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From the Sckool of Chemistry, University of Minnesota, Minneapolis. ACID PENETRATION INTO LIVING TISSUES. BY NELSON W. TAYLOR. (From the Sckool of Chemistry, University of Minnesota, Minneapolis.) (Accepted for publication, May 23, 1927.) Downloaded from http://rupress.org/jgp/article-pdf/11/3/207/1217203/207.pdf by guest on 28 September 2021 The factors governing acid penetration into living tissues have never been dearly stated and we are therefore in ignorance as to the under- lying physicochemical basis of the sour taste and allied phenomena. In this paper the writer presents quantitative evidence which he believes, shows: first, that acid penetration through a membrane occurs either in the form of the undissociated molecule or by the simultaneous passage of H + ion and anion; secondly, that the sourest acids are those which penetrate most rapidly; and finally that the relative rates of penetration of a series of acids are determined by the well known physicochemical laws of adsorption. It was early realized that solutions of equal stoichiometrical acid concentration were not equally sour. Thus, for example, the threshold concentrations of HC1 and CI-I~COOH which just excite the sour taste are 0.001 r; and 0.003 N respectively. Becker and Herzog (1907) state that the intensity of the taste sensation occasioned by acids of equal normality decreases in the following order: hydrochloric, nitric, trichloroacetic, formic, lactic, acetic and butyric. Paul (1922) in a series of papers on the physical chemistry of foodstuffs has deter- mined the concentrations of acids which are equally sour with certain standard HC1 solutions. Taking 0.005 N HC1 as a reference he finds the order of increasing acid concentration is as follows: tartaric acid (lowest concentration therefore most sour), hydrochloric, acetyl lactic, lactic, acetic, carbonic. It is noteworthy that the two acids: I tartaric and acetyl lactic, which have practically identical dissociation constants (9.7 × 10 -4) are equally sour only when the former is 0.004 N and the latter 0.0059 N. Data such as this show that some other factor beside acid concentration plays a part in determining sourness intensity. 2O7 The Journal of General Physiology 208 ACID PENETRATION INTO LIVING TISSUES That this factor is not purely hydrogen ion concentration (or activity) must be evident from the data which I have obtained from experiments carried out in this laboratory. My experimental pro- cedure consisted at first in giving 5 cc. samples of a number of concentrations of formic acid in the vicinity of the threshold concentration to fifteen people and thus determining the minimum concentration which was definitely sour. Astringent sensations are detectible in weaker solutions. A 5 cc. sample placed in the mouth is better than a few drops, as used by some investigators, because the Downloaded from http://rupress.org/jgp/article-pdf/11/3/207/1217203/207.pdf by guest on 28 September 2021 former represents more nearly the average condition in the mouth rather than that of a single spot on the tongue. The mouth was rinsed with boiled distilled water between each test. Having determined the sour threshold concentration for formic acid, solutions of other acids were taken and that concentration of each acid chosen which was indistinguishable from the standard formic acid solution. On the average ten people passed judgement on each of these solutions. In cases where the acid had an odor the nose was held during each test. This effectively excluded the odor and restricted the judgment purely to the question of sourness. All solutions were at room temperature (22°-25 °) and had been prepared from stock acid solutions of known concentration. The hydrogen ion activities of these solutions were determined electrochemically by means of cells of the type Hg I HgC1, KC1 saturated [ acid [ H~, measurements being made at 20°C. Assuming that in these dilute solutions the activity coefficient of the H + ion is unity, we may write "concentration" instead of "activity." The data are recorded in Table I. Since the values in Column 2 of this table differ from acid to acid it is evident that sourness is not purely a function of the stoichiomet- rical acid ~oncentration. Further, since the hydrogen ion concen- trations are not the same for the several acids we may conclude that this is not the determining factor. What then is the explanation of these results? It occurred to the writer to compute the ratio of the concentrations of the undissociated part of each acid outside and inside the taste bud. The former concentration can readily be calculated from the data above, and are recorded in Column 2 of Table II. In the case of the NELSON W. TAYLOR 209 dibasic acids having anions A- and HA- the concentration of the undissociated acid "outside" (H~A) has been computed from the ex- K1 K1K~'~ pression: Total acid concentration in mols = H~ 1 -k H-g q- (H+)2] TABLE I. Threshold Concelclralions for Sour Taste. Total acid concentration H + ion concentration Downloaded from http://rupress.org/jgp/article-pdf/11/3/207/1217203/207.pdf by guest on 28 September 2021 Formic ....................... 0.0018 N -I- 3 per cent 0.00055 Acetic ....................... 0.0028N 4- 6 " " 0.00028 Butyric ..................... 0.0035N-4- 6 " " 0.00027 Valeric ...................... 0.0037N 4- 5 " " 0.00015 Oxalic ....................... 0.0020N q-8 " " 0.00116 Succinic ..................... 0.0032N q- 7 " " 0.00034 Glutaric ..................... 0.0045N'4- 5 " " 0.00034 Lactic ....................... 0.0028N 4- 5 " " 0.00177 Tartaric ..................... 0.0022N q- 7 " " 0.OOO70 TABLE II. Concentration Gradients of Equally Sour Acids (from Threshold Concentration). Concentration of Relative concentra- Acid undissociated acid tion of undissociated Ratio a/b when Outside acid formic m 1 Inside (a) (b) Formic ....................... • 00125 1 1 Acetic ........................ • 00252 11.8 0.17 Butyric ..................... .00323 14.75 0.17 Valeric ....................... • 00355 13.4 0.21 Lactic ........................ .00103 1.53 0.54 Oxalic ........................ .0000285 0.0036 6.3 Succinic ..................... .00134 3.2 .34 Glutaric ..................... .00196 4.3 .38 Tartaric ....................... .000444 0.22 1.60 where K1 and K, are the first and second dissociation constants respectively. The inside concentration can be calculated, relatively but not absolutely, from the dissociation constants of the acids if we assume that the production of a given degree of sourness is due to the establishment within the cells of the taste bud of a definite hydrogen 210 ACID PENETRATION INTO LIVING TISSUES ion concentration. Thus, for example, if two solutions of acetic acid and formic acid have the same hydrogen ion concentration; the concentrations of the undissociated parts stand together in the ratio (CHsCOOH) K~ormic 2.14 X 10 -4 (HCOOH) - Kacetic 1.8 X 10 -5 - 11.8. In the case of dibasic acids the first dissociation constant K1 was used for this computation and the effect of Ks neglected, since it is impossible with the data available to take its effect into account quantitatively. The error will, however, be fairly small. These values are listed in Column 3 of Downloaded from http://rupress.org/jgp/article-pdf/11/3/207/1217203/207.pdf by guest on 28 September 2021 TABLE III. Concentration Gradients of Equally Sour Acids, from Data by Paul and Boknen. Concentration Relative con- acid H ÷ ion of undissociated centration of Ratio a/b Acid Total undlssociated when concentration concentration acid acid formic m 1 Outside Inside (o) (b) X 10--4X X lO-~ X 10-~K Formic ............ 7 3 4.0 1.00 1 Acetic ............. 22 1.9 20.1 11.8 0.43 Propionic .......... 33 2.56 30.4 14.75 0.53 Butyric ........... 5 0.96 4.0 14.75 0.068 Monochloroacetic.. 12 8.0 4.0 0.138 7.3 Monobromoacetlc.. ll 7.2 3.8 0.155 6.1 Lactic ............. 23 5.0 18 1.55 2.9 Succinic ........... 8 2 6 3.2 0.48 Malic ............. 8 4 4 .53 1.9 Tartaric ........... 4 3 1 0.22 1.1 Carbonic .......... 53 0.4 52.6 705. 0.019 Table II under the heading '~Relative concentration of undissociated acid inside." The ratios of the numbers in Columns 2 and 3 are found in Column 4, and these ratios may be taken to represent the relative gradients or driving forces which are necessary to cause the several acids to penetrate the tissues to a comparable degree (i.e. producing equal H + ion concentration inside). Paul and Bohnen (see Paul, 1922) have obtained limiting concen- trations for various acids which taste sour. Their results are given in Table III, Columns 2 and 3. I have calculated the quantities in the other columns from their experimental data. NELSON W. TAYLOR 211 Before discussing in detail the physicochemical basis for the different behavior of these acids in regard to taste excitation, I wish to draw attention to some experiments on the rate of acid penetration. Studies on rates have been made by Haas, Jacobs, Collett, Brooks, Crozier, Osterhout, Compel and others. Crozier (1916) has measured the rate of penetration of various acids into the mantle fold of Chromodoris zebra. This tissue contains a natural indicator which changes color TABLE IV. Downloaded from http://rupress.org/jgp/article-pdf/11/3/207/1217203/207.pdf by guest on 28 September 2021 Concentration Gradients of Acids, Penetrating Chromodoris Tissue and _Producing a pH = 5.6 in 20 Minutes. Concentration Concentration Acid Total acid H + ion of undissoclate, of undissoclsted Ratio concentration concentration acid acid a/b when Outside Inside formic -- 1 {a) (b) X 10 "41 X lO-,x X 10~x X 10~x Formic ............. 33.3 7.5 25.8 2.95 1.00 Acetic.............. 188.8 5.8 183 34.7 0.6O Propionic..........
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