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Structural and Functional Relationships of the Enterocyte

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Structural and Functional Relationships of the Enterocyte 12 Postgraduate Medical Journal Postgrad Med J: first published as 10.1136/pgmj.44.507.12 on 1 January 1968. Downloaded from return to the patient and determine to what DANES, B.S. & BEARN, A.G. (1967) The effect of retinol extent the findings derived from in vitro studies (vitamin A alcohol) on urinary excretion of mucopoly- saccharides in the Hurler syndrome. Lancet, i, 1029. are directly applicable to the human condition. HENDERSON, J.L. (1940) Gargoylism; review of principal features with report of five cases. Arch. Dis. Child. 15, 215. McKUSICK, V.A. Heritable Disorders of Connective Tissue. References Mosby, St Louis. DANES, B.S. & BEARN, A.G. (1966a) Hurler's syndrome. A SATO, A. (1955) Chediak and Higashi's disease. Probable genetic study in cell culture. J. exp. Med. 123, 1. identity of 'a new leucocytal anomaly (Chediak)' and DANES, B.S. & BEARN, A.G. (1966b) Hurler's syndrome. 'congenital gigantism of peroxidase granules (Higashi).' Effect of retinol (vitamin A alcohol) on cellular mucopoly- Tohoku J. exp. Med. 61, 201. saccharides in cultured human skin fibroblasts. J. exp. WHITE, J.G. (1966) The Chediak-Higashi syndrome: A Med. 124, 1181. possible lysosomal disease. Blood, 28, 143. Enteropoiesis: structural and functional relationships of the enterocyte C. C. BOOTH Royal Postgraduate Medical School, London Protected by copyright. THE REMARKABLE turnover of the cells of the layer of the small intestine is markedly increased, intestinal mucosa was first suspected by Bizzozero 'the situation is comparable with anaemia in as long ago as 1888 and has been firmly estab- which there is reduction of the peripheral red lished by modern techniques during the past 20 cells (villous cells) and compensatory hyper- years (Leblond & Stevens, 1948; Leblond & trophy of the red bone marrow (glandular Messier, 1958; Creamer, Shorter & Bamforth, mucosa)'. 1961). The absorbing cells of the small intestine This paper explores further the theoretical have, as Smyth has pointed out, a 'short life and possibilities raised by Doniach's haematological a merry one ... if merriment is the equivalent analogy and suggests a new nomenclature for in cellular terms of a remarkable range of acti- the intestinal mucosal cells. The functional im- vities possessed by few other cells in the body' plications of this classification are also discussed. (Smyth, 1967). Formed in the crypts of Lieber- kuhn, the intestinal cells are extruded on to the Nomenclature of the intestinal absorbing cells surface of an adjacent villus whose side they There are a variety of different cell types pre- http://pmj.bmj.com/ climb until they reach the villus tip, from where sent in the intestinal mucosa. Paneth cells, argen- they are shed into the intestinal lumen. The total taffin cells and goblet cells mingle with the turnover time for the cells of the jejunum is columnar cells which make up the major part known to be as rapid as 2 or 3 days in the of the absorptive epithelium and which are the mouse and perhaps 5 days in man (MacDonald, concern of this paper. These columnar cells have Trier & Everett, 1964; Shorter et al., 1964), so been given a variety of names such as villous that the body only uses the extraordinary bio- cells, absorptive cells, mucosal surface cells and on September 30, 2021 by guest. logical activities of these cells for a very brief adult epithelial cells, and the cells in the crypts period. of Lieberkuhn have been variously called the Doniach & Shiner (1960) were the first to draw glandular cells, germinative zone or production the analogy between the intestinal epithelial cells zone. The use of a haematological analogy sug- and the haemopoietic system. They compared gests a more satisfactory nomenclature which at the germinative or crypt zone, where the cells the same time links the formative cells of the are manufactured, with the erythroblasts of the crypts with their adult counterparts. If the ab- bone marrow and the adult absorbing cells of sorptive cell on the surface of the villus is given the villi with the erythrocytes of the peripheral the term 'enterocyte', then the germinative cell blood. Doniach & Shiner pointed out that in in the glandular mucosa should be called an coeliac disease, where the jejunal mucosa is 'enteroblast', the function of this cell being devoid of villi but the glandular or germinative 'enteropoiesis'. The further implications of this Festschrift for Sir John McMichael 13 Postgrad Med J: first published as 10.1136/pgmj.44.507.12 on 1 January 1968. Downloaded from nomenclature are shown schematically in Fig. 1. not hitherto been encountered in this situation. The enterocyte may be enlarged (a macroentero- cyte), normal, or reduced in size (a microentero- The normocytic enterocyte cyte). The numbers of the enterocytes may be As indicated in Fig. 1, the normocytic entero- increased (enterocytic hyperplasia), normal or re- cyte may be normoplastic, hyperplastic or hypo- duced (enterocytic hypoplasia). Furthermore, the plastic, and its function may be increased, function of the enterocyte is not static for under normal or decreased. certain circumstances there may be either in- creased or decreased function of this cell. Defin- (i) Normoplasia ing the intestinal cell in this way poses a num- Clearly the normal situation in the small in- ber of questions which can only be resolved testine is established when there are normocytic by future research, but recent studies have al- enterocytes which are present in 'normal' num- ready demonstrated many of the situations bers and which function 'normally'. As with all shown in Fig. 1. other biological phenomena, however, there is a considerable range of normality in the small The macrocytic enterocyte intestine. There are marked variations in the In pernicious anaemia it is well known that number of enterocytes which constitute norm- the cells of the buccal mucosa may be larger ality and these variations are reflected by differ- than normal. Similarly in the small intestine a ences in the appearance of the surface mucosa recent report has described macrocytic entero- in different individuals and in different popula- cytes and a megaloblastic enteroblast (Foroozan tions throughout the world. Under the dissecting & Trier, 1967). This appears to be associated microscope, the shapes of the villi of the small with enterocytic hypoplasia since there is also intestine vary widely, some individuals showing shortening of the villi. How such cells function villi which are entirely finger-shaped, whereas Protected by copyright. is not known. The cause of the intestinal ab- others have a pattern of leaf- and tongue- normality in pernicious anaemia is presumably a shaped villi. Creamer (1964) has estimated that similar 'maturation arrest', involving the entero- the number of enterocytes required to cover blast, to that seen in the bone marrow. leaf-shaped villi are only 25% of those necessary As might be expected an arrest of cell turn- when the villi are finger-shaped. The variation over in the small intestine also occurs after in the number of enterocytes in different normal X-irradiation. A recent study of jejunal biopsies individuals is not surprising in an organ where obtained from patients undergoing therapeutic cell turnover is extremely rapid and where the X-irradiation (Trier & Browning, 1966) has de- balance between enteropoiesis and cell loss must monstrated that the mitoses in the enteroblastic often be precarious. layer of the small intestine are reduced within Increased function of normocytic enterocytes 12 hr of starting X-ray therapy. Thereafter, there may occur in a variety of situations of stress. is enterocytic hypoplasia, causing marked short- The effects of bulk-feeding in the rat provide ening of the villi and at the same time megalo- an interesting example of hyperfunction (Dowl- http://pmj.bmj.com/ blastosis and megalocytosis of the epithelial cells ing et al., 1967a). When the diet of the animal develops. As in pernicious anaemia, this situation contains a large amount of unabsorbed bulk, is rapidly reversible. there is an increase in the absorptive capacity of The question must also be asked whether the the small intestine without any change in the enterocyte may become macrocytic when there macroscopic appearance of the bowel and with- is compensatory hypertrophy following partial out enterocytic hyperplasia. Using the everted resection of the small intestine. Although there sac technique Dowling et al. (1967a) showed that on September 30, 2021 by guest. is clear evidence of enterocytic hyperplasia after glucose absorption by the jejunum of rats whose intestinal resection, macrocytic enterocytes have diets contained 60 % or 80 % by weight of Macrocytic Hyperplastic Hyperfunction Enteroblast-----)' Enterocyte Normocytic \--- Normoplastic Normofunction Microcytic Hypoplastic Hypofunction Enteropoiesis FIG. 1 14 Postgraduate Medical Journal Postgrad Med J: first published as 10.1136/pgmj.44.507.12 on 1 January 1968. Downloaded from powdered kaolin was 4-81 (± 0-99 SEM) mg and in control subjects and in patients with intestinal 9-96 (+053 SEM) mg respectively per 100 mg resection. Eight of the patients in whom resec- dried intestinal tissue per hour, compared with tion had been performed absorbed 60-9 (±2.20 2-52 (+0-66 SEM) mg in control rats. Detailed SEM) mg of glucose per 25 cm of intestine per anatomical and histological studies showed that minute, as compared with 48.3 (± 2-07 SEM) this functional response was not due to any in- mg in the control subjects (Dowling & Booth, crease in absorptive surface area. However, 1966). there was an increased activity of certain enzymes More direct observations on the relationship in the enterocytes (Riecken et al., 1965) suggest- between enterocytic hyperplasia and increased ing that the enhanced absorption may be intestinal function have been made in the rat related to increased activity of the individual (Dowling & Booth, 1967). In this animal, there mucosal cells.
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