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Gastrointestinal Physiology, 1895–1975: Motility

Horace W. Davenport

10.1002/cphy.cp060101

Source: Supplement 16: Handbook of Physiology, The Gastrointestinal System, Motility and Circulation

Originally published: 1989

Published online: January 2011

Full Article on Wiley Online Library



Abstract

The sections in this article are:

1 Swallowing
1.1 Cannon's First Roentgenological Observations
1.2 Swallowing Reflex
1.3 New Methods for Studying the Human Esophagus
1.4 Muscles and Nerves of the Esophagus
1.5 Sphincters of the Esophagus
1.6 Esophageal Peristalsis
1.7 Lower Esophageal Sphincter
2 Gastric Motility and Emptying
2.1 Peristalsis
2.2 Receptive Relaxation
2.3 Gastric Mixing and Digestion
2.4 Signs of Anxiety, Rage, or Distress
2.5 Periodic Activity
2.6 Pacemaker Potentials and Action Potentials
2.7 Pyloric Sphincter as Keeper of the Gate
2.8 Gastric Emptying
2.9 Mechanisms of Control of Gastric Emptying
3 Movements of the Small Intestine
3.1 Description
3.2 Boldyreff Revisited: Periodic Activity
3.3 Slow Waves, Spikes, and Contractions
3.4 Electrophysiology of Slow Waves and Spikes
3.5 Intestinal Gradient
3.6 Enteric Nervous System and Its Autonomic Connections
3.7 Peristaltic Reflex
4 Movements of the Colon
4.1 Ileocolic Junction and Antiperistalsis
4.2 Types of Movements of the Colon
4.3 Nervous Control of Movements of the Colon
4.4 Defecation and Continence
4.5 Aganglionosis: Megacolon
4.6 Psychosomatic Factors
Figure 1. Figure 1.

Scheme of the neural organization of swallowing.

From Sumi, T. Jpn. J. Physiol. 22: 295–314, 1972
Figure 2. Figure 2.

Summary of outflow from the swallowing center of a dog recorded by electromyography. Height of line for each muscle indicates relative intensity of activity, but contours of rise and fall are schematic.

From Doty, R. W., and J. F. Bosma. J. Neurophysiol. 19: 44–60, 1956
Figure 3. Figure 3.

Code's early apparatus. a, Syringes to inflate balloons; b, reservoirs and sidearms; c, syringes for standardizing manometers; d, bulbs for damping respiratory excursions; e, timer light; f, optical manometers; g, camera; h, transformer. [.]

From Posey, E. L., Jr., W. H. Dearing, W. G. Sauer, J. A. Bargen, and C. F. Code. Proc. Staff Meet. Mayo Clin. 23: 297–301, 1948
Figure 4. Figure 4.

Cohen and Wolf's summary of resting and swallowing pressures and bolus movement in the normal human subject. Vertical lines through curves, times selected to demonstrate bolus transport and roentgen configurations shown at the bottom. Bar below the pressure curve, period during which barium is present at each level; time in seconds is indicated below each frame. P, level of the hard palate; T, tongue; L, entrance of larynx. PRR, point of respiratory pressure reversal. H, level of top of hiatus; Fu, fundus of the stomach.

From Cohen, B. H., and B. S. Wolf. In: Handbook of Physiology. Alimentary Canal, edited by C. F. Code. Washington, DC: Am. Physiol. Soc., 1968, sect. 6, vol. IV, chapt. 91, p. 1841–1860
Figure 5. Figure 5.

Resting pressures at pharyngoesophageal junction. As the Gauer transducer at the catheter tip was withdrawn it transmitted increasingly higher pressures until, at 16.5 cm from the incisors, the upper limit of the junction was reached.

From Fyke, F. E., and C. F. Code. Gastroenterology 29: 24–34, © by Williams & Wilkins, 1955
Figure 6. Figure 6.

Mean resting pressures and major changes in pressure with deglutition. Top panels, band of high pressure between pharynx and esophagus. Bottom panels, sweep of high pressure wave through pharynx, junction, and esophagus.

From Fyke, F. E., and C. F. Code. Gastroenterology 29: 24–34, © by Williams & Wilkins, 1955
Figure 7. Figure 7.

Pressures in the normal human esophagus during swallowing.

Adapted from figure provided by C. F. Code. Reproduced from Davenport, H. W. Physiology of the Digestive Tract (5th ed.). Copyright 1982 by Year Book Medical Publishers, Inc., Chicago
Figure 8. Figure 8.

Pressures in the lower normal human esophagus during swallowing.

Adapted from Fyke, F. E., C. F. Code, and J. F. Schlegel. Gastroenterologia 86: 135–150, 1956. Reproduced from Davenport, H. W. Physiology of the Digestive Tract (5th ed.). Copyright 1982 by Year Book Medical Publishers, Inc., Chicago
Figure 9. Figure 9.

Resting pressures at the gastric fundus and gastroesophageal region and their relation to ambient pressure. At the time the figure was made Code equated the hiatus with the pressure inversion point. E‐E, end expiratory; E‐I, end inspiratory.

From Code, C. F., B. Creamer, and J. F. Schlegel. An Atlas of Esophageal Motility in Health, and Disease, © 1958. Courtesy of Charles C Thomas, Publisher, Springfield, IL
Figure 10. Figure 10.

Distal tip of recording catheter is in the high‐pressure zone. Inflation of the balloon produces a peristaltic wave and early relaxation of the lower esophageal sphincter.

From Fleshler, B., T. R. Hendrix, P. Kramer, and F. J. Ingelfinger. J. Appl. Physiol. 12: 339–342, 1958
Figure 11. Figure 11.

Openchowski's scheme of nervous control of the stomach. His legend reads in part: VSR, vagus trunks (right behind) with fibers for dilator and constrictor cardiae; ND, nerv. dilator cardiae; NC, nervi constrictores; a, Auerbach's plexus; G, bundles of ganglion cells; S S, sympathetic nerves to Auerbach's plexus.

From Openchowski, T. Zentralbl. Physiol. 1–10, 1889
Figure 12. Figure 12.

pH in the lower esophagus of a normal person recorded over 15 h. Top, demonstration that acid is rapidly cleared by 9 swallows.

From Stanciu, C., and J. R. Bennett. Gut 15: 852–857, 1974
Figure 13. Figure 13.

Cannon's names for the parts of the stomach. “At C is the cardia; F, fundus; IA, incisura angularis; B, body; PC, pyloric canal; P, pylorus.”

From Cannon, W. B. The Mechanical Factors of Digestion. New York: Longmans Green, 1911
Figure 14. Figure 14.

Cannon's tracings of the shadow cast by the bismuthfilled stomach of a cat, showing changes in shape at 1‐h intervals during digestion of a meal.

From Cannon, W. B. The Mechanical Factors of Digestion. New York: Longmans Green, 1911
Figure 15. Figure 15.

Waves passing over a small balloon in stomach of a normal man. Lower right‐hand panel record of pressure is marked with letters designating the waves illustrated.

Adapted from Smith, A. W. M., C. F. Code, and J. F. Schlegel. J. Appl. Physiol. 11: 12–16, 1957. Reproduced from Davenport, H. W. Physiology of the Digestive Tract (1st ed.). Copyright 1961 by Year Book Medical Publishers, Inc., Chicago
Figure 16. Figure 16.

X‐ray photographs of stomach of a cat. Cat was fed 20 g of meat cut into lumps and was then given milk mixed with barium sulfate. A: immediately after drinking milk. B: 10 min later. C: 20 min later.

From Gianturco, C. Am. J. Roentgenol. 31: 735–744, © by Williams & Wilkins, 1931
Figure 17. Figure 17.

Entry in Cannon's notebook for 15 July 1897.

Courtesy of R. J. Wolfe, The Francis A. Countway Library of Medicine, Boston, MA
Figure 18. Figure 18.

Gastric secretion and motility in a patient with duodenal ulcer during a period of sustained resentment, guilt, and anxiety.

From Mittelman, B., and H. G. Wolff. Psychosom. Med. 4: 5–61, 1942
Figure 19. Figure 19.

Record of Boldyreff's observations of 16 March 1902. Top line of each pair, pressure recorded from balloon in stomach of a fasting dog. Bottom line, time in minutes. Record is to be read from right to left and from top to bottom. Color is reversed from the white on black of the original.

From Boldyreff, W. N. Arch. Soc. Biol. St. Pétersbourg 11: 1–157, 1905
Figure 20. Figure 20.

Cannon's legend: “One half the original size. The top record represents intragastric pressure (the small oscillations due to respiration, the large to contractions of the stomach); the second record is time in minutes (ten minutes); the third record is W's report of hunger pangs; the lowest record is respiration registered by means of a pneumograph about the abdomen.”

From Cannon, W. B., and A. L. Washburn. Am. J. Physiol. 29: 441–454, 1912
Figure 21. Figure 21.

Carlson's legend: “One third the original size. Record showing final fourth of a period of strong contractions of the empty stomach, ending in nearly complete tetanus, 11 A.M., August 6. Light breakfast at 6 A.M. Bromoform manometer. A, stomach contractions; B respiration.”

From Carlson, A. J. Am. J. Physiol. 31: 151–173, 1912
Figure 22. Figure 22.

Shadow of a bismuth‐coated condom in a dog's stomach 30 h after a meal, with corresponding pressure tracings below. A, between contractions; B, during a hunger contraction.

From Rogers, F. T., and L. L. J. Hardt. Am. J. Physiol. 38: 274–284, 1915
Figure 23. Figure 23.

Silver‐silver chloride electrodes to be implanted on canine digestive tract used by C. F. Code and colleagues.

From Kelly, K. A., C. F. Code, and L. R. Elveback. Am. J. Physiol. 217: 461–470, 1969
Figure 24. Figure 24.

Pacesetter potentials recorded from 6 positions on a dog's stomach and from 1 position on duodenum. Small pips are dog's electrocardiogram.

From Code, C. F., J. H. Szurszewski, K. A. Kelly, and I. B. Smith. In: Handbook of Physiology. Alimentary Canal, edited by C. F. Code. Washington, DC: Am. Physiol. Soc. 1968, sect. 6, vol. V, chapt. 139, p. 2881–2896
Figure 25. Figure 25.

Electrical activity and pressure in gastric antrum of a human subject. Arrow, beginning of basic electrical rhythm, which is followed by action potentials that are accompanied by an increase in pressure.

From Monges, H., J. Salducci, and C. Roman. Arch. Fr. Mal. Appl. Dig. 58: 517–530, 1969
Figure 26. Figure 26.

Tracings of shadows of bismuth‐filled contents of stomach and intestines of a cat made 2 h after feeding boiled beef (A) or boiled rice (B).

From Cannon, W. B. The Mechanical Factors of Digestion. New York: Longmans Green, 1911
Figure 27. Figure 27.

Aggregate length of bismuth‐containing food masses in small intestine of a cat at designated intervals after feeding. Dashed line, fat; heavy line, protein; light line, carbohydrate. Each line represents mean of observations on 16 animals.

From Cannon, W. B. The Mechanical Factor of Digestion. New York: Longmans Green, 1911
Figure 28. Figure 28.

Average relation of pyloric sphincter to antrum and duodenum. PS, pyloric sphincter curve; DP, duodenal peristaltic curve; DS, duodenal segmental contractions; A, antral curve. First segmental contraction (A) reaches its peak synchronously with the sphincter (Z) at time of relaxation of antrum (P). Greatest degree of relaxation of duodenum (X) occurs after antrum has stopped contracting and during contraction of sphincter. C1, C2, C3, diagrammatic representations of movement of chyme from antrum to duodenum. Time in seconds.

From Wheelon, H., and J. E. Thomas. Am. J. Physiol. 59: 72–96, 1922
Figure 29. Figure 29.

Tracings from a cineradiographic study of motions of a dog's antrum during digestion of a meal made opaque with barium sulfate. Frames are at 1‐s intervals over a complete cycle occupying 12 s. A, antrum; TA, terminal antrum; PC, pyloric canal; DC, duodenal cap; D, duodenum; PW, peristaltic wave; P, propulsion of contents orthograde; TAC, terminal antral contraction; R, retropulsion of contents; B, bolus.

Tracings made from film by H. C. Carlson, for Ph.D. thesis, Univ. of Minnesota, 1962. Reproduced from Davenport, H. W. Physiology of the Digestive Tract (5th ed.). Copyright 1982 by Year Book Medical Publishers, Inc., Chicago
Figure 30. Figure 30.

Analysis of cycle of events shown in Fig. 29. Designation open or closed applied to pyloric sphincter refers to whether the canal could or could not be seen to contain barium‐impregnated fluid.

Adapted from data contained in Carlson, H. C., C. F. Code, and R. A. Nelson. Am. J. Dig. Dis. 11:155–172, 1966. Reproduced from Davenport, H. W. Physiology of the Digestive Tract (5th ed.). Copyright 1982 by Year Book Medical Publishers, Inc., Chicago
Figure 31. Figure 31.

Square root of volume of gastric contents of a normal man after injestion of 750 ml of liquid meals plotted against time. Data for osmotic pressure of 2 meals are given. Those meals consisted of 20 g of citrus pectin per liter and either 35 or 200 g of sucrose per liter. Complete liquid meal was a mixture of milk products, cream, and sugar, containing 1.28 cal/ml; fat 40%, protein 15%, and carbohydrate 45%.

Adapted with additional data supplied by J. N. Hunt from Hunt, J. N., and W. R. Spurrel. J. Physiol. Lond. 113: 157–168, 1951. Reproduced from Davenport, H. W. Physiology of the Digestive Tract (5th ed.). Copyright 1982 by Year Book Medical Publishers, Inc., Chicago
Figure 32. Figure 32.

Idealized representation of relationship between volumes of a 750‐ml meal recovered 20 min after instillation into the stomach of a normal man and milliosmolar concentration of non‐penetrant solutes such as glucose, freely penetrant solutes such as ammonium chloride, and specifically translocated solutes such as sodium ion, glycerol, and urea. Term penetrant refers to a hypothetical osmoreceptor with properties similar to those of the erythrocyte.

From Hunt, J. N. and M. T. Knox. In: Handbook of Physiology. Alimentary Canal, edited by C. F. Code. Washington, D.C.: Am. Physiol. Soc, 1968, sect. 6, vol. IV, chapt. 94, p. 1917–1935
Figure 33. Figure 33.

Rates of emptying of 99mTc‐labeled liver (Tc‐L) and polyethylene glycol (PEG) from stomach of a normal human subject. 99mTc‐labeled liver emptied in linear fashion (slope = −11.1%/h) after lag period of 30 min, while polyethylene glycol solution left in an exponential fashion (slope on semilog plot = −1.18%/h).

From Meyer, J. H., I. L. MacGregor, R. Gueller, P. Martin, and R. Cavalieri. Am. J. Dig. Dis. 21: 296–304, 1976
Figure 34. Figure 34.

Simultaneous measurement of pressure in antrum, pylorus, and duodenum of normal human subject. During antral contraction, pyloric pressure falls.

From Fisher, R., and S. Cohen. Gastroenterology 64: 67–75, © by Williams & Wilkins, 1973
Figure 35. Figure 35.

Cannon's diagram of segmentation.

From Cannon, W. B. The Mechanical Factors of Digestion. New York: Longmans Green, 1911
Figure 36. Figure 36.

Tracings made from one of Cannon's tissue‐paper records of a cat's stomach, intestine, and colon during digestion of starch mixed with contrast medium. Ovals, parts undergoing segmentation.

From record in possession of H. W. Davenport
Figure 37. Figure 37.

Segmentation in jejunum of a dog. Dog had been fed a meal mixed with contrast medium, and continuous cinefluorographic pictures were taken as contrast medium entered upper jejunum. Tracings were made from film at approximately 1‐s intervals. Arrows, direction in which jejunal contents were moving.

Tracings made from film by H. C. Carlson and C. F. Code. Reproduced from Davenport, H. W. Physiology of the Digestive Tract (5th ed.) Copyright 1982 by Year Book Medical Publishers, Inc., Chicago
Figure 38. Figure 38.

Migrating complexes recorded from antrum, duodenum, jejunum, and ileum of a fasting dog. Phase I, pacemaker potential moves along segment of intestine, but there are few action potentials and no contraction. Bowel contents are stagnant. Phase II, random action potentials and contractions occur in step with pacemaker potential. Bowel contents move back and forth but not downward. Phase III, sudden onset of action potentials occurring at peak of each pacemaker potential; strong progressive waves of peristaltic contraction rapidly sweep bowel contents downward. Phase III ends as abruptly as it begins and is succeeded by phase IV, in which there is decreased occurrence of action potentials and contraction. Phase IV is succeeded by another phase I. At jejunum‐1 level, phase I lasted between 83rd and 130th min; phase II lasted between 130th min and 170th min; phase III lasted between 170th and 180th min; phase IV lasted between 180th and 190th min; new phase I lasted between 190th and 240th min.

Adapted from Code, C. F., and J. A. Marlett. J. Physiol. Lond. 246: 289–309, 1975. Reproduced from Davenport, H. W. Physiology of the Digestive Tract (5th ed.). Copyright 1982 by Year Book Medical Publishers, Inc., Chicago
Figure 39. Figure 39.

A: electrical activity and pressure changes in duodenum of dog and rabbit. In the dog pressure changes occur only during spiking, but in the rabbit pressure oscillations occur in the absence of spiking. B: magnitude of pressure changes is related to intensity of spiking in both species.

From Givel, M.‐L., and Y. Ruckebusch. J. Physiol. Lond. 227: 611–625, 1972
Figure 40. Figure 40.

Top, patterns of flow of digesta associated with migrating complexes. Bottom, simultaneously recorded electrical activity.

From Bueno, L., J. Fioramonte, and Y. Ruckebusch. J. Physiol. Lond. 249: 69–85, 1975
Figure 41. Figure 41.

Biebl loop with concentric electrodes inserted.

From Bass, P., C. F. Code, and E. H. Lambert. Am. J. Physiol. 201: 287–291, 1961
Figure 42. Figure 42.

Slow wave lettering according to Bass, Code, and Lambert.

From Bass, P., C. F. Code, and E. H. Lambert. Am. J. Physiol. 201: 287–291, 1961
Figure 43. Figure 43.

Electrical activity and pressure recorded from a dog's Biebl loop, as shown in Fig. 41. Dog was given morphine by slow intravenous infusion. Left half of record, slow waves with some spiking and pressure increases. Right half, prolonged spiking and accompanying pressure increases.

From Bass, P., C. F. Code, and E. H. Lambert. Am. J. Physiol. 201: 287–291, 1961
Figure 44. Figure 44.

Electrical activity and pressure in human duodenum as recorded by a pair of tandem balloons and electrodes 3.5 cm apart. Slow waves and peristalsis are traveling at about 3 cm/s.

From Monges, H., and J. Salducci. Arch. Fr. Mal. Appl. Dig. 59: 19–28, 1970
Figure 45. Figure 45.

Transmembrane potential of single cell of longitudinal muscle of rabbit's jejunum. Each slow wave consists of a prolonged depolarization (D), which often gives rise to a few generator potentials (B, C), which in turn may give rise to spikes (A).

Adapted from Bortoff, A. Am. J. Physiol. 201: 203–208, 1961
Figure 46. Figure 46.

Left: A, theoretical configuration of propagated, transient change in membrane potential recorded intracellularly from a core conductor (ordinate, voltage); B, bipolar recording with external electrodes close together (ordinate, 1st derivative of voltage); C, monopolarity recorded in a volume conductor (ordinate, 2nd derivative of voltage). Right: A, transmembrane potentials recorded from cat's jejunum; B, bipolar recording from 2 closely spaced electrodes on same tissue; C, monopolar recording with electrode about 1 mm from the tissue.

From Bortoff, A. Am. J. Physiol. 201: 203–208, 1961
Figure 47. Figure 47.

Simultaneous records of volume‐conducted potential (A) and transmembrane potential (B) from a cat's isolated gastric antrum.

From Bortoff, A., and N. Weg. Am. J. Physiol. 208: 531–536, 1965
Figure 48. Figure 48.

Dewey and Barr's schematic diagram of nexus showing fusion of outer dark lines of membrane of 2 smooth muscle cells. ECS, extracellular space; Cyt, cytoplasm; M1 and M2, cell membranes.

From Dewey, M. M., and L. Barr. Science Wash. DC 137: 670–672, 1962. Copyright 1962 by the Am. Assoc. Adv. Sci
Figure 49. Figure 49.

Electrical activity simultaneously recorded from 4 points on greater curvature of stomach near gastroduodenal junction of an anesthetized cat. A: above junction; B: at junction; C, D: below junction. Records were made before (left) and after (right) an incision was made through musculature just orad to junction.

From Bortoff, A., and N. Weg. Am. J. Physiol. 208: 531–536, 1965
Figure 50. Figure 50.

Lack of effect of functional denervation on in vitro transmission of slow waves across gastroduodenal junction of a cat. Antral electrode A was 4 mm above junction; electrode J was on junction; electrode D was 4 mm below junction. Myenteric plexus had been inactivated by anoxia.

From Bortoff, A., and R. S. Davis. Am. J. Physiol. 215: 889–897, 1968
Figure 51. Figure 51.

Frequency of contraction of a rabbit's duodenum when stomach was very active. B, bend of duodenum.

From Alvarez, W. C. Am. J. Physiol. 37: 267–281, 1915
Figure 52. Figure 52.

Frequency of contraction of a rabbit's small intestine. A and B are from an intact animal; A is average seen in a single animal; B is made up from 723 records of “nearly” 30 animals. C was obtained from excised segments.

From Alvarez, W. C. Am. J. Physiol. 37: 267–281, 1915
Figure 53. Figure 53.

Frequency of slow waves in intestine of an anesthetized cat. Step‐wise heavy line, observed frequency; segments of records from which frequencies were obtained are given above. Lower dotted line, intrinsic frequency of isolated segments of intestine. When intestine was sectioned at point marked CUT, frequency distal to the cut fell to that of intrinsic frequency at site of the cut.

Adapted from Diamant, N. E., and A. Bortoff. Am. J. Physiol. 216: 734–743, 1969, with additional data supplied by A. Bortoff. Reproduced from Davenport, H. W. Physiology of the Digestive Tract (5th ed.). Copyright 1982 by Year Book Medical Publishers, Inc., Chicago
Figure 54. Figure 54.

Hill's “model of the supposed connexions of the nervous system of the small bowel.” Muc, mucosa; mm, muscularis mucosae; sm, submucosa; Pl sm, submucosal plexus; cm, circular muscle; Pl sym, sympathetic fibers; Pl m, myenteric plexus; lm, longitudinal muscle; ss, serosa; sspl, serosal plexus; f, fiber.

From Hill, C. Proc. R. Soc. Lond. Biol. Sci. 215: 355–387, 1927
Figure 55. Figure 55.

Schofield's diagram of connections of plexuses of the gut.

From Schofield, G. C. In: Handbook of Physiology. Alimentary Canal, edited by C. F. Code. Washington, DC: Am. Physiol. Soc., 1968, sect. 6, vol. IV, chapt. 80, p. 1579–1627
Figure 56. Figure 56.

Trendelenberg's method for studying the peristaltic reflex.

From Trendelenberg, P. Naunyn‐Schmiedeberg's Arch. Pharmacol. 81: 55–129, 1917
Figure 57. Figure 57.

T configuration of 2 transducers for measurement of longitudinal and circular muscle contraction at same level of a dog's small intestine. Circular muscle transducer is sutured with 2 deep sutures, and longitudinal transducer is sutured with 4 shallow sutures.

From Reinke, D. A., A. H. Rosenbaum, and D. R. Bennett. Am. J. Dig. Dis. 12: 113–141, 1967
Figure 58. Figure 58.

Plot of distance change between markers in transverse (AD and BC) and longitudinal (DC) directions. Upper right diagram, location of markers on bowel. Longitudinal (DC) tracing is out of phase by an average of 164° with both sets of transverse movements.

From Tasaka, K., and J. T. Farrar. Am. J. Physiol. 217: 1224–1229, 1969
Figure 59. Figure 59.

Preparation of intestinal segment with attached flap of longitudinal muscle and myenteric plexus. Individual neuron or muscle cell was observed with a compound microscope and impaled with a microelectrode.

From Hirst, G. D. S., and H. C. McKirdy. J. Physiol. Lond. 238: 129–143, 1975
Figure 60. Figure 60.

Average number of contractions in successive 10‐min periods 20 cm from anal margin in 12 patients with diarrhea and 14 patients with constipation. Mean was 2 eggs, bread, fruit, and 2 cups of tea.

Adapted from Waller, S. L., J. J. Misiewicz, and N. Kiley. Gut 13: 805–810, 1972
Figure 61. Figure 61.

Original description of a deliberately evoked mass movement read: “A normal man had an ordinary breakfast with two ounces of barium sulphate at 7 am. At 12 noon the shadow of the end of the ileum, the caecum and the ascending colon was visible (A). He then had an ordinary luncheon consisting of meat, vegetables and pudding. During the meal the caecum and ascending colon became more filled owing to the rapid emptying of the end of the ileum, and toward the end of the meal a large, round mass at the hepatic flexure became cut off from the rest of the ascending colon (B). Immediately after the mean was finished some of this was seen to move slowly round the hepatic flexure (C); the diameter of the separated portion then became suddenly much smaller, and the large round shadow being replaced by a long narrow one which extended from the hepatic flexure almost to the splenic flexure (D). The shadow was at first uniform, but in a few seconds haustral segmentation developed (E). About 5 minutes later the shadow suddenly became still more prolonged and passed round the splenic flexure (F), down the descending colon (G) to the beginning of the pelvic colon (H).”

From Hertz, A. F., and A. Newton. J. Physiol. Lond. 47: 57–65, 1913
Figure 62. Figure 62.

Multihaustral propulsion. At zero time, ileal contents were entering the cecum (a). Three minutes later cecal contents distended the hepatic flexure (b). At 5 min contraction distributed cecal contents over the whole transverse colon with some narrowing (c). At 7 min as haustration was returning, most of the colon between b and c contracted. Then all contents of this segment were expelled into the distal half of the transverse colon. Conical outline at c in the 9‐min frame is typical of multihaustral contraction.

Adapted from Ritchie, J. A. Gut 9: 442–456, 1968. Reproduced from Davenport, H. W. The Physiology of the Digestive Tract (5th ed.). Copyright 1982 by Year Book Medical Publishers, Inc., Chicago
Figure 63. Figure 63.

Basis of peripheral autonomic reflex in the colon. ACh, acetylcholine; NA, noradrenaline (norepinephrine); CNS, central nervous system.

From Crowfoot, P. J., M. Holman, and J. H. Szurzsewski. J. Physiol. Lond. 219: 443–461,1971
Figure 64. Figure 64.

Schematic representation of innervation of distal colon and anal canal in the cat.

From Bishop, B., R. C. Garry, T. D. M. Roberts, and J. K. Todd. J. Physiol. Lond. 134: 229–240, 1956
Figure 65. Figure 65.

Comparison of pressure in anal sphincters in normal human subjects at rest and during distension.

From Duthie, H. L., and R. C. Bennett. Gut 4: 179–182, 1963
Figure 66. Figure 66.

Electromyogram of external anal sphincter of a man lying on his left side, showing sensitivity to very light finger touch on anal margin.

From Floyd, W. F., and E. W. Walls. J. Physiol. Lond. 122: 599–609,1953
Figure 67. Figure 67.

Diagram of recording technique used by Schuster and colleagues.

From Schuster, M. M., P. Hookman, T. R. Hendrix, and A. F. Mendeloff. Bull. Johns Hopkins Hosp. 116: 79–88, 1965
Figure 68. Figure 68.

Responses of rectum and internal and external anal sphincters to transient and repeated distension.

From Schuster, M. M., P. Hookman, T. R. Hendrix, and A. F. Mendeloff. Bull. Johns Hopkins Hosp. 116: 79–88, 1965
Figure 69. Figure 69.

Responses of rectum and internal and external anal sphincters to successive distensions of rectum.

Adapted from Denny‐Brown, D., and E. G. Robertson. Brain 58: 256–310, 1935, and Schuster, M. M., P. Hookman, T. R. Hendrix, and A. F. Mendeloff. Bull. Johns Hopkins Hosp. 116: 79–88,1965. Reproduced from Davenport, H. W. Physiology of the Digestive Tract (5th ed.). Copyright 1982 by Year Book Medical Publishers, Inc., Chicago
Figure 70. Figure 70.

A. F. Hertz's sketch of the shadow of barium in the colon of a normal man, immediately before, immediately after, and 1 h 45 min after defecation.

From Hertz, A. F. Guy's Hosp. Rep. 61: 389–427,1904
Figure 71. Figure 71.

Left: heightened motility in sigmoid colon of a 62‐yr‐old domestic servant, which coincided with discussion of life stresses. Shaded area, range of intraluminal pressures during that part of the tracing not reproduced. Right: reduction of motility in sigmoid colon of a 19‐yr‐old unemployed male with diarrhea during unsympathetic discussion of life problems.

From Almy, T. P. Am. J. Med. 10: 60–67,1951


Figure 1.

Scheme of the neural organization of swallowing.

From Sumi, T. Jpn. J. Physiol. 22: 295–314, 1972


Figure 2.

Summary of outflow from the swallowing center of a dog recorded by electromyography. Height of line for each muscle indicates relative intensity of activity, but contours of rise and fall are schematic.

From Doty, R. W., and J. F. Bosma. J. Neurophysiol. 19: 44–60, 1956


Figure 3.

Code's early apparatus. a, Syringes to inflate balloons; b, reservoirs and sidearms; c, syringes for standardizing manometers; d, bulbs for damping respiratory excursions; e, timer light; f, optical manometers; g, camera; h, transformer. [.]

From Posey, E. L., Jr., W. H. Dearing, W. G. Sauer, J. A. Bargen, and C. F. Code. Proc. Staff Meet. Mayo Clin. 23: 297–301, 1948


Figure 4.

Cohen and Wolf's summary of resting and swallowing pressures and bolus movement in the normal human subject. Vertical lines through curves, times selected to demonstrate bolus transport and roentgen configurations shown at the bottom. Bar below the pressure curve, period during which barium is present at each level; time in seconds is indicated below each frame. P, level of the hard palate; T, tongue; L, entrance of larynx. PRR, point of respiratory pressure reversal. H, level of top of hiatus; Fu, fundus of the stomach.

From Cohen, B. H., and B. S. Wolf. In: Handbook of Physiology. Alimentary Canal, edited by C. F. Code. Washington, DC: Am. Physiol. Soc., 1968, sect. 6, vol. IV, chapt. 91, p. 1841–1860


Figure 5.

Resting pressures at pharyngoesophageal junction. As the Gauer transducer at the catheter tip was withdrawn it transmitted increasingly higher pressures until, at 16.5 cm from the incisors, the upper limit of the junction was reached.

From Fyke, F. E., and C. F. Code. Gastroenterology 29: 24–34, © by Williams & Wilkins, 1955


Figure 6.

Mean resting pressures and major changes in pressure with deglutition. Top panels, band of high pressure between pharynx and esophagus. Bottom panels, sweep of high pressure wave through pharynx, junction, and esophagus.

From Fyke, F. E., and C. F. Code. Gastroenterology 29: 24–34, © by Williams & Wilkins, 1955


Figure 7.

Pressures in the normal human esophagus during swallowing.

Adapted from figure provided by C. F. Code. Reproduced from Davenport, H. W. Physiology of the Digestive Tract (5th ed.). Copyright 1982 by Year Book Medical Publishers, Inc., Chicago


Figure 8.

Pressures in the lower normal human esophagus during swallowing.

Adapted from Fyke, F. E., C. F. Code, and J. F. Schlegel. Gastroenterologia 86: 135–150, 1956. Reproduced from Davenport, H. W. Physiology of the Digestive Tract (5th ed.). Copyright 1982 by Year Book Medical Publishers, Inc., Chicago


Figure 9.

Resting pressures at the gastric fundus and gastroesophageal region and their relation to ambient pressure. At the time the figure was made Code equated the hiatus with the pressure inversion point. E‐E, end expiratory; E‐I, end inspiratory.

From Code, C. F., B. Creamer, and J. F. Schlegel. An Atlas of Esophageal Motility in Health, and Disease, © 1958. Courtesy of Charles C Thomas, Publisher, Springfield, IL


Figure 10.

Distal tip of recording catheter is in the high‐pressure zone. Inflation of the balloon produces a peristaltic wave and early relaxation of the lower esophageal sphincter.

From Fleshler, B., T. R. Hendrix, P. Kramer, and F. J. Ingelfinger. J. Appl. Physiol. 12: 339–342, 1958


Figure 11.

Openchowski's scheme of nervous control of the stomach. His legend reads in part: VSR, vagus trunks (right behind) with fibers for dilator and constrictor cardiae; ND, nerv. dilator cardiae; NC, nervi constrictores; a, Auerbach's plexus; G, bundles of ganglion cells; S S, sympathetic nerves to Auerbach's plexus.

From Openchowski, T. Zentralbl. Physiol. 1–10, 1889


Figure 12.

pH in the lower esophagus of a normal person recorded over 15 h. Top, demonstration that acid is rapidly cleared by 9 swallows.

From Stanciu, C., and J. R. Bennett. Gut 15: 852–857, 1974


Figure 13.

Cannon's names for the parts of the stomach. “At C is the cardia; F, fundus; IA, incisura angularis; B, body; PC, pyloric canal; P, pylorus.”

From Cannon, W. B. The Mechanical Factors of Digestion. New York: Longmans Green, 1911


Figure 14.

Cannon's tracings of the shadow cast by the bismuthfilled stomach of a cat, showing changes in shape at 1‐h intervals during digestion of a meal.

From Cannon, W. B. The Mechanical Factors of Digestion. New York: Longmans Green, 1911


Figure 15.

Waves passing over a small balloon in stomach of a normal man. Lower right‐hand panel record of pressure is marked with letters designating the waves illustrated.

Adapted from Smith, A. W. M., C. F. Code, and J. F. Schlegel. J. Appl. Physiol. 11: 12–16, 1957. Reproduced from Davenport, H. W. Physiology of the Digestive Tract (1st ed.). Copyright 1961 by Year Book Medical Publishers, Inc., Chicago


Figure 16.

X‐ray photographs of stomach of a cat. Cat was fed 20 g of meat cut into lumps and was then given milk mixed with barium sulfate. A: immediately after drinking milk. B: 10 min later. C: 20 min later.

From Gianturco, C. Am. J. Roentgenol. 31: 735–744, © by Williams & Wilkins, 1931


Figure 17.

Entry in Cannon's notebook for 15 July 1897.

Courtesy of R. J. Wolfe, The Francis A. Countway Library of Medicine, Boston, MA


Figure 18.

Gastric secretion and motility in a patient with duodenal ulcer during a period of sustained resentment, guilt, and anxiety.

From Mittelman, B., and H. G. Wolff. Psychosom. Med. 4: 5–61, 1942


Figure 19.

Record of Boldyreff's observations of 16 March 1902. Top line of each pair, pressure recorded from balloon in stomach of a fasting dog. Bottom line, time in minutes. Record is to be read from right to left and from top to bottom. Color is reversed from the white on black of the original.

From Boldyreff, W. N. Arch. Soc. Biol. St. Pétersbourg 11: 1–157, 1905


Figure 20.

Cannon's legend: “One half the original size. The top record represents intragastric pressure (the small oscillations due to respiration, the large to contractions of the stomach); the second record is time in minutes (ten minutes); the third record is W's report of hunger pangs; the lowest record is respiration registered by means of a pneumograph about the abdomen.”

From Cannon, W. B., and A. L. Washburn. Am. J. Physiol. 29: 441–454, 1912


Figure 21.

Carlson's legend: “One third the original size. Record showing final fourth of a period of strong contractions of the empty stomach, ending in nearly complete tetanus, 11 A.M., August 6. Light breakfast at 6 A.M. Bromoform manometer. A, stomach contractions; B respiration.”

From Carlson, A. J. Am. J. Physiol. 31: 151–173, 1912


Figure 22.

Shadow of a bismuth‐coated condom in a dog's stomach 30 h after a meal, with corresponding pressure tracings below. A, between contractions; B, during a hunger contraction.

From Rogers, F. T., and L. L. J. Hardt. Am. J. Physiol. 38: 274–284, 1915


Figure 23.

Silver‐silver chloride electrodes to be implanted on canine digestive tract used by C. F. Code and colleagues.

From Kelly, K. A., C. F. Code, and L. R. Elveback. Am. J. Physiol. 217: 461–470, 1969


Figure 24.

Pacesetter potentials recorded from 6 positions on a dog's stomach and from 1 position on duodenum. Small pips are dog's electrocardiogram.

From Code, C. F., J. H. Szurszewski, K. A. Kelly, and I. B. Smith. In: Handbook of Physiology. Alimentary Canal, edited by C. F. Code. Washington, DC: Am. Physiol. Soc. 1968, sect. 6, vol. V, chapt. 139, p. 2881–2896


Figure 25.

Electrical activity and pressure in gastric antrum of a human subject. Arrow, beginning of basic electrical rhythm, which is followed by action potentials that are accompanied by an increase in pressure.

From Monges, H., J. Salducci, and C. Roman. Arch. Fr. Mal. Appl. Dig. 58: 517–530, 1969


Figure 26.

Tracings of shadows of bismuth‐filled contents of stomach and intestines of a cat made 2 h after feeding boiled beef (A) or boiled rice (B).

From Cannon, W. B. The Mechanical Factors of Digestion. New York: Longmans Green, 1911


Figure 27.

Aggregate length of bismuth‐containing food masses in small intestine of a cat at designated intervals after feeding. Dashed line, fat; heavy line, protein; light line, carbohydrate. Each line represents mean of observations on 16 animals.

From Cannon, W. B. The Mechanical Factor of Digestion. New York: Longmans Green, 1911


Figure 28.

Average relation of pyloric sphincter to antrum and duodenum. PS, pyloric sphincter curve; DP, duodenal peristaltic curve; DS, duodenal segmental contractions; A, antral curve. First segmental contraction (A) reaches its peak synchronously with the sphincter (Z) at time of relaxation of antrum (P). Greatest degree of relaxation of duodenum (X) occurs after antrum has stopped contracting and during contraction of sphincter. C1, C2, C3, diagrammatic representations of movement of chyme from antrum to duodenum. Time in seconds.

From Wheelon, H., and J. E. Thomas. Am. J. Physiol. 59: 72–96, 1922


Figure 29.

Tracings from a cineradiographic study of motions of a dog's antrum during digestion of a meal made opaque with barium sulfate. Frames are at 1‐s intervals over a complete cycle occupying 12 s. A, antrum; TA, terminal antrum; PC, pyloric canal; DC, duodenal cap; D, duodenum; PW, peristaltic wave; P, propulsion of contents orthograde; TAC, terminal antral contraction; R, retropulsion of contents; B, bolus.

Tracings made from film by H. C. Carlson, for Ph.D. thesis, Univ. of Minnesota, 1962. Reproduced from Davenport, H. W. Physiology of the Digestive Tract (5th ed.). Copyright 1982 by Year Book Medical Publishers, Inc., Chicago


Figure 30.

Analysis of cycle of events shown in Fig. 29. Designation open or closed applied to pyloric sphincter refers to whether the canal could or could not be seen to contain barium‐impregnated fluid.

Adapted from data contained in Carlson, H. C., C. F. Code, and R. A. Nelson. Am. J. Dig. Dis. 11:155–172, 1966. Reproduced from Davenport, H. W. Physiology of the Digestive Tract (5th ed.). Copyright 1982 by Year Book Medical Publishers, Inc., Chicago


Figure 31.

Square root of volume of gastric contents of a normal man after injestion of 750 ml of liquid meals plotted against time. Data for osmotic pressure of 2 meals are given. Those meals consisted of 20 g of citrus pectin per liter and either 35 or 200 g of sucrose per liter. Complete liquid meal was a mixture of milk products, cream, and sugar, containing 1.28 cal/ml; fat 40%, protein 15%, and carbohydrate 45%.

Adapted with additional data supplied by J. N. Hunt from Hunt, J. N., and W. R. Spurrel. J. Physiol. Lond. 113: 157–168, 1951. Reproduced from Davenport, H. W. Physiology of the Digestive Tract (5th ed.). Copyright 1982 by Year Book Medical Publishers, Inc., Chicago


Figure 32.

Idealized representation of relationship between volumes of a 750‐ml meal recovered 20 min after instillation into the stomach of a normal man and milliosmolar concentration of non‐penetrant solutes such as glucose, freely penetrant solutes such as ammonium chloride, and specifically translocated solutes such as sodium ion, glycerol, and urea. Term penetrant refers to a hypothetical osmoreceptor with properties similar to those of the erythrocyte.

From Hunt, J. N. and M. T. Knox. In: Handbook of Physiology. Alimentary Canal, edited by C. F. Code. Washington, D.C.: Am. Physiol. Soc, 1968, sect. 6, vol. IV, chapt. 94, p. 1917–1935


Figure 33.

Rates of emptying of 99mTc‐labeled liver (Tc‐L) and polyethylene glycol (PEG) from stomach of a normal human subject. 99mTc‐labeled liver emptied in linear fashion (slope = −11.1%/h) after lag period of 30 min, while polyethylene glycol solution left in an exponential fashion (slope on semilog plot = −1.18%/h).

From Meyer, J. H., I. L. MacGregor, R. Gueller, P. Martin, and R. Cavalieri. Am. J. Dig. Dis. 21: 296–304, 1976


Figure 34.

Simultaneous measurement of pressure in antrum, pylorus, and duodenum of normal human subject. During antral contraction, pyloric pressure falls.

From Fisher, R., and S. Cohen. Gastroenterology 64: 67–75, © by Williams & Wilkins, 1973


Figure 35.

Cannon's diagram of segmentation.

From Cannon, W. B. The Mechanical Factors of Digestion. New York: Longmans Green, 1911


Figure 36.

Tracings made from one of Cannon's tissue‐paper records of a cat's stomach, intestine, and colon during digestion of starch mixed with contrast medium. Ovals, parts undergoing segmentation.

From record in possession of H. W. Davenport


Figure 37.

Segmentation in jejunum of a dog. Dog had been fed a meal mixed with contrast medium, and continuous cinefluorographic pictures were taken as contrast medium entered upper jejunum. Tracings were made from film at approximately 1‐s intervals. Arrows, direction in which jejunal contents were moving.

Tracings made from film by H. C. Carlson and C. F. Code. Reproduced from Davenport, H. W. Physiology of the Digestive Tract (5th ed.) Copyright 1982 by Year Book Medical Publishers, Inc., Chicago


Figure 38.

Migrating complexes recorded from antrum, duodenum, jejunum, and ileum of a fasting dog. Phase I, pacemaker potential moves along segment of intestine, but there are few action potentials and no contraction. Bowel contents are stagnant. Phase II, random action potentials and contractions occur in step with pacemaker potential. Bowel contents move back and forth but not downward. Phase III, sudden onset of action potentials occurring at peak of each pacemaker potential; strong progressive waves of peristaltic contraction rapidly sweep bowel contents downward. Phase III ends as abruptly as it begins and is succeeded by phase IV, in which there is decreased occurrence of action potentials and contraction. Phase IV is succeeded by another phase I. At jejunum‐1 level, phase I lasted between 83rd and 130th min; phase II lasted between 130th min and 170th min; phase III lasted between 170th and 180th min; phase IV lasted between 180th and 190th min; new phase I lasted between 190th and 240th min.

Adapted from Code, C. F., and J. A. Marlett. J. Physiol. Lond. 246: 289–309, 1975. Reproduced from Davenport, H. W. Physiology of the Digestive Tract (5th ed.). Copyright 1982 by Year Book Medical Publishers, Inc., Chicago


Figure 39.

A: electrical activity and pressure changes in duodenum of dog and rabbit. In the dog pressure changes occur only during spiking, but in the rabbit pressure oscillations occur in the absence of spiking. B: magnitude of pressure changes is related to intensity of spiking in both species.

From Givel, M.‐L., and Y. Ruckebusch. J. Physiol. Lond. 227: 611–625, 1972


Figure 40.

Top, patterns of flow of digesta associated with migrating complexes. Bottom, simultaneously recorded electrical activity.

From Bueno, L., J. Fioramonte, and Y. Ruckebusch. J. Physiol. Lond. 249: 69–85, 1975


Figure 41.

Biebl loop with concentric electrodes inserted.

From Bass, P., C. F. Code, and E. H. Lambert. Am. J. Physiol. 201: 287–291, 1961


Figure 42.

Slow wave lettering according to Bass, Code, and Lambert.

From Bass, P., C. F. Code, and E. H. Lambert. Am. J. Physiol. 201: 287–291, 1961


Figure 43.

Electrical activity and pressure recorded from a dog's Biebl loop, as shown in Fig. 41. Dog was given morphine by slow intravenous infusion. Left half of record, slow waves with some spiking and pressure increases. Right half, prolonged spiking and accompanying pressure increases.

From Bass, P., C. F. Code, and E. H. Lambert. Am. J. Physiol. 201: 287–291, 1961


Figure 44.

Electrical activity and pressure in human duodenum as recorded by a pair of tandem balloons and electrodes 3.5 cm apart. Slow waves and peristalsis are traveling at about 3 cm/s.

From Monges, H., and J. Salducci. Arch. Fr. Mal. Appl. Dig. 59: 19–28, 1970


Figure 45.

Transmembrane potential of single cell of longitudinal muscle of rabbit's jejunum. Each slow wave consists of a prolonged depolarization (D), which often gives rise to a few generator potentials (B, C), which in turn may give rise to spikes (A).

Adapted from Bortoff, A. Am. J. Physiol. 201: 203–208, 1961


Figure 46.

Left: A, theoretical configuration of propagated, transient change in membrane potential recorded intracellularly from a core conductor (ordinate, voltage); B, bipolar recording with external electrodes close together (ordinate, 1st derivative of voltage); C, monopolarity recorded in a volume conductor (ordinate, 2nd derivative of voltage). Right: A, transmembrane potentials recorded from cat's jejunum; B, bipolar recording from 2 closely spaced electrodes on same tissue; C, monopolar recording with electrode about 1 mm from the tissue.

From Bortoff, A. Am. J. Physiol. 201: 203–208, 1961


Figure 47.

Simultaneous records of volume‐conducted potential (A) and transmembrane potential (B) from a cat's isolated gastric antrum.

From Bortoff, A., and N. Weg. Am. J. Physiol. 208: 531–536, 1965


Figure 48.

Dewey and Barr's schematic diagram of nexus showing fusion of outer dark lines of membrane of 2 smooth muscle cells. ECS, extracellular space; Cyt, cytoplasm; M1 and M2, cell membranes.

From Dewey, M. M., and L. Barr. Science Wash. DC 137: 670–672, 1962. Copyright 1962 by the Am. Assoc. Adv. Sci


Figure 49.

Electrical activity simultaneously recorded from 4 points on greater curvature of stomach near gastroduodenal junction of an anesthetized cat. A: above junction; B: at junction; C, D: below junction. Records were made before (left) and after (right) an incision was made through musculature just orad to junction.

From Bortoff, A., and N. Weg. Am. J. Physiol. 208: 531–536, 1965


Figure 50.

Lack of effect of functional denervation on in vitro transmission of slow waves across gastroduodenal junction of a cat. Antral electrode A was 4 mm above junction; electrode J was on junction; electrode D was 4 mm below junction. Myenteric plexus had been inactivated by anoxia.

From Bortoff, A., and R. S. Davis. Am. J. Physiol. 215: 889–897, 1968


Figure 51.

Frequency of contraction of a rabbit's duodenum when stomach was very active. B, bend of duodenum.

From Alvarez, W. C. Am. J. Physiol. 37: 267–281, 1915


Figure 52.

Frequency of contraction of a rabbit's small intestine. A and B are from an intact animal; A is average seen in a single animal; B is made up from 723 records of “nearly” 30 animals. C was obtained from excised segments.

From Alvarez, W. C. Am. J. Physiol. 37: 267–281, 1915


Figure 53.

Frequency of slow waves in intestine of an anesthetized cat. Step‐wise heavy line, observed frequency; segments of records from which frequencies were obtained are given above. Lower dotted line, intrinsic frequency of isolated segments of intestine. When intestine was sectioned at point marked CUT, frequency distal to the cut fell to that of intrinsic frequency at site of the cut.

Adapted from Diamant, N. E., and A. Bortoff. Am. J. Physiol. 216: 734–743, 1969, with additional data supplied by A. Bortoff. Reproduced from Davenport, H. W. Physiology of the Digestive Tract (5th ed.). Copyright 1982 by Year Book Medical Publishers, Inc., Chicago


Figure 54.

Hill's “model of the supposed connexions of the nervous system of the small bowel.” Muc, mucosa; mm, muscularis mucosae; sm, submucosa; Pl sm, submucosal plexus; cm, circular muscle; Pl sym, sympathetic fibers; Pl m, myenteric plexus; lm, longitudinal muscle; ss, serosa; sspl, serosal plexus; f, fiber.

From Hill, C. Proc. R. Soc. Lond. Biol. Sci. 215: 355–387, 1927


Figure 55.

Schofield's diagram of connections of plexuses of the gut.

From Schofield, G. C. In: Handbook of Physiology. Alimentary Canal, edited by C. F. Code. Washington, DC: Am. Physiol. Soc., 1968, sect. 6, vol. IV, chapt. 80, p. 1579–1627


Figure 56.

Trendelenberg's method for studying the peristaltic reflex.

From Trendelenberg, P. Naunyn‐Schmiedeberg's Arch. Pharmacol. 81: 55–129, 1917


Figure 57.

T configuration of 2 transducers for measurement of longitudinal and circular muscle contraction at same level of a dog's small intestine. Circular muscle transducer is sutured with 2 deep sutures, and longitudinal transducer is sutured with 4 shallow sutures.

From Reinke, D. A., A. H. Rosenbaum, and D. R. Bennett. Am. J. Dig. Dis. 12: 113–141, 1967


Figure 58.

Plot of distance change between markers in transverse (AD and BC) and longitudinal (DC) directions. Upper right diagram, location of markers on bowel. Longitudinal (DC) tracing is out of phase by an average of 164° with both sets of transverse movements.

From Tasaka, K., and J. T. Farrar. Am. J. Physiol. 217: 1224–1229, 1969


Figure 59.

Preparation of intestinal segment with attached flap of longitudinal muscle and myenteric plexus. Individual neuron or muscle cell was observed with a compound microscope and impaled with a microelectrode.

From Hirst, G. D. S., and H. C. McKirdy. J. Physiol. Lond. 238: 129–143, 1975


Figure 60.

Average number of contractions in successive 10‐min periods 20 cm from anal margin in 12 patients with diarrhea and 14 patients with constipation. Mean was 2 eggs, bread, fruit, and 2 cups of tea.

Adapted from Waller, S. L., J. J. Misiewicz, and N. Kiley. Gut 13: 805–810, 1972


Figure 61.

Original description of a deliberately evoked mass movement read: “A normal man had an ordinary breakfast with two ounces of barium sulphate at 7 am. At 12 noon the shadow of the end of the ileum, the caecum and the ascending colon was visible (A). He then had an ordinary luncheon consisting of meat, vegetables and pudding. During the meal the caecum and ascending colon became more filled owing to the rapid emptying of the end of the ileum, and toward the end of the meal a large, round mass at the hepatic flexure became cut off from the rest of the ascending colon (B). Immediately after the mean was finished some of this was seen to move slowly round the hepatic flexure (C); the diameter of the separated portion then became suddenly much smaller, and the large round shadow being replaced by a long narrow one which extended from the hepatic flexure almost to the splenic flexure (D). The shadow was at first uniform, but in a few seconds haustral segmentation developed (E). About 5 minutes later the shadow suddenly became still more prolonged and passed round the splenic flexure (F), down the descending colon (G) to the beginning of the pelvic colon (H).”

From Hertz, A. F., and A. Newton. J. Physiol. Lond. 47: 57–65, 1913


Figure 62.

Multihaustral propulsion. At zero time, ileal contents were entering the cecum (a). Three minutes later cecal contents distended the hepatic flexure (b). At 5 min contraction distributed cecal contents over the whole transverse colon with some narrowing (c). At 7 min as haustration was returning, most of the colon between b and c contracted. Then all contents of this segment were expelled into the distal half of the transverse colon. Conical outline at c in the 9‐min frame is typical of multihaustral contraction.

Adapted from Ritchie, J. A. Gut 9: 442–456, 1968. Reproduced from Davenport, H. W. The Physiology of the Digestive Tract (5th ed.). Copyright 1982 by Year Book Medical Publishers, Inc., Chicago


Figure 63.

Basis of peripheral autonomic reflex in the colon. ACh, acetylcholine; NA, noradrenaline (norepinephrine); CNS, central nervous system.

From Crowfoot, P. J., M. Holman, and J. H. Szurzsewski. J. Physiol. Lond. 219: 443–461,1971


Figure 64.

Schematic representation of innervation of distal colon and anal canal in the cat.

From Bishop, B., R. C. Garry, T. D. M. Roberts, and J. K. Todd. J. Physiol. Lond. 134: 229–240, 1956


Figure 65.

Comparison of pressure in anal sphincters in normal human subjects at rest and during distension.

From Duthie, H. L., and R. C. Bennett. Gut 4: 179–182, 1963


Figure 66.

Electromyogram of external anal sphincter of a man lying on his left side, showing sensitivity to very light finger touch on anal margin.

From Floyd, W. F., and E. W. Walls. J. Physiol. Lond. 122: 599–609,1953


Figure 67.

Diagram of recording technique used by Schuster and colleagues.

From Schuster, M. M., P. Hookman, T. R. Hendrix, and A. F. Mendeloff. Bull. Johns Hopkins Hosp. 116: 79–88, 1965


Figure 68.

Responses of rectum and internal and external anal sphincters to transient and repeated distension.

From Schuster, M. M., P. Hookman, T. R. Hendrix, and A. F. Mendeloff. Bull. Johns Hopkins Hosp. 116: 79–88, 1965


Figure 69.

Responses of rectum and internal and external anal sphincters to successive distensions of rectum.

Adapted from Denny‐Brown, D., and E. G. Robertson. Brain 58: 256–310, 1935, and Schuster, M. M., P. Hookman, T. R. Hendrix, and A. F. Mendeloff. Bull. Johns Hopkins Hosp. 116: 79–88,1965. Reproduced from Davenport, H. W. Physiology of the Digestive Tract (5th ed.). Copyright 1982 by Year Book Medical Publishers, Inc., Chicago


Figure 70.

A. F. Hertz's sketch of the shadow of barium in the colon of a normal man, immediately before, immediately after, and 1 h 45 min after defecation.

From Hertz, A. F. Guy's Hosp. Rep. 61: 389–427,1904


Figure 71.

Left: heightened motility in sigmoid colon of a 62‐yr‐old domestic servant, which coincided with discussion of life stresses. Shaded area, range of intraluminal pressures during that part of the tracing not reproduced. Right: reduction of motility in sigmoid colon of a 19‐yr‐old unemployed male with diarrhea during unsympathetic discussion of life problems.

From Almy, T. P. Am. J. Med. 10: 60–67,1951
References
 1. The beginning of Cannon's research with X‐rays is described in Davenport, H. W. An Eagle‐Feather: The Short Life of Albert Moser, M.D., A Footnote to the Life of Walter B. Cannon. Boston, MA: Francis A. Countway Libr. Med., 1974;
 2. and in Benison, S., A. C. Barger, and E. L. Wolfe. Walter B. Cannon. Cambridge, MA: Harvard Univ. Press, 1987, p. 50–73.
 3. Kronecker, H., and S. Meltzer. Der Schluckmechanismus, seine Erregnung und seine Hemmung. Arch. Anat. Physiol. Physiol. Abt. Suppl. Bd. 328–360, 1883. Because the work was done in Berlin in 1882 it is probably correct to assume that their “kohlensäuerreiche Getränke” were beer rather than Coca‐Cola. The “ja” is a direct quote from the paper.
 4. Letter from W. B. C. to John F. Fulton, 16 April 1942.
 5. The letter is partially quoted in Barger, A. C. New technology for a new century: Walter B. Cannon and the invisible rays. Am. J. Roentgenol. 136: 87–95, 1981. Clifford Barger gave me a photocopy of the letter, for which much thanks.
 6. I have used a typescript copy of the notebook, but I have seen the crucial passages in the original, thanks to Richard J. Wolfe.
 7. The English version is Röntgen, W. K. On a new kind of ray. Science Wash. DC 3: 227–231, 1886.
 8. Brunton, T. L. A Text‐book of Pharmacology, Therapeutics and Materia Medica, Adapted to the United States Pharmacopoeia (3rd ed.), edited by F. H. Williams. Philadelphia, PA: Lea Brothers, 1889, p. 731–734.
 9. Cannon, W. B. The passage of different food‐stuffs from the stomach and through the small intestine. Am. J. Physiol. 12: 387–418, 1904.
 10. Cannon did say so in Cannon, W. B. The Mechanical Factors of Digestion. New York: Longmans, Green, 1911, p. 5.
 11. Bachem, D., and H. Günther. Bariumsulfat als schattenbildenes Kontrastmittel bei Röntgenuntersuchungen. Z. Röntgenk. 12: 369–376, 1910.
 12. The warning is Bachem, C. Baryumsulfat als Diagnosticum in der Röntgenkunde. Ber. Klin. Wochenschr. 49: 1425–1426, 1912.
 13. Rieder, H. Beiträge zur Topographie des Magens‐Darmkanals beim lebenden Menschen nebst Untersuchungen über den zeitlichen Ablauf der Verdauung. Fortschr. Geb. Roentgenstr. Nuklearmed. 8: 141–172, 1905.
 14. Albert Moser's collaboration with Cannon is described in Davenport (n.1).
 15. Cannon, W. B., and A. Moser. The movements of the food in the oesophagus. Am. J. Physiol. 1: 435–444, 1898; and Cannon (n. 8).
 16. Cannon n. 8, p. 19; and his notebook entry 6 Feb. 1896.
 17. Doty, R. W. Influence of stimulus pattern on reflex deglutition. Am. J. Physiol. 166: 142–158, 1951.
 18. For a thorough summary of the neural organization of swallowing up to about 1965, see Doty, R. W. Neural organization of deglutition. In: Handbook of Physiology. Alimentary Canal, edited by C. F. Code. Washington, DC: Am. Physiol. Soc, 1968, sect. 6, vol. IV, chapt. 92, p. 1861–1902.
 19. Other summaries are Bosma, J. F. Deglutition: pharyngeal stage. Physiol. Rev. 37: 275–300, 1957;
 20. Ingelfinger, F. J. Esophageal motility. Physiol. Rev. 38: 533–584, 1968;
 21. Wood, J. D. Neurophysiology of Auerbach's plexus and control of intestinal motility. Physiol. Rev. 55: 307–324, 1975;
 22. Brooks, F. P., and P. W. Evers editors. Nerves of the Gut. Thorofare, NJ: Slack, 1977;
 23. Diamant, N. E., and T. Y. El‐Sharkawy. Neural control of esophageal propulsion. Gastroenterology 72: 546–556, 1977;
 24. Christensen, J. editor. Gastrointestinal Motility. New York: Raven, 1979;
 25. Gabella, G. Structure of muscles and nerves in the gastrointestinal tract. In: Physiology of the Gastrointestinal Tract (1st ed.), edited by L. R. Johnson. New York: Raven, 1981, vol. 1, p. 197–246;
 26. Goyal, R. K., and B. W. Cobb. Motility of the pharynx, esophagus, and esophageal sphincter. Gastroenterology, vol. 1, p. 359–391;
 27. Roman, C., and J. Gonella. Extrinsic control of digestive tract motility. Gastroenterology, p. 289–333;
 28. Roman, C. editor. Gastrointestinal Motility. Dordrecht, Netherlands: MTP, 1984.
 29. Waller, A., and J.‐L. Prevost. Étude relative aux nerfs sensitifs qui président aux phenomènes réflexes de la déglutition. Arch. Physiol. Norm. Pathol. 3: 185–197, 343–354, 1870.
 30. Wassileff, N. Wo wird der Schlukreflex ausgelöst? Z. Biol. 24: 29–46, 1888.
 31. Storey, A. T. A Functional Analysis of Laryngeal Sensory Units in the Cat. University of Michigan, 1964. PhD dissertation;
 32. Ann Arbor: Storey, A. T. Laryngeal initiation of swallowing. Exp. Neurol. 20: 359–365, 1968;
 33. Storey, A. T. A functional analysis of sensory units innervating epiglottis and larynx. Exp. Neurol. 20: 366–383, 1968.
 34. Meltzer, S. J. On the causes of the orderly progress of the peristaltic movements in the oesophagus. Am. J. Physiol. 2: 266–272, 1899.
 35. Doty, R. W., and J. F. Bosma. An electromyographic analysis of deglutition. J. Neurophysiol. 19: 44–60, 1956.
 36. Doty (n. 14).
 37. Doty, R. W., W. H. Richmond, and A. T. Storey. Effects of medullary lesions on coordination of deglutition. Exp. Neurol. 17: 91–106, 1967. References and analysis of earlier work are in Doty's Handbook article (n. 15).
 38. Doty, Effects, (n. 22).
 39. Jean, A. Localization et activité des neurones déglutieurs bulbaries. J. Physiol. Paris 64: 227–268, 1972;
 40. Jean, A. Effect du lesions localisées du bulbe rachidien sur le stade oesophagièn de la déglutition. J. Physiol. Paris 64: 507–516, 1972.
 41. Sumi, T. Role of pontine reticular formation in the neural organization of deglutition. Jpn. J. Physiol. 22: 295–314, 1972.
 42. Lueck, W., J. Galliger, and J. F. Bosma. Persistent sequelae of bulbar poliomyelitis. J. Pediatr. 41: 549–554, 1952;
 43. Bosma, J. F. A correlated study of the anatomy and motor activity of the upper pharynx by cadaver dissection and by cinematic study of patients after maxillo‐facial surgery. Ann. Otol. Rhinol. Laryngol. 62: 51–72, 1953;
 44. Bosma, J. F. Studies of disability of the pharynx resultant from poliomyelitis. Ann. Otol. Rhinol. Laryngol. 62: 529–547, 1953.
 45. Doty and Bosma (n. 20).
 46. Mårtinson, A. Proprioceptive impulse patterns during contraction of intrinsic laryngeal muscles. Acta Physiol. Scand. 62: 176–194, 1964. There is a detailed account of the action of pharyngeal muscles in swallowing in Bosma (n. 15) and in Doty (n. 2).
 47. For a description of Code's wartime work, see Wood, E. H. Charlie Code, reminiscences. Mayo Clin. Proc. 50: 497–506, 1975.
 48. Code, C. F., and J. F. Schlegel. Motor action of the esophagus and its sphincters. In: Handbook of Physiology. Alimentary Canal, edited by C. F. Code. Washington, DC: Am. Physiol. Soc, 1968, sect. 6, vol. IV, chapt. 90, p. 1821–1839.
 49. Brody, D. A., J. M. Werle, I. Meschan, and J. P. Quigley. Intraluminal pressures of the digestive tract, especially the pyloric region. Am. J. Physiol. 130: 791–801, 1940.
 50. The glass spoon is described in Kubicek, W. G., F. R. Sedgwick, and M. B. Visscher. Adaptation of the glass spoon manometer to physiological studies. Rev. Sci. Inst. 12: 101–103, 1941.
 51. The spoon had been devised by W. G. Gibson. On a method of determining vapor densities at high temperatures and on a new form of quartz manometer. Proc. R. Soc. Edinb. 33: 1–8, 1911, but it is not described in Gibson's paper because someone who later gave Gibson credit had described it first.
 52. Code's early methods are described in Posey, E. L., W. H. Dearing, J. A. Bargen, and C. F. Code. The recording of intestinal motility. Proc. Staff Meet. Mayo Clin. 23: 297–301, 1948, and in
 53. Hightower, N. C., Jr., C. F. Code, and F. T. Maher. A method for the study of gastro‐intestinal motor activity in human beings. Proc. Staff Meet. Mayo Clin. 24: 453–462, 1949.
 54. The Gauer transducer is described in Gauer, O., and E. Grenapp. A miniature pressure recording device. Science Wash. DC 112: 404–405, 1950;
 55. and its use in Ellis, F. H., Jr., O. H. Gauer, and E. H. Wood. An intrathoracic manometer: its evaluation and application. Circulation 3: 390–398, 1951.
 56. Code, C. F., N. C. Hightower, and C. G. Morlock. Motility of the alimentary canal in man. Am. J. Med. 13: 328–351, 1952;
 57. Butin, J. W., A. M. Olsen, H. J. Moersch, and C. F. Code. A study of esophageal pressure in normal persons and in patients with cardiospasm. Gastroenterology 23: 278–291, 1953;
 58. Code, C. F., and J. F. Schlegel. The pressure profile of the gastroesophageal sphincter in man. Proc. Staff Meet. Mayo Clin. 33: 406–414, 1958.
 59. Frank, O. Kritik der elastischen Manometer. Z. Biol. 44: 445–613, 1903;
 60. Brody, D. A., and J. P. Quigley. Registration of digestive tract intraluminal pressure. In: Methods in Medical Research. Chicago, IL: Year Book, 1951, vol. 4, p. 109–123;
 61. Quigley, J. P., and D. A. Brody. A physiological and clinical consideration of pressures developed in the digestive tract. Am. J. Med. 13: 73–81, 1952.
 62. Lorber, S. H., and H. Shay. Technical and physiological considerations in measuring gastrointestinal pressures in man. Gastroenterology 27: 478–487, 1954 [2–3 ml/h];
 63. Texter, E. C., Jr., H. W. Smith, H. C. Moeller, and C. J. Barborka. Correlation of motor activity in normal subjects and patients with esophageal disorders. Gastroenterology 32: 1013–1024, 1957 [6–10 ml/h];
 64. and Stef, J. J., W. J. Dodds, W. J. Hogan, J. H. Linehan, and E. T. Stewart. Intraluminal esophageal monometry. An analysis of variables affecting recording fidelity of peristaltic pressure. Gastroenterology 67: 221–230, 1974.
 65. Ingelfinger, F. J., and W. O. Abbott. The diagnostic significance of motor disturbances. Am. J. Dig. Dis. 7: 468–474, 1940;
 66. Ingelfinger, F. J., and R. E. Moss. The activity of the descending duodenum during nausea. Am. J. Physiol. 136: 561–566, 1942.
 67. Ingelfinger gave the latter paper at the February 1942 meeting of the Philadelphia Physiological Society. The next paper on the program was later published as Davenport, H. W. The mechanism of secretion of sulfonamide drugs in gastric juice. Yale J. Biol. Med. 14: 589–597, 1942. The last paper reported the cleverest work Davenport ever did.
 68. There is a list of Ingelfinger's 57 fellows in Farrar, J. T. Preface (to an Ingelfinger Festschrift]. Am. J. Dig. Dis. 22: 276–278, 1977. In the early days Ingelfinger's fellows were supported on a research grant from the United States Public Health Service. That was long before training grants became available. Davenport takes the credit for seeing that the physiology study section approved Ingelfinger's first application.
 69. This is the story I heard Ingelfinger tell when he gave a lecture at the University of Utah College of Medicine.
 70. As always happens when someone does original work, there was a tremendous amount of me‐tooism in the next 10 years that resulted in a lot of niggling and i‐dotting. There must be a hundred papers of that kind for every one I cite.
 71. Barclay, A. E. The Digestive Tract. London: Cambridge Univ. Press, 1933.
 72. Barclay, A. E. The normal mechanism of swallowing. Br. J. Radiol. 3: 534–546, 1930.
 73. The assertion and the same evidence are repeated in Barclay, A. E.. The mechanics of the digestive tract. Lancet 1: 11–15, 1934.
 74. Example: Vantrappen, G., M. D. Liemer, J. Ikeya, E. C. Texter, Jr., and C. J. Barborka. Simultaneous fluorocinematography and intraluminal pressure measurements in the study of esophageal motility. Gastroenterology 35: 592–602, 1958.
 75. The methods are described in Cohen, B. R., and B. S. Wolf. Roentgen localization of physiologically determined esophageal hiatus. Gastroenterology 43: 43–50, 1962.
 76. Their work is summarized in Cohen, B. R., and B. S. Wolf. Cineradiographic and intraluminal pressure correlations in the pharynx and esophagus. In: Handbook of Physiology. Alimentary Canal, edited by C. F. Code. Washington, DC: Am. Physiol. Soc., 1968, sect. 6, vol. IV, chapt. 91, p. 1841–1860.
 77. For a summary of earlier work, see Code, C. F., B. Creamer, and J. F. Schlegel. An Atlas of Esophageal Motility in Health and Disease. Springfield, IL: Thomas, 1958.
 78. Cannon (n. 8);
 79. Meltzer (n. 19).
 80. Hwang, K., and M. I. Grossman. A note on the innervation of the cervical portion of the human esophagus. Gastroenterology 25: 375–377, 1953;
 81. Hwang, K., M. I. Grossman, and A. C. Ivy. Nervous control of the cervical portion of the esophagus. Am. J. Physiol. 154: 343–357, 1948.
 82. Kantrowitz, P. A., C. I. Siegel, and T. R. Hendrix. Differences in motility of the upper and lower esophagus in man and its alteration by atropine. Bull. Johns Hopkins Hosp. 118: 476–491, 1966.
 83. Tieffenbach, J., and C. Roman. Rôle de l'innervation extrinsèque vagale dans la motricité de l'esophage à musculeuse lisse: étude electromyographique chez le chat et al babouin. J. Physiol. Paris 64: 193–226, 1972.
 84. A later account of the characteristics of secondary peristalsis is Creamer, B., and J. Schlegel. Motor responses of the esophagus to distention. J. Appl. Physiol. 10: 498–504, 1957.
 85. Lumsden, K., and W. S. Holden. The act of vomiting in man. Gut 10: 173–178, 1969.
 86. McNally, E. K., J. E. Kelly, Jr., and F. J. Ingelfinger. Mechanism of belching: effect of gastric distention with air. Gastroenterology 46: 254–259, 1964.
 87. Roman, C. Contrôl nerveux due perstaltisme cesophagien. J. Physiol. Paris 58: 79–108, 1966;
 88. Roman, C., and J. Tieffenbach. Enrigistemènt de l'activité unitaire des fibres motrices vagales destinées a l'osophage du babouin. J. Physiol. Paris 64: 479–506, 1972.
 89. Winship, D. H., and F. F. Zboralske. The esophageal propulsive force: esophageal response to obstruction. J. Clin Invest. 46: 1391–1401, 1967;
 90. Dodds, W. J., W. J. Hogan, D. P. Reid, E. T. Stewart, and R. C. Arndorfer. A comparison between primary esophageal peristalsis following wet and dry swallows. J. Appl. Physiol. 35: 851–857, 1973.
 91. Siegel, C. F., and T. R. Hendrix. Evidence for the central mediation of secondary peristalsis in the esophagus. Bull. Johns Hopkins Hosp. 108: 297–307, 1961.
 92. For reviews, see Wood, J. D. Neurophysiology of Auerbach's plexus and control of intestinal motility. Physiol. Rev. 55: 307–324, 1975;
 93. Diamant, N. E., and T. Y. El‐Sharkawy. Neural control of esophageal peristalsis. Gastroenterology 72: 546–551, 1977.
 94. Cannon (n. 8), p. 23–26;
 95. Diamant, N. E., and T. Y. El‐Sharkawy. Oesophageal peristalsis after bilateral vagotomy. Am. J. Physiol. 19: 436–444, 1907.
 96. Christensen, J., and G. F. Lund. Esophageal responses to distention and electrical stimulation. J. Clin. Invest. 48: 409–419, 1969.
 97. Mukhopadhyay, A. K., and N. W. Weisbrodt. Neural organization of esophageal peristalsis: role of the vagus nerve. Gastroenterology 68: 444–447, 1975;
 98. Weisbrodt, N. W. Neuromuscular organization of esophageal motility. Arch. Intern. Med. 136: 524–531, 1976.
 99. Tieffenbach and Roman (n. 47); Kantrowitz et al. (n. 46).
 100. Roman, C., and L. Tieffenbach. Motricité de l'oesophage à musculeuse lisse après bivagotomie. J. Physiol. Paris 63: 733–762, 1971.
 101. Luciani, L. Delle oscillazione della pressione intratoracica e intraabdominale. Arch. Sci. Med. 2: 177–224, 301–351, 1878.
 102. Luciani credited Ceradini with “la felice idea di sosstituiere alla vesicicola elastica intradda nella cavita pleurale un semplice budello, legato all'esterno di tubo pertgiato, e introdotto nel canale esofago, previa aperturo di questo, praticata nel collo.” Respiratory physiologists ever since have used the method. See Mead, J., M. B. McIlroy, J. N. Selverstone, and B. C. Kriete. Measurement of intraesophageal pressure. J. Appl. Physiol. 7: 491–495, 1955.
 103. Schlippe, P. Physicalische Untersuchungen bei der Anwendung der Magenschlauches. Arch. Klin. Med. 76: 450–473, 1903.
 104. Killian, G. Uber den Mund der Speiseröhr. Z. Ohrenheilk. 55: 1–41, 1908.
 105. Kindermann, J. G. Bijdrag tot de Physiologie ven het Slikken. Nederl. Tijdschr. Geneesk. 2: 1182–1202, 1903.
 106. Fyke, F. E., and C. F. Code. Resting and deglutition pressures in the pharyngoesophageal junction. Gastroenterology 29: 24–34, 1955.
 107. Doty, Neural organization, (n. 15), p. 1873–1874.
 108. Doty and Bosma (n. 20).
 109. Clark, G. H. Deglutition apnea. J. Physiol. Lond. 54: lix, 1920.
 110. Summary and references in Code et al., Atlas (n. 43).
 111. Kodicek, J., and B. Creamer. A study of pharyngeal pouches. J. Laryngol. 75: 406–411, 1961.
 112. Kramer, P., M. Atkinson, S. M. Wyman, and F. J. Ingelfinger. The dynamics of swallowing. II. Neuromuscular dysphagia of the pharynx. J. Clin. Invest. 36: 589–595, 1957.
 113. There is a speculative discussion of the problem of fibrosis in Siegler, R. Dysphagia—an unusual presenting syndrome of the “stiff‐man syndrome.” Arch. Pathol. 70: 520–525, 1960.
 114. Bosma, J. F. Poliomyelitic disabilities of the pharynx. J. Pediatr. 19: 881–907, 1957.
 115. Arloing, S. Application de la méthode graphique à l'étude de quelques points de la déglutition. C. R. Acad. Sci. Paris 79: 1009–1013, 1874;
 116. Carlet, G. Sur le mechanisme de le déglutition. C. R. Acad. Sci. Paris 79: 1013–1014, 1874.
 117. Barclay (n. 41).
 118. Butin et al. (n. 33) said they saw the drop one‐third of the time, but Ingelfinger in discussion of the paper said he saw it less frequently. Arloing and Carlet attributed it to a brief inspiratory effort, but Code thought it is caused by pull on the esophagus as the larynx rises.
 119. The first comprehensive description of the wave from Code's laboratory is in Butin et al. (n. 33). Those records were made with a single transducer. Better and apparently otherwise unpublished records were made with multiple transducers and are in Code et al., Atlas, (n. 43). Descriptions in Butin et al. do not agree with those in the Atlas. For example, Butin's figures show respiration suspended throughout swallowing, but the Atlas asserts that respiration is only briefly inhibited. The paper asserts that the duration of the wave in the distal esophagus averages 12.6 s, whereas what is probably meant is that it ends about 12 s after swallowing begins.
 120. See also Atkinson, M., P. Kramer, S. M. Wyman, and F. J. Ingelfinger. The dynamics of swallowing. II. Normal pharyngeal mechanisms. J. Clin. Invest. 36: 581–588, 1957.
 121. Example: Dornhorst, A. C., K. Harrison, and J. W. Pierce. Observations on the normal esophagus and cardia. Lancet 1: 695–698, 1954.
 122. Sanchez, G. C., P. Kramer, and F. J. Ingelfinger. Motor mechanisms of the esophagus, particularly its distal portion. Gastroenterology 25: 321–332, 1953.
 123. Examples: Schatski, R. Reliefstudien an der normalen und krankhafte veränderten Speiseröhre. Acta Radiol. Suppl. 18, 1933;
 124. Moersch, H. J., and J. D. Camp. Diffuse spasm of the lower part of the esophagus. Ann. Otol. Rhinol. Laryngol. 43: 1165–1173, 1934.
 125. Schmidt, H. E. Diffuse spasm of the lower half of the esophagus. Am. J. Dig. Dis. 6: 693–700, 1939.
 126. Creamer, B., F. E. Donoghue, and C. F. Code. Pattern of esophageal motility in diffuse spasm. Gastroenterology 34: 782–796, 1958.
 127. Ehrmann, S. Ueber die Beziehung der Sklerodermie zu den autotoxischen Erythemen. Wien. Klin. Wochenschr. 53: 1098–1102, 1155–1159, 1903.
 128. Examples: Fessler, A., and R. Pohl. Stenosierenden Process des Ösophagus bei Sklerodermie. Dermat. Z. 63: 164–169, 1932;
 129. Kuré, K., K. Yamagata, S. Tsukada, and J. Hiyoshi. Passagestörung des Oesophagus bei Sklerodermie und Dystrophia Musculorum Progressiva. Klin. Wochenschr. 15: 516–520, 1936;
 130. Lindsay, J. R., F. E. Templeton, and S. Rothman. Lesions of the esophagus in generalized progressive scleroderma. J. Am. Med. Assoc. 123: 745–750, 1943.
 131. Olsen, A. W., P. A. O'Leary, and B. R. Kirklin. Esophageal lesions associated with acrosclerosis and scleroderma. Arch. Intern. Med. 76: 189–200, 1945.
 132. Creamer, B., H. A. Andersen, and C. F. Code. Esophageal motility in patients with scleroderma and related diseases. Gastroenterologia 86: 763–775, 1956.
 133. Lundrum, F. C. Anatomic features of the cardiac orifice of the stomach. Arch. Intern. Med. 59: 474–511, 1937.
 134. For a comprehensive review, see Botha, G. S. M. The Gastrooesophageal Junction. Boston, MA: Little, Brown, 1963.
 135. When Ingelfinger reviewed this book, he said: “G. S. Muller Botha is a big, handsome, and confident man, and he has produced a big, handsome, and confident book.” Ingelfinger's judgment on three major points was “not so good.” Ingelfinger, F. J. Review. Gastroenterology 46: 622–623, 1964.
 136. Representative negative reports are Arey, L. B., and M. J. Tremaine. The muscle coat of the lower esophagus in man. Anat. Rec. 56: 315–320, 1933;
 137. and Lundrum, F. C. Anatomic features of the cardiac orifice of the stomach. Arch. Intern. Med. 59: 474–511, 1937.
 138. The fullest description of Cannon's work on the cardiac sphincter is in Cannon (n. 8), p. 32–44.
 139. Cannon was controverted by Carlson, A. J., T. E. Boyd, and J. F. Pearse. The innervation of the cardia and the lower end of the esophagus in mammals. Arch. Intern. Med. 33: 281–291, 1924; but more modern work seems to show that Cannon was right.
 140. On account of multiple publication of essentially the same results, I make no attempt to assign priority. A representative paper by Ingelfinger's group is Ingelfinger, F. J., P. Kramer, and G. C. Sanchez. The gastroesophageal vestibule, its normal function and its role in cardiospasm and gastroesophageal reflux. Am. J. Med. Sci. 228: 417–425, 1954.
 141. One from Texter's group is Vantrappen, G., M. D. Liemer, J. Ikeya, E. C. Texter, Jr., and C. J. Barborka. Simultaneous fluorocinematography and intraluminal pressure measurements in the study of esophageal motility. Gastroenterology 35: 592–602, 1958.
 142. A roughly contemporaneous review with 163 references is Mann, C. V., R. K. Greenwood, and F. H. Ellis, Jr. The esophagastric junction. Surg. Gynecol. Obstet. 118: 853–862, 1964.
 143. There is an enormous literature on this subject. See Lerche, W. The Esophagus and Pharynx in Action. Springfield, IL: Thomas, 1950. See Fig. (this chapter) for the appearance of a bolus like an inverted arrowhead just above the diaphragm.
 144. Creamer, B., and J. W. Pierce. Observations on the gastroesophageal junction during swallowing and drinking. Lancet 2: 1309–1312, 1957;
 145. Creamer, B., G. K. Harrison, and J. W. Pierce. Further observations on the gastroesophageal junction. Thorax 14: 132–137, 1959.
 146. Harris, L. D., and C. E. Pope. II. The pressure inversion point: its genesis and reliability. Gastroenterology 51: 641–648, 1966.
 147. Cohen, B. R., and B. S. Wolf. Roentgen localization of the physiologically determined esophageal hiatus. Gastroenterology 43: 43–50, 1962.
 148. Fleshler, B., T. R. Hendrix, P. Kramer, and F. J. Ingelfinger. Resistance and reflex function of the lower esophageal sphincter. J. Appl. Physiol. 12: 339–342, 1958.
 149. Burget, G. E., and W. E. Zeller. A study of the cardia in unanesthetized dogs. Proc. Soc. Exp. Biol. Med. 34: 433–434, 1932.
 150. Cannon, W. B. The acid closure of the cardia. Am. J. Physiol. 23: 105–114, 1908;
 151. summarized in Cannon, W. B. (n. 8), p. 32–44.
 152. The results as of 1975 are briefly summarized in Castell, D. O. The lower esophageal sphincter. Physiologic and clinical results. Ann. Intern. Med. 83: 390–401, 1975;
 153. and in Snape, W. J., Jr., and S. Cohen. Hormonal control of esophageal function. Arch. Intern. Med. 136: 538–542, 1976.
 154. Giles, G. R., M. C. Mason, C. Humphries, and G. C. Clark. Action of gastrin on the lower oesophageal sphincter in man. Gut 10: 730–734, 1969.
 155. Castell, D. O., and L. D. Harris. Hormonal control of gastroesophageal sphincter strength. N. Engl. J. Med. 282: 886–889, 1970.
 156. Cohen, S., and W. Lipschutz. Hormonal regulation of human lower esophageal sphincter competence: interaction of gastrin and secretin. J. Clin. Invest. 50: 449–454, 1971;
 157. Nebel, O. T., and D. O. Castell. Lower esophageal sphincter pressure changes after food ingestion. Gastroenterology 63: 778–783, 1972.
 158. Sturdevant, R. L., and T. Kun. Interaction of pentagastrin and the octapeptide of cholecystokinin on the human lower oesophageal sphincter. Gut 15: 700–702, 1974.
 159. Giles, G. R., C. Humphries, M. C. Mason, and G. C. Clark. Effect of pH on the cardiac sphincter. Gut 10: 852–856, 1969.
 160. Openchowski, T. von. Ueber Centren und Leitungsbahnen für die Musculatur des Magens. Arch. Anat. Physiol. Physiol. Abt. 549–556, 1889.
 161. The work is described in a bit more detail in Openchowski, T. von. Ueber die nervösen Vorrichtungen des Magens. Zentralbl. Physiol. 1–10, 1889.
 162. Openchowski was working in Dorpat, the university for German‐speaking Russians in Estonia after touring Berlin, Paris, and London. There is no evidence in his papers, but he may have done his neurophysiological and anatomical work under Friedrich Bidder, who after publication of the famous Bidder and Schmidt book in 1852 did very little except some studies on the nervous system. See Bing, F. C. Friedrich Bidder (1810–1894) and Carl Schmidt (1822–1894)—a biographical sketch. J. Nutr. 103: 637–648, 1973.
 163. In 1891 Openchowski became professor of therapeutics in Kharkov. He died in Berlin in 1914 after an operation for cancer of the stomach. For a biographical sketch, see his obituary notice in Kharkov. Med. J. 27: 86–87, 1914. I am grateful to a graduate student in Michigan's department of Slavic languages for a translation.
 164. Langley, J. N. On inhibitory fibres in the vagus for the end of the oesophagus and the stomach. J. Physiol. Lond. 23: 407–414, 1899.
 165. Langley (n. 104), p. 410.
 166. Page May, W. The innervation of the sphincters and musculature of the stomach. J. Physiol. Lond. 31: 260–272, 1904. Page May used the rabbit and perhaps the monkey; his description of animals used in particular experiments is unclear.
 167. Cannon, W. B., and C. W. Lieb. The receptive relaxation of the stomach. Am. J. Physiol. 29: 267–273, 1911.
 168. Fleshler et al. (n. 94).
 169. Knight, G. C. The relation of the extrinsic nerves to the functional activity of the oesophagus. Br. J. Surg. 22: 115–168, 1922. The paper contains a photograph of the postmortem appearance of a cat with cardiospasm.
 170. Hurst, A. F. The treatment of achalasia of the cardia (so‐called “cardiospasm”). Lancet 1: 618–619, 1927.
 171. Butin et al (n. 33); Creamer, B., A. M. Olsen, and C. F. Code. The esophageal sphincters in achalasia of the cardia (cardiospasm). Gastroenterology 33: 293–301, 1957.
 172. For reviews, see Ingelfinger, F. J. The esophagus. Gastroenterology 41: 264–276, 1961;
 173. Ingelfinger, F. J. The esophagus, March 1961 to February 1963. Gastroenterology 45: 241–264, 1963;
 174. Henderson, R. D. Motor Disorders of the Esophagus. Baltimore, MD: Williams & Wilkins, 1976.
 175. Dagradi, A. E., S. J. Stempien, H. W. Seifer, and J. A. Weinberg. Terminal esophageal (vestibular) spasm after vagotomy. Arch. Surg. 85: 955–968, 1962.
 176. For a summary of the evidence as of 1978, see Goyal, R. K., and S. Rattan. Neurohumoral, hormonal, and drug receptors for the lower esophageal sphincter. Gastroenterology 74: 598–619, 1978.
 177. Kramer, P., and F. J. Ingelfinger. Esophageal sensitivity to Mecholyl in cardiospasm. Gastroenterology 19: 242–253, 1951.
 178. This was confirmed and elaborated by Hightower, N. C., Jr., A. M. Olsen, and H. J. Moersch. A comparison of the effects of acetyl‐β‐methylcholine chloride (Mecholyl) on esophageal intraluminal pressure in normal persons and in patients with cardiospasm. Gastroenterology 26: 592–600, 1954.
 179. Lundrum (n. 86).
 180. Smith, B. The neurological lesion in achalasia. Gut 11: 388–391, 1970. There are many similar reports.
 181. Cannon, W. B. A law of denervation. Am. J. Med. Sci. 198: 737–750, 1939. On p. 738: “When in a series of efferent neurones a unit is destroyed, an increased irritability to chemical agents develops in the isolated structure or structures, the effect being maximal in the part directly denervated.” In the rest of the paper Cannon established that the “chemicals” are the particular neurotransmitters.
 182. Goyal and Rattan (n. 114);
 183. Goyal, R. K. Motility of the pharynx, esophagus, and esophageal sphincters. In: Physiology of the Gastrointestinal Tract, edited by L. R. Johnson. New York: Raven, 1981, vol. 1, p. 359–391.
 184. For summary of evidence and references, see Daniel, E. E. A conceptual analysis of the pharmacology of gastro‐intestinal motility. In: Pharmacology of Gastrointestinal Motility, edited by P. Holton. Oxford, UK: Pergamon, 1973, vol. II, p. 457–545. See also Goyal and Rattan (n. 114).
 185. Christensen, J., and E. E. Daniel. Electric and motor effects of autonomic drugs on circular esophageal smooth muscle. J. Pharmacol. Exp. Ther. 159: 243–249, 1968.
 186. Nickerson, M., and L. S. Goodman. Pharmacological properties of a new adrenergic blocking agent: N, N‐dibenzyl‐chloro‐ethylamine (dibenamine). J. Pharmacol. Exp. Ther. 89: 167–185, 1947.
 187. Nickerson, M., and L. S. Call. Treatment of cardiospasm with adrenergic blockade. Am. J. Med. 11: 123–127, 1951.
 188. Bingham, J. R., F. J. Ingelfinger, and R. H. Smithwick. The effects of sympathectomy on motility of the human gastric and biliary tracts. Gastroenterology 15: 6–13, 1950.
 189. Sleisenger, M. H., H. Steinberg, and T. P. Almy. The disturbance of motility in cardiospasm: studies of autonomic stimulation and autonomic blockade, including the cardia. Gastroenterology 25: 333–348, 1953.
 190. Greenwood, R. K., J. F. Schlegel, C. F. Code, and F. H. Ellis, Jr. The effects of sympathectomy, vagotomy and oesophageal interruption on the canine gastrooesophageal sphincter. Thorax 17: 310–317, 1962.
 191. Nagler, R., and H. M. Spiro. Heartburn in pregnancy. Am. J. Dig. Dis. 7: 648–655, 1962.
 192. Bernstein, L. M., and L. A. Baker. A clinical test for esophagitis. Gastroenterology 34: 760–781, 1958;
 193. Ingelfinger, F. J. The physiologic background of heartburn, esophagitis and cardiospasm. Arch. Intern. Med. 105: 770–778, 1960;
 194. Tuttle, S. G., F. Rufin, and A. Bettarello. The physiology of heartburn. Ann. Intern. Med. 55: 292–300, 1961;
 195. Tuttle, S. G., A. Bettarello, and M. I. Grossman. Gastroesophageal regurgitation. J. Am. Med. Assoc. 176: 498–500, 1961.
 196. For reviews, see Henderson (n. 112);
 197. Castell, D. O., W. C. Wu, and D. J. Oh (editors). Gastroesophageal Reflux Disease Mount Kisco, NY: Futura, 1985. The latter would be more useful if fewer of its references were incorrect.
 198. De Meester, T. R., L. F. Johnson, G. J. Joseph, M. S. Toscano, A. W. Hall, and D. B. Skinner. Patterns of gastroesophageal reflux in health and disease. Ann. Surg. 184: 459–470, 1970;
 199. Stanciu, C., and J. R. Bennett. Oesophageal acid clearing: one factor in the production of reflux oesophagitis. Gut 15: 852–857, 1974.
 200. Tuttle, S. G., and M. I. Grossman. Detection of gastroesophageal reflux by simultaneous measurement of intraluminal pressure and pH. Proc. Soc. Exp. Biol. Med. 98: 225–227, 1958; Tuttle, Bettarello, and Grossman (n. 128); Tuttle, Rufin, and Bettarello (n. 128).
 201. Nagler and Spiro (n. 127); Tuttle, S. G., and M. I. Grossman. Heartburn in later pregnancy. Manometric studies of esophageal motor function. J. Clin. Invest. 40: 954–970, 1961.
 202. Edwards, D. A. W. The anti‐reflux mechanism: manometric and radiological studies. Br. J. Radiol. 34: 474–487, 1961.
 203. Example: Atkinson, M., D. A. W. Edwards, A. J. Honour, and E. N. Rowlands. The oesophagogastric sphincter in hiatus hernia. Lancet 2: 1138–1142, 1957.
 204. Code, C. F., M. L. Kelley, Jr., J. F. Schlegel, and H. M. Olsen. Detection of hiatal hernia during esophageal motility tests. Gastroenterology 43: 521–531, 1962.
 205. Atkinson et al. (n. 134).
 206. Cohen, S., and L. D. Harris. Does hiatal hernia affect competence of the gastroesophageal sphincter?. N. Engl. J. Med. 284: 1053–1056, 1971.
 207. Ingelfinger, F. J. The sphincter that is a sphinx. N. Engl. J. Med. 284: 1095–1096, 1971. When we reach the anal sphincters we will see that one other gastroenterologist could not resist the same pun.
 208. Cannon (n. 8), p. 46.
 209. General agreement, at least for the fundus and body, followed publication of Grossman, M. I. The names of the parts of the stomach. Gastroenterology 34: 1159–1161, 1958.
 210. Lewis, F. T. The form of the stomach in human embryos with notes upon the nomenclature of the stomach. Am. J. Anat. 13: 477–503, 1912.
 211. A version of Lewis's diagram is reproduced in Alvarez, W. C. An Introduction to Gastroenterology (4th ed.). New York: Hoeber, 1948, p. 327.
 212. Cannon, W. B. The movements of the stomach studied by means of the Röntgen rays. Am. J. Physiol. 1: 359–382, 1898.
 213. I have used the typescript copy of Cannon's notes given me by A. C. Barger.
 214. Hofmeister, F., and E. Schütz. Ueber die automatische Bewegungen des Magens. Naunyn Schmiedeberg's Arch. Pharmacol. 20: 1–33, 1886, p. 19. The translation here and elsewhere is mine.
 215. Beaumont, W. Experiments and Observations on the Gastric Juice and the Physiology of Digestion. Plattsburg, 1833, p. 109–116.
 216. Cannon (n. 8), p. 64.
 217. Brinton, W. On the movements of the stomach. Lond. Med. Gaz. 8: 1024–1031, 1849.
 218. Williams, F. H. The Roentgen Rays in Medicine and Surgery (2nd ed.). New York: Macmillan, 1902, p. 360.
 219. Groedel's tracings are reproduced in Groedel, F. M. Die Röntgenuntersuchungen des Magens. In: Lehrbuch und Atlas der Röntgendiagnostik, edited by F. M. Groedel. Munich, Germany: Lehmans, 1924, 1. Hb. 547–551, and in Alvarez (n. 141), p. 330–331.
 220. See Barclay, A. E. The Digestive Tract. London: Cambridge Univ. Press, 1933.
 221. Cannon, W. B., and J. B. Blake. Gastro‐enterostomy and pyloroplasty. Ann. Surg. 41: 686–711, 1905, p. 709.
 222. See Lane, W. A. The Operative Treatment of Chronic Intestinal Stasis (3rd ed.). London: Nisbet, 1915, for a gross example.
 223. Bard‐Macleod, Best and Taylor, Lovatt Evans‐Starling and Wiggers are examples of such textbooks.
 224. Cannon (n. 8), p. 59.
 225. Cannon and Lieb (n. 107).
 226. Openchowski, T. von. Ueber Centren und Leitungsbahnen für die Musculatur des Magens. Arch. Anat. Physiol. Physiol. Abt. 549–556, 1889;
 227. Langley, J. N. On inhibitory fibres in the vagus for the end of the oesophagus and the stomach. J. Physiol. Lond. 23: 407–414, 1899;
 228. Page May, W. The innervation of the sphincters and musculature of the stomach. J. Physiol. Lond. 31: 260–271, 1904.
 229. For a review up to about 1965, see Youmans, W. B. Innervation of the gastrointestinal tract. In: Handbook of Physiology. Alimentary Canal, edited by C. F. Code. Washington, DC: Am. Physiol. Soc, 1968, sect. 6, vol. IV, chapt. 82, p. 1655–1663.
 230. Agostini, E., J. E. Chinnock, M. de Burgh Daly, and J. G. Murray. Functional and histological study of the vagus nerve and its branches to the heart, lungs, and abdominal viscera in the cat. J. Physiol. Lond. 135: 182–205, 1957.
 231. Paintal, A. S. A study of gastric stretch receptors. Their role in the peripheral mechanism of satiation of hunger and thirst. J. Physiol. Lond. 126: 255–270, 1954;
 232. Iggo, A. Tension receptors in the stomach and the urinary bladder. J. Physiol. Lond. 128: 593.
 233. Harper, A. A., C. Kidd, and T. Scratcherd. Vaso‐vagal effects on gastric and pancreatic secretion and gastro‐intestinal motility. J. Physiol. Lond. 148: 417–436, 1959.
 234. Jansson, G. Extrinsic nervous control of gastric motility. Acta Physiol. Scand. Suppl. 326: 1969;
 235. Abrahamson, H. Studies on the inhibitory nervous control of gastric motility. Acta Physiol. Scand. Suppl. 390: 1973.
 236. Babkin, B. P., and T. J. Speakman. Cortical inhibition of gastric motility. J. Neurophysiol. Bethesda 13: 55–63, 1950;
 237. Babkin, B. P., and W. C. Kite, Jr. Cortical and reflex regulation of motility of the pyloric antrum. Acta Physiol. Scand. Suppl. 13: 321–334, 1950. Babkin died on the train while returning to Montreal from Philadelphia before the second paper was published. The issue contains an obituary notice and a picure of the handsome Babkin.
 238. Miolan, J. P., and C. Roman. Décharge unitaire des fibres vagales efférentes lors de la relaxation réceptive de l'estomac du chien. J. Physiol. Paris 68: 693–704, 1974.
 239. Van Braam Houckgeest, J. P. Untersuchungen über Peristaltik des Magens und Darmkanal. Pflúegers Arch. Gesamte Physiol. Menschen Tiere 6: 266–303, 1872.
 240. Legros, C., and E.‐N.‐J. Onimus. Recherches experimentales sur les mouvements de l'intestin. J. Anat. Physiol. Paris 6: 37–69, 163–196, 1869.
 241. There is an obituary notice of Legros: Onimus, E.‐N.‐J. J. Anat. Physiol. Paris 10: 113–136, 1874.
 242. Moritz. Ueber das Verhalten des Druckes im Magen. Z. Biol. 32: 313–370, 1895.
 243. Cannon, W. B., and H. F. Day. Salivary digestion in the stomach. Am. J. Physiol. 9: 396–416, 1903.
 244. Cannon (n. 8), p. 63.
 245. Cannon (n. 8), p. 62.
 246. Gianturco, C. Some mechanical factors in gastric physiology. Am. J. Roentgenol. 31: 735–744, 1943.
 247. Cannon (n. 8), p. 6.
 248. Cannon (n. 142).
 249. Cannon, W. B. Bodily Changes in Pain, Fear, and Rage (2nd ed.). New York: Appleton, 1929.
 250. See also Davenport, H. W. Signs of anxiety, rage, or distress. Physiologist 24 (5): 1–5, 1981, for some of Cannon's work after 1929.
 251. Mittelman, B., and H. G. Wolff. Emotions and gastroduodenal function. Experimental studies in patients with gastritis, duodenitis and peptic ulcer. Psychosom. Med. 4: 5–61, 1942.
 252. Wolf, S., and H. G. Wolff. Human Gastric Function. New York: Oxford Univ. Press, 1943. Cannon's comments are in his foreword to the book.
 253. The earliest data for an experiment in Boldyreff's paper is 2 January 1902. The paper is Boldyreff, W. N. Le travail periodique de l'appariel digestif en dehors de la digestion. Arch. Soc. Biol. St. Petersbourg 11: 1–157, 1905.
 254. Many of the same facts and ideas are in Boldyreff, W. N.. Einige neue Seitung der Tätigkeit des Pankreas. Ergeb. Physiol. 11: 121–217, 1911.
 255. Boldyreff graduated from the Russian Imperial Military Medical Academy in 1898He was Pavlov's chief assistant until 1912 when he became professor in Kazan. During the First World War he was a brigadier general in the Russian poison gas corps, serving in France and England. He returned to Russia during the revolution, but he soon made his way with his family through Siberia to Japan and then to the United States. After a year at Western Reserve University, Boldyreff, now William Nicholas, founded the Pavlov Physiological Laboratory at the Battle Creek Sanitarium in 1923, and he worked there until his death in 1946. For an obituary notice, see Ivy, A. C. Gastroenterology 6: 613–614, 1946.
 256. Boldyreff (n. 176), p. 75.
 257. Boldyreff (n. 176), p. 108.
 258. Cannon, W. B. Auscultation of the rhythmic sounds produced by the stomach and intestines. Am. J. Physiol. 14: 339–353, 1905.
 259. When electrical recording became popular after the Second World War many used tape recorders for the purpose. An elaborate example is Farrar, J. T., and F. J. Ingelfinger. Gastrointestinal motility as revealed by the study of abdominal sounds. Gastroenterology 29: 789–800, 1955. In discussion of this paper T. Grier Miller rather dismissively said: “Every physician is familiar with the loud gurgling sounds of diarrheal conditions, with the tinkling musical sounds of early intestinal obstruction, and with the ‘deathly silence’ of adynamic ileus.”
 260. Cannon (n. 8), p. 176.
 261. Cannon (n. 8), p. 204.
 262. The American Medical Directory and Washburn's obituary notice in J. Am. Med. Assoc. 192: 440, 1965, say he graduated from Harvard Medical School in 1915, but the Harvard obituary notice says he graduated in 1914. A. Clifford Barger, the authority on Cannon, tells me that Washburn graduated from Harvard College in 1910 and “apparently is listed in the class of 1914 but did not receive his degree until 1915.” If he then entered the medical school in 1910 he would have had time to learn to swallow the tube so that Cannon could give his first report on 16 December 1911. According to Barger the first entry in Cannon's diary concerning Washburn is in May 1911, and the next entries are in July of the same year.
 263. Cannon, W. B. A consideration of the nature of hunger. Harvey Lect. 6: 130–152, 1911–12.
 264. Cannon, W. B., and A. L. Washburn. An explanation of hunger. Am. J. Physiol. 29: 441–454, 1912;
 265. Cannon, W. B. Bodily Changes in Pain, Hunger, Fear and Rage (1st ed.). New York: Appleton, 1915 and 1922, 1929 (2nd ed.).
 266. James, W. The Principles of Psychology. New York: Holt, 1890, vol. II, p. 442–485.
 267. Carlson, A. J. On the nervous control of the hunger mechanism. Harvey Lect. 11: 37–100, 1915–16;
 268. Carlson, A. J.. The Control of Hunger in Health and Disease. Chicago, IL: Univ. of Chicago Press, 1916. The papers to 1916 are listed in the book.
 269. Grossman, M. I., and I. F. Stein, Jr. Vagotomy and the hunger‐producing action of insulin in man. J. Appl. Physiol. 1: 263–269, 1948.
 270. Brody, D. A., et al. (n. 31).
 271. Davenport, H. W. Human voices. Physiologist 5: 265–269, 1962. On p. 267 condom is misspelled through no fault of the author.
 272. Rogers, F. T., and L. L. J. Hardt. Contributions to the physiology of the stomach. XXVI. The relation between the digestion contractions of the filled, and the hunger contractions of the “empty” stomach. Am. J. Physiol. 38: 274–284, 1915.
 273. Carlson, Control, (n. 187), p. 50.
 274. Carlson, Control, (n. 187), p. 156–160.
 275. Grossman and Stein (n. 188), p. 268.
 276. Luckhardt, A. B., and A. J. Carlson. Contributions to the physiology of the stomach. XVII. On the chemical control of the gastric hunger mechanism. Am. J. Physiol. 36: 37–46, 1914.
 277. Bulatao, E., and A. J. Carlson. Contributions to the physiology of the stomach. Influence of experimental changes in blood sugar level on gastric hunger contractions. Am. J. Physiol. 69: 107–115, 1924.
 278. Mulinos, M. G. Studies on cholin as a motor hormone for the alimentary tract. Am. J. Physiol. 77: 158–165, 1926.
 279. This paper contains the references on the supposed role of choline. Carlson, A. J., E. A. Smith, and I. Gibbons. The action of choline on the alimentary canal of intact dogs. Am. J. Physiol. 81: 431–435, 1927.
 280. Starling, E. H. The muscular and nervous mechanisms of the digestive tract. In: Schäfer, E. A. (editor). Text‐book of Physiology. Edinburgh: Pentland, 1900, vol. II, p. 313–337.
 281. Bayliss, W. M., and E. H. Starling. The movements and innervation of the small intestine. J. Physiol. Lond. 24: 99–143, 1899.
 282. See Bülbring, E., A. F. Brading, A. W. Jones, and T. Tomita (editors). Smooth Muscle: An Assessment of Current Knowledge. London: Arnold, 1981, for 90 closely printed pages of references containing more than 2,300 items.
 283. Davenport, H. W. Physiology of the Digestive Tract (5th ed.). Chicago: Year Book, 1977, p. 22–57.
 284. Richter, C. P. Action currents from the stomach. Am. J. Physiol. 67: 612–633, 1924.
 285. Bozler, E. The activity of the pacemaker previous to the discharge of a muscular impulse. Am. J. Physiol. 136: 543–552, 1942;
 286. Bozler, E. The action potentials accompanying conducted responses in visceral smooth muscles. Am. J. Physiol. 136: 553–560, 1942;
 287. Bozler, E. The action potentials of the stomach. Am. J. Physiol. 144: 693–700, 1945.
 288. Ichikawa, S., and E. Bozler. Monophasic and diphasic potentials of the stomach. Am. J. Physiol. 182: 92–96, 1955.
 289. Weber, J., S. Kohatsu, and T. S. Nelsen. Pacemaker location and electrical conduction in the canine stomach. Gastroenterology 56: 1267, 1969;
 290. Kelly, K. A., and C. F. Code. Canine gastric pacemaker. Am. J. Physiol. 220: 112–118, 1971.
 291. Kelly, K. A., C. F. Code, and L. R. Elveback. Patterns of canine gastric electrical activity. Am. J. Physiol. 217: 461–470, 1969.
 292. Kelly, K. A., and R. C. La Force. Circumferential propagation of canine gastric pacesetter potential. Am. J. Dig. Dis. 17: 339–343, 1972.
 293. Adair, G., and E. N. Goodman. A note on the potential difference across the stomach membrane in human subjects and a simple calomel electrode. J. Physiol. Lond. 87: 35P–36P, 1930.
 294. Goodman, E. N., J. A. Ginsberg, and M. A. Robinson. An improved apparatus for measuring the electrogastrogram. Science Wash. DC 113: 682–683, 1951.
 295. Morton, H. S. The potentialities of the electrogastrogram. Ann. R. Coll. Surg. Engl. 15: 351–361, 1954.
 296. Monges, H., J. Salducci, and C. Roman. Étude électromyographique de la motricité gastrique chez l'homme normal. Arch. Fr. Appl. Dig. 58: 517–530, 1969.
 297. Kwong, N. K., B. H. Brown, G. B. Whittaker, and H. L. Duthie. Electrical activity of the gastric antrum in man. Br. J. Surg. 57: 913–916, 1970;
 298. Duthie, H. L., N. K. Kwong, B. H. Brown, and G. E. Whittaker. Pacesetter potential of the human gastroduodenal junction. Gut 12: 250–256, 1971.
 299. Thomas, J. E. Mechanics and regulation of gastric emptying. Physiol. Rev. 37: 453–474, 1957.
 300. Shay, H., and J. Gershon‐Cohen. A study of pyloric control; roles of acid and alkali. Surg. Gynecol. Obstet. 58: 935–955, 1934.
 301. The full exposition of Cannon's theory is in Cannon, W. B. The acid control of the pylorus. Am. J. Physiol. 20: 283–322, 1907.
 302. Cannon wrote in Mechanical Factors (n. 8), p. 113: “We shall find still more reason for admiration of the pyloric reflex when we see how exactly its acid control can be applied in explaining the differential discharge of different foodstuffs.” This is only one example of Cannon's customary awe of the Wisdom of the Body.
 303. Cannon (n. 8), p. 100.
 304. Cohnheim, O. Die Physiologie der Verdauung und Ernährung. Berlin: Urban & Schwartzenberg, 1908, p. 168: “Auf den Reiz der Säuer schliesst sich der Pylorus, während die Peristaltik des Antrum pylori fortdauert, Fett dagegen bewirkt anscheinend ein aufhören dieser Peristaltik.”
 305. Moore, B., and D. P. Rockwood. On the mode of absorption of fats. J. Physiol. Lond. 21: 58–84, 1897.
 306. Methyl orange changes between pH 3.1 and 4.4; phenolphthalein changes between pH 8.3 and 10.0; litmus changes between pH 4.5 and 8.3. See Clark, W. M. The Determination of Hydrogen Ions. London: Baillière, Tindall & Cox, 1928, p. 78–86.
 307. Levites, S. Über die Verdauung der Fette im tierischen Organismus. Hoppe‐Seyler's Z. Physiol. Chem. 49: 273–285, 1906.
 308. Zenon Bacq commented on Cannon's lack of interest in biochemistry at the centenary celebration of Cannon's birth. His comment is recorded in part in Bacq, Z. M. Walter B. Cannon's contribution to the theory of chemical mediation of the nerve impulse. In: The Life and Contributions of Walter Bradford Cannon, 1871–1945, edited by C. M. Brooks, K. Koizumi, and J. O. Pinkston. New York: Downstate Medical Center, 1975. On the same occasion another speaker said that “if Cannon had been able to woo Folin from his devotion to the study of the composition of urine” Cannon might have had some useful help with biochemical problems.
 309. Cannon (n. 8), p. 128.
 310. Atkinson, M., E. A. W. Edwards, A. J. Honour, and E. N. Rowlands. A comparison of cardiac and pyloric sphincters. Lancet 2: 918–922, 1957.
 311. Anderssen, S., and M. I. Grossman. Profile of pH, pressure, and potential difference at gastroduodenal junction in man. Gastroenterology 49: 364–371, 1965;
 312. Brink, B. M., J. F. Schlegel, and C. F. Code. The pressure profile of the gastroduodenal junction in dogs. Gut 6: 163–171, 1965.
 313. Thomas, J. E. The mechanism of gastric evacuation. J. Am. Med. Assoc. 97: 1663–1668, 1931.
 314. Other convenient summaries of Thomas's views are Thomas, J. E., and J. O. Crider. The regulation of gastric emptying. VA Med. Mon. 64: 181–185, 1937;
 315. and Thomas, J. E., and J. O. Crider. A study of gastric emptying with the pylorus open. Am. J. Dig. Dis. 4: 295–300, 1937. The last demonstrates that fluids empty from the stomach at the same rate when the sphincter is intact and when it is held open by a tube.
 316. Wheelon began by doing undistinguished endocrinological work under Roy Hoskins at Nortwestern University, but the gastrointestinal work he did at St. Louis University between 1914 and 1921 was as good as any at the time. For Thomas, see Code, C. F. Presentation of the Julius Friedenwald Medal to Jacob Earl Thomas. Gastroenterology 63: 725–727, 1972.
 317. Wheelon, H., and J. E. Thomas. Observations on the motility of the antrum and the relation of rhythmic activity of the pyloric sphincter to that of the antrum. J. Lab. Clin. Med. 6: 124–143, 1920;
 318. Wheelon, H., and J. E. Thomas. Rhythmicity of the pyloric sphincter. Am. J. Physiol. 54: 460–473, 1920–21;
 319. Wheelon, H. Observations on gastric and duodenal motility in duodenal obstruction. J. Am. Med. Assoc. 77: 1404–1406, 1921;
 320. Wheelon, H., and J. E. Thomas. Observations on the motility of the duodenum and the relation of duodenal activity to that of the pars pylorica. Am. J. Physiol. 59: 72–96, 1922;
 321. Wheelon, H. Duodenal motility. NY J. Med. 117: 652–655, 1923.
 322. Thomas, J. E., and J. O. Crider. Rhythmic changes in duodenal motility associated with gastric peristalsis. Am. J. Physiol. 111: 124–129, 1935.
 323. Quigley, J. P. A modern explanation of the gastric emptying mechanism. Am. J. Dig. Dis. 10: 418–421, 1943.
 324. Quigley, J. P., and M. R. Read. The spontaneous motility of the pyloric sphincter and its relation to gastric evacuation: the “pyloric diagraph.” Am. J. Physiol. 137: 234–237, 1942.
 325. Thomas (n. 226);
 326. Thomas. Differential manometer records of gastric and duodenal pressures. Am. J. Physiol. 123: 201–202, 1938.
 327. Ewald, A. A., and J. Boas. Beiträge zur Physiologie und Pathologie der Verdauung. Virchows Arch. Pathol. Physiol. Anat. 101: 325–375, 1883;
 328. Virchows Arch. Pathol. Physiol. Anat. 104: 271–305, 1886.
 329. The observations on oil are at the end of the seond, which is a continuation of the first. See Waugh, J. M. Effect of fat introduced into the jejunum by fistula on motility and emptying of the stomach. Arch. Surg. 33: 451–466, 1936, for a historical introduction doubtless assembled by the dutiful librarians at the Mayo Clinic.
 330. Marbaix, O. Le passage pylorique. Cellule 14: 249–332, 1898.
 331. Cited by Marbaix (n. 234), p. 262.
 332. Carnot, P., and A. Chassevant. Modification subies dans l'estomac et le duodénum par les solutions salines suivant leur concentration moléculaire. Le réflex regulation du sphincter pylorique. C. R. Soc. Biol. Paris 58: 173–176, 1905.
 333. Morgan, C. J., and J. E. Thomas. The effect on the pyloric sphincter of acid in the stomach and in the duodenum. Am. J. Physiol. 97: 546–547, 1931;
 334. Quigley, J. P., M. R. Read, K. H. Radzow, I. Meschan, and J. M. Werle. The effect of hydrochloric acid on the pyloric sphincter, the adjacent portions of the digestive tract, and on the process of gastric evacuation. Am. J. Physiol. 137: 153–159, 1942.
 335. Quigley, J. P., J. Werle, E. W. Ligon, Jr., M. R. Read, K. H. Radzow, and I. Meschan. The influence of fats on the motor activity of the pyloric sphincter region and on the process of gastric evacuation studied by the balloon‐water manometer and by the optical manometer‐fluroscopic technics. Am. J. Physiol. 134: 132–140, 1941;
 336. Quigley, J. P., and I. Meschan. Inhibition of the pyloric sphincter region by the digestion products of fat. Am. J. Physiol. 134: 803–807, 1941;
 337. Thomas, J. E. Gastric inhibition caused by amino acids in the small intestine. Am. J. Physiol. 135: 609–613, 1942.
 338. Thomas, J. E., and J. O. Crider. The effect of fat on the pH of the contents of the duodenum. Am. J. Physiol. 114: 603–608, 1936;
 339. Thomas, J. E. The maximal acidity of the intestinal contents during digestion. Am. J. Dig. Dis. 7: 195–197, 1940.
 340. Bircher, J., C. V. Mann, H. C. Carlson, C. F. Code, and R. A. Rovelstad. Intraluminal and juxtamucosal duodenal pH. Gastroenterology 48: 472–477, 1965.
 341. See also Tomenius, J., and G. Williams. Continuously recorded pH of gastric and duodenal contents in situ with an evaluation of the efficacy of some antacids in vivo. Acta Med. Scand. 166: 25–34, 1960.
 342. Rhodes, H., and C. J. Prestich. Acidity at different sites in the proximal duodenum of normal subjects and patients with duodenal ulcer. Gut 7: 509–514, 1966.
 343. Thomas, J. E., and H. Wheelon. The nervous control of the pyloric sphincter. J. Lab. Clin. Med. 7: 375–391, 1921;
 344. Fauley, G. B., and A. C. Ivy. The effect of exclusion of pancreatic juice on gastric digestion. Am. J. Physiol. 89: 428–437, 1929;
 345. Crider, J. O., and J. E. Thomas. A further study of the inhibitory effect on gastric peristalsis of the products of protein digestion. Am. J. Physiol. 123: 44–45, 1938;
 346. Quigley, J. P., and I. Meschan. The role of the vagus in the regulation of the pyloric sphincter and adjacent portions of the gut, with special reference to the process of gastric evacuation. Am. J. Physiol. 123: 166–170, 1938.
 347. Quigley, J. P., H. S. Zettleman, and A. C. Ivy. Analysis of the factors involved in gastric motor inhibition by fats. Am. J. Physiol. 108: 643–651, 1934;
 348. Kosaka, T., R. K. S. Lim, S. M. Ling, and A. C. Liu. On the mechanism of inhibition of gastric secretion by fat. A gastric inhibitory agent obtained from the intestinal mucosa. Chin. J. Physiol. 6: 107–128, 1932;
 349. Lim, R. K. S., S. M. Ling, and A. C. Liu. Depressor substances in extracts of intestinal mucosa. Purification of enterogastrone. Chin. J. Physiol. 8: 219–236, 1943.
 350. Lim's work and spectacular career as Lieutenant General in the Army of the Republic of China are summarized in Davenport, H. W. Robert Kho‐Seng Lim. Biogr. Mem. Natl. Acad. Sci. USA 51: 280–305, 1980.
 351. Gray, J. S., W. B. Bradley, and A. C. Ivy. On the preparation and biological assay of enterogastrone. Am. J. Physiol. 118: 463–476, 1937.
 352. Daniel, E. E. Electrical and contractile activity of pyloric region in dogs and effect of drugs administered intra‐arterially. Gastroenterology 49: 403–418, 1965;
 353. Kelly et al. (n. 206).
 354. Carlson, H. C., C. F. Code, and R. A. Nelson. Motor action of the canine gastroduodenal junction: a cineradiographic, pressure, and electric study. Am. J. Dig. Dis. 11: 155–172, 1966.
 355. See also Brody, D., and J. P. Quigley. Intraluminal pressures of the stomach and duodenum in health and disease. Gastroenterology 9: 570–575, 1947.
 356. Marbaix (n. 234).
 357. Hunt, J. N., and M. T. Knox. Regulation of gastric emptying. In: Handbook of Physiology. Alimentary Canal, edited by C. F. Code. Washington, DC: Am. Physiol. Soc., 1968, sect. 6, vol. IV, chapt. 94, p. 1917–1935 (see p. 1918).
 358. See also Hunt, J. N. Gastric emptying and secretion in man. Physiol. Rev. 39: 491–533, 1959.
 359. Hunt, J. N., and W. R. Spurrell. The pattern of gastric emptying of the human stomach. J. Physiol. Lond. 113: 157–168, 1951.
 360. Hopkins, A. The pattern of gastric emptying: a new view of old results. J. Physiol. Lond. 182: 144–150, 1966.
 361. Stubbs, D. F. Models of gastric emptying. Gut 18: 202–207, 1977.
 362. Jacobs, M. H., and D. R. Stewart. The role of carbonic anhydrase in certain ion exchanges involving the erythrocyte. J. Gen. Physiol. 25: 539–552, 1942.
 363. Hunt, J. N., and J. D. Pathak. The osmotic effects of some simple molecules and ions on gastric emptying. J. Physiol. Lond. 154: 254–269, 1954.
 364. Meeroff, J. C., V. L. W. Go, and S. F. Phillips. Control of gastric emptying by osmolarity of duodenal contents in man. Gastroenterology 68: 1144–1151, 1975.
 365. For a summary, see Meyer, J. H. Gastric emptying of ordinary food: effect of antrum on particle size. Am. J. Physiol. 239 (Gastrointest. Liver Physiol. 2): G133–G135, 1980.
 366. Griffith, G. H., G. M. Owen, S. Kirkman, and R. Shields. Measurement of rate of gastric emptying using chromium‐51. Lancet 1: 1244–1245, 1966. Owen and Kirkman were physicists, not surgeons.
 367. Bromester, D., G. Carlberger, and G. Lundh. Measurement of gastric emptying using 131I‐HSA. Scand. J. Gastroenterol. 3: 641–653, 1968.
 368. Meyer, J. H., I. L. MacGregor, R. Gueller, P. Martin, and R. Cavalieri. 99mTc‐tagged chicken liver as a marker of solid food in the human stomach. Am. J. Dig. Dis. 21: 296–304, 1976.
 369. Hinder, R. A., and K. A. Kelly. Canine gastric emptying of solids and liquids. Am. J. Physiol. 233 (Endocrinol. Metab. Gastrointest. Physiol. 2): E335–E340, 1977.
 370. Schlegel, J. F., W. M. Coburn, Jr., and C. F. Code. Gastric emptying of solid and compliant spheres in dogs (Abstract). Physiologist 9: 283, 1966.
 371. Dogois, R. K., K. A. Kelly, and C. F. Code. Effect of distal antrectomy on gastric emptying of liquids and solids. Gastroenterology 61: 675–681, 1971.
 372. Cohnheim, O., and F. Best. Ueber Bewegungreflexe des Magendarmkanal. Muench. Med. Wochenschr. 57: 1858–1860, 1910, p. 1858. As usual, my translation from the German is very free.
 373. Morlock, C. G., N. C. Hightower, Jr., C. F. Code, and W. M. Craig. Effect of throacolumbar sympathectomy and splanchnicectomy on gastric motility in man. Gastroenterology 16: 117–121, 1950.
 374. Quigley, J. P., and H. Louckes. The effect of complete vagotomy on the pyloric sphincter and the gastric evacuation mechanism. Gastroenterology 19: 533–537, 1951.
 375. Kelly, K. A., and C. F. Code. Effect of transthoracic vagotomy on canine gastric contractions. Gastroenterology 57: 51–58, 1969.
 376. Mroz, C. T., and K. A. Kelly. The role of extrinsic antral nerves in the regulation of gastric emptying. Surg. Gynecol. Obstet. 145: 369–377, 1977.
 377. Stoddard, C. J., W. E. Waterfall, B. H. Brown, and H. L. Duthie. The effects of varying the extent of vagotomy on the myoelectrical and motor activity of the stomach. Gut 14: 657–664, 1973.
 378. Quigley et al. (n. 243).
 379. Fisher, R., and S. Cohen. Physiological characteristics of the human pyloric sphincter. Gastroenterology 64: 67–75, 1973.
 380. It is only fair to say that Fisher and Cohen were themselves criticized on technical grounds by Irish gastroenterologists who could not find a pressure barrier at the pyloric sphincter. See McShane, A. J., C. O'Morain, J. R. Lennon, J. B. Coakley, and B. G. Alton. Atraumatic non‐distorting pyloric sphincter pressure. Gut 21: 826–828, 1908.
 381. Fisher, R. S., and G. Boden. Gastrin inhibition of pyloric sphincter. Am. J. Dig. Dis. 21: 468–472, 1976.
 382. Fisher, R. S., W. Lipschutz, and S. Cohen. The hormonal regulation of pyloric sphincter function in man. J. Clin. Invest. 52: 1289–1296, 1973;
 383. Fisher and Boden (n. 270).
 384. For reports of a similar increase in dogs, see Isenberg, J. I., and A. Csendes. Effect of octapeptide of cholecystokinin on canine pyloric pressure. Am. J. Physiol. 222: 428–431, 1972.
 385. Ludwig, C. Lehrbuch der Physiologie des Menschens. Heidelberg, Germany: Winter, 1861, vol. II, p. 615.
 386. Van Braam Houckgeest, [J. P.]. Untersuchungen über Peristaltik des Magens und Darmkanal. Pfluegers Arch. 6: 266–302, 1872.
 387. Cannon, W. B. The movements of the intestines studied by means of the Röntgen rays. Am. J. Physiol. 6: 251–277, 1902;
 388. Cannon, W. B. (n. 8), p. 130–163.
 389. Cannon (n. 180).
 390. Hertz, A. F. The passage of food along the human alimentary canal. Guy's Hosp. Reports 61: 389–427, 1907.
 391. There is a biographical sketch in Hurst, A. F. Selected Writings of Sir Arthur Hurst. London: Br. Soc. Gastroenterol., 1970, p. xi–xiv. Like many other patriotic Englishmen with German names, Hertz changed his during the First World War.
 392. Hukuhara, T. Die normale Dünndarmbewegungen. Pfluegers Arch. 226: 518–524, 1931.
 393. Cannon (n. 8), p. 130–163.
 394. Alvarez, W. C. An Introduction to Gastro‐enterology (4th ed.). New York: Hoeber, 1948.
 395. Starling, E. H. (n. 198).
 396. Bayliss, W. M., and E. H. Starling. The movements and innervation of the small intestine. J. Physiol. Lond. 24: 99–143, 1899, p. 110.
 397. Bayliss and Starling (n. 281), p. 114.
 398. Hukuhara, T., K. Masuda, and S. Kinose. Über das “Gesetz des Darmes.” Pfluegers Arch. 237: 619–630, 1936.
 399. Alvarez (n. 279), p. 39–58.
 400. See Davenport, H. W. Physiology of the Digestive Tract (5th ed.). Chicago, IL: Year Book, 1982, p. 98–100.
 401. Bayliss and Starling (n. 281), p. 106.
 402. Ingelfinger, F. J., and R. E. Moss. The activity of the descending duodenum during nausea. Am. J. Physiol. 136: 561–566, 1942.
 403. Smith, C. C., and K. R. Brizzee. Cineradiographic analysis of vomiting in the cat. Gastroenterology 40: 654–664, 1961.
 404. Weisbrodt, N. W., and J. Christensen. Electrical activity of the cat duodenum in fasting and vomiting. Gastroenterology 63: 1004–1010, 1972.
 405. Code, C. F. The interdigestive housekeeper of the gastrointestinal tract. Perspect. Biol. Med. 22: S49–S55, 1979, p. S49.
 406. Szurszewski, J. H., and C. F. Code. Electrical slow wave gradient of canine small bowel (Abstract). Federation Proc. 27: 449, 1968.
 407. The work was reported in full in Szurszewski, J. H., L. R. Elveback, and C. F. Code. Configuration and frequency gradient of electrical slow wave over canine small bowel. Am. J. Physiol. 218: 1468–1473, 1970.
 408. Reinke, D. A., A. H. Rosenbaum, and D. Bennett. Patterns of dog gastrointestinal contractile activity monitored in vivo with extraluminal force transducers. Am. J. Dig. Dis. 12: 113–141, 1967.
 409. Letter of 8 October 1987 from J. H. Szurszewski to H. W. D., for which much thanks. (Szurszewski's typist misspelled Reinke's name.) The abstract is Szurszewski, J. H., and C. F. Code. Activity fronts of canine small intestine. Gastroenterology 54: 1304, 1968.
 410. For Boldyreff, Cannon and Washburn, and Carlson, see gastric motility and emptying, p. 000. Others: Peustow, C. B. Intestinal Motility in the Dog and Man. Urbana: Univ. of Illinois Press, 1940;
 411. Douglas, D. M., and F. C. Mann. An experimental study of the rhythmic contractions of the small intestine of the dog. Am. J. Dig. Dis. 6: 318–322, 1934;
 412. Douglas, D. M. The activity of the duodenum. J. Physiol. Lond. 107: 472–478, 1941;
 413. Douglas, D. M. The decrease in frequency of contraction of the duodenum after transplantation to the ileum. J. Physiol. Lond. 110: 66–75, 1949;
 414. Foulk, W. T., C. F. Code, C. G. Morlock, and J. A. Bargen. A study of the motility patterns and basic rhythm of the duodenum and upper part of the jejunum in human beings. Gastroenterology 26: 601–611, 1954;
 415. Hiatt, R. B., I. Goodman, and A. Alvai. Hormonal control of intestinal motility. Ann. Surg. 166: 704–711, 1967;
 416. Reinke, D. A., A. H. Rosenbaum, and D. R. Bennett. Patterns of dog gastrointestinal activity monitored in vivo with extraluminal force transducers. Am. J. Dig. Dis. 12: 131–141, 1967;
 417. McCoy, E. J., and R. D. Baker. Effect of feeding on electrical activity of dog's small intestine. Am. J. Physiol. 214: 1291–1295, 1968.
 418. Code (n. 290).
 419. The last sentence of Szurszewski and Code (n. 291).
 420. Code, C. F., and J. F. Schlegel. The gastrointestinal interdigestive housekeeper: motor correlates of the interdigestive myoelectric complex in the dog. In: Fourth International Symposium on Gastrointestinal motility, edited by E. E. Daniel, K. Bowes, J. A. L. Gilbert, B. Schofield, T. K. Schnitka, and G. Scott. Vancouver, Canada: Mitchell, 1974.
 421. Szurszewski, J. H. A migrating electric complex of the canine small intestine. Am. J. Physiol. 217: 1757–1763, 1969.
 422. Marlett, J. A., and C. F. Code. The interdigestive gastrointestinal electric complex (Abstract). Federation Proc. 30: 609, 1971.
 423. Code, C. F., and J. A. Marlett. The interdigestive myoelectric complex of the stomach and small bowel of dogs. J. Physiol. Lond. 246: 289–309, 1975. I am grateful to Professor Marlett for a letter giving some details and a warm appreciation of Charlie Code as a mentor.
 424. Morse, L. M., R. A. Nelson, and C. F. Code. Graduate program in nutrition with clinical emphasis. J. Nutr. Ed. 7: 72–73, 1975.
 425. Example: Stefanik, P. A., M. K. Lavers, H. Smith, and C. F. Code. Effect of thiamine deficiency on secretion of acid by vagally denervated and vagally innervated gastric mucosa (Abstract). Federation Proc. 16: 124, 1957.
 426. See also Buéno, L., and M. [sic] Ruckebusch. Insulin and jejunal electrical activity in dogs and sheep. Am. J. Physiol. 230: 1538–1544, 1976, for illustrations of the transition from the fasting to the fed pattern.
 427. Example: Vantrappen, G. The migrating myoelectric complex. In: Motility of the Digestive Tract, edited by M. Wienbeck. New York: Raven, 1982, p. 157–167.
 428. Examples: Ruckebusch, Y. The electrical activity of the digestive tract as an indication of mechanical events. J. Physiol. Lond. 210: 857–882, 1970;
 429. Ruckebusch, Y., and L. Buéno. Effect of weaning on the motility of the small intestine of the calf. Br. J. Nutr. 30: 41–49, 1973.
 430. Buéno, L., and M. [sic] Ruckebusch (n. 303). When Ruckebusch published in J. Physiol. Lond. he had to quiet the anxieties of the British Home Office by adding a footnote declaring the work had been done in France.
 431. Givel, M.‐L. and Y. Ruckebusch. The propagation of segmental contractions in the small intestine. J. Physiol. Lond. 227: 611–625, 1972.
 432. Buéno, L., J. Fioramonti, and Y. Ruckebusch. Rate of flow of digesta and electrical activity of the small intestine of dogs and sheep. J. Physiol. Lond. 249: 69–85, 1975.
 433. Code and Schlegel (n. 297).
 434. Marik, F., and C. F. Code. Vagal control of the interdigestive myoelectric complex (Abstract). Physiologist 15: 208, 1972.
 435. These observations were more or less, chiefly less, confirmed by Weisbrodt, N. W., E. M. Copeland, E. P. Moore, R. W. Kearly, and L. R. Johnson. Effect of vagotomy on electrical activity of the small intestine of the dog. Am. J. Physiol. 228: 650–654, 1975.
 436. Carlson, G. M., B. S. Bedi, and C. F. Code. Mechanism of propagation of intestinal interdigestive myoelectric complex. Am. J. Physiol 222: 1027–1030, 1972.
 437. Marik, J. G., and C. F. Code. Control of the interdigestive myoelectric activity in dogs by the vagus nerves and pentagastrin. Gastroenterology 69: 387–395, 1975.
 438. Brown, J. C., L. P. Johnson, and D. F. Magee. Effect of duodenal alkalinization on gastric motility. Gastroenterology 50: 333–339, 1966.
 439. Brown, J. C. The presence of a gastric motor stimulating property in duodenal mucosa. Gastroenterology 52: 225–229, 1967;
 440. Brown, J. C., and C. Parkes. The separation of fundic pouch motor activity stimulatory and inhibitory fractions. Gastroenterology 53: 731–736, 1967;
 441. Brown, J. C., V. Mutt, and J. R. Dryburgh. The further purification of motilin, a gastric activity stimulating polypeptide from the mucosa of the small intestine of the hog. Can. J. Physiol. Pharmacol. 49: 399–405, 1971;
 442. Brown, J. C., M. A. Cook, and J. R. Dryburgh. Motilin, a gastric motor activity stimulating polypeptide: the complete amino acid sequence. Can. J. Biochem. 51: 533–537, 1973.
 443. Itoh, Z., S. Takeuchi, I. Aizawa, and R. Takayanagi. Effect of synthetic motilin on gastric motor activity in conscious dogs. Am. J. Dig. Dis. 22: 813–819, 1977.
 444. Ambache, N. The electrical activity of isolated mammalian intestine. J. Physiol. Lond. 106: 139–153, 1947.
 445. Biebl, M. Die Autointoxikation durch die Phenil‐Indolkörpers. Dtsch. Z. Chir. 218: 135–230, 1929, p. 164, where the preparation is called Darmhautschlauchplastik.
 446. Biebl, M. Graphische Darstellung der Darmbewegungen. Klin. Wochenschr. 9: 1674–1675, 1930. The photographs in the latter are labeled in English and were probably made at the Mayo Clinic.
 447. Armstrong, H. I. O., G. W. Milton, and A. W. M. Smith. Electrical changes in the small intestine. J. Physiol. Lond. 131: 147–153, 1956.
 448. Bass, P., C. F. Code, and E. H. Lambert. Motor and electric activity of the duodenum. Am. J. Physiol. 201: 287–291, 1961. Lambert was the clinic's electromyographer who provided the initial electrophysiological expertise.
 449. Monges, H., and J. Salducci. Étude électromyographique de la motricité duodénale chez l'homme normale. Arch. Mal. Appl. Dig. 59: 19–28, 1970.
 450. Waterfall, W. E., H. L. Duthie, and B. H. Brown. The electrical and motor actions of gastrointestinal hormones on the duodenum in man. Gut 14: 689–696, 1973.
 451. See Waterfall, W. E., B. H. Brown, H. L. Duthie, and G. E. Whittaker. The effects of humoral agents on the myoelectrical activity of the terminal ileum. Gut 13: 528–534, 1972, for similar results lower down.
 452. Ling, G., and R. W. Gerard. The normal membrane potential of frog sartorius fibers. J. Cell Comp. Physiol. 34: 383–396, 1949.
 453. Woodbury, L. A., H. H. Hecht, and A. R. Christopherson. Membrane resting and action potentials of single cardiac muscle fibers of the frog ventricle. Am. J. Physiol. 164: 307–318, 1951. The paper describes a method of immobilizing the heart with pressure from a plastic ring. That was used after the unpublished gelatin method was abandoned.
 454. Woodbury, J. W., and A. J. Brady. Intracellular recording from moving tissues with a flexibly mounted ultramicroelectrode. Science Wash. DC 123: 100–101, 1956.
 455. Bortoff, A. Slow potential variations of small intestine. Am. J. Physiol. 201: 203–208, 1961.
 456. Bortoff (n. 324);
 457. Bortoff, A., and N. Weg. Transmission of electrical activity through the gastroduodenal junction. Am. J. Physiol. 208: 531–536, 1965.
 458. Bortoff (n. 324), p. 207.
 459. Bortoff, A. Electrical activity of intestine recorded with pressure electrode. Am. J. Physiol. 201: 209–212, 1961;
 460. Bülbring, E., and H. Kuriyama. Effect of changes in ionic environment on the action of Ach and adrenalin on the smooth muscle cells of the guinea‐pig taenia coli. J. Physiol. Lond. 166: 59–74, 1963.
 461. Bülbring, E., and H. Kuriyama. The effect of adrenalin on the smooth muscle of the guinea‐pig taenia coli in relation to the degree of stretch. J. Physiol. Lond. 169: 198–212, 1963.
 462. For an exhaustive summary of the state of knowledge at the end of our period, see Bülbring, E., and M. F. Shuba (editors). Physiology of Smooth Muscle. New York: Raven, 1976.
 463. Davenport, H. W., and F. Alzamora. Sodium, potassium, chloride, and water in frog gastric mucosa. Am. J. Physiol. 202: 711–715, 1962;
 464. Barr, L., and R. L. Malvin. Estimation of extracellular space of smooth muscle using different sized molecules. Am. J. Physiol. 208: 1042–1045, 1965.
 465. Bülbring, E. et al. (n. 200), p. 119.
 466. Burnstock, G. The action of adrenaline on the excitability and membrane potential in the taenia coli of the guinea pig and the effect of DNP on this action and on the action of acetylcholine. J. Physiol. Lond. 143: 183–194, 1958. The word Adrenalin is a registered trademark of Parke, Davis, and in the United States the generic name almost always used in scientific publications is epinephrine. European authors frequently call the compound adrenalin or adrenaline. Except when quoting them directly, I have translated the European into the American term.
 467. Axelsson, J., E. Beuding, and E. Bülbring. The action of adrenaline on phosphorylase activity and membrane potential in smooth muscle. J. Physiol. Lond. 148: 62P–63P, 1959;
 468. Axelsson, J., and E. Bülbring. Metabolic factors affecting electrical activity of intestinal smooth muscle. J. Physiol. Lond. 156: 344–356, 1961;
 469. Axelsson, J., E. Beuding, and E. Bülbring. The inhibitory action of adrenaline on intestinal smooth muscle in relation to its action on phosphorylase activity. J. Physiol. Lond. 156: 357–374, 1961.
 470. Dewey, M. M., and L. Barr. Intercellular connection between smooth muscle cells: the nexus. Science Wash. DC 137: 670–672, 1962;
 471. Dewey, M. M., and L. Barr. Anat. Rec. 151: 343, 1965;
 472. Dewey, M. M., and L. Barr. A study of the structure and distribution of the nexus. J. Cell Biol. 23: 553–585, 1964;
 473. Dewey, M. M., and L. Barr. Structure of vertebrate intestinal smooth muscle. In: Handbook of Physiology. Alimentary Canal, edited by C. F. Code. Washington, DC: Am. Physiol. Soc., 1968, sect. 6, vol. IV, chapt. 81, p. 1629–1654.
 474. Lane, B., and J. A. G. Rhodin. Cellular interrelationships and electrical activity in two types of smooth muscle. J. Ultrastruct. Res. 10: 470–488, 1964.
 475. Barr, L. Propagation in vertebrate smooth muscle. J. Theor. Biol. 4: 73–85, 1963.
 476. Henderson, R. M., G. Duchon, and E. E. Daniel. Cell contacts in duodenal smooth muscle layers. Am. J. Physiol. 221: 564–574, 1971;
 477. Gabella, G. Intercellular junctions between circular and longitudinal intestinal muscle layers. Cell Tissue Res. 125: 191–195, 1972.
 478. For discussion of cable conduction in smooth muscle without low‐resistance pathways, see Abe, Y., and T. Tomita. Cable properties of smooth muscle. J. Physiol. Lond. 196: 87–100, 1968.
 479. Kobayashi, M., T. Nagai, and C. L. Prosser. Electrical interaction between muscle layers of cat intestine. Am. J. Physiol. 211: 1281–1291, 1966.
 480. Burnstock, G., and C. L. Prosser. Conduction in smooth muscles: comparative electrical properties. Am. J. Physiol. 199: 553–559, 1960.
 481. Nagai, T., and C. L. Prosser. Patterns of conduction in smooth muscle. Am. J. Physiol. 204: 910–914, 1963;
 482. Nagai, T., and C. L. Prosser. Electrical parameters of smooth muscle cells. Am. J. Physiol. 204: 915–924, 1963;
 483. Kobayashi, M., C. L. Prosser, and T. Nagai. Electrical properties of intestinal muscle as measured intracellularly and extracellularly. Am. J. Physiol. 213: 275–286, 1967.
 484. Brune, H. F., and H. Kolowski. Die Erregungsleitung in der glatten muskulatur des Meerschwinschen‐Dickdarmes. Pfluegers Arch. 262: 484–493, 1951.
 485. Bortoff, A. Electrical transmission of slow waves from longitudinal to circular intestinal muscle. Am. J. Physiol. 209: 1254–1260, 1965.
 486. Kobayashi et al. (n. 341).
 487. Daniel, E. E., D. R. Carlow, B. T. Wachter, W. H. Sutherland, and A. Bogoch. Electrical activity in the small intestine. Gastroenterology 37: 268–280, 1959.
 488. Daniel, E. E., and K. M. Chapman. Electrical activity of the gastrointestinal tract as an indicator of mechanical activity. Am. J. Dig. Dis. 8: 54–102, 1963, p. 73; Fig. , p. 75.
 489. Bortoff, A., and N. Weg. Transmission of electrical activity through the gastroduodenal junction. Am. J. Physiol. 208: 531–536, 1965.
 490. Bortoff, A., and R. S. Davis. Myogenic transmission of antral slow waves across the gastroduodenal junction in situ. Am. J. Physiol. 215: 889–897, 1968.
 491. Horton, B. T. Pyloric block, with special reference to the musculature, myenteric plexus and lymphatic vessels. Arch. Surg. 22: 438–462, 1931.
 492. See also Rash, R. M., and M. D. Thomas. The intrinsic innervation of the gastro‐oesophageal and pyloro‐duodenal junction. J. Anat. 96: 389–391, 1962.
 493. Bortoff and Davis (n. 348).
 494. Bass et al. (n. 318).
 495. Bedi, B. S., and C. F. Code. Pathway of coordination of postprandial, antral, and duodenal action potentials. Am. J. Physiol. 222: 1295–1298, 1972.
 496. For the sake of completeness it must be said that when Code recorded antral and duodenal activity by means of chronically implanted electrodes he could find no temporal relation between the slow waves of the antrum and those of the duodenum. See Allen, G. L., E. W. Poole, and C. F. Code. Relationship between electrical activity of antrum and duodenum. Am. J. Physiol. 207: 906–910, 1964. Daniel's and Code's contradictions might have been caused by relative insensitivity of their electrodes.
 497. Waterfal et al. (n. 320);
 498. Duthie, H. L., B. H. Brown, B. Robertson‐Dunn, N. H. Kwong, G. E. Whittaker, and W. Waterfall. Electrical activity in gastroduodenal area—slow waves in proximal duodenum. A comparison of man and dog. Am. J. Dig. Dis. 17: 344–351, 1972.
 499. Alvarez, W. C. Functional variations in contractions of different parts of the small intestine. Am. J. Physiol. 35: 177–193, 1914.
 500. Alvarez, W. C., and E. Starkweather. XI. The metabolic gradient underlying intestinal peristalsis. Am. J. Physiol. 46: 186–208, 1918.
 501. Alvarez (n. 279), p. 1–175.
 502. Luciani, L. Human Physiology (transl. by F. A. Welby). London: Macmillan, 1913, vol. II, p. 236–237.
 503. Figures from Alvarez, W. C. Further studies on intestinal rhythm. Am. J. Physiol. 37: 267–281, 1915.
 504. Castleton, K. B. An experimental study of the movements of the small intestine. Am. J. Physiol. 107: 641–646, 1934.
 505. Douglas and Mann (n. 294).
 506. Alvarez, W. C., and L. J. Mahoney. Action currents in stomach and intestine. Am. J. Physiol. 58: 476–493, 1922.
 507. Diamant, N. E., and A. Bortoff. Effects of transection on the intestinal slow‐wave frequency gradient. Am. J. Physiol. 216: 734–743, 1969.
 508. Szurszewski, Elveback, and Code (n. 291);
 509. Code, C. F., and J. H. Szurszewski. The effect of duodenal and small bowel transection on the frequency gradient of the pacesetter potential in the canine intestine. J. Physiol. Lond. 207: 281–289, 1970;
 510. Hermon‐Taylor, J., and C. F. Code. Localization of the duodenal pacemaker and its role in the organization of duodenal myoelectric activity. Gut 12: 40–47, 1971.
 511. Examples of the large literature are Nelsen, T. S., and J. C. Becker. Simulation of the electrical and mechanical gradient of the small intestine. Am. J. Physiol. 214: 749–757, 1968;
 512. Sarna, S. K., E. E. Daniel, and Y. Kingman. Simulation of slow‐wave electrical activity of small intestine. Am. J. Physiol. 221: 166–175, 1971;
 513. Brown, B. H., H. L. Duthie, A. R. Horn, and R. H. A. Smallwood. A linked oscillator model of electrical activity of human small intestine. Am. J. Physiol. 229: 384–388, 1975.
 514. Langley, J. N. Observations on the physiological action of extracts of the suprarenal bodies. J. Physiol. Lond. 27: 237–256, 1901.
 515. For the central nervous system, see text and figures in Gerlach, J. Von dem Rückenmark. In: Handbuch der Lehre von den Geweben, edited by S. Stricker. Leipzig, Germany: Wilhelm Engelmann, 1872, vol. II, p. 665–693.
 516. Meissner, G. Über die Nerven der Darmwand. Z. Rat. Med. 8: 364–396, 1857;
 517. Auerbach, L. Fernere vorläufe Mitteilung über der Nervenapparat des Darmes. Virchow's Arch. 30: 456–460, 1864. The earlier Mitteilung published in pamphlet form in Breslau in 1862 was not available to me.
 518. Billroth, T. Einige Beobachtungen über ausgedehnte Vorkommen von Nervenanastomosen im Tractus intestinalis. Arch. Anat. Physiol. Wissensch. Med. 148–158, 1858.
 519. For a summary, see the historical introduction to Hillarp, N.‐Å. Structure of the synapse and the peripheral innervation of the autonomic nervous system. Acta Anat. Suppl. IV, 1946.
 520. Mitchell, G. A. G. Anatomy of the Autonomic Nervous System. Edinburgh, UK: Livingstone, 1953, p. 53.
 521. For a summary, see Schofield, G. C. Anatomy of muscular and neural tissues in the alimentary canal. In: Handbook of Physiology. Alimentary Canal, edited by C. F. Code. Washington, DC: Am. Physiol. Soc., 1968, sect. 6, vol. IV, chapt. 80, p. 1579–1627.
 522. For enunciation of the neuron doctrine, see Waldeyer, W. Ueber einige neure Forschung im Gebiete der Anatomie des Centralnervensystems. Dtsch. Med. Wochenschr. 17: 1213–1218, 1244–1246, 1267–1289, 1331–1332, 1352–1356, 1891. See in particular the diagrams in the last section of the paper.
 523. Sherrington, C. S. Tremor, “tendon phenomenon,” and spasm. In: A System of Medicine, edited by T. C. Allbutt. New York: Macmillan, 1899, vol. VI, p. 512.
 524. Wood, J. D. Electrical discharge of single enteric neurons of guinea pig small intestine. Am. J. Physiol. 225: 1107–1113, 1973.
 525. Gaskell, W. H. On the structure, distribution and function of the nerves which innervate the visceral and vascular systems. J. Physiol. Lond. 7: 1–80, 1886.
 526. Fletcher, W. M. John Newport Langley, in memoriam. J. Physiol. Lond. 61: 1–15, 1926. Fletcher's sympathetic appreciation contains a useful bibliography.
 527. Langley, J. N. The Autonomic Nervous System. Pt. 1. Cambridge, UK: Heffer, 1921. There is no Part 2. Many persons did not like Langley, and alluding to his position as an international expert on figure skating they said he exhibited perfect technique on an icy background.
 528. Langley, J. N., and W. L. Dickinson. Pituri and nicotin. J. Physiol. Lond. 11: 265–306, 1890.
 529. Langley, J. N., and W. L. Dickinson. On the local paralysis in peripheral ganglia and on the connexion of different classes of fibres with them. Proc. R. Soc. Lond. 46: 423–431, 1889;
 530. Langley, J. N. On the physiology of salivary secretion. J. Physiol. Lond. 11: 123–133, 1890.
 531. Langley, J. N. On the stimulation and paralysis of nerve cells and of nerve endings. J. Physiol. Lond. 11: 224–236, 1901.
 532. Langley, J. N., and H. K. Anderson. Position of nerve cells on the course of efferent nerve fibres. J. Physiol. Lond. 19: 131–139, 1895.
 533. Examples: Kuntz, A. On the occurrence of reflex arcs in the myenteric and submucous plexuses. Anat. Rec. 24: 193–210, 1922, p. 200;
 534. McSwiney, B. A. Innervation of the stomach. Physiol. Rev. 11: 478–514, 1931;
 535. Ranson, S. W. The Anatomy of the Nervous System (9th ed.). Philadelphia, PA: Saunders, 1953, p. 384.
 536. Langley, J. N. Das sympatische und verwandt nervöse System. Ergeb. Physiol. 2: 818–872, 1903.
 537. Kerwenter, J. The vagal control of jejunal and ileal motility and blood flow. Acta. Physiol. Scand. Suppl. 251, 1965. Kerwenter did find high‐threshold inhibitory fibers in the subdiaphragmatic branches of the vagus (cf., Openchowski), and these are mentioned later.
 538. Pflüger, E. F. W. Ueber das Hemmungs‐Nervensystem für die peristaltischen Bewegungen der Gedärme. Berlin: Hirsch‐wald, 1857. I have not seen this; it is quoted by Starling (n. 198).
 539. Lister, J. Preliminary account of an inquiry into the functions of the visceral nerves, with special reference to the so‐called “inhibitory system.” Proc. R. Soc. Lond. 9: 367–388, 1858. Lister's paper is a confused exposition of his idea that “the intestines possess an intrinsic ganglionic apparatus which is in all cases essential to the peristaltic movement, and, while capable of independent action, is liable to be stimulated or checked by the other parts of the nervous system.…” Lister thought that “the same nerve‐fibres, while working more mildly, produce an increase of function” and are the inhibitors. He deliberately did not use chloroform in his acute experiments.
 540. Bayliss and Starling (n. 281).
 541. Cannon, W. B. Some practical applicaitons of recent studies in the physiology of the digestive system. Wis. Med. J. 7: 223–242, 1908.
 542. Cannon, W. B., and F. T. Murphy. The movements of the stomach and intestines in some surgical conditions. Ann. Surg. 43: 512–536, 1906;
 543. Cannon, W. B., and F. T. Murphy. Physiologic observations on experimentally produced ileus. J. Am. Med. Assoc. 49: 840–843, 1907.
 544. Pearcy, J. F., and E. J. Van Liere. Studies on the visceral nervous system. XVII. Reflexes from the colon. 1. Reflexes to the stomach. Am. J. Physiol. 78: 64–73, 1926.
 545. Herrin, R. C., and W. J. Meek. Distention as a factor in intestinal obstruction. Arch. Intern. Med. 51: 152–168, 1933.
 546. Hermann, H., and G. Morin. Mise en évidence d'un réflex inhibiteur intestino‐intestinale. C. R. Soc. Biol. Paris 115: 529–531, 1934.
 547. Hermann and Morin (n. 392);
 548. Morin, G., and J. Vial. Sur les voies et les centres du réflex inhibiteur intestino‐intestinale. C. R. Soc. Biol. Paris 116: 536–538, 1934.
 549. Youmans, W. B., W. J. Meek, and R. C. Herrin. Extrinsic and intrinsic pathways concerned with intestinal inhibition during intestinal distention. Am. J. Physiol. 124: 470–477, 1938;
 550. Youmans, W. B., A. I. Karstens, and K. W. Aumann. Nervous pathways for the reflex regulation of intestinal pressure. Am. J. Physiol. 135: 619–627, 1942.
 551. Hill, C. J. A contribution to our knowledge of the enteric plexuses. Philos. Trans. R. Soc. Lond. B. Biol. Sci. 215: 355–387, 1927. The paper was contributed by “Prof. J. P. Hill, F. R. S.” Professor James P. Hill was professor of embryology at University College London, and he is listed in Who's Who as having “2 ds.”
 552. Schofield, G. C. Experimental studies on the myenteric plexus in mammals. J. Comp. Neurol. 119: 159–185, 1962.
 553. Dale, H. H. Nomenclature of fibres in the autonomic system and their effects. Nature Lond. 80: 10–11, 1933.
 554. Koelle, G. B., and J. Friedenwald. A histochemical method for localizing cholinesterase activity. Proc. Soc. Exp. Biol. Med. 70: 617–620, 1949.
 555. Example: Taxi, J. La distribution des cholinesterases dans divers ganglions du système nerveux autonome des vertébrés. In: Histochemistry of Cholinesterase, edited by H. G. Schwarzacher. Basel: Karger, 1961, vol. II, p. 73–89.
 556. The status as of 1974 is reviewed in Furness, J. B., and M. Costa. The adrenergic innervation of the gastrointestinal tract. Ergeb. Physiol. 69: 1–51, 1974.
 557. Coujard, R. Récherches sur les plexuse nerveux de l'intestine. Arch. Anat. Microsc. Morphol. Exp. 39: 110–151, 1950.
 558. Example: Euler, U. S. von. The presence of the adrenergic neurotransmitter in the intra‐axonal structure. Acta. Physiol. Scand. 57: 468–480, 1963.
 559. Marks, B. H., T. Samoyaski, and E. J. Webster. Radioautographic localization of norepinephrine‐H3 in the tissues of mice. J. Pharmacol. Exp. Ther. 138: 376–381, 1962.
 560. Falck, B. Observations on the possibilities of the cellular localization of monoamines by the fluorescence method. Acta Physiol. Scand Suppl. 197, 1962.
 561. Norberg, K.‐A. Adrenergic innervation of the intestinal wall studied by means of fluorescence microscopy. Neuropharmacology 3: 379–382, 1964;
 562. Jacobowitz, D. Histochemical studies of the autonomic innervation of the gut. J. Pharmacol. Exp. Ther. 149: 358–364, 1965.
 563. For a personal account, see Page, I. H. Serotonin. Chicago, IL: Year Book, 1968.
 564. Erspamer, V., and B. Asero. Identification of enteramine, specific hormone of enterochromaffine cell system, as 5‐hydroxytryptamine. Nature Lond. 169: 800–801, 1952.
 565. Haverback, B. J., C. A. M. Hogben, M. C. Moran, and L. L. Terry. Effect of serotonin (5‐hydroxytryptamine) and related compounds on gastric secretion and intestinal motility in dogs. Gastroenterology 32: 1058–1060, 1957.
 566. Bülbring, E., and A. Crema. The release of 5‐hydroxytryptamine in relation to pressure exerted on the intestinal mucosa. J. Physiol. Lond. 146: 18–28, 1959.
 567. Bülbring, E., and A. Crema. Observations concerning the action of 5‐hydroxytryptamine on the peristaltic reflex. Br. J. Pharmacol. 13: 444–457, 1958.
 568. Gershon, M. D., and L. L. Ross. Radioisotopic studies of the localization of 5‐hydroxytryptamine (Abstract). J. Histochem. Cytochem. 12: 19, 1964;
 569. Gershon, M. D., A. B. Drakontides, and L. L. Ross. Serotonin: synthesis and release from the myenteric plexus of the mouse intestine. Science Wash. DC 149: 197–199, 1965.
 570. Jacobj, C. Beiträge zur physiologischen und pharmakologischen Kenntniss der Darmbewegungen mit besonderer Berüksichtung der Beziehung der Nebenniere zu deselben. Naunyn‐Schmiedebergs Arch. Pharmacol. Pathol. 29: 171–211, 1891.
 571. Langley (n. 365).
 572. Kock, N. G. An experimental study of the mechanisms engaged in reflex inhibition of intestinal motility. Acta Physiol Scand. Suppl. 164, 1938.
 573. For review, see Folkow, B. Nervous control of the blood vessels. Physiol. Rev. 35: 629–663, 1955.
 574. Example: Celander, O. Are there any centrally controlled sympathetic inhibitory fibres to the musculature of the intestine? Acta Physiol. Scand. 47: 299–309, 1959.
 575. Furness, J. B., and G. Burnstock. Role of circulating catecholamines in the gastrointestinal tract. In: Handbook of Physiology. Endocrinology, edited by R. O. Greep and E. B. Astwood. Washington, DC: Am. Physiol. Soc., 1975, sect. 7, vol. VI, chapt. 33, p. 515–536.
 576. Gershon, M. D. Effects of tetrodotoxin on innervated smooth muscle preparations. Br. J. Pharmacol. 29: 259–279, 1967;
 577. Kao, C. Y. Tetrodotoxin, saxitoxin and their significance in the study of excitation phenomena. Pharmacol. Rev. 18: 997–1049, 1968.
 578. Burnstock, G., G. Campbell, D. G. Satchell, and A. Smythe. Evidence that adenosine triphosphate or related nucleotide is the transmitter substance released by non‐adrenergic inhibitory nerves in the gut. Br. J. Pharmacol. 40: 668–686, 1970.
 579. Magnus, R. Wirkungweise und Angriffspunkt einiger Gifte am Katzendarm. Pfluegers Arch. 102: 349–363, 1908.
 580. Magnus's work with references is summarized in Magnus, R.. Die Bewegungen der Verdauunsrhores. Ergeb. Physiol. 7: 27–64, 1908.
 581. Alvarez (n. 279), p. 185.
 582. Gunn, J. A., and S. W. F. Underhill. Surviving mammalian intestine. Q. J. Exp. Physiol. 8: 275–296, 1914.
 583. Bayliss and Starling (n. 281), p. 115.
 584. Cannon, W. B., and I. R. Burket. The endurance of anemia by nerve cells in the myenteric plexus. Am. J. Physiol. 32: 347–357, 1913.
 585. Hukuhara, T., T. Sumi, and S. Kotani. The role of ganglion cells of the small intestine taken in the intestinal intrinsic reflex. Jpn. J. Physiol. 11: 281–288, 1961.
 586. Trendelenberg, P. Physiologische und pharmakologische Versuche über die Dünndarmperistaltik. Naunyn‐Schmiedeberg's Arch. Pharmacol. 81: 55–129, 1917.
 587. For an introduction to Kosterlitz's work, see Kosterlitz, H. W. Intrinsic and extrinsic nervous control of motility of the stomach and the intestines. In: Handbook of Physiology. Alimentary Canal, edited by C. F. Code. Washington, DC: Am. Physiol. Soc., 1968, sect. 6, vol. IV, chapt. 104, p. 2147–2171.
 588. Hukuhara, T., and H. Fukuda. The motility of isolated guinea‐pig small intestine. Jpn. J. Physiol. 15: 125–139, 1965;
 589. Bortoff, A., and F. Sachs. Electrotonic spread of slow waves in circular muscle of small intestine. Am. J. Physiol. 218: 576–581, 1970;
 590. Bortoff, A., and G. Ghalib. Temporal relationship between electrical and mechanical activity of longitudinal and circular muscle during intestinal peristalsis. Am. J. Dig. Dis. 17: 317–325, 1972.
 591. McKirdy, H. C. Functional relationship of longitudinal and circular layers of muscularis externa of the rabbit large intestine. J. Physiol. Lond. 227: 839–853, 1972.
 592. Jacoby, H. T., P. Bass, and D. B. Bennett. In vivo extraluminal contractile force transducer for gastrointestinal muscle. J. Appl. Physiol. 18: 658–665, 1967.
 593. Tasaka, K., and J. T. Farrar. Mechanics of small intestinal muscle function in the dog. Am. J. Physiol. 217: 1224–1229, 1969.
 594. Bülbring, E., R. C. Y. Lin, and G. C. Schofield. An investigation of the peristaltic reflex in relation to anatomical observations. Q. J. Exp. Physiol. 43: 26–37, 1958.
 595. Bülbring et al. (n. 432);
 596. see also Schofield (n. 371).
 597. Berkely, H. J. The nerves and nerve endings of the mucous layer of the ileum, as shown by the rapid Golgi method. Anat. Anz. 8: 12–19, 1893.
 598. Yokoyoma, S. Aktionpotentiale der Ganglionzeller der Auerbachschen plexus in kanisnschendünndarm. Pfluegers Arch. 288: 95–102, 1966.
 599. Wood, J. D. Electrical activity from single neurons in Auerbach's plexus. Am. J. Physiol. 219: 159–169, 1970;
 600. Wood, J. D. Electrical discharge of single enteric neurons of guinea pig small intestine. Am. J. Physiol. 225: 1107–1113, 1973;
 601. Wood, J. D., and C. J. Mayer. Pattern of discharges of six different neurones of the intestine of the cat, dog, and guinea pig. Pfluegers Arch. 338: 247–256, 1973.
 602. Wood's work is summarized in Wood, J. D. Neurophysiology of ganglion of Auerbach's plexus. Am. Zool. 14: 973–989, 1974;
 603. and Wood, J. D., Physiol. Rev. (n. 54). Wood was a student of Prosser, but because his most important work was done after leaving Urbana I have cited him separately.
 604. See also Ohkawa, H., and C. L. Prosser. Electrical activity in myenteric and submucous plexuses of cat intestine. Am. J. Physiol. 222: 1412–1419, 1972;
 605. and Ohkawa, H., and C. L. Prosser. Functions of neurons in enteric plexuses of cat intestine. Am. J. Physiol. 1420–1426.
 606. Wood, Physiol. Rev. (n. 436).
 607. Jansson, G., and J. Mårtinson. Studies on the ganglionic site of action of sympathetic outflow to the stomach. Acta Physiol. Scand. 68: 184–192, 1966.
 608. Hirst, G. D. S., and H. C. McKirdy. Presynaptic inhibition at mammalian peripheral synapse? Nature Lond. 250: 430–431, 1974;
 609. Hirst, G. D. S., and H. C. McKirdy. Synaptic potentials recorded from neurones of the submucous plexus of guinea‐pig small intestine. J. Physiol. Lond. 249: 369–385, 1975.
 610. Hirst, G. D. S., and H. L. McKirdy. A nervous mechanism for descending inhibition in guinea‐pig small intestine. J. Physiol. Lond. 238: 129–143, 1974.
 611. Hirst, G. D. H., M. E. Holman, and H. C. McKirdy. Two descending nerve pathways activated by distention of guinea‐pig small intestine. J. Physiol. Lond. 244: 113–127, 1975.
 612. Paton, W. D. M., and E. S. Vizi. The inhibitory action of noradrenaline and adrenaline of acetylcholine output by guinea‐pig ileum longitudinal strips. Br. J. Pharmacol. 35: 10–28, 1969.
 613. Beani, C., C. Bianchi, and A. Creama. The effect of catecholamines and sympathetic stimulation on the release of acetylcholine by the guinea‐pig colon. Br. J. Pharmacol. 36: 1–17, 1969.
 614. Schlesinger, M. L. Snatched From Oblivion; A Cambridge Memoir. Boston, MA: Little, Brown, 1979. This memoir by Cannon's daughter gives a somewhat appalling view of his home life.
 615. Alvarez, W. C. An Introduction to Gastro‐enterology. New York: Hoeber, 1948, p. 569.
 616. Cannon, W. B. Am. J. Physiol. (n. 274).
 617. See Lane, W. A. The Operative Treatment of Chronic Intestinal Stasis (3rd ed.). London: Nisbet, 1915, p. 51. This is not the place to discuss Lane's irresistible urge to excise every colon in Great Britain, but it is worth recording that Lane's biographer quotes a letter from Bernard Shaw categorically denying the widespread rumor that Lane was the model for the surgeon in The Doctor's Dilemma.
 618. See Layton, W. B. Sir William Arbuthnot Lane, Bt., C.B., M.S. Edinburgh: Livingstone, 1956, p. 97–98.
 619. Elliott, T. R., and E. Barclay‐Smith. Antiperistalsis and the muscular activities of the colon. J. Physiol. Lond. 31: 272–304, 1904. Barclay‐Smith's horror of the human colon is recorded in Lane (n. 446).
 620. Jacobj, C. Phamakologische Untersuchungen über des Colchigumgift. Naunyn‐Schmiedebergs Arch. Pharmacol. Pathol. 27: 119–157, 1890, p. 147.
 621. Barcroft, J., and F. R. Steggerda. Observations on the proximal portion of exteriorized colon. J. Physiol. Lond. 76: 460–471, 1932.
 622. Hertz, A. F. The passage of food along the human alimentary canal. Guy's Hosp. Rep. 61: 389–427, 1907;
 623. Hurst, A. F. The sphincters of the alimentary canal and their clinical significance. Br. Med. J. i: 145–151, 1925.
 624. Lawson, L. M., and J. A. Bargen. Physiology of the colon. Arch. Surg. 27: 1–50, 1933. This review would be more useful if its bibliographic citations were more complete.
 625. Elliott, T. R. On the innervation of the ileocolic sphincter. J. Physiol. Lond. 31: 157–168, 1904.
 626. Elliott did not cite the just‐published book describing the junction in 112 species of fishes, amphibia, reptile, birds, and mammals: Huntington, G. S. The Anatomy of the Human Peritoneum and Abdominal Cavity. Philadelphia, PA: Lea Bros., 1903.
 627. Bauhin, C. Theatrum Anatomicum. Frankfurt, Germany: Becker, 1605.
 628. Crooke, H. Mikrokosmographia. London: Jaggard, 1615, p. 166–167.
 629. Modern historians do not have Crooke's confidence in Laurentius. Cecelia Mettler said Laurentius's book published in 1595 was poorly written, badly illustrated, and ignorant of the work of Laurentius's predecessors. Mettler, C. C. History of Medicine. Philadelphia, PA: Blakiston, 1947, p. 49–50.
 630. Grützner, P. Ueber die Bewegungen des Darminhaltes. Pfluegers Arch. 71: 492–522, 1895.
 631. Truelove, S. C. Movements of the large intestine. Physiol. Rev. 46: 457–512, 1966.
 632. Langley, J. N., and H. K. Anderson. On the innervation of the pelvic and abdominal viscera. J. Physiol. Lond. 18: 67–105, 1895.
 633. Elliott, T. R. On the action of adrenalin. J. Physiol. Lond. 31: xx–xxi, 1904.
 634. Elliott (n. 452).
 635. Hinrichsen, J., and A. C. Ivy. Studies on the ileo‐cecal sphincter of the dog. Am. J. Physiol. 96: 494–507, 1931.
 636. Examples: White, H. L., W. R. Rainey, B. Monoghan, and A. S. Harris. Observations on the nervous control of the ileocolic sphincter and on intestinal movements in an unanesthetized human subject. Am. J. Physiol. 108: 449–457, 1934;
 637. Gazet, J. C., and R. J. Jarrett. The ileocoloic sphincter. Studies in vitro in man, monkey, cat, and dog. Br. J. Surg. 51: 368–370, 1964;
 638. Singleton, A. O., D. C. Redmond, and J. E. McMurray. Ileocecal resection and small bowel transit and absorption. Ann Surg. 159: 690–694, 1954;
 639. Kelley, M. L., Jr., E. A. Gordon, and J. A. De Weese. Pressure studies of the ileocolonic junctional zone of dogs. Am. J. Physiol. 209: 333–339, 1965;
 640. Kelley, M. L., Jr., E. A. Gordon, and J. A. De Weese. Pressure responses of canine ileocolonic junctional zone to intestinal distention. Am. J. Physiol. 211: 614–618, 1966;
 641. Jarrett, R. J., and J. C. Gazet. Studies in vivo of the ileocaecal sphincter in the cat and dog. Gut 7: 271–275, 1966;
 642. Cohen, S., L. D. Harris, and R. Levitan. Manometric characteristics of the human ileocecal junctional zone. Gastroenterology 54: 72–75, 1968;
 643. Kelley, M. A., Jr., and J. A. De Weese. Effects of eating and intraluminal filling on ileocolonic junctional zone pressures. Am. J. Physiol. 216: 1491–1495, 1969;
 644. Conklin, J. L., and J. Christensen. Local specialization at ileocecal junction of the cat and opossum. Am. J. Physiol. 228: 1075–1081, 1975;
 645. Pahlin, P.‐E., and J. Kewenter. Reflexogenic contraction of the ileocolic sphincter in the cat following small or large intestinal distention. Acta Physiol. Scand. 95: 126–132, 1975;
 646. Pahlin, P.‐E., and J. Kewenter. The vagal control of the ileocaecal sphincter in the cat. Acta Physiol. Scand. 96: 433–442, 1975;
 647. Pahlin, P.‐E., and J. Kewenter. Sympathetic nervous control of cat ileocecal sphincter. Am. J. Physiol. 231: 296–305, 1976. 462.
 648. Starling, E. H. (n. 198).
 649. Bayliss, W. M., and E. H. Starling. The movements and innervation of the large intestine. J. Physiol. Lond. 26: 107–118, 1900.
 650. Templeton, R. D., and H. Lawson. Studies in the motor activity of the large intestine. I. Normal motility in the dog, recorded by the tandem balloon method. Am. J. Physiol. 96: 667–676, 1931.
 651. Adler, H. F., A. J. Atkinson, and A. C. Ivy. A study of the motility of the human colon. Am. J. Dig. Dis. 8: 197–204, 1941;
 652. Adler, H. F., A. J. Atkinson, and A. C. Ivy. Motility of the human colon. J. Am. Med. Assoc. 121: 646–651, 1943.
 653. Spriggs, E. A., C. F. Code, J. A. Bargen, R. K. Curtiss, and N. C. Hightower, Jr. Motility of the pelvic colon and rectum of normal persons and patients with ulcerative colitis. Gastroenterology 19: 480–487, 1951;
 654. Posey, E. L., Jr., and J. A. Bargen. Observations of normal and abnormal intestinal function. Am. J. Med. Sci. 221: 10–17, 1951.
 655. For a summary as of 1966, see Truelove (n. 456).
 656. Connell, A. M. Motility of the pelvic colon. II. Paradoxical motility in diarrhoea and constipation. Gut 3: 342–348, 1962.
 657. Waller, S. L., J. J. Misiewicz, and N. Kiley. Effect of eating on motility of the pelvic colon in constipation and diarrhoea. Gut 13: 805–810, 1972.
 658. Aub, J. C., B. Wolbach, B. J. Kennedy, and O. T. Bailey. Mycosis fungoides followed for 14 years;
 659. the case of W. B. Cannon. AMA Arch. Pathol. 60: 535–547, 1955.
 660. Cannon (n. 445).
 661. Holzknecht, G. Die normale Persistatlik des Kolon. Muench. Med. Wochenschr. 47: 2401–2403, 1909.
 662. Barclay, A. E. Note on the movements of the large intestine. Br. J. Roentgenol. 16: 422–424, 1912.
 663. Hertz, A. F. The ileo‐caecal sphincter. J. Physiol. Lond. 47: 54–56, 1913;
 664. Hertz, A. F., and A. Newton. The normal movements of the colon in man. J. Physiol. Lond. 47: 57–65, 1913.
 665. Groedel, F. M. (n. 149), p. 500–640, which contains figures reproduced from a book published by Groedel in 1912.
 666. Scharz, G. Zur Physiologie und Pathologie der menschlichen Dickdarmbewegungen. Muench. Med. Wochenschr. 58: 1489–1494, 1911.
 667. Luboshez, B. E. Une méthode pratique de cinéradiographie. Paris Med. 71: 117–118, 1929;
 668. Luboshez, B. E. Cineradiographia. Radiol. Med. 18: 450–457, 1931. The quotation is from the second paper.
 669. Apparently the only description of this project is in Gianturco, C., and W. C. Alvarez. Roentgen motion pictures of the stomach. Proc. Staff Meet. Mayo Clin. 7: 669–671, 1932.
 670. See Gianturco, C. Some mechanical factors in gastric physiology. Am. J. Roentgenol. 31: 735–744, 745–750, 1934.
 671. Barclay, A. E. Direct x‐ray cinematography with a preliminary note of non‐propulsive movements of the large intestine. Br. J. Radiol. 8: 652–658, 1935.
 672. Chaudhary, N. A., and S. C. Truelove. Human colonic motility: a comparative study of normal subjects, patients with ulcerative colitis and patients with irritable colon syndrome. I. Resting patterns of motility. Gastroenterology 40: 1–17, 1966.
 673. Truelove, S. C. Movements of the large intestine. Physiol. Rev. 46: 457–512, 1966.
 674. Ritchie, J. A., G. M. Ardran, and S. C. Truelove. Motor activity of the sigmoid colon in humans. Gastroenterology 43: 642–668, 1962.
 675. Ritchie, J. A. Colonic activity and bowel function. Pt. I. Normal movement of contents. Gut 9: 442–456, 1968.
 676. Garry, R. C. The movements of the large intestine. Physiol. Rev. 14: 103–132, 1934.
 677. Cannon (n. 8), p. 276.
 678. Ludwig, C. Lehrbuch des Physiologie des Menschen. Leipzig, Germany: Winder'sche Verlaghandlung, 1861, vol. II, p. 617.
 679. Gabella, G. Structure of the Autonomic Nervous System. London: Chapman & Hall, 1976, p. 155.
 680. Furness, J. B. The presence of inhibitory nerves in the colon after sympathetic denervation. Eur. J. Pharmacol. 6: 349–352, 1969.
 681. Schmidt, C. A. Distribution of vagus and sacral nerves to the large intestine. Proc. Soc. Exp. Biol. Med. 30: 739–740, 1933;
 682. Lannon, J., and E. Weller. The parasympathetic supply to the distal colon. Br. J. Surg. 34: 373–378, 1946–1947.
 683. Hultén, L. Extrinsic nervous control of colonic motility and blood flow. Acta Physiol. Scand. Suppl. 335: 1969.
 684. Langley and Anderson (n. 457).
 685. Wallentin, I. Studies on intestinal circulation. Acta Physiol. Scand. Suppl. 279: 1960.
 686. Reviews: Christensen, J. Myoelectric control of the colon. Gastroenterology 68: 601–609, 1975;
 687. Daniel, E. E. Electrophysiology of the colon. Gut 16: 298–329, 1975.
 688. Christensen, J., R. Caprilli, and G. F. Lund. Electric slow waves in circular muscle of cat colon. Am. J. Physiol. 217: 771–776, 1969;
 689. Christensen, J., and R. L. Hauser. Longitudinal axial coupling of slow waves in proximal cat colon. Am. J. Physiol. 221: 246–250, 1971;
 690. Christensen, J., and R. L. Hauser. Circumferential coupling of electric slow waves in circular muscle of cat colon. Am. J. Physiol. 221: 1033–1037, 1971;
 691. Christensen, J., and B. F. Freeman. Circular muscle electromyogram in the cat colon. Gastroenterology 63: 1011–1015, 1972;
 692. Christensen, J., and S. C. Rasmus. Colon slow waves: size of oscillators and rates of spread. Am. J. Physiol. 223: 1330–1333, 1972.
 693. Wienbeck, M., and J. Christensen. Electromyography of the colon in the unanesthetized cat. Am. J. Dig. Dis. 7: 356–362, 1972;
 694. Wienbeck, M. The normal electrical activity and its relationship to contractile activity. Res. Exp. Med. 158: 268–279, 1972.
 695. Wankling, W. J., B. H. Brown, C. D. Collins, and H. L. Duthie. Basal electrical activity in the anal canal in man. Gut 9: 457–460, 1968;
 696. Taylor, I., H. L. Duthie, B. K. Small‐wood, and D. Linkens. The effect of stimulation on the myoelectrical action of the rectosigmoid in man. Gut 15: 599–607, 1974.
 697. French workers made similar but less complete observations at the same time: Coutoutier, D., C. Roze, M. H. Coutourier‐Turpin, and C. Debray. Electromyography of the colon in situ: an experimental study in man and in the rabbit. Gastroenterology 56: 317–322, 1969.
 698. Szurszewski, J. H. Toward a new view of prevertebral ganglion. In: Nerves and the Gut, edited by F. P. Brooks and P. Evers. Thorofare, NJ: Slack, 1977.
 699. Kuntz, A., and R. J. Mosel. An experimental analysis of the pelvic autonomic ganglia in the cat. J. Comp. Neurol. 64: 63–75, 1936;
 700. Kuntz, A., and G. Saccomanno. Reflex inhibition of intestinal motility mediated through decentralized prevertebral ganglia. J. Neurophysiol. Bethesda 7: 163–170, 1944.
 701. Crowfoot, P. J., and J. H. Szurszewski. A study of the inferior and pelvic ganglia of guinea‐pigs with intracellular electrode. J. Physiol. Lond. 219: 421–441, 1971;
 702. Crowfoot, P. J., M. Holman, and J. H. Szurszewski. Excitatory input from the distal colon to the inferior mesenteric ganglion in the guinea‐pig. J. Physiol. Lond. 219: 443–461, 1971.
 703. Schuster, M. M. The riddle of the sphincters. Gastroenterology 69: 249–262, 1975.
 704. Fellner, L. Weitere Mittheilungen über die Bewegungs‐ und Hemmungsnerven des Rectums. Pfluegers Arch. 56: 542–557, 1894.
 705. Langley and Anderson (n. 457).
 706. Starling (n. 462), p. 313–337.
 707. Sherrington, C. S. The spinal cord. In: Text‐book of Physiology, edited by E. A. Schäfer. Edinburgh: Pentland, 1900, vol. II, p. 782–883, p. 851.
 708. Garry, R. C. The nervous control of the caudal region of the large bowel in the cat. J. Physiol. Lond. 77: 422–431, 1933.
 709. Garry, R. C. The response to stimulation of the caudal end of the large bowel in the cat. J. Physiol. Lond. 78: 208–224, 1933;
 710. see also Garry (n. 485) for review as of 1934.
 711. This may be true in the cat, but it is not true in the human. See Floyd, W. F., and E. W. Walls. Electromyography of the sphincter ani externis in man. J. Physiol. Lond. 122: 599–609, 1953.
 712. Bishop, B., R. C. Garry, T. D. M. Roberts, and J. K. Todd. Control of the external sphincter of the anus in the cat. J. Physiol. Lond. 134: 229–240, 1956.
 713. Hill, J. R., M. L. Kelley, Jr., J. F. Schlegel, and C. F. Code. Pressure profile of the rectum and anus in healthy persons. Dis. Colon Rectum 3: 203–209, 1960.
 714. Duthie, H. L., and F. W. Gairns. Sensory nerve‐endings and sensations in the anal region of man. Br. J. Surg. 47: 585–595, 1960.
 715. Duthie, H. L., and R. C. Bennett. The relation of sensation in the anal canal to the functional anal sphincter: a possible factor in anal continence. Gut 4: 179–182, 1963.
 716. Beck, A. Electromyograpische Untersuchungen am Sphincter ani. Pfluegers Arch. 224: 278–292, 1930.
 717. Floyd and Walls (n. 508); Kawakami, M. Electromyographic investigation on the human external sphincter muscle of the anus. Jpn. J. Physiol. 4: 196–204, 1954;
 718. Frenckner, B., and C. von Euler. Influence of pudendal block on the function of the anal sphincter. Gut 16: 482–489, 1975.
 719. Schuster, M. M., P. Hookman, T. R. Hendrix, and A. F. Mendeloff. Simultaneous manometric recording of internal and external anal sphincter response. Bull. Johns Hopkins Hosp. 116: 79–88, 1965.
 720. Head, H., and G. Riddoch. The automatic bladder, excessive sweating and other reflex conditions in gross injuries of the spinal cord. Brain 40: 188–263, 1917.
 721. Denny‐Brown, D., and E. G. Robertson. An investigation of the nervous control of defaecation. Brain 58: 256–310, 1935.
 722. Goltz, F., and J. R. Ewald. Der Hund mit verkürzten Rückmark. Pfluegers Arch. 63: 362–400, 1896.
 723. Hatcher, R. A., and S. Weiss. Studies on vomiting. J. Pharmacol. Exp. Ther. 22: 139–193, 1923.
 724. Koppani, T. Studies on defecation with special reference to a medullary defecation center. J. Lab. Clin. Med. 16: 225–238, 1930.
 725. Rostad, H. Colonic motility in the cat. Acta Physiol. Scand. Suppl. 89: 1973.
 726. I quote from Hurst, A. F. Constipation and Allied Disorders (2nd ed.). London: Frowde, 1919. I have not seen the first edition of 1909, and I am assuming the second simply follows the first edition.
 727. I quote O'Beirne, J. New Views of the Process of Defecation. Washington, DC: Duff Green, 1834.
 728. This is apparently a pirated edition. The Surgeon General's catalog says the first edition was published in Dublin in 1833 by Hodges & Smith. Hurst gives the spelling of the operant word in the title as Defecation, whereas it is always otherwise spelled Defaecation. Therefore it is probably safe to assume that the Irish spelling was the same as the American, as indeed the Surgeon General's catalog cites it. There is an anonymous obituary of O'Beirne in Lancet 1: 114, 1862.
 729. Cannon, W. B. The movements of the intestines studied by means of the Röntgen rays. Am. J. Physiol. 6: 251–277, 1902.
 730. Hertz, A. F. The passage of food along the alimentary canal. Guy's Hosp. Rep. 61: 384–427, 1904, p. 423–424.
 731. Rendich, R. A., and L. A. Harrington. The roentgen kymography of the normal colon. Defecation in man. Am. J. Roentgenol. 40: 173–179, 1938.
 732. Schuster, M. M. Motor action of the rectum and sphincters in continence and defecation. In: Handbook of Physiology. Alimentary Canal, edited by C. F. Code. Washington, DC: Am. Physiol. Soc., 1968, sect. 6, vol. IV, chapt. 103, p. 2121–2140.
 733. Reviews: Duthie, H. L. Anal continence. Gut 12: 844–852, 1971;
 734. Garry (n. 485);
 735. Dickinson, V. A. Maintenance of continence: a review of pelvic physiology. Gut 19: 1163–1174, 1978.
 736. See Lipkin, M., T. P. Almy, and B. M. Ball. Pressure volume characteristics of the human colon. J. Clin. Invest. 41: 1831–1839, 1962. Observations reported in this paper show the colonometrogram of the sigmoid colon has the same characteristics as the cystometrogram. One can, perhaps, assume the rectum would behave in the same mammer if the reflexes were suppressed.
 737. Harris, L. D., and C. E. Pope II. “Squeeze” vs. resistance: an evaluation of the mechanism of sphincter competence. J. Clin. Invest. 43: 2272–2278, 1965;
 738. Harris, L. D., C. S. Winans, and C. E. Pope II. Determination of yield pressure: a method for measuring anal sphincter competence. Gastroenterology 50: 754–760, 1966.
 739. Phillips, S. F., and D. A. W. Edwards. Some aspects of anal competence and defaecation. Gut 6: 396–406, 1965.
 740. Hirschsprung, H. Stuhlträgheit neugeborenes in Folge von Dilatation und Hypertrophie des Kolons. Ann. Paediatr. 27: 1–7, 1886.
 741. Concetti, L. Ueber einige angeborene, bei Kindern die habituelle Verstopfung hervorrubenden Missbildung des Colons. Klin. Paediatr. 27: 318–366, 1899.
 742. Tittle, K. Ueber eine angeborene Missbildung des Dickdarmes. Wien. Klin. Wochenschr. 14: 903–907, 1901.
 743. For reviews, see Eherenpreis, T. Megacolon in the newborn, a clinical and roentgenological study with special reference to pathogenesis. Acta Chir. Scand. Suppl. 112: 1946;
 744. Whitehouse, F. R., and J. W. Kernohan. Myenteric plexus in congenital megacolon: a study of 11 cases. Arch. Intern. Med. 82: 75–83, 1948.
 745. Gannon, B. J., H. R. Noblet, and G. Burnstock. Adrenergic innervation of bowel in Hirschsprung's disease. Br. J. Med. 3: 338–340, 1969.
 746. Davidson, M., M. H. Sleisenger, H. Steiberg, and T. P. Almy. Studies of distal colonic motility. III. The abnormal physiology of congenital megacolon. Gastroenterology 29: 803–823, 1955.
 747. Wade, R. B., and N. D. Royle. The operative treatment of Hirschsprung's disease: a new method. Med. J. Aust. 1: 137–141, 1927. The report of the procedure is by Wade; the “explanation” is by Royle.
 748. Judd, E. S., and A. W. Adson. Lumbar ganglionectomy and ramisection for congenital idiopathic dilatation of the colon. Ann. Surg. 88: 479–498, 1928.
 749. A slightly later paper from the Mayo Clinic is Learmouth, F. W., and J. R. Rankin. Section of the sympathetic nerves of the distal part of the colon and the rectum in the treatment of Hirschsprung's disease and certain types of constipation. Ann. Surg. 92: 710–720, 1930.
 750. Adamson, W. A. D., and I. Aird. Megacolon: evidence in favor of a neurogenic origin. Br. J. Surg. 20: 220–223, 1932.
 751. Report of 178 cases: Penick, R. M., Jr. Problems in surgical treatment of congenital megacolon. J. Am. Med. Assoc. 128: 423–427, 1945.
 752. The “pull‐through” operation: Swenson, O., and A. H. Bill, Jr. Resection of rectum and rectosigmoid with preservation of sphincter for benign spastic lesions produring megacolon. Surgery St. Louis 24: 212–220, 1948.
 753. Derrick, E. H., and B. M. St. George‐Grambauer. Megacolon in mice. J. Pathol. Bacteriol. 73: 569–571, 1957.
 754. Beilchowsky, M., and G. C. Schofield. Studies on megacolon in piebald mice. Aust. J. Exp. Biol. Med. Sci. 40: 395–400, 1962.
 755. Wood, J. D. Electrical activity of the intestine of mice with hereditary megacolon and absence of enteric ganglion cells. Am. J. Dig. Dis. 18: 477–488, 1973.
 756. Cannon (n. 8), p. 220.
 757. Alvarez, W. C. Nervous Indigestion. New York: Hoeber, 1930.
 758. White, B. V., and C. M. Jones. The effect of irritants and drugs affecting the autonomic nervous system upon the mucosa of the normal rectum and rectosigmoid, with especial reference to “mucous colitis.” New Engl. J. Med. 218: 791–797, 1938.
 759. Chi Kim, I., and G. J. Barbero. The pattern of rectosigmoid motility in children. Gastroenterology 45: 57–66, 1963.
 760. Deller, D. J., and G. Wangel. Intestinal motility in man. I. A study combining intraluminal pressure recording with cineradiography. Gastroenterology 48: 45–57, 1965.
 761. Connell, A. M. The motility of the pelvic colon. I. Motility in normals and in patients with asymptomatic duodenal ulcer. Gut 2: 175–186, 1961.
 762. Ritchie et al. (n. 483).
 763. For appreciations of Wolff, see Almy, T. P. Harold G. Wolff, 1898–1962. Trans. Assoc. Am. Physicians. 75: 45–47, 1962;
 764. and a longer one: Soriano, V. Harold G. Wolff, 1898–1962. Int. J. Neurol. 3: 279–286, 1962.
 765. Grace, W. J., S. Wolf, and H. G. Wolff. The Human Colon. New York: Hoeber, 1951, p. 113–114.
 766. Almy's ideas are summarized in Almy, T. P. Experimental studies on the “irritable colon.” Am. J. Med. 10: 60–67, 1951;
 767. and Almy, T. P. What is the “irritable colon?” Am. J. Dig. Dis. 2: 93–97, 1957.
 768. Almy, T. P., F. Kern, Jr., and M. Tulin. Experimental production of sigmoid spasm in healthy persons. Gastroenterology 12: 425–436, 1949.
 769. Essentially similar results are reported in Almy, T. P., and M. Tulin. Alterations in colonic function in man under stress: experimental production of changes simulating the “irritable colon.” Gastroenterology 8: 616–626, 1947.
 770. Almy, T. P., L. E. Hinkle, Jr., B. Berle, and F. Kern, Jr. Experimental production of sigmoid spasm in patients with spastic constipation. Gastroenterology 12: 437–449, 1949.
 771. Almy, Am. J. Med. (n. 554), p. 64.

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Article Title: Gastrointestinal Physiology, 1895–1975: Motility
 

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Horace W. Davenport. Gastrointestinal Physiology, 1895–1975: Motility. Compr Physiol 2011, Supplement 16: Handbook of Physiology, The Gastrointestinal System, Motility and Circulation: 1-101. First published in print 1989. doi: 10.1002/cphy.cp060101