Cross-modal interactions between olfaction and touch

We report two experiments designed to investigate the nature of any cross-modal interactions between olfactory and tactile information processing. In Experiment 1, we assessed the influence of olfactory cues on the tactile perception of fabric softness using computer-controlled stimulus presentation. The results showed that participants rated fabric swatches as feeling significantly softer when presented with a lemon odor than when presented with an animal-like odor, demonstrating that olfactory cues can modulate tactile perception. In Experiment 2, we assessed whether this modulatory effect varied as a function of the particular odors being used and/or of the spatial coincidence between the olfactory and tactile stimuli. The results replicated those reported in Experiment 1 thus further supporting the claim that people's rating of tactile stimuli can be modulated by the presence of an odor. Taken together, the results of the two experiments reported here support the existence of a cross-modal interaction between olfaction and touch.     

Action without awareness: reaching to an object you do not remember seeing

Background

Previous work by our group has shown that the scaling of reach trajectories to target size is independent of obligatory awareness of that target property and that “action without awareness” can persist for up to 2000 ms of visual delay. In the present investigation we sought to determine if the ability to scale reaching trajectories to target size following a delay is related to the pre-computing of movement parameters during initial stimulus presentation or the maintenance of a sensory (i.e., visual) representation for on-demand response parameterization.

Methodology/Principal Findings

Participants completed immediate or delayed (i.e., 2000 ms) perceptual reports and reaching responses to different sized targets under non-masked and masked target conditions. For the reaching task, the limb associated with a trial (i.e., left or right) was not specified until the time of response cuing: a manipulation that prevented participants from pre-computing the effector-related parameters of their response. In terms of the immediate and delayed perceptual tasks, target size was accurately reported during non-masked trials; however, for masked trials only a chance level of accuracy was observed. For the immediate and delayed reaching tasks, movement time as well as other temporal kinematic measures (e.g., times to peak acceleration, velocity and deceleration) increased in relation to decreasing target size across non-masked and masked trials.

Conclusions/Significance

Our results demonstrate that speed-accuracy relations were observed regardless of whether participants were aware (i.e., non-masked trials) or unaware (i.e., masked trials) of target size. Moreover, the equivalent scaling of immediate and delayed reaches during masked trials indicates that a persistent sensory-based representation supports the unconscious and metrical scaling of memory-guided reaching.     

Crossmodal visual input for odor tracking during fly flight

Flies generate robust and high-performance olfactory and visual behaviors. Adult fruit flies can distinguish small differences in odor concentration across antennae separated by less than 1 mm [1], and a single olfactory sensory neuron is sufficient for near-normal gradient tracking in larvae [2]. During flight a male housefly chasing a female executes a corrective turn within 40 ms after a course deviation by its target [3]. The challenges imposed by flying apparently benefit from the tight integration of unimodal sensory cues. Crossmodal interactions reduce the discrimination threshold for unimodal memory retrieval by enhancing stimulus salience [4], and dynamic crossmodal processing is required for odor search during free flight because animals fail to locate an odor source in the absence of rich visual feedback [5]. The visual requirements for odor localization are unknown. We tethered a hungry fly in a magnetic field, allowing it to yaw freely, presented odor plumes, and examined how visual cues influence odor tracking. We show that flies are unable to use a small-field object or landmark to assist plume tracking, whereas odor activates wide-field optomotor course control to enable accurate orientation toward an attractive food odor.     

The allocation of attention on a perpendicular visual display during reaching movements in depth

We investigated how visual attentional resources are allocated during reaching movements. Particularly, this study examined whether or not the direction of the reaching movement affected visual attention resource allocation. Participants held a stylus pen to reach their hand toward a target stimulus on a graphics tablet as quickly and accurately as possible. The direction of the hand movement was either from near to far space or the reverse. They observed visual stimuli and a cursor, which represented the hand position, on a perpendicularly positioned display, instead of directly seeing their hand movements. Regardless of the movement direction, the participants tended with quickly responding to the target stimuli located far from the start position as compared with those located near to the start position. These results led us to conclude that attentional resources were preferentially allocated in the areas far from the start position of reaching movements. These findings may provide important information for basic research on attention, but also contribute to a decrease of human errors in manipulation tasks performed with visual feedback on visual display terminals.     

The grasping side of odours

Background

Research on multisensory integration during natural tasks such as reach-to-grasp is still in its infancy. Crossmodal links between vision, proprioception and audition have been identified, but how olfaction contributes to plan and control reach-to-grasp movements has not been decisively shown. We used kinematics to explicitly test the influence of olfactory stimuli on reach-to-grasp movements.

Methodology/Principal Findings

Subjects were requested to reach towards and grasp a small or a large visual target (i.e., precision grip, involving the opposition of index finger and thumb for a small size target and a power grip, involving the flexion of all digits around the object for a large target) in the absence or in the presence of an odour evoking either a small or a large object that if grasped would require a precision grip and a whole hand grasp, respectively. When the type of grasp evoked by the odour did not coincide with that for the visual target, interference effects were evident on the kinematics of hand shaping and the level of synergies amongst fingers decreased. When the visual target and the object evoked by the odour required the same type of grasp, facilitation emerged and the intrinsic relations amongst individual fingers were maintained.

Conclusions/Significance

This study demonstrates that olfactory information contains highly detailed information able to elicit the planning for a reach-to-grasp movement suited to interact with the evoked object. The findings offer a substantial contribution to the current debate about the multisensory nature of the sensorimotor transformations underlying grasping.     

Orientation responses of the grasshopper, Melanoplus sanguinipes, to visual, olfactory and wind stimuli and their combinations

Prereproductive adults of the grasshopper, Melanoplus sanguinipes (F.) (Orthoptera, Acrididae), demonstrated orientation and movement towards both visual and olfactory stimulus sources in a still-air chamber. Visual stimuli (wheat and lima bean foliage, vertical black or yellow-green stripes, and a yellow-green broad leaf pattern) were approached more frequently than the control white background surface. Olfactory stimuli (chopped wheat foliage and a four-component, synthetic, grass odor blend of volatiles) elicited an even greater positive response than the visual stimuli. Changing the proportions of the four volatiles in the blend significantly reduced positive orientation responses to the stimulus source. Visual cues of wheat foliage and olfactory cues of either chopped wheat odor or the grass odor blend gave greater responses when combined than when presented separately.In flowing air or wind, nearly all insects demonstrated a rapid positive response to odors of chopped wheat and the grass odor blend, significantly greater than the response to the same stimuli in still air. However, positive responses to visual cues were not significantly greater in wind than in still air. When combined with the olfactory stimuli in flowing air, visual cues did not increase the incidence of response. Grasshoppers responding to grass odors in wind moved more rapidly and directly toward the source, and stopped less often and for shorter durations than insects responding to odor in still air or to visual cues.We conclude from these studies that M. sanguinipes adults show orientation behavior to both visual and olfactory stimuli from food plant sources, although leaf odors elicit a stronger positive response particularly when carried by wind.     

Olfactory Orientation and Navigation in Humans

Although predicted by theory, there is no direct evidence that an animal can define an arbitrary location in space as a coordinate location on an odor grid. Here we show that humans can do so. Using a spatial match-to-sample procedure, humans were led to a random location within a room diffused with two odors. After brief sampling and spatial disorientation, they had to return to this location. Over three conditions, participants had access to different sensory stimuli: olfactory only, visual only, and a final control condition with no olfactory, visual, or auditory stimuli. Humans located the target with higher accuracy in the olfaction-only condition than in the control condition and showed higher accuracy than chance. Thus a mechanism long proposed for the homing pigeon, the ability to define a location on a map constructed from chemical stimuli, may also be a navigational mechanism used by humans.     

Auditory-olfactory integration: congruent or pleasant sounds amplify odor pleasantness

Even though we often perceive odors while hearing auditory stimuli, surprisingly little is known about auditory-olfactory integration. This study aimed to investigate the influence of auditory cues on ratings of odor intensity and/or pleasantness, with a focus on 2 factors: "congruency" (Experiment 1) and the "halo/horns effect" of auditory pleasantness (Experiment 2). First, in Experiment 1, participants were presented with congruent, incongruent, or neutral sounds before and during the presentation of odor. Participants rated the odors as being more pleasant while listening to a congruent sound than while listening to an incongruent sound. In Experiment 2, participants received pleasant or unpleasant sounds before and during the presentation of either a pleasant or unpleasant odor. The hedonic valence of the sounds transferred to the odors, irrespective of the hedonic tone of the odor itself. The more the participants liked the preceding sound, the more pleasant the subsequent odor became. In contrast, the ratings of odor intensity appeared to be little or not at all influenced by the congruency or hedonic valence of the auditory cue. In conclusion, the present study for the first time provides an empirical demonstration that auditory cues can modulate odor pleasantness.     

Role of visual and olfactory cues from agricultural hedgerows in the orientation behavior of multicolored Asian lady beetle (Coleoptera: Coccinellidae)

Harmonia axyridis Pallas is an introduced lady beetle common in eastern North American agroecosystems. Two-choice behavioral bioassays were performed to determine whether visual and olfactory stimuli from prey and host habitats could elicit taxis in wild-collected H. axyridis adults and whether beetles exhibit a preference among stimuli. Soybean aphid (Aphis glycines Matsumura) spends much of the year in agricultural hedgerows residing on buckthorn (Rhamnus cathartica L), and H. axyridis is frequently observed feeding on aphids in this habitat. Olfactory bioassays were performed in a Y-tube olfactometer and tested the response of beetles to the odor of buckthorn leaves, apple leaves (Malus domestica Borkh.), and buckthorn leaves both naturally and artificially infested with A. glycines. No differences were observed between the numbers of beetles moving toward the odor of buckthorn artificially infested with A. glycines and uninfested buckthorn, but more beetles preferred naturally infested buckthorn over uninfested buckthorn. Visual bioassays were performed in an acrylic tube arena,and tested beetle response to silhouettes and to apple and buckhorn leaves. Beetles were significantly more likely to choose silhouettes over blank space in visual trials. Significantly more beetles moved toward buckthorn leaves than blank space, but beetles did not discern between apple and buckthorn until olfactory cues were also included. This study lays the foundation for future work examining the response of H. axyridis to visual and olfactory cues in Ontario agroecosystems, which could help enhance effectiveness of H. axyridis as a biological control and mitigate its impacts as a pest species.     

Toward a framework for the evaluation of feature binding in pigeons

Pigeons were trained in a new procedure to test for visual binding errors between the dimensions of color and shape. In Experiment 1, pigeons learned to discriminate a target compound from 15 non-target compounds (constructed from four colors and shapes) by choosing one of two hoppers in a two-hopper choice task. The similarity of the target to non-target stimuli influenced choice responding. In Experiment 2, pigeons learned to detect a target compound when presented with a non-target compound within the same trial under conditions of simultaneity and sequentiality. Non-target trials were arranged to allow for the testing of binding errors (i.e., false identifications of the target on certain non-target trials). Transient evidence for binding errors in two of the birds occurred at the start of two-item training, but decreased with training. The experiments represent an important step toward developing a framework for the evaluation of visual feature binding in nonhumans.     

A simple and reliable test of olfactory learning and memory in mice

The present paper describes a quick and efficient method for assessing olfactory discrimination learning in mice. In training mice received trials in which one odor (CS+) was paired with sugar and another odor (CS-) was paired with no sugar. When the mice were subsequently placed in a chamber with CS+ odor at one end and CS- odor at the other, they spent more time digging in CS+ than in CS- odor. In Experiment 2 mice trained with this procedure and tested after 60 days also spent more time digging in CS+ than CS- in the test phase, indicating that this olfactory discrimination task is effective for assessing long-term memory. In addition to the outbred strain of CD1 mice used in Experiments 1 and 2, C57Bl/6NCr/BR and DBA/2NCr/BR mice used in Experiment 3 also acquired this learned odor discrimination. Moreover, Experiment 4 showed that DBA animals were capable of acquiring this odor discrimination after receiving only two training trials (one exposure each to CS+ and CS-) per day for 4 days.     

Grasping the past. delay can improve visuomotor performance.

"Optic ataxia" is caused by damage to the human posterior parietal cortex (PPC). It disrupts all components of a visually guided prehension movement, not only the transport of the hand toward an object's location, but also the in-flight finger movements pretailored to the metric properties of the object. Like previous cases, our patient (I.G.) was quite unable to open her handgrip appropriately when directly reaching out to pick up objects of different sizes. When first tested, she failed to do this even when she had previewed the target object 5 s earlier. Yet despite this deficit in "real" grasping, we found, counterintuitively, that I.G. showed good grip scaling when "pantomiming" a grasp for an object seen earlier but no longer present. We then found that, after practice, I.G. became able to scale her handgrip when grasping a real target object that she had previewed earlier. By interposing catch trials in which a different object was covertly substituted for the original object during the delay between preview and grasp, we found that I.G. was now using memorized visual information to calibrate her real grasping movements. These results provide new evidence that "off-line" visuomotor guidance can be provided by networks independent of the PPC.     

Colour Terms Affect Detection of Colour and Colour-Associated Objects Suppressed from Visual Awareness

The idea that language can affect how we see the world continues to create controversy. A potentially important study in this field has shown that when an object is suppressed from visual awareness using continuous flash suppression (a form of binocular rivalry), detection of the object is differently affected by a preceding word prime depending on whether the prime matches or does not match the object. This may suggest that language can affect early stages of vision. We replicated this paradigm and further investigated whether colour terms likewise influence the detection of colours or colour-associated object images suppressed from visual awareness by continuous flash suppression. This method presents rapidly changing visual noise to one eye while the target stimulus is presented to the other. It has been shown to delay conscious perception of a target for up to several minutes. In Experiment 1 we presented greyscale photos of objects. They were either preceded by a congruent object label, an incongruent label, or white noise. Detection sensitivity (d’) and hit rates were significantly poorer for suppressed objects preceded by an incongruent label compared to a congruent label or noise. In Experiment 2, targets were coloured discs preceded by a colour term. Detection sensitivity was significantly worse for suppressed colour patches preceded by an incongruent colour term as compared to a congruent term or white noise. In Experiment 3 targets were suppressed greyscale object images preceded by an auditory presentation of a colour term. On congruent trials the colour term matched the object’s stereotypical colour and on incongruent trials the colour term mismatched. Detection sensitivity was significantly poorer on incongruent trials than congruent trials. Overall, these findings suggest that colour terms affect awareness of coloured stimuli and colour- associated objects, and provide new evidence for language-perception interaction in the brain.     

Olfactory and visual stimuli affecting host plant detection in Homalodisca coagulata (Hemiptera: Cicadellidae)

The relative effects of visual and olfactory stimuli on host plant detection in immature and adult Homalodisca coagulata Say (Homoptera: Cicadellidae) were studied using a novel olfactometer and factorial experimental designs. Colored, gray, and white cards were used as visual targets. Each card was attached to a glass thistle tube from which host-plant odor (from Vigna unguiculata L.) or blank, humidified air was dispensed. Visual + odor stimuli combinations were presented in no-choice tests. Nymphs were released onto a perch stick downwind from the target. Nymph response to color + odor treatments was measured by the duration of orientation behavior, residence time on the perch, and percentage of individuals that jumped to the target. The assay was modified so that adults crawled from the perch onto the target. Adult response was measured by the duration of individual behaviors (e.g., foraging) and by their position and residence time on the target. Both main effects and interactive effects of the stimuli were observed. Nymphs showed a decrease in orientation and residence times in the colored target + host odor treatments and increased jumping response in the gray + host odor treatment. When adults were exposed to host odor, the duration of foraging behavior increased, whereas crawling and phototactic behaviors decreased. Although nymphs and adults responded to visual stimuli + blank air treatments, host odor enhanced their responses. The primary effect of host odor on host detection behavior may be to enhance H. coagulata responsiveness to visual cues.     

Variation in female guppy preference for male olfactory and visual traits

Animals often use different sensory systems to assess different sexually selected signals from potential mates. However, the relative importance of different signals on mate choice is not well understood in many animal species. In this study, we examined the relative importance of male olfactory and visual cues on female preference in the guppy Poecilia reticulata. We used digitally modified male images to standardize visual stimuli. We found that, regardless of whether females were presented without male visual stimuli or with identical male visual stimuli, they preferred stimuli with the odor of males to those without. However, when females were allowed to choose between dull male visual stimuli with male odor, and brightly colored male visual stimuli without male odor, there was no clear preference for either. Some females preferred the dull male visual stimuli with male odor, whereas some other females preferred the brightly colored male visual stimuli without male odor. These results indicate that the relative importance of olfactory and visual cues in female mate preference varied between individuals.     

Social Groups Prioritize Selective Attention to Faces: How Social Identity Shapes Distractor Interference

Human faces automatically attract visual attention and this process appears to be guided by social group memberships. In two experiments, we examined how social groups guide selective attention toward in-group and out-group faces. Black and White participants detected a target letter among letter strings superimposed on faces (Experiment 1). White participants were less accurate on trials with racial out-group (Black) compared to in-group (White) distractor faces. Likewise, Black participants were less accurate on trials with racial out-group (White) compared to in-group (Black) distractor faces. However, this pattern of out-group bias was only evident under high perceptual load—when the task was visually difficult. To examine the malleability of this pattern of racial bias, a separate sample of participants were assigned to mixed-race minimal groups (Experiment 2). Participants assigned to groups were less accurate on trials with their minimal in-group members compared to minimal out-group distractor faces, regardless of race. Again, this pattern of out-group bias was only evident under high perceptual load. Taken together, these results suggest that social identity guides selective attention toward motivationally relevant social groups—shifting from out-group bias in the domain of race to in-group bias in the domain of minimal groups—when perceptual resources are scarce.     

High hunger state increases olfactory sensitivity to neutral but not food odors

Understanding how hunger state relates to olfactory sensitivity has become more urgent due to their possible role in obesity. In 2 studies (within-subjects: n = 24, between-subjects: n = 40), participants were provided with lunch before (satiated state) or after (nonsatiated state) testing and completed a standardized olfactory threshold test to a neutral odor (Experiments 1 and 2) and discrimination test to a food odor (Experiment 2). Experiment 1 revealed that olfactory sensitivity was greater in the nonsatiated versus satiated state, with additionally increased sensitivity for the low body mass index (BMI) compared with high BMI group. Experiment 2 replicated this effect for neutral odors, but in the case of food odors, those in a satiated state had greater acuity. Additionally, whereas the high BMI group had higher acuity to food odors in the satiated versus nonsatiated state, no such differences were found for the low BMI group. The research here is the first to demonstrate how olfactory acuity changes as a function of hunger state and relatedness of odor to food and that BMI can predict differences in olfactory sensitivity.     

Effects of Haptic Information on the Perception of Dynamic 3-D Movement

This study examined effects of hand movement on visual perception of 3-D movement. I used an apparatus in which a cursor position in a simulated 3-D space and the position of a stylus on a haptic device could coincide using a mirror. In three experiments, participants touched the center of a rectangle in the visual display with the stylus of the force-feedback device. Then the rectangle''s surface stereoscopically either protruded toward a participant or indented away from the participant. Simultaneously, the stylus either pushed back participant''s hand, pulled away, or remained static. Visual and haptic information were independently manipulated. Participants judged whether the rectangle visually protruded or dented. Results showed that when the hand was pulled away, subjects were biased to perceive rectangles indented; however, when the hand was pushed back, no effect of haptic information was observed (Experiment 1). This effect persisted even when the cursor position was spatially separated from the hand position (Experiment 2). But, when participants touched an object different from the visual stimulus, this effect disappeared (Experiment 3). These results suggest that the visual system tried to integrate the dynamic visual and haptic information when they coincided cognitively, and the effect of haptic information on visually perceived depth was direction-dependent.     

Implicit learning and implicit memory for odors: the influence of odor identification and retention time

One hundred and fifty-two subjects, divided into eight groups, were exposed to a room with a low concentration of either orange or lavender and to an odorless room. In a careful double-blind procedure, neither the subjects nor the experimenters were made aware of the presence of the odors in the experimental conditions. Later they were asked to indicate how well each of 12 odor stimuli, including the experimental and control odors, befitted each of 12 visual contexts, including the exposure rooms. At the end of this session they rated the pleasantness and the familiarity of the odors, and identified them by name. Finally they were debriefed and asked specifically whether they had perceived the experimental odors anywhere in the building. The results of four subjects who answered positively to the latter question were omitted. The results confirm the earlier finding that non-identifiers implicitly link odor and exposure room, whereas identifiers do not show such a link. It is suggested that episodic information is an essential constituent of olfactory memory and that its function is comparable to that of form and structure in visual and auditory memory systems.