Arta anal. 92: 411- 416 (1975)
Anatomy Department, Abbassia Medical Faculty, Ein-Shams University, Cairo, Egypt
Intrinsic innervation of the gall bladder in the dog N.N.Y. N awar and I. K amel Key Words. Gall bladder • Innervation • Dog • Gall bladder innervation
Introduction Reviewing the literature, it was found that several workers have studied the light-microscopic structure of the gall bladder in different animal species [Oppel . 1900: J ubisch, 1909: Hoyden, 1926; D awson, 1943: K amel and N awab, 1975], The submicroscopic structure of the gall bladder epithelium received the attention of the other workers [YaMada. 1955: J ohnson el at.. 1962: H ayward, 1962, 1965, 1966: E vett el at., 1964: Bader, 1965, 1966: Chapman et al.. 1966], However, few workers have studied the intrinsic innervation of the gall bladder. H ebmaNN |19511 reported the intrinsic innervation of the human gall bladder. Sutherland 11966, 1967] described that of the monkey and guinea pig. Therefore, it became the aim of the present work to investigate the intrinsic innervation of the gall bladder of the dog and compare the results with the findings of other workers especially since some of the information was divergent.
Material and methods The gall bladders studied were taken from eight dogs 3 months old. The animals were killed by an overdose of anaest hetic, inhaled or administered inlraperitoneally. The abdomen was incised by a right subcostal incision and the gall bladder was removed as soon as possible after death and fixed in Bodian solution for 1 week. The materials were dehydrated, cleared and embedded in paraffin. Serial sections 15 /an thick were cut and stained by Holme’s silver technique and modified GrossBielschowskv stain.
As the gall bladder is a derivative of the gut, it is not surprising that it resembles other derivatives from this structure. Therefore, the nervous arrangem ent might accordingly he described in a similar way to th at of the intestine, i.e. (1) plexus subserus, (2) plexus m yentericus, (3) plexus tnuscularis profundus, (4) plexus intericus interims, and (5) plexus submucosus. However, it was necessary to describe the
Downloaded by: Nagoya University 133.6.82.173 - 1/14/2019 3:54:04 PM
Observations
Downloaded by: Nagoya University 133.6.82.173 - 1/14/2019 3:54:04 PM
412 N awar and K am Kt.
Intrinsic innervation of the gall bladder
413
normal histological appearance of the gall bladder of the dog in order to loeate these plexuses. A broad perivascular connective tissue coat rich in blood vessels, lym phatic vessels and elastic fibres surrounded the muscularis. As the nerve fibres reached the gall bladder, they divided and spread out around the organ to form the plexus subserosus. The plexus myentericus was situated in the outer layer of the gall bladder ju st ex ternal to the muscle layer. Cells with vesicular nuclei with one or two large nucleoli were found in relation with this plexus (fig. 1). The smooth muscle fibres of the gall bladder were disposed in longitudinal, circular and oblique directions. Reticular and collagen fibres filled the interstices between the muscle bundles. Undulating fasciculi of fine nerve fibres were observed parallel to the muscle fibres; these constituted the plexus muscularis profundus. Some of these nerve fibres ended in knob-like term inals on the muscle fibres (fig.2) and in a circular type of nerve ending (fig. 3). The mucous m em brane was thrown into folds. The epithelial cells were similar tall colum nar and were supported by a fine basement membrane and a delicate connective tissue (lamina propria). The latter contained fine undulating fasciculi of nerve fibres, which ap peared of equal thickness and were different from the reticular fibres in silver preparations. These formed the plexus entericus interims (fig.4). However, no ganglion cells were detected along the course of its fibres. Other nerve fibres were observed below the basem ent mem brane; these constituted the plexus submucosus (fig.5).
Discussion
Fig. 1. Photomicrograph of a nerve cell in relation with the myenteric plexus and undulating nerve fibres constituting the plexus muscularis profundus. Holme’s stain, x 380. Fig. 2. Photomicrograph of a knob-like terminal on the muscle fibres. Holme’s stain. X 380. Fig. 3. Photomicrograph of a circular nerve ending on the muscle fibres. Holme’s stain. X 380. Fig. 4. Photomicrograph of fasciculi of nerve fibres constituting the plexus entericus interims. Gross-Biclschowsky stain. X 188. Fig. 5. Photomicrograph of nerve fibres running below the basement membrane forming the plexus submucosus. Holme’s stain. X 380.
Downloaded by: Nagoya University 133.6.82.173 - 1/14/2019 3:54:04 PM
The gall bladder possesses a dual innervation, sym pathetic and parasym pathetic, derived from the coeliac plexus and passing along
N awar and K amel
the hepatic artery and its branches. Fibres from the right phrenic nerve through the com m unication between the phrenic plexus and the coeliac plexus also appear to reach the gall bladder in the hepaticplexus [W arwick and W illiams , 1973). For descriptive purposes the arrangem ent of the nerve libres within the wall of the organ were described as being divided into five plexuses, but this was rather artificial as the nerve plexuses were so interconnected th at they formed an extensive network in the wall of the gall bladder. Gerlach |1873| was the first to report th at there was more than one nerve plexus situated within the wall of the gall bladder. H ermann [1951] reported that the outer ganglionated plexus, like that described by A uerbach |18(>4| in the gut divided into prim ary, secondary and tertiary plexuses and the submucous plexus could be also divided, like th at in the intestine. In the dog, however, the division of the myenteric plexus was not clear, but the submucous plexus could be subdivided into an outer part, similar to the plexus entericus in terims described by H enle | 18711, and a true plexus submucosus, like th at reported by M eissner [1857|. In the present study, nerve cells were found in relation with the myenteric plexus. Ganglia in associ ation with this plexus were described in man [H ermann , 1951], monkey and guinea pig [S utherland , 1966]. Fine nerve fibres from the m yenteric plexus constituted the plexus muscularis profundus. The nerve fibres ran parallel to the muscle fibres. Fine nerve fibres were noticed to form a term inal reticulum in man | H ermann , 1951] while in the dog. knob-like term inals and circular type of nerve endings were observed. In the dog, no ganglia were met with in association with the plexus muscularis profundus. H ermann [1951] described ganglia in this plexus in man. S utherland [1966] did not find any ganglia in this situation in the monkey nor the guinea pig and suggested th a t what H ermann had observed was part of the myenteric plexus which was closely adherent and thus on section might appear to belong to the muscular layer. S utherland [1967| did not find ganglion cells in the submucous plexus of the guinea pig gall bladder, but he described sparse ganglia scattered throughout the mucosa in the monkey. H ermann [1951] described several forms of smaller ganglia in the submucous plexus in man. In the dog, no ganglia were met with in this situation and it probable th a t what H ermann described were neurilemma! or inter stitial cells, which he confused w ith ganglion cells. S utherland 119671 observed that neurilemma! cells lying at the intersection or bifurcation
Downloaded by: Nagoya University 133.6.82.173 - 1/14/2019 3:54:04 PM
414
Intrinsic innervation of the gall Madder
IIS
of a nerve bundle, where an excessive deposit of silver might occur, could sim ulate a small ganglion cell, while in choline esterase prepara tions, these cells did not stain. Although ganglion cells in the wall of the gall bladder were described as early as I860 by Mans [quoted in H arting , 1930|, the significance of the intrinsic innervation of the gall bladder is still not clear. D ogiel [ 18951 regarded the ganglion cells as sym pathetic in origin. A lexan der 11940] described the nerve cells as being com ponents of vagal efferent chains. Z abusov [ 1958] considered that the giant cells re present structures com parable to mechanoreeeptors registering disten sion and initiating reflex peristaltic contractions. Although textbooks of physiology describe the humoral mechanism as being responsible for evacuation of the gall bladder, it can nevertheless be assumed th at both humoral and nervous mechanisms are probably responsible in causing the gall bladder to em pty. This assumption is based on the histological findings of the rich network of nerve fibres and cells within the wall of the organ. However. B loom and F awcett |1969| emphasized the greater clinical im portance of the sensory nerve end ings as overdistension or spasm of the extrahepatic biliary tract leads to respiratory inhibition and reflex disturbance in the gastrointestinal tract. Summary The gall bladder of eight dogs 3 months old were extraeted and studied by the Holme’s and Gross-Biclschowsky techniques. The nerves were found to form an extensive network within the wall of the gall bladder. However. 5 plexuses were identified and their arrangement was similar to that in the wall of the intestine. Nerve cells were only found in relation with the myenteric plexus. Moreover, knob-like terminals and circular type of nerve endings were noticed on the muscularis. The findings of the intrinsic innervation of the gall bladder of the dog were compared with that of man. monkey and guinea pig and the significance of that innervation was discussed. Alexander. W.F. : The innervation of the biliary system. J. comp. Neurol. 72: 357-370 (1940). Auerbach, I..: Fernere vorläufige Mittheilung über den Nervenapparat des Darmes. Virchows Arch. path. Anal. 30: 457-460 (1864). Bader. G.: The submieroscopical structure of the epithelium of the gall bladder and its regeneration. 1. Carp and Frog. Z. mikr.-anat. Forsch. 74: 92-107 (1965). Bader. G.: The ultra-microscopical structure of the epithelium of the gall bladder and its regeneration. II. The chicken and various mammals. Z. mikr.-anat. Forsch. 74: 303-320 (1966).
Downloaded by: Nagoya University 133.6.82.173 - 1/14/2019 3:54:04 PM
References
N a ii and K amel \VA
Bloom. W. and F awcett. D.W.: A text book of histology; 9th ed.. pp. 582-613 (Saunders, Philadelphia 1968). Boyden, E. A.: The problem of pancreatic bladder. A critical survey of six cases based on new histological and cmbryological observations. Amcr. J. Anal. 36: 151-184- (1926). Chapman, G.B.: Cuiardo, A.I.; Coffey, R. 1... and Weineke , k.: The fine struc ture of the human gall bladder. Anal. Rec. 154: 579-615 (1966). Dawson, II. L.: A study of certain cells observed in the gall bladder mucosa of cats. Anal. Rec. 85: 135-155 (1943). DociEL, A.S.: Zur Frage über den feineren Bau des sympathischen Nervensystems bei den Säugetieren. Arch. mikr. Anal. 46: 305-344 (1895). Evett. R.D.: H iggins, J. A., and Brown, A.J.. jr.: The fine structure of normal mucosa in human gall bladder. Gastroenterology 47: 49-60 (1964). G erla CH, L.: Über die Nerven der Gallenblase. Zbl. med. Vt iss. 36: 562 563 (1873). Harting, K.: Über die feinere Innervation der extrahepatischen Gallenwege. I.Über die mikroskopische Innervation der Gallenblase. Z. Zellforsch. 12: 518- 548 (1930). H ayward, A. F .: Electron microscopic observations on absorption in epithelium of the guinea pig gall bladder. Z. Zellforsch. 56: 197-202 (1962). H ayward. A .F.: The fine structure of the gall bladder epithelium of the sheep. Z. Zellforsch. 65: 331-339 (1965). Hayward, A. F .: An electron microscopic study of developing gall bladder epi thelium in the rabbit. J. Anat.. Lond. 100: 245-259 (1966). II enle, J .; Handbuch der Nervcnlehre des Menschen (Vieweg. Braunschweig 1871). Hermann. H.: Das Nervensystem der menschlichen Gallenblase und seine Verän derungen bei Cholelithiasis. Virchows Arch. path. Anat. 322: 17-48 (1951). J ohnson, F. R.: McMin .n, R.M. II., and Birchenougii. R.F.: The ullrastructure of the gall bladder epithelium of the dog. J. Anal.. Lond. 96: 477-487 (1962). J irisch . A.: Beiträge zur mikroskopischen Anatomic und Histologie der Gallen blase. Anat. H. 39: 395-467 (1909). Kamel, 1. and N awar, N.N.Y.: Comparative microscopical study of the gall bladder mucosa. Acta anat. 91: 104-109 (1975). Meissner, G.: Über die Nerven der Darmwand. Z. rat. Med. (new series) 8: 364-366 (1857). Oppel , A.: Lehrbuch der vergleichenden mikroskopischen Anatomie, vol.3, pp.972 980 (Fischer. Jena 1900). Sutherland, S. I).: The intrinsic innervation of the gall bladder in Macaca rhesus and Cavia porcellus. J. Anat., Lond. 100: 216-268 (1966). Sutherland, S. D.: The neurons of the gall bladder and gut. J. Anat., Lond. 101: 701-709 (1967). Warwick, R. and W illiams, P. L.: Gray’s anatomy: 35th ed., pp. 1311-1312 (Long man’s Green, London 1973). Y amada. E .; The fine structure of the gall bladder epithelium of the mouse. J. biophys. bioehem. Cytol. 1: 455-458 (1955). Zabusov, G. I.: The significance of giant cells of the nerve plexus of the gall bladder in mammals. Abstract. Vop. morf. Chuvst intervast. 5: 80-89 (1958). Received May 27, 1974 Dr. N .N .Y .N awar, 11. Aziz Osman St., Zamalck. Cairo (Egypt)
Downloaded by: Nagoya University 133.6.82.173 - 1/14/2019 3:54:04 PM
+16
Anatomy Department, Abbassia Medical Faculty, Ein-Shams University, Cairo, Egypt
Intrinsic innervation of the gall bladder in the dog N.N.Y. N awar and I. K amel Key Words. Gall bladder • Innervation • Dog • Gall bladder innervation
Introduction Reviewing the literature, it was found that several workers have studied the light-microscopic structure of the gall bladder in different animal species [Oppel . 1900: J ubisch, 1909: Hoyden, 1926; D awson, 1943: K amel and N awab, 1975], The submicroscopic structure of the gall bladder epithelium received the attention of the other workers [YaMada. 1955: J ohnson el at.. 1962: H ayward, 1962, 1965, 1966: E vett el at., 1964: Bader, 1965, 1966: Chapman et al.. 1966], However, few workers have studied the intrinsic innervation of the gall bladder. H ebmaNN |19511 reported the intrinsic innervation of the human gall bladder. Sutherland 11966, 1967] described that of the monkey and guinea pig. Therefore, it became the aim of the present work to investigate the intrinsic innervation of the gall bladder of the dog and compare the results with the findings of other workers especially since some of the information was divergent.
Material and methods The gall bladders studied were taken from eight dogs 3 months old. The animals were killed by an overdose of anaest hetic, inhaled or administered inlraperitoneally. The abdomen was incised by a right subcostal incision and the gall bladder was removed as soon as possible after death and fixed in Bodian solution for 1 week. The materials were dehydrated, cleared and embedded in paraffin. Serial sections 15 /an thick were cut and stained by Holme’s silver technique and modified GrossBielschowskv stain.
As the gall bladder is a derivative of the gut, it is not surprising that it resembles other derivatives from this structure. Therefore, the nervous arrangem ent might accordingly he described in a similar way to th at of the intestine, i.e. (1) plexus subserus, (2) plexus m yentericus, (3) plexus tnuscularis profundus, (4) plexus intericus interims, and (5) plexus submucosus. However, it was necessary to describe the
Downloaded by: Nagoya University 133.6.82.173 - 1/14/2019 3:54:04 PM
Observations
Downloaded by: Nagoya University 133.6.82.173 - 1/14/2019 3:54:04 PM
412 N awar and K am Kt.
Intrinsic innervation of the gall bladder
413
normal histological appearance of the gall bladder of the dog in order to loeate these plexuses. A broad perivascular connective tissue coat rich in blood vessels, lym phatic vessels and elastic fibres surrounded the muscularis. As the nerve fibres reached the gall bladder, they divided and spread out around the organ to form the plexus subserosus. The plexus myentericus was situated in the outer layer of the gall bladder ju st ex ternal to the muscle layer. Cells with vesicular nuclei with one or two large nucleoli were found in relation with this plexus (fig. 1). The smooth muscle fibres of the gall bladder were disposed in longitudinal, circular and oblique directions. Reticular and collagen fibres filled the interstices between the muscle bundles. Undulating fasciculi of fine nerve fibres were observed parallel to the muscle fibres; these constituted the plexus muscularis profundus. Some of these nerve fibres ended in knob-like term inals on the muscle fibres (fig.2) and in a circular type of nerve ending (fig. 3). The mucous m em brane was thrown into folds. The epithelial cells were similar tall colum nar and were supported by a fine basement membrane and a delicate connective tissue (lamina propria). The latter contained fine undulating fasciculi of nerve fibres, which ap peared of equal thickness and were different from the reticular fibres in silver preparations. These formed the plexus entericus interims (fig.4). However, no ganglion cells were detected along the course of its fibres. Other nerve fibres were observed below the basem ent mem brane; these constituted the plexus submucosus (fig.5).
Discussion
Fig. 1. Photomicrograph of a nerve cell in relation with the myenteric plexus and undulating nerve fibres constituting the plexus muscularis profundus. Holme’s stain, x 380. Fig. 2. Photomicrograph of a knob-like terminal on the muscle fibres. Holme’s stain. X 380. Fig. 3. Photomicrograph of a circular nerve ending on the muscle fibres. Holme’s stain. X 380. Fig. 4. Photomicrograph of fasciculi of nerve fibres constituting the plexus entericus interims. Gross-Biclschowsky stain. X 188. Fig. 5. Photomicrograph of nerve fibres running below the basement membrane forming the plexus submucosus. Holme’s stain. X 380.
Downloaded by: Nagoya University 133.6.82.173 - 1/14/2019 3:54:04 PM
The gall bladder possesses a dual innervation, sym pathetic and parasym pathetic, derived from the coeliac plexus and passing along
N awar and K amel
the hepatic artery and its branches. Fibres from the right phrenic nerve through the com m unication between the phrenic plexus and the coeliac plexus also appear to reach the gall bladder in the hepaticplexus [W arwick and W illiams , 1973). For descriptive purposes the arrangem ent of the nerve libres within the wall of the organ were described as being divided into five plexuses, but this was rather artificial as the nerve plexuses were so interconnected th at they formed an extensive network in the wall of the gall bladder. Gerlach |1873| was the first to report th at there was more than one nerve plexus situated within the wall of the gall bladder. H ermann [1951] reported that the outer ganglionated plexus, like that described by A uerbach |18(>4| in the gut divided into prim ary, secondary and tertiary plexuses and the submucous plexus could be also divided, like th at in the intestine. In the dog, however, the division of the myenteric plexus was not clear, but the submucous plexus could be subdivided into an outer part, similar to the plexus entericus in terims described by H enle | 18711, and a true plexus submucosus, like th at reported by M eissner [1857|. In the present study, nerve cells were found in relation with the myenteric plexus. Ganglia in associ ation with this plexus were described in man [H ermann , 1951], monkey and guinea pig [S utherland , 1966]. Fine nerve fibres from the m yenteric plexus constituted the plexus muscularis profundus. The nerve fibres ran parallel to the muscle fibres. Fine nerve fibres were noticed to form a term inal reticulum in man | H ermann , 1951] while in the dog. knob-like term inals and circular type of nerve endings were observed. In the dog, no ganglia were met with in association with the plexus muscularis profundus. H ermann [1951] described ganglia in this plexus in man. S utherland [1966] did not find any ganglia in this situation in the monkey nor the guinea pig and suggested th a t what H ermann had observed was part of the myenteric plexus which was closely adherent and thus on section might appear to belong to the muscular layer. S utherland [1967| did not find ganglion cells in the submucous plexus of the guinea pig gall bladder, but he described sparse ganglia scattered throughout the mucosa in the monkey. H ermann [1951] described several forms of smaller ganglia in the submucous plexus in man. In the dog, no ganglia were met with in this situation and it probable th a t what H ermann described were neurilemma! or inter stitial cells, which he confused w ith ganglion cells. S utherland 119671 observed that neurilemma! cells lying at the intersection or bifurcation
Downloaded by: Nagoya University 133.6.82.173 - 1/14/2019 3:54:04 PM
414
Intrinsic innervation of the gall Madder
IIS
of a nerve bundle, where an excessive deposit of silver might occur, could sim ulate a small ganglion cell, while in choline esterase prepara tions, these cells did not stain. Although ganglion cells in the wall of the gall bladder were described as early as I860 by Mans [quoted in H arting , 1930|, the significance of the intrinsic innervation of the gall bladder is still not clear. D ogiel [ 18951 regarded the ganglion cells as sym pathetic in origin. A lexan der 11940] described the nerve cells as being com ponents of vagal efferent chains. Z abusov [ 1958] considered that the giant cells re present structures com parable to mechanoreeeptors registering disten sion and initiating reflex peristaltic contractions. Although textbooks of physiology describe the humoral mechanism as being responsible for evacuation of the gall bladder, it can nevertheless be assumed th at both humoral and nervous mechanisms are probably responsible in causing the gall bladder to em pty. This assumption is based on the histological findings of the rich network of nerve fibres and cells within the wall of the organ. However. B loom and F awcett |1969| emphasized the greater clinical im portance of the sensory nerve end ings as overdistension or spasm of the extrahepatic biliary tract leads to respiratory inhibition and reflex disturbance in the gastrointestinal tract. Summary The gall bladder of eight dogs 3 months old were extraeted and studied by the Holme’s and Gross-Biclschowsky techniques. The nerves were found to form an extensive network within the wall of the gall bladder. However. 5 plexuses were identified and their arrangement was similar to that in the wall of the intestine. Nerve cells were only found in relation with the myenteric plexus. Moreover, knob-like terminals and circular type of nerve endings were noticed on the muscularis. The findings of the intrinsic innervation of the gall bladder of the dog were compared with that of man. monkey and guinea pig and the significance of that innervation was discussed. Alexander. W.F. : The innervation of the biliary system. J. comp. Neurol. 72: 357-370 (1940). Auerbach, I..: Fernere vorläufige Mittheilung über den Nervenapparat des Darmes. Virchows Arch. path. Anal. 30: 457-460 (1864). Bader. G.: The submieroscopical structure of the epithelium of the gall bladder and its regeneration. 1. Carp and Frog. Z. mikr.-anat. Forsch. 74: 92-107 (1965). Bader. G.: The ultra-microscopical structure of the epithelium of the gall bladder and its regeneration. II. The chicken and various mammals. Z. mikr.-anat. Forsch. 74: 303-320 (1966).
Downloaded by: Nagoya University 133.6.82.173 - 1/14/2019 3:54:04 PM
References
N a ii and K amel \VA
Bloom. W. and F awcett. D.W.: A text book of histology; 9th ed.. pp. 582-613 (Saunders, Philadelphia 1968). Boyden, E. A.: The problem of pancreatic bladder. A critical survey of six cases based on new histological and cmbryological observations. Amcr. J. Anal. 36: 151-184- (1926). Chapman, G.B.: Cuiardo, A.I.; Coffey, R. 1... and Weineke , k.: The fine struc ture of the human gall bladder. Anal. Rec. 154: 579-615 (1966). Dawson, II. L.: A study of certain cells observed in the gall bladder mucosa of cats. Anal. Rec. 85: 135-155 (1943). DociEL, A.S.: Zur Frage über den feineren Bau des sympathischen Nervensystems bei den Säugetieren. Arch. mikr. Anal. 46: 305-344 (1895). Evett. R.D.: H iggins, J. A., and Brown, A.J.. jr.: The fine structure of normal mucosa in human gall bladder. Gastroenterology 47: 49-60 (1964). G erla CH, L.: Über die Nerven der Gallenblase. Zbl. med. Vt iss. 36: 562 563 (1873). Harting, K.: Über die feinere Innervation der extrahepatischen Gallenwege. I.Über die mikroskopische Innervation der Gallenblase. Z. Zellforsch. 12: 518- 548 (1930). H ayward, A. F .: Electron microscopic observations on absorption in epithelium of the guinea pig gall bladder. Z. Zellforsch. 56: 197-202 (1962). H ayward. A .F.: The fine structure of the gall bladder epithelium of the sheep. Z. Zellforsch. 65: 331-339 (1965). Hayward, A. F .: An electron microscopic study of developing gall bladder epi thelium in the rabbit. J. Anat.. Lond. 100: 245-259 (1966). II enle, J .; Handbuch der Nervcnlehre des Menschen (Vieweg. Braunschweig 1871). Hermann. H.: Das Nervensystem der menschlichen Gallenblase und seine Verän derungen bei Cholelithiasis. Virchows Arch. path. Anat. 322: 17-48 (1951). J ohnson, F. R.: McMin .n, R.M. II., and Birchenougii. R.F.: The ullrastructure of the gall bladder epithelium of the dog. J. Anal.. Lond. 96: 477-487 (1962). J irisch . A.: Beiträge zur mikroskopischen Anatomic und Histologie der Gallen blase. Anat. H. 39: 395-467 (1909). Kamel, 1. and N awar, N.N.Y.: Comparative microscopical study of the gall bladder mucosa. Acta anat. 91: 104-109 (1975). Meissner, G.: Über die Nerven der Darmwand. Z. rat. Med. (new series) 8: 364-366 (1857). Oppel , A.: Lehrbuch der vergleichenden mikroskopischen Anatomie, vol.3, pp.972 980 (Fischer. Jena 1900). Sutherland, S. I).: The intrinsic innervation of the gall bladder in Macaca rhesus and Cavia porcellus. J. Anat., Lond. 100: 216-268 (1966). Sutherland, S. D.: The neurons of the gall bladder and gut. J. Anat., Lond. 101: 701-709 (1967). Warwick, R. and W illiams, P. L.: Gray’s anatomy: 35th ed., pp. 1311-1312 (Long man’s Green, London 1973). Y amada. E .; The fine structure of the gall bladder epithelium of the mouse. J. biophys. bioehem. Cytol. 1: 455-458 (1955). Zabusov, G. I.: The significance of giant cells of the nerve plexus of the gall bladder in mammals. Abstract. Vop. morf. Chuvst intervast. 5: 80-89 (1958). Received May 27, 1974 Dr. N .N .Y .N awar, 11. Aziz Osman St., Zamalck. Cairo (Egypt)
Downloaded by: Nagoya University 133.6.82.173 - 1/14/2019 3:54:04 PM
+16
