Ultrastructure of the Kidney PDF
Preview Ultrastructure of the Kidney
ULTRASTRUCTURE IN BIOLOGICAL SYSTEMS Edited by Albert J. Dalton Francoise Haguenau VOLUME 1 / Tumors Induced by Viruses: Ultrastructure Studies, 1962 VOLUME 2 / Ultrastructure of the Kidney, 1967 IN PREPARATION: The Nucleus The Membranes ULTRASTRUCTURE OF THE KIDNEY Edited by Albert J. Dalton Francoise Haguenau PUBLIC HEALTH SERVICE LABORATOIRE DE MEDECINE EXPERIMENTALE NATIONAL CANCER INSTITUTE COLLEGE DE FRANCE BETHESDA, MARYLAND PARIS, FRANCE 1967 ACADEMIC PRESS NEW YORK and LONDON COPYRIGHT © 1967, BY ACADEMIC PRESS INC. ALL RIGHTS RESERVED. NO PART OF THIS BOOK MAY BE REPRODUCED IN ANY FORM, BY PHOTOSTAT, MICROFILM, OR ANY OTHER MEANS, WITHOUT WRITTEN PERMISSION FROM THE PUBLISHERS. ACADEMIC PRESS INC. Ill Fifth Avenue, New York, New York 10003 United Kingdom Edition published by ACADEMIC PRESS INC. (LONDON) LTD. Berkeley Square House, London W. 1 LIBRARY OF CONGRESS CATALOG CARD NUMBER : 66-16437 PRINTED BY THE ST. CATHERINE PRESS LTD., BRUGES, BELGIUM CONTRIBUTORS Numbers in parentheses indicate the pages on which the authors' contributions begin. Sergio A. Bencosme (143), Departments of Pathology and Medicine, Queen's Univ- ersity and the Kingston General Hospital, Kingston, Ontario, Canada Julia Creemers (57), Department of Internal Medicine, Universiteitsklinieken Sint Rafael, Louvain, Belgium, and the Department of Anatomy, University of Louvain, Belgium Pierre-Yves Hatt (101), Centre Hospitalier Emile Roux, Limeil-Brevannes, Seine et Oise, France and Laboratoire de Medecine experimentale du College de France, Paris, France Harrison Latta (1), Department of Pathology, School of Medicine, University of California, Los Angeles, California Arvid B. Maunsbach (1), Department of Pathology, Center for the Health Sciences, University of California, Los Angeles, California Max G. Menefee (73), Department of Anatomy, State University of New York, Upstate Medical Center, Syracuse, New York1 Paul A. Michielson (57), Department of Internal Medicine, Universiteitsklinieken, Sint Rafael, Louvain, Belgium, and the Department of Anatomy, University of Louvain, Belgium Peter A. F. Morrin (143), Departments of Pathology and Medicine, Queen's Univ- ersity and the Kingston General Hospital, Kingston, Ontario, Canada C. Barber Mueller (73), State University of New York, Upstate Medical Center? Syracuse, New York Lydia Osvaldo (1), Department of Pathology, Center for the Health Sciences, University of California, Los Angeles, California2 1 Present address: Department of Pathology, University of Cincinnati Medical Center, Cincinnati, Ohio. 2 Present address: Karolinska Stukhusets, Patologiska Avdelning, Stockholm, Sweden. v PREFACE The monograph series on "Ultrastructure in Biological Systems" was begun not simply to start another group of reviews on ultrastructure but to fill what was felt to be a need for summary statements of the current status of particular areas of ultrastructural research in which significant and rapid advances are being made and in which considerable controversy and discussion are continuing. The ultra- structure of the mammalian kidney is certainly one of these areas. One of the first applications of the electron microscope to problems in human pathology was to the kidney, and resulted in the demonstration of the hitherto undetected early thickening of the basement membrane of glomerular capillaries in glomerulonephritis. Yet many problems remain, particularly in relation to the correlation between function and the ultrastructure of components of the kidney— mesangium, glomerulus, juxtaglomerular apparatus, and the renal tubules. It is only very recently that the mesangium has come to be accepted as real, and many questions remain as to the function of its cells. The existence of true membranes between foot processes of the epithelial cells of glomeruli is a newly established fact; but what this has to do with glomerular filtration is not known at present. Granules apparently secretory in nature have been identified in cells of the juxtaglomerular apparatus, but so far their presence has not been correlated with specific functional change. Artifacts introduced at fixation are now known to have considerable relevance in interpreting the ultrastructure of the normal nephron. Partly because evidence for this is quite recent, the correlation between fine structure and function of parts of the nephron is still in its infancy. Contributions of electron microscopy to an understanding pathological conditions have been several, but it would appear that the most valuable future contributions will come from the application of some of the more sophisticated techniques such as radioautography and ferritin-labeled antibody. These are paraphrased views of the contributors to this monograph who, while acquainting the reader with the research being carried on in these areas, have also brought into focus the many problems still awaiting solution. ALBERT J. DALTON November, 1966 FRANCHISE HAGUENAU vii THE FINE STRUCTURE OF RENAL TUBULES IN CORTEX AND MEDULLA1 HARRISON LATTA, ARVID B. MAUNSBACH, AND LYDIA OSVALDO2 Department of Pathology, Center for the Health Sciences, and Department of Zoology, University of California, Los Angeles, California I. Introduction 2 II. Anatomy 3 III. Renal Physiology 7 A. Glomerular Filtration 7 B. Tubular Resorption 7 C. Tubular Secretion and Synthesis 9 IV. Proximal Tubule 9 A. Fixation 9 B. Plasma Membrane 11 C. Brush Border and Cilia 13 D. Attachment Zones 14 E. Plasma Membrane Invaginations and Apical Vacuoles 15 F. Cytomembranes (Endoplasmic Reticulum or Ergastoplasmic Membranes). . 16 G. Cytoplasmic Bodies 18 H. Mitochondria 22 I. Nuclei 23 J. Segmentation of the Proximal Tubule 23 V. Thin Segment of Loop of Henle 25 A. Thin Segments with Elaborate Basal Processes (Descending Type) 27 B. Thin Segments without Elaborate Basal Processes (Ascending Type) . . .. 28 VI. Distal Tubule 28 A. Ascending Part of Distal Tubule 28 B. Convoluted Part of Distal Tubule 30 1 The work from the authors' laboratories which is mentioned in this review has been supported in part by U.S. Public Health Service research grants A-06074, RG 5762, A 1702, and AM-07731. 2 While each author has contributed to each section, primary responsibility is as follows: H. Latta, Sections I, II, IX, XII; A. B. Maunsbach, Sections IV, VI, X, XT; L. Osvaldo, Sections III, V, VII, VIII. 1 2 HARRISON LATTA, ARVID B. MAUNSBACH, AND LYDIA OSVALDO VII. Collecting Tubule 31 A. Cortical Collecting Tubule 31 B. Collecting Tubule in Medulla 31 C. Papillary Duct 33 VIII. Other Structures in Cortex and Medulla 33 A. Vessels 33 B. Nerves 35 C. Interstitial Cells 37 IX. Embryology 38 X. Renal Phylogenetic Differences 39 XI. Other Tissues Transporting Salts or Fluids 40 XII. Physiological and Pathological Reactions 40 A. Fixation of the Kidney as a Whole 40 B. Autolysis, Ischemia, and Shock 41 C. Poisoning 41 D. Ions, Hormones, and Vitamins 42 E. Renal Homotransplantation 43 F. Various Disease Processes 44 G. Renal Tumors 45 XIII. Conclusion 45 References 45 I. Introduction The kidneys play a leading role in maintaining the internal environment of the body, the "milieu interieur," in the optimal state for cellular function. In the course of evolution the kidney of the early chordates developed the ability to conserve sodium, enabling these chordates to leave the ocean and invade freshwater streams. Later the kidney became able to conserve water also and the chordates moved into a drier environment on land (H. W. Smith, 1951, 1959). Of course, many other substances have come to be selectively retained or excreted as necessary. The localization of sites and the elucidation of mechanisms for handling each substance are challenges to investigators who must try to explain the chemical identification, analysis, and work performed by the kidney, as well as the alterations produced by disease—all in terms of the chemical composition and finer structures of the different renal cell types and their macromolecular components. Such an analysis will require clarification of the factors involved in filtration in the glomerulus, permeability, selective resorption, active and passive transport, and biochemical synthesis and secretion, whether into the bloodstream or into the urine. It is apparent that such an analysis will involve different fields including anatomy, histology, biochemistry, physiology, pharmacology, and pathology, as well as presenting challenges in the application and development of physical and chemical techniques at the macromolecular level. While renal processes cannot be fully understood until these intracellular processes are worked out, the high degree of structural and FINE STRUCTURE OF RENAL TUBULES IN CORTEX AND MEDULLA 3 functional differentiation in different parts of the nephron suggest that the kidney may well be the first organ in which will be clarified many of the secrets of cellular activity in transferring chemical energy of catabolism into the chemical work of active transport. More electron microscopic work has been done on the glomerulus than on the tubules for two apparent reasons. First, the present electron microscopic techniques give better pictures of the glomerulus in health and disease because the glomerulus is less altered by present preparative techniques and by the trauma of needle biopsy, if one is performed. Second, most human renal disease seems to affect the glomerulus primarily. Although the etiological processes in the glomerulus are manifold and obscure, the functional alterations seem relatively simple. Passive glomerular filtration is affected either by changes in the permeability of the glomerular capillary wall or by changes in the capillary blood pressure. Although the tubules are secondarily affected in most renal diseases, they have much more complex physiological functions which accomplish most of the work of the kidney. Moreover, it is apparent that they are relatively labile structures, rapidly changing in response to interference with the blood supply and to many of the procedures necessary to study them. This biological sensitivity has made the interpretation of many electron microscopic studies quite difficult and demands stringent efforts to control conditions and discover the more sensitive reaction mechanisms of the cell. This is why studies of technical variations necessarily precede an analysis of any but the grossest physiological and pathological alterations. On the other hand, this sensitivity might well be expected of cells required to perform constantly so much chemical and osmotic work. Moreover, reactions to changes in the environment during study and fixation are clues to cellular changes under physiological and pathological conditions. Certain elementary points are summarized in this review because some readers will be familiar with the kidney but not with electron microscopy, whereas others will be familiar with electron microscopy but not with the kidney. In order to keep within manageable proportions, the bibliography is not complete, but emphasizes reviews and recent references rather than the historical development of our knowledge. II. Anatomy The main anatomical features are given in histology and renal textbooks (von Mollendorff, 1930; Allen, 1962; Coupland, 1962). Only a few points will be mentioned here. The different parts of the nephron are listed in Table I, which has been modified from von Mollendorff (1930), Sjostrand (1944), and Rhodin (1958). The corresponding names in several languages are given in order to facilitate translation of foreign papers. The same part sometimes has different names in the same language; some of the equivalents are given in parentheses. In English, "part," "portion," and "segment" are used interchangeably. The division of the nephron given in Table I refers primarily 4 TABLE I DIFFERENT PARTS OF THE NEPHRON AND CORRESPONDING NAMES IN FOREIGN LANGUAGES H A R R IS English German Latin French Spanish O N L A T T Glomerulus Nierenkorperchen Corpusculum renis Glomerule Glomerulo A , (renal corpuscle) (corpusculo renal) A R V (Malpighian corpuscle) (corpusculo de Malpighi) ID B . M Neck Hals Col Cuello A U N S B A C Proximal tubule Hauptstuck Portio principalis Tube contourne proximal Tubo contorneado proxi- H , (I) mal A N Convoluted (proximal) gewundener Teil pars convoluta la portion contournee D part porcion contorneada LY D Terminal part (straight gestreckter Teil pars recta (pars medul- la portion rectiligne IA part) laris) porcion recta OS V A L D O Thin segment (limb) of Uberleitungsstuck (diinner Pars conducens La branche grele de l'anse Porcion delgada del asa de Henle's loop Teil der Henleschen de Henle (segment grele) Henle Schleife) Distal tubule Mittelstuck Portio intermedia Tube contourne distale (II) Tubo contorneado distal (segment intermediare) Ascending thick segment gestreckter Teil (auf- pars recta la branche large (as- porcion gruesa de la F (limb) of Henle's loop steigender Schenkel der cendante) de l'anse de rama ascendente del asa IN E Henleschen Schleife, Henle (portion epaisse, de Henle S T breiter Teil) anse large) RU C Medullary part in Markstrahl pars medullaris T U Cortical part in der Aussenzone pars corticalis R E O Convoluted part gewundener Teil pars convoluta segment contourne porcion contorneada F R Macula densa Zwischenstiick pars maculata macula densa macula densa E N Intercalated part Schaltstiick pars intercalata piece intermediare porcion intercalar A L Connecting part Verbindungsstiick pars reuniens canal d'union T U B U L E Collecting tubule (excre- Sammelrohrsystem Pars colligens Tube collecteur (excreteur) Tubo collector S tory duct) IN C Arched collecting tubule proximaler Teil — tubo collector arqueado O R (proximal part, connec- — (porcion cortical) TE X ting tubule, cortical col- — A ting tubule) — N D Straight collecting tubule distaler Teil — tube droit tubo collector recto M E D Part in cortex — — porcion cortical U L Part in outer medulla — — porcion medular LA Papillary duct of — ductus papillaris canal papillaire conductos de Bellini Bellini (Bellini) 5
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