Chemical senses and aging: taste versus smell

Earlier studies have suggested that aging may adversely affectthe sense of smell more than the sense of taste. Although bothsense modalities lose some absolute (threshold) sensitivity,agerelated losses of suprathreshold magnitude seem to occurcommonly in the sense of smell, less commonly in the sense oftaste. This apparent difference between taste and smell wasput to direct test in the same subjects in the same test session.Young (20–25 years) and two groups of elderly people (65–78and 80–95 years) estimated the taste intensity of variousconcentrations of NaCl and the odor intensity of various levelsof iso-amyl butyrate, under instruction to make magnitude estimationsof both kinds of stimuli on a common scale. Relative to thetaste estimates, the old gave lower odor estimates than theyoung. Also, the 80–95 year group showed, on average,a greater odor deficit than the 65–78 year group. Thisoutcome supports the main hypothesis that as age increases smelldeficits present a more serious problem than do taste deficits.     

Decline in taste and odor discrimination abilities with age, and relationship between gustation and olfaction

It is important to learn about changes in both taste and odor perceptions with increasing age, because the taste of foods we encounter in our daily life is strongly affected by their smell. This study discusses the difference in qualitative taste and odor discrimination between the elderly and the young. Tastants and odorants used in this study were presented not as single stimuli but as a taste mixture (sucrose and tartaric acid) and an odor mixture (beta-phenylethyl alcohol and gamma-undecalactone). The results showed that quality discrimination abilities of the elderly subjects for both taste and odor were significantly lower than those of the young subjects, indicating a decline in quality discrimination abilities related to age. Also, a moderate but significant correlation was observed between the taste discrimination ability and the odor discrimination ability. We measured thresholds for single-taste and odor components in mixtures and compared them between the elderly and the young to investigate the cause for these findings.     

Speeding up reaction time with invisible stimuli

Normal subjects react more quickly to a pair of visual stimuli than to a stimulus alone. This phenomenon is known as the redundant signal effect (RSE) and represents an example of divided visual attention in which signal processing is carried out in parallel to the advantage of response speed. A most interesting aspect of this phenomenon is that it can occur when one stimulus in a pair cannot be consciously detected because of hemianopia or unilateral extinction resulting from brain damage. Here, we report that a similar dissociation between visual awareness and visually guided behavior is present in normal subjects who show an RSE even when the luminance of one of a pair of stimuli is below detection threshold. The observed RSE cannot be attributed to probability summation because it violates Miller's race inequality and is likely to be related to neural summation between supra- and subthreshold stimuli. Given that a similar implicit RSE is present in hemispherectomy patients, we hypothesize that the site of this summation might be the superior colliculus (SC).     

Modality-specific neural effects of selective attention to taste and odor

The insular cortex is implicated in general attention and in taste perception. The effect of selective attention to taste on insular responses may therefore reflect a general effect of attention or it may be (taste) modality specific. To distinguish between these 2 possibilities, we used functional magnetic resonance imaging to evaluate brain response to tastes and odors while subjects passively sampled the stimuli or performed a detection task. We found that trying to detect a taste (attention to taste) resulted in activation of the primary taste cortex (anterior and mid-dorsal insula) but not in the primary olfactory cortex (piriform). In contrast, trying to detect an odor (attention to odor) increased activity in primary olfactory but not primary gustatory cortex. However, we did identify a region of far anterior insular cortex that responded to both taste and odor "searches." These results demonstrate modality-specific activation of primary taste cortex by attention to taste and primary olfactory cortex by attention to odor and rule out the possibility that either response reflects a general effect of attentional deployment. The findings also support the existence of a multimodal region in far anterior insular cortex that is sensitive to directed attention to taste and smell.     

Chemosensory cross-modal stroop effects: congruent odors facilitate taste identification

In order to explore the cross-modal cognitive associations between smell and taste, a chemosensory analogue of the Stroop task (Stroop 1935) was developed. Fourteen participants were presented with an odorant and a tastant and asked to identify the tastant as "sweet" or "sour" by pressing 1 of 2 buttons as quickly as possible. Participants were faster to name the taste when it was presented with an odor that was congruent (e.g., strawberry/sweet) than with an incongruent odor (e.g., strawberry/sour). These results support the concept of a high level of cognitive integration between the senses of smell and taste and illustrates occasions of interference between information arising from different sensory systems.     

A general rule for sensory cue summation: evidence from photographic, musical, phonetic and cross-modal stimuli

The Euclidean and MAX metrics have been widely used to model cue summation psychophysically and computationally. Both rules happen to be special cases of a more general Minkowski summation rule , where m = 2 and ∞, respectively. In vision research, Minkowski summation with power m = 3-4 has been shown to be a superior model of how subthreshold components sum to give an overall detection threshold. Recently, we have previously reported that Minkowski summation with power m = 2.84 accurately models summation of suprathreshold visual cues in photographs. In four suprathreshold discrimination experiments, we confirm the previous findings with new visual stimuli and extend the applicability of this rule to cue combination in auditory stimuli (musical sequences and phonetic utterances, where m = 2.95 and 2.54, respectively) and cross-modal stimuli (m = 2.56). In all cases, Minkowski summation with power m = 2.5-3 outperforms the Euclidean and MAX operator models. We propose that this reflects the summation of neuronal responses that are not entirely independent but which show some correlation in their magnitudes. Our findings are consistent with electrophysiological research that demonstrates signal correlations (r = 0.1-0.2) between sensory neurons when these are presented with natural stimuli.     

Suggestions for the functional relationship between differently located taste buds and vomeronasal olfaction in several mammals

In almost all mammals a well developed, paired and blind ending vomeronasal Organ (VNO) situated within the basement of the nasal septum, communicates with the oral cavity. This contact is established by two nasopalatine ducts, which penetrate the rostral palate close to the incisors. These ducts open orally into the sulcus which moulds the palatine papilla. In several mammals taste buds were found in the epithelium of the patatine papilla located within the nasopalatine ducts or close to their oral openings. Presumably these taste buds interact with the vomeronasal olfaction. It is likely that they are leading to a chemosensory sensation comparable to the combination of normal taste and smell. As not all mammals with a functionable VNO possess taste buds in this position, an inspection of the rostral part of the tongue which touches the palatine papilla presented an interesting situation concerning the distribution of taste buds. This region of the tongue is almost completely free of taste buds in species like Tupaia glis and Didelphis marsupialis virginiana, which have taste buds in the epithelium of their palatine papilla. In Lemur catta however, where the palatine papilla is lacking taste buds, the respective tongue part is densely covered with them. In this case it appears likely that they in a way of substitution functionally are connected with vomeronasal olfaction.     

Integration of subthreshold and suprathreshold excitatory barrages along the somatodendritic axis of pyramidal neurons

Neurons integrate inputs arriving in different cellular compartments to produce action potentials that are transmitted to other neurons. Because of the voltage- and time-dependent conductances in the dendrites and soma, summation of synaptic inputs is complex. To examine summation of membrane potentials and firing rates, we performed whole-cell recordings from layer 5 cortical pyramidal neurons in acute slices of the rat's somatosensory cortex. We delivered subthreshold and suprathreshold stimuli at the soma and several sites on the apical dendrite, and injected inputs that mimic synaptic barrages at individual or distributed sites. We found that summation of subthreshold potentials differed from that of firing rates. Subthreshold summation was linear when barrages were small but became supralinear as barrages increased. When neurons were discharging repetitively the rules were more diverse. At the soma and proximal apical dendrite summation of the evoked firing rates was predominantly sublinear whereas in the distal dendrite summation ranged from supralinear to sublinear. In addition, the integration of inputs delivered at a single location differed from that of distributed inputs only for suprathreshold responses. These results indicate that convergent inputs onto the apical dendrite and soma do not simply summate linearly, as suggested previously, and that distinct presynaptic afferents that target specific sites on the dendritic tree may perform unique sets of computations.     

A new method to assess the influence of odor on food selection in dogs

Previous research provides evidence that odor is a key driver in food selection in dogs. Dogs' flavor preferences are generally assessed through paired comparison tests based on food intake. Methods for evaluating odor preference in canines are lacking. In this study, the paired comparison test was modified by replacing standard bowls with false‐bottom bowls (FBBs). Made of two compartments separated by a drilled, stainless‐steel plate, FBBs enable odorant compounds to be placed under the food that is presented to the dogs. Several paired comparison trials were conducted on a trained canine panel with FBBs containing various odorant substances under the kibbles. Results showed that dogs were able to perceive the hidden substances and to distinguish between the bowls accordingly. These results demonstrate that the false‐bottom bowl paired comparison method could be helpful in evaluating the role of odor in dogs' food preferences, thus, also as a way of assessing food odor performance.

Practical applications

The false‐bottom bowl method is an adaptation of the paired comparison test that enables the influence of odor on dog behavior to be isolated from that stimulated by vision, taste or textural parameters. The odor impact of a hidden substance is tested under pet meal conditions. This new method could be useful in pet food industry to measure the odor potential of a new ingredient, or to understand the key food selection drivers for dogs and cats. In addition, as the olfactory stimulus is not eaten by the animal, the influence of odor in non‐food products for dogs, such as pet care and pet medicines, could also be evaluated using this method.     

Multimodal chemosensory integration through the maxillary palp in Drosophila

Drosophila melanogaster has an olfactory organ called the maxillary palp. It is smaller and numerically simpler than the antenna, and its specific role in behavior has long been unclear. Because of its proximity to the mouthparts, I explored the possibility of a role in taste behavior. Maxillary palp was tuned to mediate odor-induced taste enhancement: a sucrose solution was more appealing when simultaneously presented with the odorant 4-methylphenol. The same result was observed with other odors that stimulate other types of olfactory receptor neuron in the maxillary palp. When an antennal olfactory receptor was genetically introduced in the maxillary palp, the fly interpreted a new odor as a sweet-enhancing smell. These results all point to taste enhancement as a function of the maxillary palp. It also opens the door for studying integration of multiple senses in a model organism.     

Insectivorous birds can see and smell systemically herbivore‐induced pines

Several studies have shown that insectivorous birds are attracted to herbivore‐damaged trees even when they cannot see or smell the actual herbivores or their feces. However, it often remained an open question whether birds are attracted by herbivore‐induced changes in leaf odor or in leaf light reflectance or by both types of changes. Our study addressed this question by investigating the response of great tits (Parus major) and blue tits (Cyanistes caeruleus) to Scots pine (Pinus sylvestris) damaged by pine sawfly larvae (Diprion pini). We released the birds individually to a study booth, where they were simultaneously offered a systemically herbivore‐induced and a noninfested control pine branch. In the first experiment, the birds could see the branches, but could not smell them, because each branch was kept inside a transparent, airtight cylinder. In the second experiment, the birds could smell the branches, but could not see them, because each branch was placed inside a nontransparent cylinder with a mesh lid. The results show that the birds were more attracted to the herbivore‐induced branch in both experiments. Hence, either type of the tested cues, the herbivore‐induced visual plant cue alone as well as the olfactory cues per se, is attractive to the birds.     

Taste and Smell Perception in Aging1

The taste and smell experience of other human beings cannot be known directly. Thus our understanding of age-related changes in the perception of taste and smell is derived inferentially. Inferences based on verbal reports and on the performance of tasks involving taste and smell stimuli suggest that some older individuals are impaired. The perceptual disadvantage of older adults appears to be more marked or more easily measured for complex than for simple stimuli. Some difference between older and younger adults may not be primarily sensory; others may arise from differences only incidentally associated with aging. The study of taste and smell perception in aging continues to challenge the psychophysical investigator to define the nature and extent of age-related change and to demonstrate its underlying mechanisms.     

Odor preference in honeybees as a function of amount of reward: Tests of two explanations

How does a foraging honeybee come to prefer a color or odor paired with the large of two amounts of sucrose solution to a color or odor paired with the smaller amount? One hypothesis is that the attractiveness of a color or odor is based on the strength of its association with the taste of sucrose, which increases with the duration of concurrent color-taste or odor-taste stimulation. Another hypothesis is that the attractiveness of a color or odor is based on association with the afferent consequences of feeding, which are different for the two amounts. Both hypotheses were tested in experiments on proboscis-extension conditioning, a technique that provides better control of stimulation than is possible in work with free-flying foragers. In Experiments 1–3, which were designed to test the first hypothesis, an odor that accompanied the ingestion of sucrose on training trials was found to elicit extension of the proboscis when subsequently it was presented alone, but the duration of concurrent stimulation had no significant effect on the probability or persistence of the response. In Experiments 4 and 5, which were designed to test the second hypothesis, an odor that immediately followed the ingestion of sucrose on training trials failed to elicit extension of the proboscis when subsequently it was presented alone. The results support neither hypothesis.     

The independence and integration of olfaction and taste

To study the influences of smell on the scaling of taste, andvice versa, an unobtrusive device was developed which permitsthe independent varying of the olfactory and gustatory componentsof a flavor compound. With instant coffee and the method ofmagnitude estimation, the slopes of the olfactory, taste, andflavor curves produced by this device were similar to thoseproduced by more traditional delivery systems. To study theinteraction of taste on smell and vice versa, subjects scaledolfactory stimuli against different taste backgrounds and tastestimuli against different odorant backgrounds. Taste backgroundsdid not significantly alter the scaling of olfaction. Similarly,olfactory backgrounds did not significantly alter the scalingof taste. Finally, subjects scaled overall intensity (flavor)when presented with all combinations of four concentrationsof odorants and tastants. Smell and taste were shown to contributeindependently to the estimation of overall intensity.     

Individual differences in the chemical senses: is there a common sensitivity?

Taste, smell, and chemical irritation (so-called trigeminal sensation) combine in our daily experience to produce the supramodal sensation of flavor, are processed by partly overlapping neural mechanisms, and show functional interconnectivity in experiments. Given their collaboration in flavor formation and the well-established connections between these senses, it is plausible that polymodal detection mechanisms might contribute to individual differences in measured sensitivity. One would expect the existence of a general chemosensory sensitivity factor to result in associations among taste, smell, and trigeminal stimulation thresholds. Measures of 5 detection thresholds from all the chemical senses were assessed in the same group of young healthy subjects (n=57). An unbiased principal components analysis (PCA) yielded a 2-component solution. Component 1, on which taste thresholds loaded strongly, accounted for 29.4% of the total variance. Component 2, on which the odor and trigeminal lateralization thresholds loaded strongly, accounted for 26.9% of the total variance. A subsequent PCA restricted to a 3-component solution cleanly separated the 3 sensory modalities and accounted for 75% of the total variance. Thus, though there may be a common underlying factor that determines some individual differences in odor and trigeminal lateralization thresholds, a general chemical sensitivity that spans chemosensory modalities seems unlikely.     

Comparison of hyperalgesia induced by capsaicin injection and controlled heat injury: effect on temporal summation

The relationship between induction of central sensitization and facilitation of temporal summation to repetitive stimulation is still unclear. The aim of this study was to investigate temporal summation before and after the induction of secondary hyperalgesia by two different experimental methods: capsaicin injection and controlled heat injury. The effect of each injury model was assessed on a separate day with an interval of at least 5 days. Twelve healthy volunteers participated. Each experiment was performed using electrical, radiant heat, mechanical impact, and punctuate stimuli consecutively. The pain threshold (PT) to a single stimulus and the summation threshold to five repetitive stimuli for electrical (2?Hz) and radiant heat (0.83?Hz) were assessed within the secondary hyperalgesic area. The degree of temporal summation for stimulus intensities of 0.8, 1.0, and 1.2 times the baseline pain thresholds were evaluated by the increase in visual analogue scale (VAS) scores from the first to the fifth stimulus of the train. Further, the degrees of temporal summation were assessed for mechanical impact and punctuate stimuli within the primary and secondary hyperalgesic areas. The contra-lateral forearm served as control (no injury). The pain threshold and the summation threshold to electrical and heat stimuli decreased significantly within the secondary hyperalgesic area after the injury induced by both heat injury or capsaicin injection. However, there was no temporal summation for heat and electrical stimuli in either model. In contrast, for the mechanical impact and punctuate mechanical stimuli the degree of temporal summation was significantly facilitated in the secondary hyperalgesic areas compared with the baseline and the control arm in both models. In the primary hyperalgesic area, the degree of temporal summation was facilitated to mechanical impact and punctuate stimuli but only following the capsaicin injection. In conclusion, the temporal summation mechanism for mechanical stimuli was facilitated in the secondary hyperalgesic area.     

Comparison of hyperalgesia induced by capsaicin injection and controlled heat injury: effect on temporal summation

The relationship between induction of central sensitization and facilitation of temporal summation to repetitive stimulation is still unclear. The aim of this study was to investigate temporal summation before and after the induction of secondary hyperalgesia by two different experimental methods: capsaicin injection and controlled heat injury. The effect of each injury model was assessed on a separate day with an interval of at least 5 days. Twelve healthy volunteers participated. Each experiment was performed using electrical, radiant heat, mechanical impact, and punctuate stimuli consecutively. The pain threshold (PT) to a single stimulus and the summation threshold to five repetitive stimuli for electrical (2 Hz) and radiant heat (0.83 Hz) were assessed within the secondary hyperalgesic area. The degree of temporal summation for stimulus intensities of 0.8, 1.0, and 1.2 times the baseline pain thresholds were evaluated by the increase in visual analogue scale (VAS) scores from the first to the fifth stimulus of the train. Further, the degrees of temporal summation were assessed for mechanical impact and punctuate stimuli within the primary and secondary hyperalgesic areas. The contra-lateral forearm served as control (no injury). The pain threshold and the summation threshold to electrical and heat stimuli decreased significantly within the secondary hyperalgesic area after the injury induced by both heat injury or capsaicin injection. However, there was no temporal summation for heat and electrical stimuli in either model. In contrast, for the mechanical impact and punctuate mechanical stimuli the degree of temporal summation was significantly facilitated in the secondary hyperalgesic areas compared with the baseline and the control arm in both models. In the primary hyperalgesic area, the degree of temporal summation was facilitated to mechanical impact and punctuate stimuli but only following the capsaicin injection. In conclusion, the temporal summation mechanism for mechanical stimuli was facilitated in the secondary hyperalgesic area.     

Can Honey Bees Learn the Removal of a Stimulus as a Conditioning Cue?

Experiments investigated a Pavlovian conditioning situation where the presence and absence of the stimulus are reversed temporally with respect to the presentation of a reward. Instead of a conditioned stimulus (e.g. odor) signaling the presence of a reward, the stimulus (e.g. odor) is present in the environment except just prior to the presence of the reward. Thus, the absence of the stimulus, or offset of the stimulus (e.g. absence of odor), serves as a conditioned stimulus and is the reward cue. Honey bees (Apis mellifera) were used as a model invertebrate system, and the proboscis‐conditioning paradigm was used as the test procedure. Using both simple Pavlovian conditioning and discrimination‐learning protocols, animals learned to associate the onset of an odor as conditioned stimuli when paired with a sucrose reward. They could also learn to associate the onset of a puff of air with a sucrose reward. However, bees could not associate the offset of an order stimulus with the presentation of a sucrose reward in either a simple conditioning or a discrimination‐learning situation. These results support the model that a very different cognitive architecture is used by invertebrates to deal with certain environmental situations, including signaled avoidance.     

fMRI activation in response to odorants orally delivered in aqueous solutions

During food intake flavor perception results from simultaneous stimulation of the gustatory, olfactory and trigeminal systems. Olfactory stimulation occurs mainly through the retronasal pathway and the resulting perception is often interpreted as a taste perception, thus leading to the well-known sensory confusion between taste and olfaction. The present experiment was designed to study, with functional magnetic resonance imaging (fMRI), the cortical representation of olfactory perception in humans in response to retronasal stimulation by odorants delivered in aqueous solution. Psychophysical evaluation confirmed that the stimuli acted as pure olfactory stimuli through the retronasal pathway and did not present any taste component. Results showed activation in all brain regions previously described with neuroimaging techniques using olfactory stimulation with an odorized air flow. Piriform and orbitofrontal cortex were found activated as well as the hippocampal region, the amygdala, the insular lobe, the cingulate gyrus and the cerebellum. These results demonstrate the feasibility of efficiently stimulating the olfactory system in an fMRI scanner through the retronasal pathway with liquids delivered to the oral cavity. The presentation of olfactory stimuli in liquids to the mouth is a realistic model for the study of food-related flavor perception. This stimulation protocol furthermore allows presenting taste and olfactory stimuli separately or combined, thus allowing for direct comparisons between single modality representation, taste or olfaction, and representation of multi-modality mixtures.     

Repeated Recent Aggressive Encounters Do Not Affect Behavioral Consistency in Male Siamese Fighting Fish

Consistent individual differences in behavior suggest that individuals respond in a predictable and repeatable manner in a specific situation while differing from other individuals. Male Siamese fighting fish exhibit consistent individual differences in decision‐making strategies when they encounter a receptive female and a rival male simultaneously. However, whether these differences are altered by recent experience is unknown. We examined the influence of repeated aggressive encounters on behavioral consistency and decision‐making. Males were presented with paired female–male dummies prior to any aggressive experiences to obtain a baseline measure. Next, males either won or lost three consecutive contests against rivals and then received the paired female–male dummies after each of these encounters. Overall levels of highly aggressive behaviors were affected by contest outcome, while levels of female‐directed were not. Not surprisingly, winning a fight led to an increase in male‐directed bites, an overtly aggressive behavior that only occurs after fights have escalated. Fighting a male before encountering the dummies caused males to perform more tail beats to the dummy male, perhaps as a result of increased motivation. Males exhibited similar levels of repeatability and used the same strategies when faced with conflicting stimuli regardless of fighting experience. Thus, while winning or losing a fight impacts overall aggression, it does not influence behavioral consistency. This study demonstrates that consistent individual differences and decision‐making strategies may be resistant to recent aggressive experiences, even over a period of days.