titration. Thus, Neumeister(ly states that the reaction of bile is

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1 ON THE REACTION OF BILE. BY SEIZABURO OKADA (Tokyo). (From the Institute of Physiology, University College, London.) WITH regard to the chemical reaction of bile from the gall-bladder there are various contradictory statements. This is probably due to the variety of methods used. Most investigators have made use of titration. Thus, Neumeister(ly states that the reaction of bile is alkaline and that it contains about 02 p.c. sodium carbonate and about the same amount of alkaline-reacting sodium phosphate. He does not state whether he refers to the bile from the gall-bladder or from the liver or to both of them. Jolles(2) observed that the bile from the gall-bladder of oxen, pigs, dogs, as well as of men, reacts always acid to phenolphthalein and that the acidity of human bile is much stronger than that of oxen. pigs and dogs. Chittenden and Albro(3) measured the amount of alkalinity of bile, using lacmoid as an indicator, and also its acidity, using phenolphthalein as an indicator. They found this so-called alkalinity and acidity in various kinds of bile except rabbit's bile, in which no measurable amount of acidity was found, Pisenti (4) found bile from a fistula to have an alkaline reaction, B rand (5) a neutral or alkaline reaction. Quagliariello(6) measured the bile from the gall-bladder of various animals (dog, rabbit, peacock, ox and sheep) with the hydrogen-electrode method and found that the H-ion concentration of it is x As to the reaction of bile from the liver I have found no exact data. Methods. I measured with the hydrogen-electrode the reaction of bile from permanent fistulae in two dogs under various conditions. Dog A was provided with a complete fistula (the same dog as that called Dog E in my paper "On the Secretion of Bile"(7)), Dog B with an incomplete fistula (the same dog as that called Dog B in my paper " On the Contractile Movement of the Gall-bladder " (8)). I also measured the bile from the gall-bladders of various animals. The following apparatus was used: (a) Luther's capillary electrometer (closed form). (b) Weston-standard cell. (c) Accumulator,

REACTION OF BILE. 115 partly discharged, as the working cell. (d) Measuring wire, consisting of a one metre long nickelin wire, graduated in millimetres and provided with a sliding contact.

2 REACTION OF BILE. 115 partly discharged, as the working cell. (d) Measuring wire, consisting of a one metre long nickelin wire, graduated in millimetres and provided with a sliding contact. (e) Gas-chains. (i) Gas electrode, similar in principle to that of Hasselbalch(9), with a capacity of 10 c.c. of which the liquid fills about 7 c.c. As electrode was used platinum wire instead of platinum plate, which was platinized in the usual way. (ii) Calomel electrode, with N/10 KCI. (iii) Intermediate liquid consisting of saturated KCI solution. In measuring the H'-ion concentration hydrogen gas was first passed through the electrode for 30 minutes. The gas was produced in a Kipp's apparatus and washed with 2 0/0 permanganate solution and saturated mercuric chloride solution. Then the bile was introduced into it by means of a funnel, avoiding every possibility of mixing with air. After closing all taps, the electrode was turned over some 200 times, and then its siphon was dipped in the saturated KCI solution. After the potential had become constant, the liquid was renewed two or three times, without changing the gas, and the measurement repeated. But all the values agreed so well with each other that the change of the liquid was afterwards omitted. This is perhaps owing to the fact that the electrode was filled with liquid as far as possible, so that the gas-space was relatively small. Probably also bile does not deliver much carbon dioxide into the gas-space. The correctness of the electrodes was often tested with standard solution. The bile was kept closed from air and was measured as fresh as possible, otherwise the value of Hi-ion concentration was found to be subject to change. 1. H%-ion concentration of the bile from the gall-bladder. I measured the values of H-ion concentration of the bile from the gall-bladder in a series of animals (dogs, oxen, cat, sheep) and obtained variable results. The hydrogen ion exponents were between 5-33 (cat) and 7.47 (ox), the Hi-ion concentration therefore being between 4-7 x 10- and 3.4 x 10-8, that is to say, the bile from the gall-bladder may be acid, neutral or alkaline. This variability also exists in different individuals of the same species, and would, no doubt, be more considerable if it were measured in more animals. It seems to be connected with the condition of the animals; in fasting the reaction is more acid than after feeding. Whether this variability has any relation to the species of animal should be determined by testing various animals under the same conditions. The dogs and cat which I used were almost without exception in a fasting condition.

3 116 S. OKADA. The details of results are given in the following table: TABLE L Bile from gal-bladder. Number Animal T. C. P.H. Number Animal T. C. a b 1 Dog Ox ,, '74 2,, 22* ,, 25* ,, ,, *32 4,, , , (after 5 hours) 7,, Cat ,, 23* ,, 210 6*97 1 Sheep 22' ,, ,, 22* ,, 22* ,, 22* ,, 25* ,, *61 5,, 22*0 6*22 6*22 14,, *39 6,, *98 P.H. 2. H--iAn concentration of bile from liver during fasting. In the two dogs the hydrogen ion exponents were between 7-41 and 8-14, that is, the H-ion concentration is between 3-9 x 1o-8 and 7-2 x The average H-ion concentration is 1-5 x 10-8 at C. TABLE 1I. Bile from liver of fasting dogs. Number Animal T. C. P.H. Number Animal T., C. PH. 1 Dog A Dog A Dog B 23' ,, 24' Dog A ,, *15 12,, * Dog B 240 7X Dog A 24* Dog B Dog B Average H--ion concentration of bile from liver during digestion. The hydrogen ion exponents of the bile from the liver after feeding with meat, biscuits and fat were between 7-54 and 8-01, that is the Hi-ion concentration is between 2-9 and 1-0 x The average H--ion concentration is 1-5 x 10-8 at C. No perceptible difference is seen with various kinds of food.

4 REACTION OF BILE. 117 TABlE III. Bile from liver during digestion. Number Animal Food T. C. P.H. Number Animal Food T. C. P.H. 1 Dog A Meat *91 13 Dog A Meat ,, (400 gm.) 22* ,, (400 gm.) '70 3,,,, ,,,, ,,,, ,,,, 23* ,,,, 21*0 7*68 17,,,, *86 6,,,, Dog B,, ,,,, Dog A,, ,,,, ,, Biscuits ,,,, ,,,, ,,,, '94 22,, Butter Dog B,, ,, Olive oil Dog A,, (100 C.C.) Average TABLE IV. Bile from liver after introduction of 200 c.c. solution into the stomach. 0-4 /0 HCI T. C. P.H. Dog A Dog B Discussion. From the above results it is evident that the reaction of the bile from the gall-bladder and from the liver is different; the reaction of the former is more variable and always inclining more to the acid side than that of the latter. To test this fact on the same animal by estimating each kind of bile separately, the cystic duct was first ligated and the gall-bladder taken out. A cannula was then inserted in the common bile-duct and after introducing 0 40/0 HCI solution into the duodenum, the outflowing bile was collected in an india-rubber bag, avoiding exposure to the air. The result is as follows: Bile from T." C. P.H. 1. Gall-bladder Liver Gall-bladder Liver The bile from the liver changed the red lacmoid paper into blue before the introduction of acid into the intestine, while the bile from the gall-bladder merely changed it into yellow. The fact that bile is, in so many animals, poured into the intestine by an orifice common to it and the pancreatic juice and that these two fluids cooperate in their actions on the ingested food-stuffs, suggest that they must have a reaction which is optimal for their action in the

118 S. OKADA. processes of digestion. Auerbach and Pick(lot found the H-ion concentration of pancreatic juice to be 0-2-5 x 10-8, that of intestinal juice 05-5 x 10-8.

5 118 S. OKADA. processes of digestion. Auerbach and Pick(lot found the H-ion concentration of pancreatic juice to be x 10-8, that of intestinal juice 05-5 x But these authors used a method which neglects the escape of carbon dioxide, so that the value should be a little higher, i.e. nearly 2 x 10-8 for both fluids. Kurt Meyer(1i) found the optimal point for trypsin to be 1.0 x 10-8, Michaelis and Davidsohn(12) 2-0 x 10-8 at 370 C., Rona and Arnheim(l3) that of erepsin 2-0 x 10-8, Davidsohn(14) that of pancreatic lipase 1.0 x The result of my research on the bile from the liver during digestion is on the average 1 5 x 10-8 at C., which may be estimated to become 2 x 10-8 at 370 C. Therefore it corresponds exactly with the reaction of pancreatic and intestinal juices and with the optimal point for trypsin, pancreatic lipase and erepsin. The bile from the gall-bladder, which has usually a higher Hi-ion concentration, might be poured out into the intestine at the beginning of digestion, so that afterwards at the height of digestion the freshly secreted bile may be mainly in action. SUMMARY. 1. The reaction of bile from the gall-bladder is variable; it may be acid, neutral or alkaline. 2. The reaction of bile from the liver is alkaline, its HP-ion concentration is 1-5 x 10-8 at 230 C., a reaction which turns red lacmoid blue but does not turn phenolphthalein red. 3. The reaction of bile from the liver during digestion corresponds to that of pancreatic and intestinal juices and with the optimal poinlt for trypsin, pancreatic lipase and erepsin. REFERENCES. (1) Neumeister. Lhrb. d. Physiol. Chem. 2te Auf. S (2) Jolles. Arch. f. Physiol. LVII. S (3) Chittenden and Albro. Amer. Journ. of Physiol. i. p (4) Pisenti. Arch. Med. Ital. xiv. p. 13. (Quoted from Brand.) (5) Brand. Arch. f Physiol. xc. S (6) Quagliariello. Rendiconti Acc. dei Lincei. xx. Ser. 5a, p (7) Okada. This Jourmal, xlmx. p (8) Ibid. This Journal, L. p (9) Hasselbalch. Biochem. Ztsch. xxx. S (10) Auerbach and Pick. Arb. a. d. k. Gesundheitsamt. Berlin, xlm. S (Quoted by H6ber in Physikal. Chem. d. Zelle u. d. Gew. 2te Auf. S ) (11) Meyer. Biochem. Ztsch. XxXII. S (12) Michaelis and Davidsohn. Biochem. Ztsch. xxxvi. S (13) Rona and Arnheim. Biochem. Ztsch. LVII. S (14) Davidsohn. Biochem. Ztsch. XLV. S

Babkin, Savitsch) that pancreatic secretion is due, in part, to reflex

Babkin, Savitsch) that pancreatic secretion is due, in part, to reflex THE MECHANISM OF PANCREATIC DIGESTION-THE FUNCTION OF SECRETIN. BY J. MELLANBY. (From the Physiological Laboratory, St Thomas's Hospital, London.) A SECRETION of pancreatic juice may be evoked by appropriate