(BQ) Part 1 book Histology text and atlas presents the following contents: Methods, cell cytoplasm, the cell nucleus, tissues - Concept and classification, epithelial tissue, connective tissue, adipose tissue, nerve tissue, cardiovascular system,...
72006_FM 15/07/10 6:59 PM Page vi This page intentionally left blank 72006_FM 15/07/10 6:59 PM Page i Sixth Edition HISTOLOGY A Text and Atlas with Correlated Cell and Molecular Biology 72006_FM 15/07/10 6:59 PM Page ii Michael H Ross (1930–2009) 72006_FM 15/07/10 6:59 PM Page iii Sixth Edition HISTOLOGY A Text and Atlas with Correlated Cell and Molecular Biology Michael H Ross, PhD (deceased) Professor and Chairman Emeritus Department of Anatomy and Cell Biology University of Florida College of Medicine Gainesville, Florida Wojciech Pawlina, MD Professor and Chair Department of Anatomy Department of Obstetrics and Gynecology Assistant Dean for Curriculum Development and Innovation Mayo Medical School College of Medicine, Mayo Clinic Rochester, Minnesota 72006_FM 15/07/10 6:59 PM Page iv Acquisitions Editor: Crystal Taylor Product Manager: Jennifer Verbiar Designer: Doug Smock Compositor: MPS Limited, A Macmillan Company Sixth Edition Copyright © 2011 Lippincott Williams & Wilkins, a Wolters Kluwer business 351 West Camden Street Baltimore, MD 21201 Two Commerce Square 2001 Market Street Philadelphia, PA 19103 All rights reserved This book is protected by copyright No part of this book may be reproduced or transmitted in any form or by any means, including as photocopies or scanned-in or other electronic copies, or utilized by any information storage and retrieval system without written permission from the copyright owner, except for brief quotations embodied in critical articles and reviews Materials appearing in this book prepared by individuals as part of their official duties as U.S government employees are not covered by the above-mentioned copyright To request permission, please contact Lippincott Williams & Wilkins at 530 Walnut Street, Philadelphia, PA 19106, via email at permissions@lww.com, or via website at lww.com (products and services) Printed in China Library of Congress Cataloging-in-Publication Data Ross, Michael H Histology: a text and atlas: with correlated cell and molecular biology/Michael H Ross, Wojciech Pawlina.—6th ed p ; cm Includes bibliographical references and index ISBN 978-0-7817-7200-6 (alk paper) Histology Histology—Atlases I Pawlina, Wojciech II Title [DNLM: Histology—Atlases QS 517 R825h 2011] QM551.R67 2011 611’.018—dc22 2010024700 DISCLAIMER Care has been taken to confirm the accuracy of the information present and to describe generally accepted practices However, the authors, editors, and publisher are not responsible for errors or omissions or for any consequences from application of the information in this book and make no warranty, expressed or implied, with respect to the currency, completeness, or accuracy of the contents of the publication Application of this information in a particular situation remains the professional responsibility of the practitioner; the clinical treatments described and recommended may not be considered absolute and universal recommendations The authors, editors, and publisher have exerted every effort to ensure that drug selection and dosage set forth in this text are in accordance with the current recommendations and practice at the time of publication However, in view of ongoing research, changes in government regulations, and the constant flow of information relating to drug therapy and drug reactions, the reader is urged to check the package insert for each drug for any change in indications and dosage and for added warnings and precautions This is particularly important when the recommended agent is a new or infrequently employed drug Some drugs and medical devices presented in this publication have Food and Drug Administration (FDA) clearance for limited use in restricted research settings It is the responsibility of the health care provider to ascertain the FDA status of each drug or device planned for use in their clinical practice To purchase additional copies of this book, call our customer service department at (800) 638-3030 or fax orders to (301) 223-2320 International customers should call (301) 223-2300 Visit Lippincott Williams & Wilkins on the Internet: http://www.lww.com Lippincott Williams & Wilkins customer service representatives are available from 8:30 am to 6:00 pm, EST 72006_FM 15/07/10 6:59 PM Page v This edition is dedicated to my wife Teresa Pawlina whose love, patience, and endurance created safe havens for working on this project and to my children Conrad Pawlina and Stephanie Pawlina whose stimulation and excitement have always kept my catecholamine levels high 72006_FM 15/07/10 6:59 PM Page vi This page intentionally left blank 72006_FM 15/07/10 7:00 PM Page vii Preface This sixth edition of Histology: A Text and Atlas with Correlated Cell and Molecular Biology continues a tradition of providing medical, dental, and allied health science students with a textual and visual introduction to histology with correlative cell biology As in previous editions, this book is a combination “text-atlas” in that standard textbook descriptions of histologic principles are supplemented by illustrations and photographs In addition, separate atlas sections follow each chapter and provide large-format, labeled atlas plates with detailed legends highlighting elements of microanatomy Histology: A Text and Atlas is therefore “two books in one.” Significant modifications have been made in this edition in order to create an even more useful and understandable approach to the material: Updated cellular and molecular biology Material introduced in the fifth edition has been updated to include the latest advancements in cellular and molecular biology The sixth edition focuses on selected information to help students with overall comprehension of the subject matter To accommodate reviewers’ suggestions, the sixth edition also integrates new cell biology information into several chapters For instance, the cell biology of endothelial cells has been added to the discussion of the cardiovascular system; a section on primary cilia, including their structure and function, was added to the epithelial tissue chapter; a new clinical nomenclature for cells involved in hemopoiesis and a detailed description of the respiratory burst reaction in neutrophils were added to the chapter on blood; new information and diagrams of nerve fiber regeneration were added to the nerve tissue chapter; and the cell biology of taste receptors was incorporated into the chapter on the digestive system Reader-friendly innovations The book has been redesigned in an attempt to provide more ready access to important concepts and essential information Additional color font is used in the body of the text Important concepts are listed as sentence headings Features of cells, tissues, and organs and their functions, locations, and other relevant short phrases are formatted as bulleted lists that are clearly identifiable in the body of the text by oversized color bullets Essential terms within each specific section are introduced in the text in an eye-catching oversized red bolded font that clearly stands out from the remaining black text Text containing clinical information or the latest research findings is presented in blue, with terminology pertaining to diseases, conditions, symptoms, or causative mechanisms in oversized bolded blue The clinical sections of the text are easily found within each chapter Emphasis on features Many of the pedagogic features from the last edition have been refined, and some new features have been added: • More summary tables are included to aid students in learning and reviewing material without having to rely on strict memorization of data These include a review table of the specializations in the apical domains of epithelial cells and a table of features of adipose tissue Many tables have been updated and modified • Previous clinical and functional correlations boxes have been replaced with Clinical Correlation and Functional Consideration Folders More new folders have been added to each chapter, and existing folders have been redesigned, updated, enhanced, and illustrated with new diagrams and images of clinical specimens New folders contain clinical information related to the symptoms, photomicrographs of diseased tissues or organs, short histopathological descriptions, and treatment of specific diseases Important terms have been highlighted with oversized bolded text While the information in these folders might be considered ancillary material, it demonstrates the functional impact and clinical significance of histology • More Atlas Plates have been added to the atlas section at the end of each chapter Several orientation micrographs were added to the summary box in the atlas section Atlas plates for the blood chapters have been completely redesigned so as to show both mature forms of blood cells and the stages through which they pass during hemopoiesis Many plates have been replaced with vibrant digital images • More new figures and illustrations have also been added, and about one-third of all old figures have been redrawn for vii 72006_FM 15/07/10 7:00 PM Page viii greater clarity and conceptual focus This sixth edition incorporates many new clinical images and photomicrographs to illustrate information in the clinical correlation folders Many new high-resolution digital photomicrographs have been integrated into each chapter • New design A bright, energetic text design sets off the new illustrations and photos and makes navigation of the text even easier than in previous editions As in the last five editions, all of the changes were undertaken with student needs in mind; namely, to understand the subject matter, to become familiar with the latest information, and to be able to practically apply newfound knowledge Wojciech Pawlina 72006_ch13 15/07/10 3:18 PM Page 425 Muscular venules are distinguished from postcapillary venules by the presence of a tunica media thoroughfare channel Muscular venules are located distal to the postcapillary precapillary sphincters metarteriole arteriole venule lymphatic capillary FIGURE 13.23 anchoring filaments • Diagram of microcirculation This schematic is usually patent; that of the vein is often collapsed Many veins, especially those that convey blood against gravity, such as those of the limbs, contain valves that allow blood to flow in only one direction, back toward the heart The valves are semilunar flaps consisting of a thin connective tissue core covered by endothelial cells Venules and Small Veins Postcapillary venules collect blood from the capillary network and are characterized by the presence of pericytes Postcapillary venules possess an endothelial lining with its basal lamina and pericytes (Plate 35, page 438) The endothelium of postcapillary venules is the principal site of action of vasoactive agents such as histamine and serotonin Response to these agents results in extravasation of fluid and emigration of white blood cells from the vessel during inflammation and allergic reactions Postcapillary venules of lymph nodes also participate in the transmural migration of lymphocytes from the vascular lumen into the lymphatic tissue Pericytes form umbrella-like connections with the endothelial cells The relationship between endothelial cells and pericytes promote their mutual proliferation and survival Both synthesize and share the basal lamina (see Fig 13.21), synthesize growth factors, and communicate with each other through tight and gap junctions Pericyte coverage is more extensive in the postcapillary venules than capillaries The postcapillary venules in the lymph nodes are also called high endothelial venules (HEVs) because of the prominent cuboidal appearance of their endothelial cells and their ovoid nuclei Medium veins have a diameter of as much as 10 mm Most deep veins that accompany arteries are in this category (e.g., radial vein, tibial vein, popliteal vein) Valves are a characteristic feature of these vessels and are most numerous in the inferior portion of the body, particularly the lower limbs, to prevent retrograde movement of blood because of gravity Often deep veins of lower limbs are the site of thrombus (blood clot) formation, a condition known as deep venous thrombosis (DVT) DVT is associated with immobilization of the lower limbs due to prolonged bed rest (after surgery or hospitalization), orthopedic casts, or restricted movement during long-haul flights DVT can be a life-threatening condition because of the potential for development of pulmonary embolism (blockage of the pulmonary arteries) by a dislodged blood clot originating from deep veins The three tunics of the venous wall are most evident in medium-sized veins (Fig 13.24) The tunica intima consists of an endothelium with its basal lamina, a thin subendothelial layer with occasional smooth muscle cells scattered in the connective tissue elements, and, in some cases, a thin internal elastic membrane The tunica media of medium-sized veins is much thinner than the same layer in medium-sized arteries It contains several layers of circularly arranged smooth muscle cells with interspersed collagen and elastic fibers In addition, longitudinally arranged smooth muscle cells may be present just beneath the tunica adventitia The tunica adventitia is typically thicker than the tunica media and consists of collagen fibers and networks of elastic fibers (Fig 13.24b) • • • Large Veins In large veins, the tunica media is relatively thin, and the tunica adventitia is relatively thick Veins with a diameter greater than 10 mm are classified as large veins tunica intima of these veins (Fig 13.25 and Plate 34, • The page 436) consists of an endothelial lining with its basal lamina, a small amount of subendothelial connective tissue, and some smooth muscle cells Often the boundary between the tunica intima and tunica media is not clear, and it is not always easy to decide whether the smooth muscle cells close to the intimal endothelium belong to the tunica intima or to the tunica media 425 425 Cardiovascular System ᭹ VE I N S diagram shows a metarteriole (initial segment of a thoroughfare channel) giving rise to capillaries The precapillary sphincters of the arteriole and metarteriole control the entry of blood into the capillaries The distal segment of the thoroughfare channel receives capillaries from the microcirculatory bed, but no sphincters are present where the afferent capillaries enter the thoroughfare channels Blind-ended lymphatic vessels are shown in association with the capillary bed Note the presence of anchoring filaments and the valve system within the lymphatic capillaries Medium Veins chapter 13 venules in the returning venous network and have a diameter of as much as 0.1 mm Whereas postcapillary venules have no true tunica media, the muscular venules have one or two layers of smooth muscle that constitute a tunica media These vessels also have a thin tunica adventitia Usually, pericytes are not found in muscular venules 72006_ch13 15/07/10 3:18 PM Page 426 tunica adventitia tunica media elastic fibers tunica intima bundle of smooth muscles endothelial cells collagen fibers basal lamina smooth muscle cells macrophage fibroblast a unmyelinated nerves MEDIUM-SIZED VEIN FIGURE 13.24 • Schematic diagram and photomicrograph of a medium-sized vein a The cellular and extracellular components are labeled Note that the tunica media contains a few layers of circularly arranged smooth muscle cells with interspersed collagen and elastic fibers Also, longitudinally arranged smooth muscle cells are present at the junction with the tunica adventitia b This photomicrograph shows a section through the wall of a medium-sized vein in routine H&E preparation The tunica intima consists of endothelium and a very thin subendothelial layer of connective tissue containing some smooth muscle cells The tunica media contains a few layers of circularly and spirally arranged smooth muscle cells with collagen and elastic fibers Note that the thickest layer is the tunica adventitia, which contains an abundance of collagen and some elastic fibers The few nuclei seen in this layer belong to fibroblasts ϫ360 tunica media is relatively thin and contains circum• The ferentially arranged smooth muscle cells, collagen fibers, • and some fibroblasts In some animals, but not in humans, cardiac muscle cells extend into the tunica media of the both superior and inferior vena cava and the pulmonary veins, near their junction with the heart The tunica adventitia of large veins (e.g., the subclavian veins, portal vein, and the venae cavae) is the thickest layer of the vessel wall Along with the usual collagen and elastic fibers and fibroblasts, the tunica adventitia also contains longitudinally disposed smooth muscle cells (Fig 13.26) ATYPICAL BLOOD VESSELS In several locations in the body, blood vessels—both arteries and veins with an atypical structure are present These include the following Coronary arteries, considered to be medium-sized muscular arteries, originate from the proximal part of the ascending aorta and lie on the surface of the heart in • 426 • • the epicardium surrounded by adipose tissue The walls of coronary arteries are usually thicker than those of comparable arteries on the upper or lower limb because of the large amounts of circular smooth muscle layers in the tunica media In routine H&E preparation, the subendothelial layer of the tunica intima of younger people is inconspicuous, but it progressively thickens by increasing amounts of smooth muscle cell and fibroelastic tissue with aging (Fig 13.27) The internal elastic membrane is well developed, although it may be fragmented, duplicated, or focally lost in older individuals The relatively “loose” consistency of the tunica adventitia is reinforced by the longitudinal bundles of collagen fibers that allow for continuous changes of the vascular diameter Atherosclerotic changes in coronary arteries that restrict blood flow and oxygen supply to cardiac muscle lead to ischemic heart disease (see Folder 13.3) Dural venous sinuses represent venous channels in the cranial cavity They are essentially broad spaces within the dura mater that are lined with endothelial cells and devoid of smooth muscles The great saphenous vein represents a long subcutaneous vein of the lower limb that originates in the foot and 72006_ch13 15/07/10 3:18 PM Page 427 tunica adventitia tunica media collagen fibers tunica intima chapter 13 endothelial cells basal lamina bundles of smooth muscle cells 427 a fibroblast LARGE VEIN FIGURE 13.25 • Schematic diagram and photomicrograph of a large vein a The cellular and extracellular components are labeled Note a thin layer of circumferentially arranged smooth muscles of tunica media and the tunica adventitia with a large amount of longitudinally arranged smooth muscle bundles b This photomicrograph shows a section through the wall of a human portal vein in a routine H&E preparation The tunica intima is indiscernible at this magnification The tunica media contains a layer of circumferentially arranged smooth muscle cells with collagen and elastic fibers Note the thickest layer of this wall is the tunica adventitia In addition to an extensive collagen and elastic fiber network, the tunica adventitia contains a broad layer of smooth muscle cells arranged in longitudinal bundles These bundles are variable in size and separated from each other by connective tissue fibers ϫ125 (Courtesy of Dr Donald J Lowrie Jr., University of Cincinnati College of Medicine.) • drains into the femoral vein just below the inguinal ligament This vein is often described as a muscular vein because of the presence of an unusual amount of smooth muscle (Fig 13.28) In addition to the thick circular arrangement of smooth muscle in its tunica media, the great saphenous vein possesses numerous longitudinal smooth muscle bundles in the intima and in the welldeveloped adventitia A thin, poorly developed internal elastic membrane separates the tunica intima from the media The great saphenous vein is frequently harvested from the lower limb and used for autotransplantation in coronary artery bypass graft (CABG) surgery when arterial grafts (usually taken from internal thoracic artery) are not available or many grafts are required for multiplebypass anastomoses CABG is one of the most commonly performed major surgical operations in the United States The central adrenomedullary vein that passes through the adrenal medulla and its tributaries have an unusual tunica media It contains several longitudinally oriented bundles of smooth muscle cells that vary in size and apperance (Fig 13.29) These irregularly arranged smooth muscle bundles (also called muscle cushions) extend into larger tributaries of the central adrenomedullary vein This unique eccentric arrangement of smooth muscle bundles results in the irregularity in the thickness of the vascular wall In areas where muscle bundles are absent, cells of the adrenal medulla or sometimes adrenal cortex are separated from the lumen of the vein only by a thin layer of the tunica intima (see Fig 13.29) Contraction of the longitudinally arranged smooth muscles in the tunica media enhances the efflux of hormones from the adrenal medulla into the circulation Veins in certain other locations (e.g., retina, placenta, trabeculae of the spleen) also have atypical walls and are discussed in the chapters that describe these organs LYMPHATIC VESSELS Lymphatic vessels convey fluids from the tissues to the bloodstream In addition to blood vessels, another set of vessels circulates fluid called lymph through most parts of the body These lymph-carrying vessels serve as adjuncts to the blood vessels Unlike the blood vessels, which convey blood to and from tissues, the lymphatic vessels are unidirectional, conveying fluid 427 Cardiovascular System ᭹ LYM P HATI C VE S S E LS unmyelinated nerves 72006_ch13 15/07/10 3:18 PM Page 428 endothelium TA tunica intima CM TM tunica media TI IEM tunica adventitia longitudinal bundles of smooth muscle cells 13.26 • Photomicrograph of a large vein This photomicrograph shows the three tunics in a section through the wall of the portal vein stained with H&E The tunica intima consists of endothelium and a thin subendothelial layer of connective tissue containing a few smooth muscle cells The tunica media contains relatively thin layer of circularly arranged smooth muscle cells Tunica adventitia is the thickest layer of this vessel It contains a thick layer of longitudinally arranged smooth muscle bundles (seen here in cross section) separated by collagen and elastic fibers Note a layer of connective tissue containing coarse collagen and elastic fibers that separates longitudinal bundles of smooth muscle in the tunica adventitia from a layer of smooth muscles of the tunica media ϫ240 (Courtesy of Dr Donald J Lowrie Jr., University of Cincinnati College of Medicine.) FIGURE only from tissues The smallest lymphatic vessels are called lymphatic capillaries They are especially numerous in the loose connective tissues under the epithelium of the skin and mucous membranes The lymphatic capillaries begin as “blind-ended” tubes in the microcapillary beds (see Fig 13.23) Lymphatic capillaries converge into increasingly larger vessels called lymphatic vessels They ultimately unite to form two main channels that empty into the blood vascular system by draining into the large veins in the base of the neck Lymph enter the vascular system at the junctions of the internal jugular and subclavian veins The largest lymphatic vessel, draining most of the body and emptying into the veins on the left side, is the thoracic duct The other main channel is the right lymphatic trunk Lymphatic capillaries are more permeable than blood capillaries and collect excess protein-rich tissue fluid 428 FIGURE 13.27 • Photomicrograph of the coronary artery This photomicrograph of a cross section of the coronary artery obtained from adult human shows all three vascular tunics similar to those in muscular arteries The subendothelial layer of the tunica intima (TI) is considerably thicker because of the aging process than a comparable muscular artery The internal elastic membrane (IEM) is visible at the border with the tunica media (TM), which is also thicker than in other muscular-type arteries Connective tissue of the tunica adventitia (TA) is loosely arranged and contains peripherally positioned longitudinal bundles of collagen fibers There is an artificial separation between cardiac muscle (CM) and tunica adventitia ϫ175 Lymphatic capillaries are a unique part of the circulatory sys- tem, forming a network of small vessels within the tissues Because of their greater permeability, lymphatic capillaries are more effective than blood capillaries in removing protein-rich fluid from the intercellular spaces Once the collected fluid enters the lymphatic vessel, it is called lymph Lymphatic vessels also serve to convey proteins and lipids that are too large to cross the fenestrations of the absorptive capillaries in the small intestine Before lymph is returned to the blood, it passes through lymph nodes, where it is exposed to the cells of the immune system Thus, the lymphatic vessels serve not only as an adjunct to the blood vascular system but also as an integral component of the immune system Lymphatic capillaries are essentially tubes of endothelium that, unlike the typical blood capillary, lack a continuous basal lamina This incomplete basal lamina can be correlated with their high permeability Anchoring filaments extend between the incomplete basal lamina and the perivascular collagen These filaments may help maintain the patency of the vessels during times of increased tissue pressure such as in inflammation 72006_ch13 15/07/10 3:18 PM Page 429 endothelium SM tunica intima SM SM SM adrenal medulla tunica media * central adrenomedullary vein chapter 13 SM SM tunica adventitia As lymphatic vessels become larger, the wall becomes thicker The increasing thickness is because of connective tissue and bundles of smooth muscle Lymphatic vessels possess valves that prevent backflow of the lymph, thus aiding unidirectional flow (Plate 35, page 438) There is no central pump in the lymphatic system Lymph moves sluggishly, driven primarily by compression of the lymphatic vessels by adjacent skeletal muscles adrenal cortex tributary vein SM FIGURE 13.29 • Photomicrograph of the central adrenomedullary vein This photomicrograph of the human adrenal gland shows a large central adrenomedullary vein with its tributary stained with H&E The wall of the vein is highly irregular, containing several longitudinally oriented bundles of smooth muscles (SM) that extend into the wall of a tributary This unique eccentric arrangement of smooth muscles, sometimes called muscle cushions, results in the irregularity in the thickness of the vascular wall Note that in the cleft between two smooth muscle bundles (asterisk), the lumen of the vein is separated from the chromaffin cells of the adrenal medulla only by the tunica intima On the opposite side of the wall, muscle bundles are absent (arrowheads) and the cells of the adrenal cortex are in direct contact with the tunica intima ϫ120 (Courtesy of Dr Donald J Lowrie Jr., University of Cincinnati College of Medicine.) • FOLDER 13.3 Clinical Correlation: Ischemic Heart Disease Ischemic heart disease or ischemic cardiomyopathy is defined as the imbalance between supply and demand of the heart for oxygenated blood Ischemic heart disease is the most common type of heart disease in the United States and affects approximately in every 100 people The most common cause of ischemic heart disease is atherosclerosis The risk of developing atherosclerosis increases with age, family history, hypertension, cigarette smoking, hypercholesterolemia, and diabetes In atherosclerosis, the lumina of the coronary arteries progressively narrow because of the accumulation of lipids, extracellular matrix, and cells, leading to the development of atheromatous plaques (Fig F13.3.1) Plaques are formed by intracellular and extracellular lipid deposition, smooth muscle proliferation, and increased synthesis of proteoglycans and collagen within the intima of the vessel wall Blood flow continued next page 429 Cardiovascular System ᭹ LYM P HATI C VE S S E LS FIGURE 13.28 • Photomicrograph of the great saphenous vein This photomicrograph shows a section through the wall of the greater saphenous vein Tunica intima is usually thicker than in the other medium size veins and is characterized by a presence of numerous longitudinal smooth muscle bundles (SM) separated by connective tissue fibers Tunica media contains relatively thick layer of circularly arranged smooth muscles Tunica adventitia is well developed and contains additional layers of smooth muscle fibers arranged in spiral, oblique, and longitudinal bundles ϫ380 (Courtesy of Dr Joseph J Maleszewski, Mayo Clinic, Rochester, MN.) 429 SM tunica intima 72006_ch13 15/07/10 3:18 PM Page 430 FOLDER 13.3 Clinical Correlation: Ischemic Heart Disease (Cont.) TM IEM TA TM TI TA TI thrombus fatty streak atheromatous plaque FIGURE F13.3.1 • Photomicrograph of an atheromatous plaque in the coronary artery This low magnification photomicrograph shows a cross section of the human coronary artery with chronic ischemic heart disease The specimen is stained with the Verhoeff van Gieson technique for elastic and connective tissue fibers The black strands represent elastic lamellae; a distinct intact internal elastic membrane (IEM) is present between the dark red-stained tunica media (TM) containing smooth muscle cells and the pathologically changed tunica intima (TI) Variable shades of pink material represent collagen fibers deposited in a thick tunica intima, which contains advanced atheromatous plaque with visible calcifications (dark pink–orange color) and accumulation of extracellular lipids (cholesterol clefts) The light pink color surrounding the lumen of the vessel represents the most recent deposition of the pathological material Note that the lumen of the vessel is occluded almost 90%, which led to inadequate coronary blood flow Tunica adventitia (TA) represents an outermost layer of the vessel ϫ34 (Courtesy of Dr William D Edwards, Mayo Clinic, Rochester, MN.) becomes critical when it is reduced by 90% or more Sudden occlusion of the narrowed lumen by a thrombus (blood clot) released from the surface of an atheromatous plaque precipitates an acute ischemic event Ischemic events are 430 FIGURE F13.3.2 • Photomicrograph of the coronary artery with a mural thrombus This photomicrograph shows a cross section of the coronary artery in a less advanced stage of the atherosclerotic disease The fibrofatty plaque is visible in the tunica intima (TI ) and developed thrombus superimposed on a plaque partially obstructing the arterial lumen Dashed line indicates the border between the tunica intima and tunica media (TM ) The tunica adventitia (TA) forms an outermost layer of the vessel ϫ40 (Courtesy of Dr William D Edwards, Mayo Clinic, Rochester, MN.) characterized by anginal pain associated with loss of oxygenated blood flow to the region of the heart supplied by the affected coronary vessel Coronary artery thrombosis usually precedes and precipitates a myocardial infarct—that is, a sudden insufficiency of blood supply that results in an area of muscle cell death Mural thrombus may develop and is usually associated with dysfunctional or ruptured endothelium overlying atheromatous plaque (Fig F13.3.2) With time, the area of the heart affected by the myocardial infarct heals A scar forms and replaces the damaged tissue, but the area of infarction loses contractile function Multiple infarctions over time can produce sufficient loss of cardiac function to cause death Infarction also commonly occurs in the brain, spleen, kidney, lung, intestine, testes, and tumors (especially of the ovaries and uterus) 72006_ch13 15/07/10 3:18 PM Page 431 This page intentionally left blank 72006_ch13 15/07/10 3:18 PM Page 432 ᭹ PLATE 32 Heart The cardiovascular system is a transport system that carries blood and lymph to and from the tissues of the body The cardiovascular system includes the heart, blood vessels, and lymphatic vessels Blood vessels provide the route by which blood circulates to and from all parts of the body The heart pumps the blood Lymphatic vessels carry tissue-derived fluid, called lymph, back to the blood vascular system The heart is a four-chambered organ consisting of a right and left atrium and a right and left ventricle Blood from the body is returned to the right atrium from which it enters the right ventricle Blood is pumped from the right ventricle to the lungs for oxygenation and returns to the left atrium Blood from the left atrium enters the left ventricle from which it is pumped to the rest of the body, i.e., the systemic circulation The heart, which differentiates from a straight vascular tube in the embryo, has the same basic three-layered structure in its wall as the blood vessels above the level of capillaries and postcapillary venules In the blood vessels, the three layers are called the tunica intima, including the vascular endothelium and its underlying connective tissue; the tunica media, a muscular layer that varies in thickness in arteries and veins; and the tunica adventitia, the outermost layer of relatively dense connective tissue In the heart, these layers are called the endocardium, the myocardium, and the epicardium, respectively Atrioventricular septum, heart, human, H&E ì45; inset ì125 P LATE 32 H E ART 432 This micrograph of the field shows portions of the atrial (A) and ventricular (V) walls at the level of the atrioventricular septum and the root of the mitral valve (MV) Both chambers and the valve are lined with the squamous endothelium of the endocardium (En) Purkinje fibers (PF) of the cardiac conduction system are seen in the atrial wall between the relatively thin subendocardial connective tissue (CT) and the underlying modified cardiac muscle cells (CM) of the atrioventricular node (AVN) Dense fibrous connective tissue (DCT) that is continuous with that of the septum and the subendocardial layers of the atrium and ventricle extends from the root of the valve into the leaflet Thin cardiac muscle fibers can also be seen extending from the wall Coronary artery and cardiac vein, heart, human, H&E ×30 This micrograph shows cross sections of a coronary artery and cardiac vein in the coronary sulcus The surrounding adipose tissue (AT) serves to cushion the blood vessels that run in the coronary sulcus The coronary artery (CA) in the lower left of this micrograph is surrounded by small bundles of small cardiac muscle cells of the atrium into the upper portion of the valve Inset This higher-magnification view of the field outlined by the rectangle (turned ~90°) shows more clearly the endothelial layer of the endocardium (En) and the dense fibrous connective tissue of the endocardium (DCT) and subendocardial layer A thin layer of smooth muscle (SM) appears between the more densely packed fibrous tissue immediately subjacent to the endothelium and the more loosely packed dense fibrous tissue of the subendocardium Particularly evident are the longitudinally sectioned Purkinje fibers (PF) of the cardiac conduction system These modified cardiac muscle cells contain the same fibrillar contractile system as their smaller counterparts in the myocardium, but the fibrils are fewer, are more loosely packed, and often surround what appear to be vacuolated areas Intercalated discs (ID), typical of cardiac muscle cell organization, are evident in some areas (CM) that are part of the atrioventricular node (AVN) A loop of the conduction bundle (CB) containing Purkinje fibers is evident to the right of the artery The darkly stained tunica intima (TI) is delimited by an internal elastic membrane (IEM) that is easily distinguished even at this relatively low magnification The thick muscular tunica media (TM) is also easily distinguished from the thinner, fibrous tunica adventitia (TA) A smaller arterial vessel (AЈ) KEY A, atrium A, small artery AT, adipose tissue AVN, atrioventricular node B, blood CA, coronary artery CB, conduction bundle CM, cardiac muscle CT, connective tissue CV, cardiac vein DCT, dense connective tissue En, endothelium ID, intercalated disc IEM, internal elastic membrane LN, lymph node MV, mitral valve PF, Purkinje fibers SM, smooth muscle TA, tunica adventitia TM, tunica media TI, tunica intima V, ventricle 72006_ch13 15/07/10 3:18 PM Page 433 En CM MV En DCT CT PF A CM AVN 433 V P LATE 32 En PF DCT SM • DCT H EART ID AT AVN CM CV B LN CA CB TI IEM TM TA TI B 72006_ch13 15/07/10 3:18 PM Page 434 ᭹ PLATE 33 Aorta The aorta, the main systemic artery of the body, is an elastic artery The presence of numerous fenestrated elastic lamellae allows it to resist the pressure variations caused by rhythmic contraction of the left ventricle The intima is comparatively much thicker than that seen in muscular arteries The subendothelial layer of the intima consists of connective tissue with both collagen and elastic fibers The cellular component consists of smooth muscle cells and fibroblasts The external border of the intima is bounded by an internal elastic membrane that represents the first layer of the many concentric fenestrated laminae in the media of the vessel The media constitutes the bulk of the wall Between the elastic laminae are collagen fibers and smooth muscle cells The latter are responsible for the synthesis of collagen and elastic fibers With age, the number and thickness of elastic laminae in the wall increases By 35 years as many as 60 laminae are found in the thoracic aorta At approximately 50 years, individual laminae begin to show signs of degeneration and gradually become replaced by collagen leading to a gradual loss of elasticity of the aortic wall The adventitia consists of irregular dense connective tissue with intermixed elastic fibers that tend to be organized in a circumferential pattern It also contains small blood vessels that supply the outer portion of the media They are the vasa vasorum of the aorta Also present in the adventitia are lymphatic capillaries ORIENTATION MICROGRAPHS: The upper micrograph shows a cross section of an H&E stained human aorta from a child The intima (I) stains considerably lighter than the adjacent media (M) The adventitia (A) contains an abundance of collagenous fibers and stains more densely than that of either the media or intima The lower micrograph is from an adult and has been stained to reveal the elastic component of the vessel wall The intima (I) is very lightly stained, in this case, due to the paucity of elastic material The media (M) is heavily stained due to the presence of large amounts of elastic laminae The adventitia (A) contains in addition to the dense connective tissue a moderate amount of elastic fibers M A I I 434 M P LATE 33 AORTA A Aorta, human, H&E, ì365; inset ×700 This micrograph shows the layers of the aortic wall The intima consists of an endothelium (En) overlying loose connective tissue (LCT) The thickest portion of the vessel wall is the media (M) The wavy eosinophilic material is the collagenous fibers The eosin stain does not reveal the elastic laminae The nuclei are those of smooth muscle cells Fibroblasts are absent The outer Aorta, human, iron hematoxylin and aniline blue, ×255; inset ×350 The specimen shown here has been stained to distinguish collagen from elastic material The intima (I) consists mostly of collagenous fibers The endothelium (En) represented by several nuclei is just barely evident The media (M) contains numerous elastic lamellae that appear as the black wavy lines The intervening blue stained material consists of collagen fibers Careful examination of the Aorta, human, iron hematoxylin and aniline blue, ×255 This micrograph shows the outer portion of the media (M) with its elastic lamellae The major portion of the micrograph is the adventitia (A) Here, the thick collagenous fibers (CF) are readily recognized The outer portion of the adventitia layer of the vessel wall is the adventitia (A) The eosinophilic material here consists of dense connective tissue The nuclei that are evident belong to fibroblasts Also note the small blood vessel (BV) in the adventitia The inset shows the intima at higher magnification and includes part of the media Note the endothelium (En) The eosinophilic material in the intima consists of collagenous fibers (CF) The main cell type here is the smooth muscle cell (SMC) media reveals nuclei of smooth muscle cells dispersed between the elastic lamellae The inset shows the intima at higher magnification Note the nuclei of the endothelial cells (EnC) at the luminal surface The remainder of the intima consists mostly of collagenous fibers (stained blue) with occasional elastic fibers (EF) identified by their darker coloration The nuclei of the fibroblasts and occasional smooth muscle cells (SMC) appear randomly arranged contains numerous elastic fibers which appear as the black dot-like structures These elastic fibers are arranged in a circumferential pattern, thus when sectioned they appear as black, dot-like structures KEY A, adventitia BV, blood vessel CF, collagenous fibers EF, elastic fibers En, endothelium EnC, endothelial cells I, intima LCT, loose connective tissue M, media SMC, smooth muscle cells 72006_ch13 15/07/10 3:18 PM Page 435 En BV LCT M A 435 P LATE 33 SMC CF En • AORTA CF I A En M M EnC SMC EF CF 72006_ch13 15/07/10 3:18 PM Page 436 ᭹ PLATE 34 Muscular Arteries and Medium Veins Muscular arteries have more smooth muscle and less elastin in the tunica media than elastic arteries Thus, as the arterial tree is traced further from the heart, the elastic tissue is considerably reduced and smooth muscle becomes the predominant component of the tunica media The muscular arteries are characterized, however, by a refractile internal elastic membrane separating the tunica intima from the tunica media and, usually, by an external elastic membrane separating the tunica media from the tunica adventitia Muscular arteries, or arteries of medium caliber, constitute the majority of the named arteries in the body Veins usually accompany arteries as they travel in the loose connective tissue The veins have the same three layers in their walls, but the tunica media is thinner than in the accompanying artery, and the tunica adventitia is the predominant layer in the wall The veins usually have the same name as the artery they accompany Muscular artery and medium vein, monkey, H&E ×365 436 In this photomicrograph, the lumen of the artery is at the left, the lumen of the vein is at the right The arterial endothelium (AEn) is clearly seen on the corrugated surface of the tunica intima, whereas the venous endothelium (VEn) is somewhat P LATE • M U SC U L A R ARTE R I E S AN D M E DI U M VE I N S Muscular artery, monkey, H&E ×545 This is a higher-magnification micrograph of the portion of the figure above outlined by the rectangle turned 90° At this magnification, it is evident that the flattened endothelial cells (EN) follow the contours of the refractile, corrugated internal Medium vein, monkey, H&E ×600 In this higher-magnification view of a portion of the wall of the vein in the figure above, the endothelial cells (EN) are more easily recognized and are seen to be plumper than those of the arterial endothelium The margin between the tunica intima (TI) and the thin tunica media (TM) is difficult to discern, but the smooth muscle cells (SM) in the thin media are more easily recognized harder to distinguish The internal elastic membrane (IEM) is seen as a thin clear zone immediately beneath the endothelial layer, separating the tunica intima from the underlying smooth muscle (SM) of the tunica media (TM) It is evident here that the tunica media is almost twice as thick as the tunica adventitia (TAЈ) elastic membrane (IEM), which rests directly on the most luminal layer of smooth muscle cells (SM) of the thick tunica media (TM) The thinner tunica adventitia (TAЈ) than in the figure above because of the shape of their nuclei and the slight basophilia of their cytoplasm The tunica adventitia (TA) is about twice as thick as the tunica media and appears to contain only bundles of collagen fibers and fibroblasts, with the latter recognizable by their nuclei (N) The collagen bundles of the loose connective tissue beneath the tunica adventitia are larger than those of the adventitia, and there are fewer cells in this portion of the specimen KEY AEn, arterial endothelium C, collagen bundles EF, elastic fibers EN, endothelial cells IEM, internal elastic membrane N, nuclei SM, smooth muscle SSm, small smooth muscle TA' tunica adventitia of artery TA, tunica adventitia of accompanying vein TI, tunica intima TM, tunica media VEn, venous endothelium 72006_ch13 15/07/10 3:18 PM Page 437 AEn N TM SM VEn IEM 437 TA SSm P LATE EF • TI SM IEM SM N EF N N TA N TM C TM SM M USCU LAR ARTE R I E S AN D M E DI U M VE I N S EN EN 72006_ch13 15/07/10 3:18 PM Page 438 ᭹ PLATE 35 Arterioles, Venules, and Lymphatic Vessels The terminal components of the arterial tree just before a capillary bed or an arteriovenous shunt are the arterioles Arterioles have an endothelial lining and smooth muscle in the wall but the smooth muscle is limited in thickness to one or two cells There may or may not be an internal elastic membrane, according to the size of the vessel Arterioles control blood flow into capillary networks In the normal relationship between an arteriole and a capillary network, contraction of the smooth muscle of the arteriole wall reduces or shuts off the blood going to the capillaries A precapillary sphincter is formed by a slight thickening of the smooth muscle at the origin of a capillary bed from an arteriole Nerve impulses and hormonal stimulation can cause the muscle cells to contract, directing blood into capillary beds where it is most needed Arteriole, venule, and small nerve, fingertip, human, H&E ì600 P LATE ARTE R IOLE S, VE N U LE S, AN D LYM P HATIC VE S S E LS 438 This micrograph shows two cross-sectioned arterioles (A) and a venule (V) The arteriole on the left is identified as a large arteriole, based on the presence of two discrete layers of smooth muscle cells that form the tunica media of the vessel The nuclei of the muscle cells appear in longitudinal profile as a result of the circumferential arrangement of the cells The endothelial cell nuclei of the vessel appear as small round profiles surrounding the lumen These cells are elongate and oriented with their long axis in the direction of flow Thus, Arteriole, fingertip, human, H&E ×350 This micrograph shows a longitudinal section of an arteriole Because of its twisting path through the section, its wall has been cut such that the single layer of muscle cells of the tunica media is seen in different planes along its length In the segment numbered 1, at the left, the vessel wall has been cut tangentially Thus, the vessel lumen is not included in the plane of section, but the smooth muscle cell nuclei of the tunica media are seen in longitudinal Lymphatic vessel, fingertip, human, H&E ×175 The lymphatic vessel shown in this figure shows a region where the vessel is making a U-shaped turn in the plane of the section, thus disappearing at the top and bottom of the micrograph The wall of the vessel consists of an endothelial lining and a small amount of connective tissue, with one being indistinguishable from the other A valve (Val), which is characteristic of Lymphatic vessel, fingertip, human, Mallory ×375 The lymphatic vessel shown here is contained within dense irregular connective tissue (DCT) The lumen is irregular, appearing relatively narrow below the valve (Val) A few endothelial cell nuclei are evident (arrows) A thin layer of their nuclei are seen here as cross-sectioned profiles The arteriole on the right is a very small arteriole, having only a single layer of smooth muscle Again, the muscle cell nuclei are seen in longitudinal profile The endothelial cell nuclei appear as the small round profiles at the luminal surface A venule is seen in proximity to the larger arteriole, and a cross section of peripheral nerve (N) is seen in proximity to the smaller arteriole Compare the wall of the venule, consisting only of endothelium and a thin layer of connective tissue, with the arterioles Also, note the relatively large lumen of the venule profile After the arteriole makes an acute turn (segment numbered 2), the vessel wall is cut to reveal the lumen Here, the smooth muscle nuclei appear as round profiles and the nuclei of the endothelial cells lining the lumen appear in longitudinal profile In the segment numbered 3, the vessel wall is again only grazed In the segment numbered 4, the cut is deeper, again showing the lumen and some of the endothelial cells in face view (arrowheads) The structure below the vessel is a Pacinian corpuscle (P) lymphatic vessels, is seen within the vessel It is formed of a miniscule layer of connective tissue that is covered on both sides by endothelium The arrows indicate nuclei that are just barely visible at this magnification; most of them belong to endothelial cells Typically, the lumen contains precipitated lymph material (L); sometimes, lymphocytes may be present Adjacent to the vessel, on the right, is adipose tissue (AT) and on the upper left is dense irregular connective tissue (DCT) connective tissue that is present outside of the endothelium blends with the dense connective tissue beyond the wall of the vessel A venule (V) is also present; it can readily be distinguished from the lymphatic vessel by the presence of red blood cells in the lumen KEY A, arteriole Ad, adipocyte AT, adipose tissue DCT, dense irregular connective tissue L, lymph material N, nerve P, Pacinian corpuscle V, venule Val, valve arrowheads, endothelial cells arrows, endothelial cell nuclei 72006_ch13 15/07/10 3:18 PM Page 439 N V A P LATE 439 AT • DCT A L V DCT Val Val DCT DCT L AT Ad L ARTE R IOLE S, VE N U LE S, AN D LYM P HATIC VE S S E LS P ... 978-0-7 817 -7200-6 (alk paper) Histology Histology—Atlases I Pawlina, Wojciech II Title [DNLM: Histology Atlases QS 517 R825h 2 011 ] QM5 51. R67 2 011 611 ’. 018 —dc22 2 010 024700 DISCLAIMER Care has been taken...72006_FM 15 /07 /10 6:59 PM Page vi This page intentionally left blank 72006_FM 15 /07 /10 6:59 PM Page i Sixth Edition HISTOLOGY A Text and Atlas with Correlated Cell and Molecular Biology 72006_FM 15 /07 /10 ... Mineralization and Matrix Vesicles | 2 41 Physiologic Aspects of Bone | 242 xiv Atlas Plates Plate 11 Atlas Plates Plate 17 Erythrocytes and Granulocytes | 302 Plate 18 Agranulocytes and Red Marrow