The relationship between PTC taster status and taste thresholds in young adults

Ju-Hee Hong, DDS,a Jin-Woo Chung, DDS, PhD,b Young-Ku Kim, DDS, PhD,c Sung-Chang Chung, DDS, PhD,c Sung-Woo Lee, DDS, PhD,c and Hong-Seop Kho, DDS, PhD,b Seoul, Korea
SEOUL NATIONAL UNIVERSITY

Objective. The aim of this study was to compare taste detection and recognition thresholds of young males and females, and
young phenylthiocarbamide (PTC) tasters and nontasters for stimuli representing sweet, sour, bitter, salty, and umami classes of taste sensations. Study design. Thirty-eight men and 37 women (mean [SD] age = 24.5 [2.5] years) were classied as PTC tasters and nontasters according to their PTC recognition thresholds. Detection and recognition thresholds for the non-PTC stimuli were determined using a 2-alternative, forced choice procedure. Results. The detection thresholds for quinine-HCl and sucrose and the recognition threshold for quinine-HCl were signicantly higher in the PTC nontasters than in the tasters. The PTC threshold showed signicant correlations with detection or recognition thresholds for sucrose, sodium chloride, quinine-HCl, and monosodium glutamate. The sucrose recognition threshold was lower in women than in men. Conclusion. In this study, gender and PTC taster status were found to be associated with thresholds for sucrose and quinine-HCl. (Oral Surg Oral Med Oral Pathol Oral Radiol Endod 2005;99:711-5)

Taste refers to the sensation arising from the direct stimulation of the taste bud receptors by substances dissolved in saliva.1 Classically, it has been suggested that there are 4 basic tastes: sweet, sour, bitter, and salty. Such categories have been hotly debated, however, and more recently some authors have suggested that a category of umami taste be added as well.2-4 Umami is a Japanese term applied to the avor of monosodium glutamate. The methods for evaluating taste function can be classied into qualitative and quantitative taste tests. A qualitative test would be one that determines a subjects response to a tastant without quantifying the response. A quantitative test, on the other hand, provides a numerical metric of the subjects function. Of the available quantitative tests, threshold tests are perhaps the most widely used. A taste detection threshold test establishes the lowest concentration at which a substance can be distinguished from water, whereas a taste
a

recognition threshold test determines the lowest concentration at which the taste quality of a substance can be identied.5 Phenylthiocarbamide (PTC) and its chemically related compound, 6-n-propylthiouracil (PROP), provide extremely bitter taste to some subjects (tasters) but are tasteless or only slightly bitter to others (nontasters).6-9 Although approximately 15% to 30% of people are known to be genetically nontasters,1,10 the taster/nontaster frequency is different among ethnic groups.11-13 PTC and PROP perception has been known to correlate with the taste perception of various primary taste quality.14-16 However, these results were based on food aversions and taste intensity, not thresholds. Therefore, a comparison of these taste detection and recognition thresholds between the tasters and nontasters is warranted. The aim of this study was to report the taste detection and recognition thresholds of the 5 taste sensations and to compare these taste thresholds between tasters and nontasters in the young adult population. MATERIAL AND METHODS Subjects Seventy-ve subjects (38 males and 37 females, mean age: 24.562.5 years), who were students of the College of Dentistry, Seoul National University, were enrolled in the study. A questionnaire, which included questions regarding chronic sinusitis, chronic obstructive pulmonary disease, diabetes, psychological disorders, the loss of olfactory sense, and dry mouth, was used to select the 711

Graduate Student, Department of Oral Medicine and Oral Diagnosis, College of Dentistry, Seoul National University, Seoul, Korea. b Assistant Professor, Department of Oral Medicine and Oral Diagnosis, College of Dentistry, Seoul National University. c Professor, Department of Oral Medicine and Oral Diagnosis, College of Dentistry, Seoul National University. Received for publication Apr 11, 2003; returned for revision Jul 30, 2004; accepted for publication Aug 6, 2004. Available online 5 November 2004. 1079-2104/$ - see front matter 2005 Elsevier Inc. All rights reserved. doi:10.1016/j.tripleo.2004.08.004

712 Hong et al Table I. The proportion of tasters and nontasters, n(%)

Male Taster Nontaster Total 30 (79) 8 (21) 38 (100) Female 30 (81) 7 (19) 37 (100) Total 60 (80) 15 (20) 75 (100)

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subjects. The subjects had no history of serious illness and did not show any positive responses to the checklists included in the questionnaire. None of the subjects wore dentures. They were instructed to refrain from smoking, eating, or drinking anything but water for at least 1 hour prior to being tested.17-19 Preparation of taste solutions Five series of stimulus uids were prepared, 1 each for sucrose (sweet, 2.4 3 104 to 1.0 M), sodium chloride (salty, 2.4 3 104 to 1.0 M), citric acid (sour, 7.5 3 106 to 3.2 3 102 M), quinine-HCl (bitter, 7.5 3 108 to 3.2 3 104 M), and monosodium glutamate (umami, 2.4 3 104 to 1.0 M). In each series, successive solutions, which comprised a total of 30 grades that differed by 0.125 log units of the molar concentration. Fourteen grades of PTC solutions (0.50 3 105 to 4.20 3 102 M) were prepared. All solutions were prepared from reagent grade chemicals using distilled, deionized water. All uids were stored at 48C and brought to room temperature before use. Determination of taste detection threshold The experiment was performed in the order of measuring the taste detection threshold, the taste recognition threshold, and the PTC taster status in each subject. The taste thresholds were obtained by standard 2alternative forced choice trials.18,20 The taste solution and distilled water were dispensed to the subject as 5-mL samples in a 15-mL test tube according to a random schedule. The subject rinsed his or her mouth with distilled water prior to tasting each sample. The subject then indicated which tube contained the taste solution that could be distinguished from water, even though the subject could not determine taste quality. The subject was always required to choose one tube or the other but was reassured that guess was appropriate when the taste solution was so weak as to be indistinguishable from water. The concentration of an individuals rst exposure to each taste solution was determined by the detection threshold of the previous subject for that taste. Subsequent increases and decreases in concentration depended on the individuals own responses. Any incorrect response caused an increase in the stimulus concentration on the next trial. The stimulus concentration

was decreased after 2 consecutive correct responses. The stimulus at which the concentration sequence changed from decreasing to increasing or vice versa was designated a reversal. The procedure was repeated 3 times. The rst reversal was disregarded, and the taste detection threshold was determined as the mean of the last 2 reversal concentrations. The rotation among the taste solutions continued in the order of sweet, salty, sour, umami, and bitter tastes for a period of approximately 1 hour. For 7 subjects (9.3%) who did not complete the procedure within 1 hour, the experiment was continued the next day.18 Determination of taste recognition threshold The taste recognition threshold was determined according to the same method used for determining the taste detection threshold. However, the subjects had to indicate which tube contained the taste solution and had to identify what kind of taste was contained in the taste solution. The concentration of the rst exposure to each taste solution was determined by the subjects detection threshold of that taste. The subject was always required to choose 1 tube from a choice of 2. Determination of taster status The experiment was commenced with the weakest PTC solution in the order of increasing concentration. The method was the same as that of the taste recognition threshold. The tasters were classied as those individuals able to recognize the bitterness of a PTC solution with a concentration less than 1.80 3 104 M. Subjects who could not recognize the bitterness of a solution stronger than 1.80 3 104 M were classied as nontasters.9 Statistics Student t test was used to analyze differences in the taste detection and recognition thresholds between genders in the tasters. Because of a relatively small number of nontasters, the data from only the tasters were used for the analysis of gender difference. Analyses of variance (ANOVA) for each dependent variable of taste detection and recognition thresholds were used to analyze differences between the tasters and nontasters. The interactions between gender and taste status in each dependent variable were assessed. Pearsons correlation analysis was used to examine the relationships between PTC thresholds with thresholds for 5 taste sensations. A difference was considered signicant only if the P value was less than or equal to .05. RESULTS As shown in Table I, the percentage of nontasters was 20.0% of the 75 subjects in both genders, 21.0% in males and 19.0% in females.

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Table II. Taste detection thresholds in the tasters

Sucrose (3 102 M) Male (n = 30) Female (n = 30) P* Total (n = 60) 0.89 6 0.60 0.68 6 0.36 .11 0.78 6 0.50 NaCl (3 102 M) 0.33 6 0.29 0.28 6 0.14 .39 0.30 6 0.23 Citric acid (3 104 M) 0.65 6 0.31 0.62 6 0.35 .80 0.63 6 0.33 Quinine-HCl (3 106 M) 4.77 6 5.08 3.68 6 7.51 .51 4.22 6 6.38 MSG (3 102 M) 0.22 6 0.15 0.20 6 0.10 .57 0.21 6 0.12

MSG, monosodium glutamate. *P values were obtained from Student t test.

Table III. Taste recognition thresholds in the tasters

Sucrose (3 102 M) Male (n = 30) Female (n = 30) P* Total (n = 60) 2.32 6 0.95 1.83 6 0.70 .03* 2.07 6 0.86 NaCl (3 102 M) 2.37 6 1.83 2.06 6 1.23 .45 2.21 6 1.55 Citric acid (3 104 M) 2.31 6 1.29 1.75 6 1.12 .08 2.03 6 1.23 Quinine-HCl (3 106 M) 8.93 6 9.29 7.72 6 10.51 .64 8.33 6 9.85 MSG (3 102 M) 1.01 6 1.77 0.69 6 0.55 .35 0.85 6 1.31

MSG, monosodium glutamate. P values were obtained from Student t test. *P \ .05.

Table IV. Taste detection thresholds in the tasters and nontasters

Sucrose (3 102 M) Taster (n = 60) Nontaster (n = 15) P Total (n = 75) 0.78 6 0.50 1.11 6 0.80 .05* 0.85 6 0.58 NaCl (3 102 M) 0.30 6 0.23 0.35 6 0.31 .53 0.31 6 0.24 Citric acid (3 104 M) 0.63 6 0.33 0.70 6 0.38 .58 0.65 6 0.34 Quinine-HCl (3 106 M) 4.22 6 6.38 9.25 6 9.45 .02* 5.23 6 7.31 MSG (3 102 M) 0.21 6 0.12 0.26 6 0.15 .22 0.22 6 0.13

MSG, monosodium glutamate. P values were obtained from analysis of variance (ANOVA). *P \ .05.

The detection threshold for sucrose averaged 0.78 3 102 M; 0.30 3 102 M for NaCl; 0.63 3 104 M for citric acid; 4.22 3 106 M for quinine-HCl; and 0.21 3 102 M for MSG in the tasters (Table II). Although the mean value of each detection threshold for the 5 tastes in the male tasters was higher than in the female tasters, there was no statistically signicant difference. The recognition threshold for sucrose averaged 2.07 3 102 M; 2.21 3 102 M for NaCl; 2.03 3 104 M for citric acid; 8.32 3 106 M for quinine-HCl; and 0.85 3 102 M for MSG in the tasters (Table III). The mean value of each recognition threshold for the 5 tastes in the male tasters was higher than in the female tasters but only the sucrose recognition threshold of the male tasters was signicantly higher than that of the female tasters (P \ .05). As shown in Table IV, the detection thresholds for quinine-HCl and sucrose were signicantly higher in the nontasters than the tasters (P \ .05). The PTC threshold showed signicant correlations with detection

Table V. Correlations (r) between PTC threshold and taste detection thresholds
n = 75 PTC Sucrose .378* NaCl .129 Citric acid .158 Quinine-HCl .477* MSG .326*

MSG, monosodium glutamate; PTC, phenylthiocarbamide. *P \ .01.

thresholds for sucrose (r = 0.378, P \ .01), quinine-HCl (r = 0.477, P \ .01), and monosodium glutamate (r = 0.326, P \ .01) (Table V). The recognition threshold for quinine-HCl was also signicantly higher in the nontasters than the tasters (P \ .01) (Table VI). The PTC threshold showed signicant correlations with recognition thresholds for sodium chloride (r = 0.307, P \ .01) and quinine-HCl (r = 0.410, P \ .01) (Table VII). None of the interaction terms between gender and taste status reached statistical signicance both in the taste detection and recognition thresholds.

714 Hong et al Table VI. Taste recognition thresholds in the tasters and nontasters
Sucrose (3 102 M) Taster (n = 60) Nontaster (n = 15) P Total (n = 75) 2.07 6 0.86 2.33 6 0.75 .32 2.12 6 0.84 NaCl (3 102 M) 2.21 6 1.55 2.55 6 2.63 .60 2.28 6 1.80 Citric acid (3 104 M) 2.03 6 1.23 2.75 6 3.03 .15 2.18 6 1.74 Quinine-HCl (3 106 M) 8.33 6 9.85 18.00 6 12.68 .003* 10.26 6 11.09

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MSG (3 102 M) 0.85 6 1.31 0.96 6 0.85 .80 0.87 6 1.22

MSG, monosodium glutamate. P values were obtained from analysis of variance (ANOVA). *P \ .01.

Table VII. Correlations (r) between PTC threshold and taste recognition thresholds
n = 75 PTC Sucrose 0.178 NaCl 0.307* Citric acid 0.186 Quinine-HCl 0.410* MSG 0.224

MSG, monosodium glutamate; PTC, phenylthiocarbamide. *P \ .01.

DISCUSSION The percentage of nontasters is 30% to 40% in European and American Caucasians, 5% to 15% in the Japanese population, and about 5% in the American Indian and Ainu populations.11,12 In this investigation using 75 Korean dental students, the percentage of nontasters was found to be 20%, which was higher than found in Japanese and lower than observed in European and American Caucasians. This is likely explained by a combination of factors, such as (1) ethnic differences, (2) variations in the experimental procedures, and (3) the PTC concentrations used for determining taster status. Further research is needed using a larger number of subjects to establish more denitive conclusions. The results of this study showed that the taste thresholds of the nontasters were higher than those of the tasters for all 5 stimuli evaluated in this study. These ndings were consistent with the results of the previous studies using suprathreshold taste intensity scaling and food preferences. The novel nding in the present study was the difference in taste thresholds, especially detection taste thresholds, and there were signicant differences in the detection thresholds for quinine-HCl and sucrose and in the recognition threshold for quinineHCl. The PTC threshold showed signicant positive correlations with the detection thresholds for quinineHCl, sucrose, and MSG and with the recognition thresholds for quinine-HCl and NaCl. These ndings had meaningful signicance because the ability to taste PTC was suggested to be a continuously distributed variable rather than a conventional bimodal categorical one.13,21 The previous studies also showed that PTC and PROP perception correlates with the taste perception of

various primary taste quality compounds, including sweet, sour, bitter, and salty.15,16 The nontasters were known to be relatively insensitive to quinine and the PROP sensitivity weakly correlated with sucrose as well as quinine.15,16 Work on sweeteners has shown that saccharin and sucrose tasted sweeter to the tasters.22-24 The taste thresholds for PROP and quinine showed a relationship to the percentage of food aversions.14 In addition, anatomical data also support the difference between the tasters and nontasters. The relationship between the taste intensity perception and taste bud density on the tongue was reported.25 The tasters had more fungiform papillae and taste pores on their anterior tongue.15 In this study, female subjects had lower mean values of detection and recognition thresholds for all the 5 tastes than male subjects, although these results did not reach statistical signicance except for the taste recognition threshold for sucrose. The nding of higher taste sensitivity in female adult subjects was consistent with the results of the previous studies. For example, it was reported that both males and females showed a gradual increase in sensitivity up to the age of 16 to 20 years, which was followed by an exponential decline.26 The sensitivities to PROP and quinine were similar in both genders up to the age of 16 to 20 years. However, men declined at a faster rate than women in those older than 20 years of age. For HCl, females were more sensitive tasters until adolescence, where there was almost an equal maximum sensitivity for both women and men. After this, the sensitivity to HCl also decreased more rapidly for men.26 Other studies have suggested that women tend to have lower thresholds for sucrose, citric acid, and acetic acid than men.3,18 Anatomical data are seemingly in accord with the aforementioned gender difference, in that women have more fungiform papillae and more taste buds than men.15,27 The reason for this anatomical difference is not clear, however, since it could reect differing dietary habits, smoking behaviors, and alcohol consumption, as well as possibly hormonal factors.22,27-29 In addition, the pattern of gender difference can differ according to

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15. Bartoshuk LM, Duffy VB, Miller IJ. PTC/PROP tasting: anatomy, psychophysics, and sex effects. Physiol Behav 1994; 56:1165-71. 16. Pasquet P, Oberti B, El Ati J, Hladik CM. Relationships between threshold-based PROP sensitivity and food preferences of Tunisians. Appetite 2002;39:167-73. 17. Hyde RJ, Feller RP, Sharon IM. Tongue brushing, dentifrice, and age effects on taste and smell. J Dent Res 1981;60:1730-4. 18. Weiffenbach JM, Baum BJ, Burghauser R. Taste thresholds: quality specic variation with human aging. J Gerontol 1982;37: 372-7. 19. Weiffenbach JM. Taste-quality recognition and forced-choice response. Percept Psychophys 1983;33:251-4. 20. Jones FN. A forced-choice method of limits. Am J Psychol 1956; 69:672-3. 21. Kalmus H. Improvements in the classication of the taster genotypes. Ann Hum Genet 1958;22:222-30. 22. Lucchina LA, Curtis V OF, Putnam P, Drewnowski A, Prutkin JM, Bartoshuk LM. Psychophysical measurement of 6-n-propylthiouracil (PROP) taste perception. Ann N Y Acad Sci 1998;30;855:816-9. 23. Bartoshuk LM. Bitter taste of saccharin related to the genetic ability to taste the bitter substance 6-n-propylthiouracil. Science 1979;205:934-5. 24. Gent JF, Bartoshuk LM. Sweetness of sucrose, neohesperidin dihydrochalcone, and saccharin is related to genetic ability to taste the bitter substance 6-n-propylthiouracil. Chem Senses 1983;7:265-72. 25. Miller IJ, Reedy FE. Variations in human taste bud density and taste intensity perception. Physiol Behav 1990;47:1213-9. 26. Glanville EV, Kaplan AR, Fischer R. Age, sex, and taste sensitivity. J Gerontol 1964;19:474-8. 27. Prutkin J, Duffy VB, Etter L, Fast K, Gardner E, Lucchina LA, et al. Genetic variation and inferences about perceived taste intensity in mice and men. Physiol Behav 2000;69: 161-73. 28. Kaplan RA, Glanville EV, Fischer R. Cumulative effect of age and smoking on taste sensitivity in males and females. J Gerontol 1965;20:334-7. 29. Yamauchi Y, Endo S, Yoshimura I. A new whole-mouth gustatory test procedure. II. Effects of aging, gender and smoking. Acta Otolaryngol 2002;Suppl 546:49-59.

age groups. Therefore, further studies including more controlled subjects of different age groups are needed to reach a meaningful conclusion about gender difference in the taste thresholds. Overall, the taste detection threshold was different from and lower than the taste recognition threshold. Therefore, the term detection or recognition should be described when the taste threshold is mentioned. The taste detection threshold as well as taste recognition threshold was affected by the gender and taster status. For additional information on the effects of aging, gender, and taster status on taste thresholds, further studies including a large number of well-controlled subjects are essential.
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Reprint requests: Hong-Seop Kho, DDS, PhD Department of Oral Medicine and Oral Diagnosis College of Dentistry and Dental Research Institute Seoul National University 28-22 Yunkeun-Dong, Chongro-Ku Seoul 110-744, Korea (ROK) hkho@plaza.snu.ac.kr