of spatial factors had no systematic effects on accuracy and precision except that near distracters yielded higher precision. Interestingly, localization performance was unbiased in 150 ms condition when there were distracters in the display, while being biased towards the fovea in 1000 ms condition regardless of their presence or absence. These results suggest a temporal dynamics in dominance-suppression between egocentric and exocentric cues in the construction of memory for location.     

Olfactory discrimination: when vision matters?

Many previous studies have attempted to investigate the effect of visual cues on olfactory perception in humans. The majority of this research has only looked at the modulatory effect of color, which has typically been explained in terms of multisensory perceptual interactions. However, such crossmodal effects may equally well relate to interactions taking place at a higher level of information processing as well. In fact, it is well-known that semantic knowledge can have a substantial effect on people's olfactory perception. In the present study, we therefore investigated the influence of visual cues, consisting of color patches and/or shapes, on people's olfactory discrimination performance. Participants had to make speeded odor discrimination responses (lemon vs. strawberry) while viewing a red or yellow color patch, an outline drawing of a strawberry or lemon, or a combination of these color and shape cues. Even though participants were instructed to ignore the visual stimuli, our results demonstrate that the accuracy of their odor discrimination responses was influenced by visual distractors. This result shows that both color and shape information are taken into account during speeded olfactory discrimination, even when such information is completely task irrelevant, hinting at the automaticity of such higher level visual-olfactory crossmodal interactions.     

Visual event-related potentials during odor labeling

Sixteen subjects judged the accuracy of visual labels whichidentified previously occurring olfactory or visual stimuli.Labels correctly identified the previous stimuli on 75% of thetrials (frequent condition) and incorrectly identified the previouscondition in 25% of the trials (rare condition). Event-relatedpotentials were recorded from nine scalp locations during theadministration of the visual labels. As would be expected, P300bwas significantly larger during the rare non-matching trials,but was independent of the modality of the previous stimulus.Previously occurring olfactory stimuli did, however, producea significant increase in negativity recorded over the leftfrontal lobe during the visual label. The amplitude of thisnegativity was found to be correlated with subjects’ performanceon the Connecticut Clinical Chemosensory Research Center testsuch that increased left frontal negativity predicted betterolfactory performance. These data suggest the presence of aleft frontal mechanism that is important for judging the accuracyof labels for olfactory, but not necessarily visual, stimuli.     

Comparison of visual and olfactory reversal learning in pigeons

Pigeons trained to discriminate between either two differentodors or two different colored lights acquired their discriminationsat the same rate. When the discrimination problems were reversedwithin a modality, however, the birds using visual cues acquirednew discriminations more rapidly than in original learning (positivetransfer), whereas the birds using olfactory cues acquired thediscrimination reversal less rapidly. On subsequent reversals,pigeons in the visual task condition developed a successivediscrimination reversal set with repeated reversals of the stimuli,while those in the olfactory condition did not. In a secondexperiment designed to assess the acquisition of redundant cues,birds received additional training with visual and olfactorycues in compound as discriminative stimuli, and were then testedwith only visual or olfactory cues. Birds previously trainedwith odor attended to visual cues in the compound, whereas birdspreviously trained to discriminate between lights did not attendto odor cues until they were presented alone. These resultsdemonstrate that the selection of stimuli may play a crucialrole in the performance of successive discrimination reversalsand suggest that, in contrast to rats, birds selectively attendto visual over olfactory cues in discrimination learning.     

Discrimination training with multimodal stimuli changes activity in the mushroom body of the hawkmoth Manduca sexta

Background

The mushroom bodies of the insect brain play an important role in olfactory processing, associative learning and memory. The mushroom bodies show odor-specific spatial patterns of activity and are also influenced by visual stimuli.

Methodology/Principal Findings

Functional imaging was used to investigate changes in the in vivo responses of the mushroom body of the hawkmoth Manduca sexta during multimodal discrimination training. A visual and an odour stimulus were presented either together or individually. Initially, mushroom body activation patterns were identical to the odour stimulus and the multimodal stimulus. After training, however, the mushroom body response to the rewarded multimodal stimulus was significantly lower than the response to the unrewarded unimodal odour stimulus, indicating that the coding of the stimuli had changed as a result of training. The opposite pattern was seen when only the unimodal odour stimulus was rewarded. In this case, the mushroom body was more strongly activated by the multimodal stimuli after training. When no stimuli were rewarded, the mushroom body activity decreased for both the multimodal and unimodal odour stimuli. There was no measurable response to the unimodal visual stimulus in any of the experiments. These results can be explained using a connectionist model where the mushroom body is assumed to be excited by olfactory stimulus components, and suppressed by multimodal configurations.

Conclusions

Discrimination training with multimodal stimuli consisting of visual and odour cues leads to stimulus specific changes in the in vivo responses of the mushroom body of the hawkmoth.     

Odor supported place cell model and goal navigation in rodents

Experiments with rodents demonstrate that visual cues play an important role in the control of hippocampal place cells and spatial navigation. Nevertheless, rats may also rely on auditory, olfactory and somatosensory stimuli for orientation. It is also known that rats can track odors or self-generated scent marks to find a food source. Here we model odor supported place cells by using a simple feed-forward network and analyze the impact of olfactory cues on place cell formation and spatial navigation. The obtained place cells are used to solve a goal navigation task by a novel mechanism based on self-marking by odor patches combined with a Q-learning algorithm. We also analyze the impact of place cell remapping on goal directed behavior when switching between two environments. We emphasize the importance of olfactory cues in place cell formation and show that the utility of environmental and self-generated olfactory cues, together with a mixed navigation strategy, improves goal directed navigation.     

Saccade latency during probability presentation of the visual targets in man

The latent periods of saccadic eye movements in response to peripheral visual stimuli were measured in 8 right-handed healthy subjects using Posner's paradigm "COST-BENEFIT". In 6 subjects, the saccade latency in response to visual target presented in expected location in valid condition was shorter than that in neutral condition ("benefit"). Increase in saccade latency in response to the visual target presented in unexpected location in valid condition versus neutral condition took place only in 4 subjects ("cost"). A decrease in left-directed saccade latency in response to expected target presented in the left hemifield and increase in saccade latency in response to unexpected left target in comparison with analogous right-directed saccades were observed in valid condition. This phenomenon can be explained by the dominance of the right hemisphere in the processes of spatial orientation and "disengage" of attention.     

The role of the posterior parietal cortex in coordinate transformations for visual-motor integration

Lesion to the posterior parietal cortex in monkeys and humans produces spatial deficits in movement and perception. In recording experiments from area 7a, a cortical subdivision in the posterior parietal cortex in monkeys, we have found neurons whose responses are a function of both the retinal location of visual stimuli and the position of the eyes in the orbits. By combining these signals area 7 a neurons code the location of visual stimuli with respect to the head. However, these cells respond over only limited ranges of eye positions (eye-position-dependent coding). To code location in craniotopic space at all eye positions (eye-position-independent coding) an additional step in neural processing is required that uses information distributed across populations of area 7a neurons. We describe here a neural network model, based on back-propagation learning, that both demonstrates how spatial location could be derived from the population response of area 7a neurons and accurately accounts for the observed response properties of these neurons.     

Role of sensory cues on food searching behavior of a captive Manta birostris (Chondrichtyes,Mobulidae)

This study reports on the first experimental research designed specifically for Manta birostris behavior. The authors attempted to learn about the feeding behavior and environmental cues influencing this behavior, as well as general cognitive ability. The preconditioned Manta's ability to identify food, on the basis of a fraction of the ordinary food signal complex, was tested. The opening of cephalic fins was considered a good indicator of feeding motivation level. The study subject animal used its biological clock to predict time and also associated a specific location with food, suggesting an ability to build up a cognitive map of its environment. Both underwater visual stimuli and olfactory stimuli had a very intense effect on food searching behavior over a 30 m distance, in contrast to visual signs from above the water surface. In addition, although an underwater visual signal resulted in a more intense response than from an olfactory signal, the specimen did not discriminate between different objects tested on the basis of visual sensation. It could therefore be suggested that food searching behavior of Mantas are governed by triggering stimuli, including smell or visual recognition, and modulated by the cognitive spatial map stored in their long‐term memory. These findings will hopefully prove useful while devising protecting policies in the natural environment and/or while keeping these animals in captivity. Zoo Biol 27:294–304, 2008. © 2008 Wiley‐Liss, Inc.     

Beware the serpent: the advantage of ecologically-relevant stimuli in accessing visual awareness

Snakes and spiders constitute fear-relevant stimuli for humans, as many species have deleterious and even fatal effects. However, snakes provoked an older and thus stronger evolutionary pressure than spiders, shaping the vision of earliest primates toward preferential visual processing, mainly in the most complex perceptual conditions. To the best of our knowledge, no study has yet directly assessed the role of ecologically-relevant stimuli in preferentially accessing visual awareness. Using continuous flash suppression (CFS), the present study assessed the role of evolutionary pressure in gaining a preferential access to visual awareness. For this purpose, we measured the time needed for three types of stimuli - snakes, spiders (matched with snakes for rated fear levels, but for which an influence on humans but not other primates is well grounded) and birds - to break the suppression and enter visual awareness in two different suppression intensity conditions. The results showed that in the less demanding awareness access condition (stimuli presented to the participants' dominant eye) both evolutionarily relevant stimuli (snakes and spiders) showed a faster entry into visual awareness than birds, whereas in the most demanding awareness access condition (stimuli presented to the participants' non-dominant eye) only snakes showed this privileged access. Our data suggest that the privileged unconscious processing of snakes in the most complex perceptual conditions extends to visual awareness, corroborating the proposed influence of snakes in primate visual evolution.     

Evidence for separate perceptive and semantic memories for odours: a priming experiment

Sixty-four subjects participated in an olfactory priming experiment comprising separate study and test phases. Priming was measured within the olfactory modality (intramodal condition) and from the visual modality to the olfactory modality (intermodal condition). In the study phase of the intramodal condition, subjects were exposed twice to a series of odours: once performing a semantic orientation task (deciding which of seven categories odour stimuli belonged to) and once performing a perceptual orientation task (judging the intensity, the hedonicity and the familiarity of odour stimuli). Half of the odour stimuli corresponded to edible products, the other half did not. The study phase of the intermodal condition was similar, with the exception that the names of the odours (instead of the odours themselves) were presented. In the test phase, subjects were presented with primed and non-primed odour targets and had to decide as fast as possible whether the target corresponded to an edible product or not. Response times and types were recorded by a computer. The analysis of response times revealed a priming effect in the intramodal condition only. Results are discussed in terms of separate perceptual and semantic subsystems that store odour representations.     

Direct evidence from parietal extinction of enhancement of visual attention near a visible hand

Brain areas exist that appear to be specialized for the coding of visual space surrounding the body (peripersonal space). In marked contrast to neurons in earlier visual areas, cells have been reported in parietal and frontal lobes that effectively respond only when visual stimuli are located in spatial proximity to a particular body part (for example, face, arm or hand) [1-4]. Despite several single-cell studies, the representation of near visual space has scarcely been investigated in humans. Here we focus on the neuropsychological phenomenon of visual extinction following unilateral brain damage. Patients with this disorder may respond well to a single stimulus in either visual field; however, when two stimuli are presented concurrently, the contralesional stimulus is disregarded or poorly identified. Extinction is commonly thought to reflect a pathological bias in selective vision favoring the ipsilesional side under competitive conditions, as a result of the unilateral brain lesion [5-7]. We examined a parietally damaged patient (D.P.) to determine whether visual extinction is modulated by the position of the hands in peripersonal space. We measured the severity of visual extinction in a task which held constant visual and spatial information about stimuli, while varying the distance between hands and stimuli. We found that selection in the affected visual field was remarkably more efficient when visual events were presented in the space near the contralesional finger than far from it. However, the amelioration of extinction dissolved when hands were covered from view, implying that the effect of hand position was not mediated purely through proprioception. These findings illustrate the importance of the spatial relationship between hand position and object location for the internal construction of visual peripersonal space in humans.     

Contextual taste cues modulate olfactory learning in C. elegans by an occasion-setting mechanism

Manipulations of context can affect learning and memory performance across species in many associative learning paradigms. Using taste cues to create distinct contexts for olfactory adaptation assays in the nematode Caenorhabditis elegans, we now show that performance in this associative learning paradigm is sensitive to context manipulations, and we investigate the mechanism(s) used for the integration of context cues in learning. One possibility is that the taste and olfactory stimuli are perceived as a combined, blended cue that the animals then associate with the unconditioned stimulus (US) in the same manner as with any other unitary conditioned stimuli (CS). Alternatively, an occasion-setting model suggests that the taste cues only define the appropriate context for olfactory memory retrieval without directly entering into the primary association. Analysis of genetic mutants demonstrated that the olfactory and context cues are sensed by distinct primary sensory neurons and that the animals' ability to use taste cues to modulate olfactory learning is independent from their ability to utilize these same taste cues for adaptation. We interpret these results as evidence for the occasion-setting mechanism in which context cues modulate primary Pavlovian association by functioning in a hierarchical manner to define the appropriate setting for memory recall.     

Sex Differences in Spatial Memory in Brown-Headed Cowbirds: Males Outperform Females on a Touchscreen Task

Spatial cognition in females and males can differ in species in which there are sex-specific patterns in the use of space. Brown-headed cowbirds are brood parasites that show a reversal of sex-typical space use often seen in mammals. Female cowbirds, search for, revisit and parasitize hosts nests, have a larger hippocampus than males and have better memory than males for a rewarded location in an open spatial environment. In the current study, we tested female and male cowbirds in breeding and non-breeding conditions on a touchscreen delayed-match-to-sample task using both spatial and colour stimuli. Our goal was to determine whether sex differences in spatial memory in cowbirds generalizes to all spatial tasks or is task-dependant. Both sexes performed better on the spatial than on the colour touchscreen task. On the spatial task, breeding males outperformed breeding females. On the colour task, females and males did not differ, but females performed better in breeding condition than in non-breeding condition. Although female cowbirds were observed to outperform males on a previous larger-scale spatial task, males performed better than females on a task testing spatial memory in the cowbirds’ immediate visual field. Spatial abilities in cowbirds can favour males or females depending on the type of spatial task, as has been observed in mammals, including humans.     

Conceptualization of above and below relationships by an insect

Relational rules such as 'same' or 'different' are mastered by humans and non-human primates and are considered as abstract conceptual thinking as they require relational learning beyond perceptual generalization. Here, we investigated whether an insect, the honeybee (Apis mellifera), can form a conceptual representation of an above/below spatial relationship. In experiment 1, bees were trained with differential conditioning to choose a variable target located above or below a black bar that acted as constant referent throughout the experiment. In experiment 2, two visual stimuli were aligned vertically, one being the referent, which was kept constant throughout the experiment, and the other the target, which was variable. In both experiments, the distance between the target and the referent, and their location within the visual field was systematically varied. In both cases, bees succeeded in transferring the learned concept to novel stimuli, preserving the trained spatial relation, thus showing an ability to manipulate this relational concept independently of the physical nature of the stimuli. Absolute location of the referent into the visual field was not a low-level cue used by the bees to solve the task. The honeybee is thus capable of conceptual learning despite having a miniature brain, showing that such elaborated learning form is not a prerogative of vertebrates.     

Free Fatty Acid and Diacylglycerol Accumulation in the Rat Brain During Recurrent Seizures Is Related to Cortical Oxygenation

Abstract: Environmental regulation of sensory function has provided an important model of plastic mechanisms mediating neural information processing. To define potential commonalities in information processing in different systems, we investigated molecular changes elicited by sensory deprivation in the developing rat olfactory and visual systems. Protein kinase C (PKC), an intracellular messenger implicated in synaptic plasticity and memory, was analyzed. Initial, developmental studies indicated that PKC activity in the soluble and particulate fractions of the olfactory bulb increased three- to fourfold from birth to 3 months of age. Unilateral olfactory deprivation prevented the developmental increase in both soluble and particulate PKC activities in the ipsilateral olfactory bulb and piriform cortex, the second-order relay. Phorbol ester binding localized PKC to intrinsic neuronal populations and their dendrites in the control and deprived bulbs. Moreover, PKC was similarly lower in the visual cortex of dark-reared rats than in light-reared controls. The changes in PKC were region specific, as activity was unchanged by either treatment in the parietal cortex, a control area that does not process primary olfactory or visual information. Our results suggest that the important intracellular messenger, PKC, is similarly regulated in entirely different sensory systems by different environmental stimuli. Consequently, different sensory systems may use common molecular mechanisms to process information.     

Predator and heterospecific stimuli alter behaviour in cattle

Wild and domestic ungulates modify their behaviour in the presence of olfactory and visual cues of predators but investigations have not exposed a domestic species to a series of cues representing various predators and other ungulate herbivores. We used wolf (Canis lupus), mountain lion (Puma concolor), and mule deer (Odocoileus hemionus) stimuli (olfactory and visual), and a control (no stimuli) to experimentally test for differences in behaviour of cattle (Bos taurus) raised in Arizona. We measured (1) vigilance, (2) foraging rates, (3) giving up density (GUD) of high quality foods and (4) time spent in high quality forage locations in response to location of stimuli treatments. In general, we found a consistent pattern in that wolf and deer treatments caused disparate results in all 4 response variables. Wolf stimuli significantly increased cattle vigilance and decreased cattle foraging rates; conversely, deer stimuli significantly increased cattle foraging rate and increased cattle use of high quality forage areas containing stimuli. Mountain lion stimuli did not significantly impact any of the 4 response variables. Our findings suggest that domestic herbivores react to predatory stimuli, can differentiate between stimuli representing two predatory species, and suggest that cattle may reduce antipredatory behaviour when near heterospecifics.     

Spatial localization of electrotactile stimuli on the fingertip in humans

This study was designed to determine the extent to which sensations elicited by discrete electrotactile stimulation can be spatially localized, with a qualitative comparison to mechanical stimulation, in a 2 x 2 electrode array on the fingertip. Electrotactile stimulation was delivered in two modes: (1) same current to all locations (constant) or (2) current adjusted to perceptual threshold of each location (varied). For each stimulus location, subjects were asked to identify the location of the stimulus. Mechanical stimulation of the same locations on the fingerpad was delivered through von Frey hairs (0.07, 0.2 and 0.4 g). The percentage of accurate responses was computed for all stimulation modes. We found that the accuracy of discrimination of stimulus location in both the constant (46%) and varied (40%) electrotactile stimulation modes was significantly higher than chance level (25%; p < 0.01). Furthermore, subjects were significantly more accurate in discriminating electrotactile stimuli in the constant than in the varied mode (p < 0.05). We also found that the accuracy of spatial discrimination was dependent on stimulation site for mechanical, but not electrotactile stimulation. Finally, we found a significant difference in accuracy over the duration of the experiment only for mechanical modes, which may indicate that electrotactile stimuli are less biased over time. These results suggest that, although low in accuracy, human subjects are able to extract spatial information from electrotactile stimuli. Further research is needed to optimize the amount of the information that can be delivered through electrotactile stimulation.     

The roles of vision and olfaction in mate location by males of the tsetse fly Glossina morsitans morsitans

The roles of visual and/or olfactory stimuli in eliciting mating responses from male Glossina morsitans morsitans Westwood were examined, using a system for automatically recording the number and duration of mating strikes made towards decoys, under controlled conditions. The results confirm that there is no olfactory component of the female sex recognition pheromone sensed by the male antennae, and the attraction of males to females appears to be visual. The absence of male-male mating strikes was the result of the absence of female sex-pheromone, rather than the presence of a repellent mating deterrent in the male cuticle. Experiments with coloured, artificial, sex-pheromone-dosed, cotton decoys showed that colour had only weak effects on attractiveness and number of encounters with decoys, and that no colour caused significant enhancement of mating responses over those shown to decoy females.