The role of attention in the localization of odors to the mouth

Odors can be perceived as arising from the environment or as part of a flavor located in the mouth. One factor that may dictate where an odor is perceived to be is concurrent gustatory stimulation in the mouth. A taste may impair the ability to attend to an odor, especially if they are perceptually similar. Alternatively, salient mouth-based features of a flavor might command attention at the expense of smell. Experiment 1 and 2, using different stimulus sets, explored the impact of perceptually similar and dissimilar pairings of tastes in the mouth and odors at the nose. In each case, these were followed by judgments of the odor's location (mouth vs. nose). Perceptual similarity had no impact on localization judgments. Experiment 3 then manipulated the salience of the olfactory and gustatory cues and showed that each could independently shift the perceived location of an odorant-salient olfactory cues toward the nose and gustatory cues toward the mouth. These findings suggest that the salient features of a flavor may command attention at the expense of olfaction and, thereby, contribute to oral localization, with implications for flavor binding.     

Olfactory test performance and its relationship with the perceived location of odors

Odors are generally perceived as arising via the nose when sniffed and as part of an orally located flavor during ingestion. The perceived location of an odor may in part be an attentional phenomenon, with concurrent oral stimulation occurring at the expense of access to the olfactory channel. Two predictions were derived from this account: (a) tasks dependent on a capacity to attend to the olfactory channel--odor discrimination and naming--should be adversely affected by oral localization; and (b) tasks not dependent upon a capacity to attend--incidental learning/recognition memory--should not. Using a procedure to generate oral localization, in which odors were presented via the nose with concurrent oral stimulation (sucrose, a viscous fluid or water), greater reported oral localization was associated with poorer odor discrimination and naming, but not with recognition memory performance. These results support the notion that attentional processes contribute to oral localization of odors by reducing the capacity to attend to the olfactory channel.     

Acute ethanol ingestion impairs appetitive olfactory learning and odor discrimination in the honey bee

Invertebrates are valuable models for increasing our understanding of the effects of ethanol on the nervous system, but most studies on invertebrates and ethanol have focused on the effects of ethanol on locomotor behavior. In this work we investigate the influence of an acute dose of ethanol on appetitive olfactory learning in the honey bee (Apis mellifera), a model system for learning and memory. Adult worker honey bees were fed a range of doses (2.5%, 5%, 10%, or 25%) of ethanol and then conditioned to associate an odor with a sucrose reward using either a simple or differential conditioning paradigm. Consumption of ethanol before conditioning significantly reduced both the rate of acquisition and the asymptotic strength of the association. Honey bees also exhibited a dose dependent reduction in arousal/attention during conditioning. Consumption of ethanol after conditioning did not affect recall 24h later. The observed deficits in acquisition were not due to the affect of ethanol on gustatory sensitivity or motor function. However, honey bees given higher doses of ethanol had difficulty discriminating amongst different odors suggesting that ethanol consumption influences olfactory processing. Taken together, these results demonstrate that an acute dose of ethanol affects appetitive learning and olfactory perception in the honey bee.     

Failure of selective attention to phonetic segments in consonant-vowel syllables

Subjects performed a two-choice speeded classification task that required selective attention to either the consonant or the vowel in synthetic consonant-vowel (CV) syllables. When required to attend selectively to the consonant, subjects could not ignore irrelevant variation in the vowel. Similarly, when required to attend selectively to the vowel, they could not ignore irrelevant variation in the consonant. These results suggest that information about an initial stop consonant and the following vowel is processed as an integral unit.     

Potentiation and overshadowing in preweanling and adult rats

Four experiments compared the aversion acquired by 18-day-old and 60-day-old rats to a flavor that was either tasted alone or in combination, simultaneously or successively, with another flavor when paired with illness. The purpose was to study temporal variables and theoretical issues pertinent to potentiation and overshadowing while investigating ontogenetic differences in these phenomena. When either the preweanlings or the adults were presented a simultaneous compound flavor (sucrose/coffee) followed by lithium chloride-induced illness, greater sucrose aversions were found than for animals conditioned on sucrose alone--that is, potentiation. Preweanlings demonstrated greater potentiation than did adults, whether the conditioned stimulus-unconditioned stimulus (CS-US) delay was 0 or 1 hr. This potentiation was eliminated by nonreinforced presentations of the alternative CS element. Potentiation was not seen when the two flavors were presented successively as the CS; instead, overshadowing occurred. Tests of configuring by extinction procedures indicated a tendency for these animals to form a configured representation of the simultaneous compound solution. This disposition for configuring tended to be more pronounced for preweanlings than for adults. Ontogenetic differences in response to configuration, CS saliency, and generalization decrement seem consistent with at least one model of potentiation and the ontogenetic differences in potentiation seen in the present experiments.     

Attention and attribute overlap in preferential choice

Attributes that are common, or overlapping, across alternatives in two-alternative forced preferential choice tasks are often non-diagnostic. In many settings, attending to and evaluating these attributes does not help the decision maker determine which of the available alternatives is the most desirable. For this reason, many existing behavioural theories propose that decision makers ignore common attributes while deliberating. Across six experiments, we find that decision makers do direct their attention selectively and ignore attributes that are not present in or associated with either of the available alternatives. However, they are as likely to attend to common attributes as they are to attend to attributes that are unique to a single alternative. These results suggest the need for novel theories of attention in preferential choice.     

Modeling panel detection frequencies by queuing system theory: an application in gas chromatography olfactometry

In continuous vigilance tasks, the number of coincident panel responses to stimuli provides an index of stimulus detectability. To determine whether this number is due to chance, panel noise levels have been approximated by the maximum coincidence level obtained in stimulus-free conditions. This study proposes an alternative method by which to assess noise levels, derived from queuing system theory (QST). Instead of critical coincidence levels, QST modeling estimates the duration of coinciding responses in the absence of stimuli. The proposed method has the advantage over previous approaches that it yields more reliable noise estimates and allows for statistical testing. The method was applied in an olfactory detection experiment using 16 panelists in stimulus-present and stimulus-free conditions. We propose that QST may be used as an alternative to signal detection theory for analyzing data from continuous vigilance tasks.     

Sensory responsiveness and the effects of equal subjective rewards on tactile learning and memory of honeybees

In tactile learning, sucrose is the unconditioned stimulus and reward, which is usually applied to the antenna to elicit proboscis extension and which the bee can drink when it is subsequently applied to the extended proboscis. The conditioned stimulus is a tactile object that the bee can scan with its antennae. In this paper we describe the quantitative relationships between gustatory antennal stimulation, gustatory proboscis stimulation, and tactile learning and memory. Bees are 10-fold more responsive to sucrose solutions when they are applied to the antenna compared to proboscis stimulation. During tactile conditioning, the sucrose solution applied to the proboscis determines the level of acquisition, whereas antennal input is of minor importance. Bees differing in their gustatory responsiveness measured at the antenna differ strongly in their tactile acquisition and memory. We demonstrate how these differences in tactile acquisition and memory can be greatly reduced by calculating equal subjective rewards, based on individual gustatory responsiveness.     

Selective attention to the chemosensory modality

Previous studies have shown that behavioral responses to auditory, visual, and tactile stimuli are modulated by expectancies regarding the likely modality of an upcoming stimulus (see Spence & Driver, 1997). In the present study, we investigated whether people can also selectively attend to the chemosensory modality (involving responses to olfactory, chemical, and painful stimuli). Participants made speeded spatial discrimination responses (left vs. right) to an unpredictable sequence of odor and tactile targets. Odor stimuli were presented to either the left or the right nostril, embedded in a birhinally applied constant airstream. Tactile stimuli were presented to the left or the right hand. On each trial, a symbolic visual cue predicted the likely modality for the upcoming target (the cue was a valid predictor of the target modality on the majority of trials). Response latencies were faster when targets were presented in the expected modality than when they were presented in the unexpected modality, showing for the first time that behavioral responses to chemosensory stimuli can be modulated by selective attention.     

Effects of a flavor-placement reversal test after different modalities of taste aversion learning

Taste aversion learning is induced through two different behavioral procedures: a short-term or concurrent (two-daily flavors) and a long-term or sequential (one-daily flavor) procedure. For the concurrent group of animals, two gustatory/olfactory stimuli are presented separately but at the same time on a daily basis. One is paired with simultaneous intragastric administration of hypertonic NaCl and the other with physiological saline. For the sequential group, the two stimuli are presented on alternate days, one of them followed by intragastric injection of the aversive stimulus and the other by saline, both after a delay of 15 min. The two groups learned the task, but when they were subjected to a flavor-placement reversal test only the sequential group was successful in achieving it. In a second experiment, three groups of animals had to learn concurrent or sequential discrimination tasks (with either simultaneous or delayed administration of the visceral stimulus) using only spatial/proprioceptive cues. The data show that none of the groups learned them under these conditions. The results are discussed in terms of the different modalities of learning. Short-term and long-term taste aversion learning are different in the anatomical structures involved, the number of trials required for acquisition and, as shown in this paper, flexibility.     

Colour-cueing in visual search

Several studies have shown that people can selectively attend to stimulus colour, e.g., in visual search, and that preknowledge of a target colour can improve response speed/accuracy. The purpose was to use a form-identification task to determine whether valid colour precues can produce benefits and invalid cues costs. The subject had to identify the orientation of a "T"-shaped element in a ring of randomly-oriented "L"s when either two or four of the elements were differently coloured. Contrary to Moore and Egeth's (1998) recent findings, colour-based attention did affect performance under data-limited conditions: Colour cues produced benefits when processing load was high; when the load was reduced, they incurred only costs. Surprisingly, a valid colour cue succeeded in improving performance in the high-load condition even when its validity was reduced to the chance level. Overall, the results suggest that knowledge of a target colour does not facilitate the processing of the target, but makes it possible to prioritize it.     

Attention to faces modulates early face processing during low but not high face discriminability

In the present study, we investigated whether attention to faces results in sensory gain modulation. Participants were cued to attend either to faces or to scenes in superimposed face—scene images for which face discriminability was manipulated parametrically. The face-sensitive N170 event-related potential component was used as a measure of early face processing. Attention to faces modulated N170 amplitude, but only when faces were not highly discriminable. Additionally, directing attention to faces modulated later processing (~230—300 msec) for all discriminability levels. These results demonstrate that attention to faces can modulate perceptual processing of faces at multiple stages of processing, including early sensory levels. Critically, the early attentional benefit is present only when the “face signal” (i.e., the perceptual quality of the face) in the environment is suboptimal.     

Attentional modulation of electrophysiological activity in auditory cortex for unattended sounds within multistream auditory environments

In three experiments, we addressed the issue of attention effects on unattended sound processing when one auditory stream is selected from three potential streams, creating a simple model of the cocktail party situation. We recorded event-related brain potentials (ERPs) to determine the way in which unattended, task-irrelevant sounds were stored in auditory memory (i.e., as one integrated stream or as two distinct streams). Subjects were instructed to ignore all the sounds and attend to a visual task or to selectively attend to a subset of the sounds and perform a task with the sounds (Experiments 1 and 2). A third (behavioral) experiment was conducted to test whether global pattern violations (used in Experiments 1 and 2) were perceptible when the sounds were segregated. We found that the mismatch negativity ERP component, an index of auditory change detection, was evoked by infrequent pattern violations occurring in the unattended sounds when all the sounds were ignored, but not when attention was focused on a subset of the sounds. The results demonstrate that multiple unattended sound streams can segregate by frequency range but that selectively attending to a subset of the sounds can modify the extent to which the unattended sounds are processed. These results are consistent with models in animal and human studies showing that attentional control can limit the processing of unattended input in favor of attended sensory inputs, thereby facilitating the ability to achieve behavioral goals.     

Disruption of conditioned taste aversion: evidence that ECS weakens the gustatory engram

Two experiments are reported concerning the neural substrate of conditioned taste aversion (CTA) and the amnesic mechanisms involved when such learning is disrupted by electroconvulsive shock (ECS). Subjects were adult male rats. In the first experiment it is demonstrated that an aversion established to a 2.5% sucrose solution can simulate the learning loss reported in earlier studies which was caused by interpolating ECS between tasting and illness when the aversion was established using a 10% sucrose cue. It is concluded that if ECS acts to disrupt the memory of the taste cue, it possibly reduces it to only about one-quarter of its original strength, a substantial deficit not readily apparent simply from the degree of aversion displayed. In the second experiment, CTAs were established using either a 2.5% sucrose cue or a 10% cue with ECS interpolated during the taste-illness interval. Animals in the two groups were subsequently confronted with a range of sucrose stimuli and their respective aversions were compared. Near identical responses were observed under all conditions tested. These findings are consistent with the theory that ECS disrupts CTA by weakening the gustatory engram.     

Temporally selective attention modulates early perceptual processing: event-related potential evidence

Some of the most important information we encounter changes so rapidly that our perceptual systems cannot process all of it in detail. Spatially selective attention is critical for perception when more information than can be processed in detail is presented simultaneously at distinct locations. When presented with complex, rapidly changing information, listeners may need to selectively attend to specific times rather than to locations. We present evidence that listeners can direct selective attention to time points that differ by as little as 500 msec, and that doing so improves target detection, affects baseline neural activity preceding stimulus presentation, and modulates auditory evoked potentials at a perceptually early stage. These data demonstrate that attentional modulation of early perceptual processing is temporally precise and that listeners can flexibly allocate temporally selective attention over short intervals, making it a viable mechanism for preferentially processing the most relevant segments in rapidly changing streams.     

Selective attention to the color and direction of moving stimuli: Electrophysiological correlates of hierarchical feature selection

Event-related brain potentials (ERPs) were recorded from subjects who attended to pairs of adjacent colored squares that were flashed sequentially to produce a perception of movement. The task was to attend selectively to stimuli in one visual field and to detect slower moving targets that contained the critical value of the attended feature, be it color or movement direction. Attention to location was reflected by a modulation of the early P1 and N1 components of the ERP, whereas selection of the relevant stimulus feature was associated with later selection negativity components. ERP indices of feature selection were elicited only by stimuli at the attended location and had distinctive scalp distributions for features mediated by “ventral” (color) and “dorsal” (motion) cortical areas. ERP indices of target selection were also contingent on the prior selection of location but initially did not depend on the selection of the relevant feature. These ERP data reveal the timing of sequential, parallel, and contingent stages of visual processing and support early-selection theories of attention that stipulate attentional control over the initial processing of stimulus features.     

Reinforcer devaluation in palatability-based learned flavor preferences

Rats exposed to simultaneous compounds of 1 neutral flavor with dilute (2%) sucrose and a 2nd flavor with dilute (2%) maltodextrin subsequently consumed both flavors in preference to a 3rd flavor that was never paired with a palatable taste. Brief training exposure under ad lib food and water minimized the post-ingestive effects of nutrients, emphasizing the contribution of palatability to these preferences. Devaluation of sucrose or maltodextrin by pairing with illness (Experiment 1) or sensory-specific satiety (Experiment 2) selectively reduced the preference for the flavor previously paired with the devalued reinforcer. Such reinforcer-specific devaluation effects suggest that palatability-based learned flavor preferences are under-pinned by a Pavlovian process whereby the cue flavor is associated with the taste of the concurrently consumed palatable reinforcer.     

Salivary conditioning with antennal gustatory unconditioned stimulus in an insect

Classical conditioning of olfactory conditioning stimulus (CS) with gustatory unconditioned stimulus (US) in insects has been used as a pertinent model for elucidation of neural mechanisms underlying learning and memory. However, a conditioning system in which stable intracellular recordings from brain neurons are feasibly obtained while monitoring the conditioning effect has remained to be established. Recently, we found classical conditioning of salivation in cockroaches Periplaneta americana, in which an odor was associated with sucrose solution applied to the mouth, and this conditioning could be monitored by activities of salivary neurons. Application of gustatory US to the mouth, however, leads to feeding movement accompanying a movement of the brain that prevents stable recordings from brain neurons. Here we investigated whether a gustatory stimulus presented to an antenna could serve as an effective US for producing salivary conditioning. Presentation of sucrose or sodium chloride solution to an antenna induced salivation and also increased activities of salivary neurons. A single pairing trial of an odor with antennal presentation of sucrose or sodium chloride solution produced conditioning of salivation or of activities of salivary neurons. Five pairing trials led to a conditioning effect that lasted for one day. Water or tactile stimulus presented to an antenna was not effective for producing conditioning. The results demonstrate that gustatory US presented to an antenna is as effective as that presented to the mouth for producing salivary conditioning. This conditioning system provides a useful model for studying the neural basis of learning at the level of singly identifiable neurons.     

Taste aversion learning: simulation of interference with the gustatory cue during conditioning

Rats were taught an aversion to a sucrose taste cue of varying strengths. The concentration of the sucrose solution was either, 10, 7.5, 5, 2.5, 1, 0.5, or 0.25% which the animals drank for 5 min. Thirty minutes later they were poisoned with lithium chloride. On the test day all animals had access to a 10% sucrose solution regardless of the concentration they had drunk on the conditioning day. Animals conditioned with a 10, 7.5, or 5% sucrose cue subsequently displayed an identically strong aversion to the 10% cue. Only those animals conditioned with a sucrose cue which was 1% or less displayed a significantly weaker aversion to the 10% cue. The results are discussed in terms of the theory that interference with taste aversion learning by such agents as pentylenetetrazol and electroconvulsive shock may have their effect by disrupting the gustatory engram. If this assumption is correct then it suggests that the memory of the gustatory cue may be stored, at least prior to poisoning, in a quite labile state and an apparently limited disruption of taste aversion learning may in fact represent a substantial amnesic effect.     

Visual salience in the change detection paradigm: the special role of object onset

The relative efficacy with which appearance of a new object orients visual attention was investigated. At issue is whether the visual system treats onset as being of particular importance or only 1 of a number of stimulus events equally likely to summon attention. Using the 1-shot change detection paradigm, the authors compared detectability of new objects with changes occurring at already present objects--luminance change, color change, and object offset. Results showed that appearance of a new object was less susceptible to change blindness than changes that old objects could undergo. The authors also investigated whether it is onset per se that leads to enhanced detectability or onset of an object representation. Results showed that the onset advantage was eliminated for onsets that did not correspond with the appearance of a new object. These findings suggest that the visual system is particularly sensitive to the onset of a new object.