Psychophysical exponents for single papillae: A comparison with whole-mouth exponents

Taste intensity judgments in response to chemical stimulation of single human fungiform papillae were obtained by the method of magnitude estimation. Intensity scales constructed from these data were compared to those obtained from the same subjects in response to stimulation of the whole mouth. Single papilla functions conformed well to the power law governing the growth of sensation magnitude, although a minority of single papilla functions exhibited peaking of response magnitude at intermediate concentrations, followed by a decline in response magnitude at higher concentrations. Single papilla exponents were found to be lower than whole mouth exponents and were positively correlated with detection and/or recognition thresholds of the papillae for most compounds. Exponents of summated single papilla functions were greater than median single papilla exponents, but their relative magnitude vs. whole mouth exponents varied by test compound. The data were discussed in relationship to previous findings concerning flow rate effects, threshold-exponent relationships across the tongue surface, and level-dependent spatial summation.     

Bulimia and taste: possible interactions

Potential abnormalities of taste were examined in bulimic subjects who purged by vomiting and in controls. When spatial testing of the tongue and palate was performed by direct local application of sweet, salty, sour, and bitter solutions, bulimics showed a selective spatial loss on the palate. The palate may be affected by purging because vomit is directed toward the roof of the mouth where the palate receptors are located. The data suggest that the acid in vomit damages these receptors. Bulimics and controls did not differ in their basal ratings of intensity or pleasantness of sweet, salty, sour, and bitter stimuli when these were sipped rather than directly applied to the tongue. However, after ingesting a glucose load, controls found sweet taste significantly less pleasant, whereas bulimics did not. The results suggest that bulimics may also have an abnormal experience of satiety.     

Effect of presentation procedure on taste intensity functions

Magnitude estimations were made for the taste intensity of sodium chloride (NaCl) and quinine sulfate (QSO₄) presented by three different methods: sip, anterior dorsal tongue flow, and whole-mouth flow. Power functions fitted to the data indicate that, for the anterior tongue stimulus (NaCl), the two flowing procedures produced lower exponents than did the sip procedure. For the posterior tongue stimulus (QSO₄), the exponent obtained with dorsal tongue flow was lower than the exponents obtained with either of the whole-mouth procedures, sip or flow. The results are compared to previous experiments on ratio scaling of taste intensity to elucidate the effects of several procedural variables.     

The effect of stimulus delivery technique on perceived intensity functions for taste stimuli: Implications for fMRI studies

Classic psychophysical studies have provided significant information on the psychophysical functions for taste stimuli. With the advent of fMRI, studies are being conducted that provide insight into central processing of gustation in humans. However, fMRI experiments impose physical limitations on stimulus delivery. In the present study, we compared psychophysical functions relating perceived intensity to concentration, derived from previous studies that used the traditional sip-and-spit and dorsal flow delivery techniques, to psychophysical functions generated in this study using a simulated stimulus delivery technique (SSDT). The SSDT delivered minute quantities of taste stimuli to the dorsal surface of the tongue, just as in an fMRI scanner. As was hypothesized, the results indicated that slopes of intensity functions were dependent on the type of stimulus delivery technique. The SSDT resulted in slopes that were more similar to those generated by dorsal flow than by sip-andspit stimulus delivery techniques, suggesting the importance of considering the influence of stimulus delivery on psychophysical response in designing and interpreting experiments.     

Psychophysics of taste lateralization on anterior tongue

There have been very few investigations of the spatial properties of taste stimuli localized to specific areas of the oral cavity. This is surprising, since the spatial localization of taste sensations may contribute to the overall taste percept, much as do quality, intensity, and the temporal characteristics of tastes. The difficulty in eliminating the confounding factor of a tactile sensation may partially account for the paucity of such studies, since a gustatory stimulus cannot be presented as a liquid without a tactile component. As a step toward understanding the localizability of gustatory sensations, we designed a yoked stimulator and an experimental procedure to control for tactile cues. Lateral discrimination was evaluated at the tip of the tongue with four taste stimuli (sodium saccharin, sodium chloride, citric acid, and quinine hydrochloride) by presenting a taste and a blank solution simultaneously at two locations on the tongue. We found that subjects could lateralize all four taste stimuli in the absence of any discriminative tactile cues. Subjects' ability to lateralize varied as a psychometric function of the stimulus concentration. Detection thresholds, measured in a forced-choice two-interval staircase procedure with the same yoked stimulator that was used in the lateralization task, were always lower than lateralization thresholds, and both lateralization and detection thresholds were correlated within subjects. Subjects were unable to lateralize taste cues on a nongustatory surface under the upper lip at the highest tested concentrations, at which performance was 100% on a gustatory surface (dorsal anterior tongue). These results show that (1) taste compounds can be lateralized in the absence of any discriminative mechanical cue (but only on the gustatory epithelium) and (2) although the localization of a compound does not logically require conscious detection of the taste (cf. blind sight), subjects always detected a taste when they were able to lateralize.     

Comparison of taste qualities elicited by tactile,electrical, and chemical stimulation of single human taste papillae

Taste quality responses elicited by chemical, electrical, and tactile stimulation of 40 single fungiform papillae in four subjects were examined. A comparison of responses revealed differences in taste quality mediation for the three different classes of stimuli. Chemical stimulation elicited multiple taste qualities from a greater proportion of papillae than did either electrical or tactile stimulation. In addition, the chemical data revealed the presence of consistent bitter-sour and sour-salty confusions. Both tactile and electrical stimulation elicited relatively few bitter and sweet responses, as compared with either sour or salty responses. However, correct quality identification in these papillae for bitter and sweet compounds was no different from that for sour or salty compounds. In addition, electrical stimulation elicited a greater proportion of salty responses than did tactile stimulation. Comparison of the taste quality elicited by either tactile or electrical stimulation of a papilla with the taste quality exhibiting the greatest relative chemical sensitivity in the papilla also revealed independence of responding, and it was observed that both electrical and tactile stimulation elicited consistent taste quality responses from papillae in which these taste qualities could not be elicited by any concentration of any chemical test compound. The observed differences in quality judgments are discussed as possibly being the result of cross-modal gustatory associations resulting from nongustatory components of inadequate stimulation.     

Stability of magnitude production scales for oral and nonoral vibrotactile mechanoreceptive systems

The stability of oral and nonoral vibrotactile magnitude scales produced by the method of magnitude production was investigated for 11 adult subjects. Each subject was tested over four sessions separated by 1-wk. intervals. Data analyses included correlations across trials of power function slopes as well as individual adjustments of intensity. Analysis showed a lack of stability over time for the slope functions obtained on the thenar eminence but moderate stability for the functions obtained on the tongue. Individual adjustments on intensity showed consistently high correlations across trials and structures. It was concluded that the subjects' responses were guided by the imposed experimental variables and that each individual's scaling was not necessarily reflected by the correlations based on power function slopes established over the four trials.     

Spatial interactions of taste stimuli on the human tongue

Two nearby loci on the human tongue were stimulated with solutions representing the four basic taste qualities. The recognition threshold for each test stimulus was measured when a strong concentration of one of the four qualities or water was placed nearby. Decreased sensitivity for the test stimulus resulted when both stimuli were the same quality, with the exception of an enhancement of bitter sensitivity by QHCl. The only effect across qualities was a suppression of citric acid by NaC1. In contrast, classical studies reported across-quality enhancement. Stimulation of two loci on the same half of the tongue and the use of modern psychophysical techniques may explain the differences between these and earlier results.     

The duration and perceived intensity of sucrose taste

Thirty-two subjects judged the perceived intensity of each of four concentrations of sucrose over 2 min. Stimuli were either sipped and expectorated or flowed over the subject’s extended tongue. Ratio judgments on a line scale and category ratings were made. Sixteen of the subjects had had extensive training in judging the sensory attributes of food products, and another group of 16 subjects were untrained. The perceived intensity of sucrose rose to a peak within 5 or 10 sec, and then declined over 2 min. In both the sip and the flow conditions, the taste disappeared completely for 26 of 32 subjects. Stronger concentrations were perceived as having greater peak intensities and longer lasting taste. The differences between concentrations were enhanced when sipped rather than flowed over the tongue. Judgments of intensity and duration were largely unaffected by the training level of subjects and the use of different rating scales.     

Sensation magnitude scales for vibrotactile stimulation of the tongue and thenar eminence

This study represents the first attempt to generate magnitude-estimation and magnitude-balance functions for the human tongue. Vibrotactile magnitude scales for the tongue and thenar eminence of the hand were investigated for nine adult subjects. The methods of magnitude estimation and magnitude production were used to obtain magnitude-balance functions for both structures. The magnitude-estimation task resulted in magnitude functions for the tongue which were similar in slope value to those found for the thenar eminence. The magnitude-production task also gave magnitude functions which were similar in slope value for the two structures.     

Gustatory cross adaptation: Sourness and bitterness

In five experiments, Sa were presented with a variety of sour and bitter compounds after the tongue was rinsed with distilled H₂0, QHCl, urea, or citric acid. All the acids tested were significantly less sour following adaptation to citric acid than after adaptation to distilled H₂O. The taste of these acids was not affected by rinsing the tongue with QHCl or urea. QHCl adaptation markedly reduced the bitterness of some compounds, while having little effect on others, including urea and citric acid. Both urea and citric acid had smaller but reliable effects on the bitterness of QHCI. These apparently incompatible results do not seem to be the result of a simple verbal confusion between sourness and bitterness. Some compounds were not affected by any of the adapting conditions. The coding mechanisms for the sourness of acids appears to be relatively simple, while that for bitterness is more complex.     

Duration of subsegments within the intervocalic interval in stutterers and nonstutterers

Intervocalic intervals (IVI) from the contextual fluent speech of 14 stutterers and controls were examined at one-quarter speed on simultaneously prepared spectrograms and intensity x time displays. Seven subsegments within the IVI were identified and their durations compared between the two groups. Stutterers were slower than controls in transitional subsegments, corresponding to movements of the tongue and larynx, but not in steady-state subsegments. The results are interpreted as suggesting that stutterers are not able to move their laryngeal and supralaryngeal structures as quickly as nonstutterers.     

Individual psychophysical functions for 28 odorants

Individual scales of odor intensity were obtained for 28 different chemical compounds by the method of magnitude estimation. Eleven Ss participated in an experiment with 196 olfactory stimuli which differed in both quality and intensity. It was found (1) that power functions described the relationship between partial vapor pressure of the odorants and their subjective odor intensity for all Ss, (2) that all exponents were less than one but varied greatly between Ss, (3) that consistent intraindividual differences in the exponents of different odorants exist, and (4) that these are attributable to perceptual differences rather than to response bias.     

Method of stimulation, mouth movements, concentration, and viscosity: effects on the degree of taste adaptation

Although sensory adaptation, the gradual loss of sensation during prolonged stimulation, has been demonstrated in laboratory taste experiments, a comparable loss of taste intensity is not experienced in real-life eating situations. This discrepancy may be due to differences in the proximal stimuli or to differences in the ways the taste receptors are stimulated. In two experiments, the effects of four potentially relevant variables were investigated: stimulus intensity, stimulus viscosity, mouth movements, and presentation method. During the initial seconds of stimulation, adaptation to the weakest of the two solutions was faster. Although more viscous stimuli were less sweet, viscosity as such did not affect adaptation rate, nor did mouth movements. Among the three presentation methods, a sucrose-soaked filter paper on the tongue produced more adaptation than either sipping the solution or flowing it over the tongue. This suggests that even mouth movements far more subtle than those still present in the no-movement condition of a sip-and-spit experiment can disrupt the adaptation process.     

Taste aversions to mother's milk: the age-related role of nursing in acquisition and expression of a learned association.

The development of taste aversion learning to novel cues contained in mother's milk was examined in rat pups. Pups receiving distinctive mild by experimenter-delivered oral infusions followed by toxicosis formed an aversion to the dam's diet. Robust aversions were learned as early as Day 10 and were retained for at least 11 days. When the same distinctive milk was obtained directly from a foster mother through nursing, however, only weanling-age pups (over 20 days) formed an aversion. X-ray analysis of nipple location in the mouths of suckling pups suggested that pups between the ages of 10 and 21 days receive milk at a similar tongue locus. Flavored milk was then delivered at specific time intervals in controlled quantities through tongue cannulas implanted at loci corresponding to the nipple position shown by the X-rays. Cannulated preweanling pups that were attached to a nipple during milk delivery failed to associate the taste cue with illness, whereas both preweanlings off the nipple and weanlings on the nipple acquired aversions to the taste cue in the milk. The evidence obtained in these experiments suggests that pups of all ages are incapable of expressing a taste aversion in a nursing situation and that preweanling pups in particular are also dificient in acquiring aversions within a suckling context. The inability of preweanling pups to acquire taste aversions in a nursing situation appears to result from a failure to associate taste cues with illness rather than a failure to detect taste cues obtained from a nipple.     

Covert behavior as a direct electromyographic measure of mediating responses

Electromyographic (EMG) measures (in addition to eye activity, and electroencephalograms, EEG) were taken from the tongue and arms of children under conditions of verbal oral mediation, non-oral (leg and arm) mediation based on the concepts of right and left, and no-mediation. It was found that amplitude of tongue EMG (a measure of covert oral behavior) significantly increased only for the Verbal Mediation Group, and the increase was significantly greater than for the other two (control) groups. The arms were possible loci of mediational behavior for the Non-oral Mediation Group; arm EMG was relatively large under this non-oral mediation condition, the changes being significant for the left arm. Eye movements, possible indicators of right and left mediational activity, were greater under the non-oral mediation condition, too. These psychophysiological measures of covert behavior during mediation are thus consistent with verbal mediational theory that has been based on the study of overt behavior.     

Magnitude estimations of the course of gustatory adaptation

Magnitude estimations of 15-sec intervals were obtained during 2-min and 5-min adaptation of suprathreshold solutions of NaCl. quinine sulfate (QSO₄, and sucrose. Expectorating and adding fresh adapting solution at the end ofeach minute produced higher estimations than those obtained with continuous adaptation because of salivary dilution. Separation of the adaptation determinations by 45-sec rest intervals rather than 5 min had no effect on the form or magnitude of the functions for NaCl or sucrose, although QSO₄. adaptations separated by the shorter rest intervals were of greater magnitude. The absence of complete adaptation was largely attributed to the effects of tongue movements.     

Soft tissue anatomy of the tongue and floor of the mouth: An ultrasound demonstration

Ultrasound technology has not been used extensively in the study of normal and abnormal oral physiology and speech. Features such as soft tissue detail, real-time motion display, and subject safety make ultrasound ideal for imaging the tongue and the floor of the mouth. This study demonstrates visualization of the muscles of the tongue and floor of the mouth for a normal subject using ultrasound imaging. By employing submandibular transducer placement of real-time sector scanners, tongue anatomy and motion were continuously visualized in sagittal or coronal planes. In addition to the entire tongue surface, much of the intrinsic anatomy was identified including: the genioglossus, geniohyoid, mylohyoid, and digastric muscles; fascial boundaries such as the median fibrous septum, floor intermuscular septum, and paramedian septums; and the hyoid bone. A tongue excised from a human cadaver was scanned using ultrasound and dissected to confirm the anatomy seen in the live tongue. Tongue surface shape and configuration of the intrinsic tissue structures were observed and compared for the phonemes /k/, /u/, and /i/. Anatomical landmarks in the resting and speaking tongue are discussed as well as applications in the fields of speech science and speech pathology.     

Lingual vibrotactile threshold alterations in response to varying stimulation levels of intensity and duration.

Lingual vibrotactile thresholds were obtained at a frequency of 250 Hz by using an ascending-continuous series of stimulation during five sessions. Each of three adult male Ss attended a control session which included a prescribed period of constant tongue positioning, a pre-experimental session where the mean of six thresholds was obtained as a basis of further testing, and three experimental sessions. In the experimental sessions vibrotactile thresholds after exposure to vibratory stimulation of varying levels of intensity and duration were obtained. The findings of the control session reflected a slight pattern of decreased lingual sensitivity indicative of the influential factor of constant maintenance of tongue positioning. Results from the experimental sessions demonstrated shifts toward progressively decreased lingual sensitivity after exposure to vibratory signals of increased levels of intensity and duration. A recovery phase to near-normal sensitivity followed each stimulus presentation.     

The stimulus range effect: Evidence for top-down control of sensory intensity in audition

The influence of intensity range on the perceived magnitude of a stimulus is well documented and usually attributed to response biases. Recent studies, however, have suggested that the range effect might be sensory in origin. To test this notion, we had one set of subjects compare loudness intervals in three conditions : a broad-range condition (15 tones, 23–95 dB SPL), a soft shortrange condition (the lowest 10 tones from the broad-range condition), and a loud short-range condition (the highest 10 tones). Nonmetric scaling showed that the broad-range and loud short-range conditionshad identical loudness functions, However, the second derivative of the loudness function was larger for the soft short-range condition than fox the broad-range condition. This pattern of results is consistent with the notion of a nonlinear ampler whose gain and degree of nonlinearity are adjusted under top-down control, so as to prevent distortion and increase discriminabztity.