COMPARATIVE VETERINARY HISTOLOGY WITH CLINICAL CORRELATES Elizabeth Aughey Fredric L. Frye PUBLISHINGOrganized by body system, this highly illustrated volume covers the normal histological appearance of tissues a wide range animals, both domestic and exotic species, with relevant clinical correlates emphasizing the need to appreciate the normal in order to recognize the abnormal. The breadth of coverage - farm animals, dogs, cats, birds, reptiles, amphibians and fish - and the integration of normal and abnormal tissue provide a reference of lasting value to veterinary students, veterinary practitioners and pathologists. Containing 758 superb colour illustrations, this book offers comprehensive coverage of normal tissue as well as relevant disease conditions. “ Py 4 jee Ise: | olrere7alsa5ece!COMPARATIVE VETERINARY HISTOLOGY WITH CLINICAL CORRELATES Elizabeth Aughey BVMS, MRCVS Department of Veterinary Anatomy University of Glasgow Veterinary School, UK Fredric L. Frye BSc, DVM, MSc, CBiol, Fliiol, FRSM Fredric L. Frye & Associates Davis CA, USA Clinical Correlates by Fredric L. Frye BSc, DVM, MSc, CBiol, FIBiol, FRSM Hazel Johnston BVMS, MRCVS Manson Publishing/The Veterinary Pressce 1s 3 Copyright © 2001 Manson Publishing Ltd ISBN 1-874545-66-9 All rights reserved. No Part of this publication may be reproduced, stored in a permission of the copyright holder or in accordance with the provisions ofthe 4 Copyright Act 1956 (as amended), or under the terms limited copying issued by the Copyright Licensing Agency, London WC1E 7DP, UK. 33-34 Alfred Place, 's any unauthorized act in relation to this publication may able to criminal prosecution and civil claims for dam Any person who d A CIP catalogue record for this book is available from the British Library For full details of all Manson Publishing Ld, titles please write to Manson Publishing Lxd, 6. 73 Corringham R London NW11 7DL, UK. Commissioning editor: Jill Northcott Project management: John Ormiston Design and layout: Judith Campbell and Paul Be aker and Martin Maxwe Colour reproduction: Jade Reprographics Text editing and proof-reading: Andy Printed by: Grafos $A, Barcelona, SpainCONTENTS Preface Dedications Bibliography Introduction Origin of tissues Preparation of tissue sections Staining techniques Microscopy Artefacts induced by histological processing Clinical correlates The cell Epithelium, Glands Clinical correlates Connective tissue Embryonal, mesenchymal and mucoid Connective tissue proper Cartilage and bone Clinical correlates Blood Blood cells and platelets Clinical correlate Bone marrow: production of blood ells Clinical correlates Muscle Muscle types Clinical correlates Cardiovascular system Arteries Capillaries and venules Sinusoids Veins. Arteriovenous anastomoses Heart Lymphatic vessels Clinical correlates a Respiratory system ‘Conduction of air Respiration Clinical correlates Avian respiratory system Reptlian, amphibian and fish respiratory systems Clinical correlates Digestive system Oral cavity Clinical correlates ‘Alimentary canal Clinical correlates, Alimentary system of reptiles and amphibians Clinical correlates, Liver Clinical correlates Gall bladder Clinical corzelates Pancreas Clinical correlates Reptilian, amphibian and fish pancreas Clinical correlates Avian digestive system Urinary system Kidneys Renal pelvis Ureter Urinary bladder Urethra Avian urinary system Reptilian urinary system Amphibian urinary system Fish urinary system Clinical correlates10. Endocrine system Endocrine tissue Clinic Iypophysis cerebri (pituitary gland) Clinical correlates ‘Thyroid gland Clinical correlates Parathyroid g Adrenal gland Clinical correlates Epiphysis cerebri (pineal gland) Avian endocrine system Reptilian, amphibian and fi Clinical correlates 11, Male reproductive system Testis Production of spermatozoa Structure of spermatozoa Tubuli rectir Epididymis and ductus deferens Urethr: Penis Male accessory genital glands Clinical corr Reptilian, an produ phibian and fi Clinical correlates 12. Female reproductive system Uterine tubes (oviduets) Uterus Vulva, labia and clitoris Mammary gland Placentation Umbilical cord Clinical correlates Avian female reproductive system Reptilian, amphibian and fish female reproductive system Clinical correlates 149 149 149 150 153 Ls4 89 192 194 194 194 196 13, Nervous system 215 Peripheral nervous system 216 Central nervous system 219 Clinical correlates 226 The 14. Special senses 27 ion Eye esse Clinical correlates Ino Avian ey see Reptilian, amphibian and fish eyes stu Clinical correlates, of Bar the Clinical correlates his Avian ear : Reptilian and amphibian ears em (Organ of smell (olfactory organ) oft Other specialized sense organs ak int 15. Lymphatic system In Lymph nodes lyn Spleer vai Thym un Local aggregations of lymphoid tissue lyr Clinical correlate gy Avian lymphatic system re Repilian lymphatic system al Amphibian lymphatic system fu ish lymphatic system te Clinical correlates, m The immune system 16. Integumentary system Clinical correlates Avian skin Repailian skin Amphibian skin Fish skin Clinical correlates Specialized integumentary structures Hi Sinus Hooves, horns and claws Appendices Index1 Comparative Veterinary Histology with Clinical Correl Preparation of tissue sections Fixation For tissue sections to be evaluated, they must have be architecture do not decompose after cell death. Generally, a 10% neutral buffered formalin solu- tion is employed as a tissue preservative. However, for certain tissues, such as adrenal gland, brain, eyes and a few other structures, special fixati xed of preserved so that their cells and solutions such as Bouin’s or Karnovsky’s are preferable. Usually, specimens of blood and some body fluids containing cells are fixed onto the glass slide with absolute methanol before staining by one of vari us dyes. In some cases, such as when supravital staining is used, the stain is applied directly to a specimen without prior fixation Small portions (blocks) of tissue, usually less than 0.5 em in thickness, are removed from the animal as soon as possible after death and immersed in a special preservative fluid, a fixative aa degenerative cl Delay for even a minute can lead to serious ges in the tissue caused by the release of enzymes from the cells. The smaller the sample, the faster the fixative can penetrate the whole block of tissue before dege nerative changes Although many different fixatives are available for different purposes, the most commonly used. general-purpose fixative is 10% formol saline. It is, important to use adequate volumes of fixative approximately 50 volumes of fixative to one volume of specimen. Depending on the temper of the: is a minimum of 2 days. Fixation kills the cell quick pecimen and the type of fixative, fixation time enerative processes and preserves the x i Il My iTVIACHTTE HII mT MU IHMLUELLUIE 10 FE LAE structural integrity of the cellular components of the tissue. Soft specimens, such as brain, are hardened by fixation, which allows easier manipulation, By coagulating proteins, fixation prevents their leak: age from the cells jon to be identified in situ, Fixation facilitates id allows their po: staining of the tissue Paraffin embedding Once the specimen is properly fixed itis embedded in paraffin wax to support the tissue during the cut ting process without altering the morphology of the specimen. The process begins with the removal of the water-based fixative by immersion in a graded series of alcohols of increasing con the tissue is saturated with 100% ethanol (i. dehy dated). The specimen is then infiltrated with a clearing agent, such as xylene, that is miscible with both paraffin wax and alcohol. The specimen is ingilteated with warm paraffin wax to replace the the tissue firmly in place (1 dure is usually processor. There are disadvantages in paraffin embeddings itis time consuming and the clearing agents are lipid solvents, so this method cannot be used to demonstrate fats (see Freezing Excess paraffin is trimmed and the block is ready for cutting on a microtome. The block is clamped ‘onto the cutting frame of the microtome, and is moved toward the blade of the microtome using an adjustable wheel until the face of the block is against the blade. With each revolution thin slices (optimally less than 6 nm thick) of the block are cut into a rib bon and emersed (floated) in a warm water-bath (1.5). The sections flatten and are floated onto glass slides (1.6). The slides are placed on a warm plate to ¢ (1.2). The wax hardens as it cools, holding and 1.4). This proce: automatically in a tissue done 4.1 Specimens of fixed tissues are placed into disposable plastic cassettes that confine and identify leach accession during laboratory processing.sponents of the yare hardened nipulation, By nts their leak position ro be es subsequent iis embedded during the cut: phology of the he removal of main a graded atration until nol (ie. dehy trated with a miscible wit to replace the zoos, holdin ). This proc y in a tissue s in paratfi d the clearing ck is clamped jome using ant lock is agains’ 3 (optimally cut into a rib m water-bath redo warm plate t ced into i ‘ring Introduction 12 126 412 (a) Once the tissue has dehydrated, (b) a histology laboratory technician embeds it in a melted paratfin wax-plastc polymer compound, 1.3 A small piece of paper iting special staining equasts is embedded on the opposite side of the sett, In this instance, haematoxylin and eosin periodic acid-Schff (PAS) stains wil be parate du lssue sections 1a Four casettes embedded tissue ready for Sectioning by a microtome, 4.5 Using the finely honed microtome blade, the is transferred to a water bath and histology technician cuts a thin ribbon of paraffin onto the glass microzeape side with a fine tmbedsed tissue camel's hair brush, Note the matt black finish of the water bath, which facilitates Visualizing the nearlyComparative Veterinary Histology with Cli dry; and the section adheres to the glass slide (1.7). Removal of the paraffin wax by such as xylene, and rehydration allows the tissue 0 be examined unstained; this has no advantage ove the direct examination of living cells. Ie is necessa to stain the component cells and tissues selectively and make a permanent preparation for examination with the light microscope a selection of these tech niques is described later (see Staining technique). Dec fied or calcified components befo ding, otherwise the hardness ofthe tissue will cesule n difficulty in cutting the sections, causing artefacts Specimens are fixed in formalin or other chemical fixatives, and then transferred to the decalcifying solution to allow removal ofthe mineral salts. Most of these decalcfying agents contain acids such as ification is necessary for tissue with oss paraffin embed formic, malic, glacial acetic, hydrochloric or nitric. Freezing A cryostat, a microtome confined to a fr chamber, is required to cut frozen sections (1.8 -al Correlates These may be from fixed or from unfixed tissue. The advantage of this method is that the time between taking che sample and examining it under the microscope is much reduced. A biopsy may be taken and examined while the patient is still in the operating room. Fat-containing cells retain the lipid content and the tissue is often more life-like in appearance than non-frozen sections. The disad vantages of this method are tissue distortion, caused by the freezing and thawing, and thicker sectio Once tions are cut and mounted on g slides, conven ional staining techniques are used. Consequence of freezing unfixed tissues ‘When unfixed tissues are frozen and then thawed before being chemically fixed, their delicate cell membranes may become distorted or ruptured, o both, by the forces induced by the expansion and contraction of the intracellular fluid as it freezes and thaws. Therefore, if tissues are to examined histologically, unfixed specimens must not be frozen. Examples of tissues that were frozen before histological fixation and processing are illustrated in 19, 117 Slides bearing th ons are thermal table, which causes evaporation a the adhesion of the sections onto the glass su smooths out iregularites, whichis preparatory to xylene earing enhances 1.8 Frozen tissue sections are created with the use of a yostat, which is @ conventional microtome enclosed, within a freezing temperature chamber. Whereas tissue enzymes and some other cellular constituents cannot be detected in paraffin-embedded sections, specially stained frozen tissue sections reveal them. The advantages of frozen tissue sections are that they require ess time than paraffin-embedded sections to process, and stained tissue specimens obtained during surgery can be examined and nterpreted while the patient still in the operating room. Fat-containing cells retain thelr lipid contents because they are not dissolved by xylene, Tr slide 19) (am (av Coo befoi bt dlsru histo of a Such the Welfixed tissue. hat the time ning it under iopsy may be is sil inthe ain the lipid elifeike in The disad: tion, caused ker sections. don glass es are used. d tissues then thawed delicate cell ruptured, or pansion and a it freezes o examined aust not be rozen before eillustrated armed on a enhances, urfaces, and ay to lene ages of time than ained tisue mined arc rating ments The rehydrated sections of tissue are now immersed in a solution of one or more stains; any excess stain is removed during this process. ‘The slides are dried again, cleared in xylene, and per- 19 Histological sections of (a) myocardium, (0) kidney and (@ Iver from a boa constrictor (B02 constrictor) that were frozen before being fixed in 10% neutral buffered formalin solution. Note the Aisruption and distortion of the histological architecture, and the loss fall but the erythrocytic nuce Such destruction can be prevented if the tissues are fixed before freezing HAE a, 625; b, x625;¢ 2625, Introduction ently mounted beneath a glass or plastic cover- slip using a mounting medium chat is xylene mis cible. Sections that require special stains are stained and given individual coverslips, as shownfs ee ae er oe Comparative Veterinary Histology with Clinical Correlates in 1.40. Sections requiring standard haematoxylin St ing techniques and eosin (H & E) stain fe stained and cover slipped by automated machines that process the Numerous different dyes in various combinations tissues and then dispense an appropriate volume are formulated into stains that are used to impart of mounting medium, apply a coverslip and com- specific and reproducible colouration. Many of these ess the finished mounted slide to remove any dyes possess positive and negative electrical charges trapped bubbles of air (1.11). The completed andare attracted or repulsed by electrostatic charges stained microsections are placed onto the surface of that are characteristic of certain tissue constituents. a warming table beneath a fume hood, where the In order for some dyes to combine with tissue com- xylene in the mounting medium evaporates. This ponents, a merallic salt, termed a mordant, is final step fixes the coverslip firmly to the tissue and required. The combination of a dye with an appro: glass slide, forming a permanent ‘sandwich’ that can _ priate mordant forms a ‘lake’ and carries a positive be handled without dislodging any portion of the electrostatic charge. Dye-mordant combinations stained section. with positive e 4.10 Slides that have received special staining are ‘coverslipped anally sure of laboratory personnel to potentially toxic xjlene vapours is reduced by conducting the overslipping operations beneath a vacuum fume hood. 1.11 When large volumes of slides with standard H & € machine is employed, “4combinations sed to impar Many of these trical charges static charges th tissue com ith an appro: res a positive mbinations d are termed at have a staining are boratory otentially operations Lum fume ge volumes landard H & € overslipped oversipping loved. basic’ stains. These cationic basic lakes combine electrochemically with negatively charged tissue con: stituents, such as nuclear chromatin, other nucleo. proteins, and phosphate inherently basic without requiring the addition of a mordant; they carry their own positive electrostatic tharge. Basic fuchsin, toluidine blue and methylene blue are examples of natu ‘oups. Some dyes ate sic stains, Con Versely, anionic or ‘acidic’ dyes carry a negative ot anionic charge, and are called ‘acidic’ because they to and combine with tissne constituents ative electrostatic charge. Eosin imple of an acidic stain, Differential stain- ing is possible because some tissues may be acidic, basic or amphoteric. Thus, the pH of the extracel Iular fluid causes their electrostatic charge to vary and, as a result, their acceptance of acidic and basic Many special dye combinations, some requiring rare metallic salts, are used to stain certain tissue types etabolic by products and so on. Many formularies containing 4.12 Digital pad (dos), The nuclei are Pra stained deep blue (arrowed). (1) The Qtoplasm and fibres are stained varying shades of pink with eosin. 2) Fat cellsare the fat is lost during processing \ HAE. «160, y 4.13 Longitudinal ection (LS nerve (dos) ‘The myelin sheath surrounding the nerve fibre reacts with osmic acid and stains black; the supporting connective tisue is stained. Osmic acid «250. Introduction recipe-like staining formulae are available and new staining techniques are continually being developed. Examination of living he ligh scope yields very litle information, Therefore, thin ter they have been excised and processed, are stained with special dyes to enable detailed observations to be made on th ells with micro sections of tissue, a ture. The most widely used staining technique is H 8 E, Haematoxylin stains a deep purple colour and acts as a basic stain (basophilic). Eosin is pink t red in colour and acts as an acid stain (acidophilic or eosinophilic), Haematoxylin reacts with bonucleic acid and ribonucleic acid, and eosin reacts with cytoplasmic proteins and a variety of extra- cellular structures. Thus, nuclei and rough endo- plasmic reticulum stain blue to purple and cytoplasm stains pink co red depending upon the concentration of the hasic and acid components of the cell (1. Specialized staining med 1ods are used to illustrate particular features. Osmic acid reacts with fat to give 1 grey-black colour (1.13), periodic acid Schiff (PAS)Comparative Vet and alcian bine reveal glycosaminogly 1.15), silver impregnat ts of nervous tissue (1.16 and 1.17), on’s trichrome differentiates bi ion displays reticular fibres and some and Ma nective tissue and muscle (1.18 and 1.19). Some ci ‘cumstances require the combination of two or more staining methods to yield the maximum tion, There are many other metho: only the commonly used ones are mentioned here. availab) 4.14 Duodenum (dog). The mucus-secreting goblet cells react with PAS (arrowed). HaematoxylinPAS, «125. 44.16 Adrenal (horse). The reticular fibres form a fine networkin (1) the capsule and (2) asa delicate supporting framework for the adrenal secretory calls, The method of Gordon and Sweet for reticular fibees. 125, 16 inary Histology with Clinical Correlates Examination of living material withthe electron microscope has necessitated the developme: new techniques in preparation procedures to illas trate the arrangement of organelles, membranes and cell contents (1.20). Te h provide a three-dimensional picture without dis tortion (1.21). All of these techniques are now steandard tools in histology and have advanced our been further refined to understanding, 1.15 Cervix (sheep). The ep alls lining the cervix react with ether alan blue or PAS, illustrating chemical differences in the types of mucus secreted. Alcian bluesPAS, +20, Used specifically to llustrate the oj the neurons of the central nervous 5 Cajal's uranium silver. 1250 nic processes of (arrowed)the electron elopment of ures co illus: mbranes and er tefined to without dis- dvanced our ” the cervix chemical method is oceses of nue). 1.18 ‘Tongue (dog). The muscle is stained red and the 41.49 Kidney (dog). this trichrome stain the connective connective tissue is stained green, Masson's trichrome, x60, ssUe is stained a blue/green. Gomor's trichrome. x125, 1.20 Transmission electron micrograph ofa fibroblast (sheep). (1) Nucleus, (2) nuclear membrane, (3) cisternae of rough endoplasmic reticulum (RER) (@ plasmalemma, (5) mitochondria, (6) fat croplet and (7) collagen fibril +8000, 1121 Scanning electron micrograph of kidney (dog). (1 Renal tubule, (2) interstitial connective tissue, (3) free erythrocyte ~ a biconcave dic with the 'ypical indentation. 675.Comparative Vet Microscopy The examination and study of normal cells and tis- sues by microscopy is called histology or microscopic anatomy. The study of abnormal cells and tisstes is al is histopathology. An understanding of the not essential for the recogait investigative microscopes range from the simple light, high-resolution elec wide variety of spe of the abnormal, microscops fo the soph tron microscope. In becween lie cialized microscopes to meet special needs, such as phase contrast, polarizing and fluoresc pes, and the scanning electron microscope Units of istological measurement A micromerre (jim) is equal tc a metre and is the unit of measurement of the light microscope; a red blood cell is approximately 8 jim in diameter: millionth part of A nanometre (nm) is equal to a billionth part of a metre. The thickness of the basal lamina of an epithelial cell is 70 nm, which the electron microscope. an be resolved using, Light microscopy he light microscope is the instr visualization of cells and tissues. With it magnifications of up to 2000 times are pos- sible. The limit to the size of the structure that can be distinguished with the ligh by the physical nature of light. The wavelength of microscope is limited visible light ranges from 0.4 to 0.7 jim. Therefore, even with the best optical system available the res olution, or resolv power, of the light microscope wan that is limited to 0.2 um, and anything s will not be clearly distinguishes In order to hieve the best results a few preliminary checks must he made, ‘© Ensure that the glass slide is clean, free from dust and smears, ‘Ensure that the microscope condenser, objec: tives and ocular lenses are clean ~ take great care to clean the microscope with soft lens tis # Set the microscope up for critical illumination fe cach objective by inary Histology with Clinical Correlates closing the iris diaphragm (the substage con- denser diaphragm (2) adjusting the condenser until the circular area of il asa sharp edge, and. 3) making sure the condenser is centred by using the adjusting screws, Always begin with the lowest objective and increase he magnification slowly Transmission electron microscopy This microscope uses an electron beam instead of a light source and allows resolution of structures as small as 1 nm. Small pieces of tissue (cubes not more than 1 mm on a side) are fixed rapidly (to avoid artefacts induced by tissue degradation) in cold glutaraldchyde-based fixative, dehydrated and embedded in epoxy resin. Sections are cut at 0.03-0.05 ym on an ultramicrotome using a glass or a diamond knife, mounted on copper grids and ad sul c vapours of fixa stained with heavy metal solutions such as phate and uranium nitrate. T tives used for electron microscopy processing are volatile and hazardous to the eyes and mucous membranes, so an exhaust hood or adequate ven- Scanning electron microscopy Solid pieces of tissue fixed in a glutaraldehyde fix- ative, are dried, coated with gold and placed in the microscope. The electron beam scans # and a three-dimensional representation of the sur face is obtained e specimen Artefacts induced by histological processing The preparation of tissue sections involves a number of stages during fixing, dehydrating, paraffin embea: ding, sectioning, deparaffinizing,rehydracing, stain ing and coverslipping. Each of these processes necessitates the manipulation of tissue specimens and laboratory reagents, thus providing opportunities for errors to be made. J nique can spoil the final result. Some of the common illustrated in 1,221.26, one flawed laboratory tech artefacts 124tubstage con roscopy (cubes d rapidly (v0 pydrared and using er grids and has lead sul fix and mucous fequate ven opy Idehyde fix laced in the he specimer nof dl i ng fin embe e processes ecimens and nities for 128 Introduction 41.22 Ovary sheep). A knife mark, caused by a nick in the 4.23. Uterus (cat). Shrinkage of the adhesive medium microtome’ cuting edge, leaves a straight line across the used to mount the coversip to the slide captures ait and fection arowed), Messrs tcvome 25. Couses bubbles Wi E655, pees i SOG zt | i 1.24 Spleen (bird). When crystals accumulate in the stain Solutions or are not removed during standard processing, d tissue are present bec stain deposits precipitate onto the surfaces of the tissue he tissue completely, part Section. H & E925. In very thin sections, H & E125, 1126 Spleen (dog). Compression of the paraffin 5 embectied tissue cause HAE 25, parallel ‘chatter’ marks 4.23Comparative Veterinary Histology with Clinical Correlates 20 Clinical correlates In order to appreciate the often subtle alterations that accompany disease or other physical abnor- malities, itis useful to compare the characteris tic changes by which histopathological diagnoses are made and classified. To that end, clinical cor relates sections are inserted throughout this text. Itis important to note that in many instances the tissues comprising an organ of one species are similar or even identical to those found in a dis- parate species. Generally, there are fewer substantive differ- ences within a phylogenetic group of animals than between different groups of animals. For instance, the livers of sheep, cattle, horses, swine, dogs and cats are relatively quite similary the liver tissues of many fish resemble the hepatie tissue found in amphibians; and the hepatic tissues of many reptiles resemble those observed in birds. Because of these characteris- tic similarities and differences, we have selected ‘examples of tissues that are particularly instrue- tive in order to avoid showing repetitively the same tissues for every animal, irrespective of its phylogeny. However, examples from a wide variety of species are included for purposes of ‘comparison, These correlates are placed where they most readily illustrate specific, clinically significant medical conditions. Recognizing normal tissue facilitates interpreting the often subtle alterations in abnormal tissues. Where appropriate, the physiological attributes or significance, or both, of a particular organ or structure are discussed briefly so that their importance to the survival of the animal becomes apparent.mible those haracteris- ve selected ly instruc tively the ctve ofits ma wide urposes of they most significant mal tissue aherations rate, the 2 oF both, discussed survival of 2. THE CELL otein, carbohydrate, fat, nucleic complex acids and inorganic material. Cells are limited by cell membrane, the pla sma. Within the cell lies the membrane-bound nucleus. Th the nuclear membrane or envelope, and sequesters the cell complement of deoxyribonucleic acid (DNA surrounded by a double membran which consists of two long strands wound together in a double helix. The DNA is chromosomes, which carry the genetic informa: tion: the genes. Chromosomes are rarely visible except during tein-DNA complexes are s small, darkly staining bodies within the nucleus are the nucleoli. Ribonucleic acid (RNA) is present Il replication (2.1) and where pro 253 € a | D. Anaphose within the nucleoli together with a small amount of DNA, Small amounts smic organelle, the mitochondric DNA are also presen Ribosomal RNA is synthesized in the nucleoli and lines the outer nuclear membrane as it exp. plasmic reticulum (RER: CCytoplasm surrounds the nucleus and is bound by the plasmalemma. The components of the cytoplasm are divided into organelles and inclusions lying in the oplasm to become the rough endo- cytoplasmic matrix (or cytosol). The organelles ma be membranous (e.g. the cell membrane, mito: chondria, RER and smooth endoplasmic reticulum, Golgi apparatus, secretory vesicles and granules, and lysosomes) or non-membranous (e.g, stored food, f . Metaphase 2:1 The stages of mitotic division. (A) Interphase. (1) Nucleus. (2) Centromere. (8) Prophase. The granular ‘appearance of the nucleus (1) i the early condensation (©) Metaphase. The short, compact chromosomes are the nuclear chromatin in preparation fo rtanged around a central spindi, (0) Anaphase. The "tromere has divided and two chromatids for each chromosome have Separated and moved towards exch tromere (8) (E) Early telophase. Separation of the daughter chromoromes is comp and the cytoplasm begins te divide. (F) Late telophase. The nucleus (1) of each daughter cel is reconstructed with 3 nuclear membrone arc hromosomes are no longer visible, The ¢‘Comparative Vet Epithelium Epithelium is the term used for all the covering and ing membranes of the body, It is composed of contiguous cells linked by cell junctions and resting ‘on a specialized matrix, the basement membrane. All epithelia are avascular and are supported and nourished by the underlying connective tissue cap- illary bed. They are derived from all three basic germ layers (ectoderm, endoderm and mesoderm). The ectoderm provides the nervous system, the ‘outer layer of the skin and the epidermis, and the endoderm provides the lining of the respiratory and digestive tracts, Both ectoderm and endoderm ‘grow into the underlying embryonic connective tissue (mesenchyme) and form exocrine and endocrine glands. Exocrine glands secrete onto the surface of the epithelial membrane through a sys- tem of ducts. Endocrine glands are ductless; islands of secretory cells embedded in connective tissue secrete into the local capillary bed and thus directly into the blood to be carried to the target organ Epithelium of mesodermal origin forms a thin squamous membrane lining the pleural, peritoneal and pericardial cavities of the body. The meso mally derived epithelium lining the heart, blood and lymphatic vessels is called endothelium. The urogenital system is derived from mesoderm, and the epithelial membranes of most of the geni tal system, the kidneys and ureters are of mesodet mal origin. All epithelial membranes are capable of regeneration and repair. Damaged and dead cells, are replaced by adjoining cells to maintain the cover and the integrity of the membrane Epithelium may be either simple, where a single layer of cells is present, or stratified, where a vari able number of cell layers are superimposed. Simple epithelium Squamous Simple squamous epithelium is a single continuo\ layer of flattened cells, which is often so attenuated, that ivis difficult to identify the boundaries of indi vidual cells using the light microscope. The nucleus bulges from the thickest part of the cell, as in the endothelium lining blood vessels (2.2) and in the am of the body cavities (2.3). mesothel Cuboidal Simple cuboidal epithelium is a single layer of cells; each cell is square in cross-section with a central nucleus. Minor variations in proportion may occur ary Histology with Clinical Correlates to give short cuboidal and tall cuboidal cells, Examples can be found covering the ovary, thyroid gland and mammary gland (2.42.6) columnar Simple columnar epithelium is a single continuous layer of tall hexagonal cells with a basal nucleus forming a celatively thick membrane. These cells are often specialized, performing a particular function. In secretory epithelium the cells secrete mucus and have a lubricant and protective function; examples can be found in the stomach and cervical canal (2.7 and 2.8). In the small intestine the luminal surface area is markedly increased by microvillous processes to form a striated border, a functional adaptation, designed to increase the surface area for absorption. Adjoining goblet cells secrete mucus, keeping the membrane moist and protecting against digestion by the luminal contents (2,9) 2.2. Simple squamous endothelium. Uterus (ca). Arteriole. Simple attenuated squamous cell line the Tumen; the nucleus of one cells arrowed. H & E.x250. 2.3 Simple squamous mesothelium. Uterus (ct). The simple squamous cells are on the free serous surface of ‘thevuterus. H & E160.: The Cell as atest = 2.5 Simple cuboidal epithelium. Thyroid (dog). The ted Simple cuboidal epithelium on the free surface of the simple cuboidal epithelium lines the colloid filled thyrokd ovary is arrowed. H&E. x50. follicles. 8 €. x20 keeping the s¢ digestion ve thelium. Lactating mammary 2.7. Simple columnar epithelium, Gallbladder (dog) he secretory alveolus of the mammary (G) connective core of the lamina propria 2) Tall 5 simple cuboidal cells, H& €.»125. hexagonal epithelial cells witha basal nucleus. H&E. »125. 23 W Ais 4 lum. Cervix sheep). The 2.9 Simple columnar epithelium. Duodenum (dog). The ‘ BS aries sy oa he Sak a epithelium lining the cer le mucus-secreting goblet cel is wateaylinvPAS, «500,2.10 Comparative Veterinary Histology with Clinical Correlates | Pseudostratified J Pseudstatified epithelium appears co consis of more than one layer of ells All the cells ae in Contact with the basement membrane, bur not d very cell reaches the luminal surface, The nucle licat diferent levels, causing the staifed appear ance. This type of membrane is seem in the re ratory tract (2-10-2.12) and in the genital tract (2.13), where cells may be seeretory or ciliated pit 2.10 Pseudostratifed columnar ciliated epithelium with ‘goblet cells respiratory epithelium lining the nares {horse), The alia appear as a fringe on the free surface, the goblet cells, as clear rounded spaces. There are ‘Several layers of nuclei, but all the cells rest on the basement membrane. H & E100, 2.11 Respiratory epithelium. Lung : y epithelium. Lung (donkey). In this (con), The mucussecreting goblet cells scanning electron micrograph the mucus secretion is & are individually tained bulbous projection surrounded by cilia. 2500, Gomorialdehyde fuchsin. 100. 2.13 Pseudostratified epithelium. Epididymis (bul. Several rows of nuclei create the appearance of 2 stratified epithelium, but al the cells rest on the basement membrane. The stereoclia give a fringe effect tothe luminal surface. H & E.»200.ats 10 consist of the cells are in brane, but not face, The nuclei ratified appear ven in the respi he ry or ciliated mis ul the fringe Stratified epithelium Designed to withstand wear and tear, stratified ‘epithelia consist of two or more layers of cells with ‘only the basal layer resting on the basement mem: brane. Stratified columnar and cuboidal epithelia are found lining large gland ducts and the urethra (2.14 and 2.15). They are usually not suitable for absorption and require gland secretion to keep the surface moist, bur the epithelium lining the rumen is absorptive (see Chapter 8, Digestive System}, The cell Squamous Stratified squamous epithelium may be keratinized ‘or non-keratinized. The later is common on surfaces subject to wear and tear, where the secretions nec: essary to keep the surface wet come from associated glands, and is found lining the oesophagus and ‘vagina (2.16 and 2.17). The basal cell layer is mitot- ically active and the new cells are continually formed and pushed towards the surface, moving away from the nourishing capillary bed beneath the epithelium, ‘These cells are dead or dying by the time the surface 2as 2.14 Stratified columnar epithelium. Penile urethra (horse) The epithelium Is several layers deep; the Superficial layer is columnar. H&E, «200 2.15 Stratified cuboidal epithelium. Urethra (bitch). The surface layer of cells is cuboidal. H & E. x00. 216 2.16 Stratified squamous non keratinized epithelium, (esophagus (at). The polyhedral cells ofthe basal layer divide and the daughter cells are pushed towards the surface where the dead squames are shed. H & E.x125. 2.17 Stratified squamous keratinized epithelium Footpad (dos). (1) Stratum germinativum; the basal layer ‘of simple columnar cells. 2) Stratum spinosum: several layers of pear-shaped cells. (3) Stratum granulosum; layer of cells deeply stained containing keratohyalin granules, (@ Stratum corneum: muttlayered zone of anucleate squames. H&E. x100, 25es Comparative Veterinary Histology with Clinical Correlates is reached. There they lose their nuclei and become detached (desquamate); only on the surface layers are the cells squamous. In some sites, such as the epi- dermis and the tongue, the cells become ker inized and form a protective waterproof layer on the sur face (2.18 and 2.19). Urethelium (transitional epithelium) Urethelium lines most of the renal pelvis, the ureters and the urinary bladder, and is designed to allow 248 2319 [ 220 stretching of the membrane without rupture. It is My classified as pseudostratified, because in the relaxed, My unstretched state a number of layers of cells are pre end sent, The surface cells are rounded (2.20). In the and stretched state the appearance is that of stratified bas squamous epithelium. The cells have the ability to fou stretch and distore, without pulling apart, and are duc ideally suited to the demands of the bladder and my ureters. The surface is thickened and gives a water: thu proof coating. thes 2.18 Stratified squamous keratinized epithelium. Skin (cov), Relatively fewer layers of cells; the surface is covered with keratin, H & E100, sa 2.19 Stratified squamous epithelium. Teat (cow). The stratum lucidum is present in thick skin as dear translucent layer. Phosphotungstic acid haemotoxylin 100, 2.20 Urethelium, Bladder (dog). The surface cellshave 3 rounded appearance, the mide layer of cells are pear shaped and the basal layer is columnar. H & E.x125,ut rupture, I is cin the relaxed, of eels are pre- d (2.20). In the hat of stratified ve the ability to apart, and are he bladder and d gives a water- thelium. Skin cow), The 2c have a Myoepithelium Myoepithelial cells, derived from ectoderm and ‘endoderm, are found in sweat and mammary glands and lie between the secretory epithelial cell and the basement membrane. Myoepithelial cells are also found in the modified salivary (venom) glands and. ducts of venomous snakes. The cytoplasm has ‘myofilaments and the cell is capable of contraction, thus assisting the expulsion of the secretions from ructures (2.21-2.23), these 2.22. The paired venom glands of many venomous snakes [and the four pairs in the single genus of venomous lizards (Hefoderma)] are composed of ‘thin-walled, folicleike structures lined by a single layer of non-keratinized squamousto-plump cuboidal epithelial secretory calls. There dilated follicles store the pink staining, proteln-rch venrom Until itis delivered via the duct system and fangs. Contractile myoepithelial cells (arrowed) and skeletal ‘muscle aid inthe expression and delivery of venom hough the colled ducts and into the hallow fange of snakes (and to external grooves in the solid teeth of Helodermati lizards). lustrated isa section of e venom gland of a small Mexican rattlesnake nyo). H&E, xB, The Cell 2.21 Sweat gland, kin (horse). Myoepithelial cll le between the simple columnar se lium and the basement membrane, and appear as a deep pink line (arrowed). H & E.x125, 2.23 The much colled venom duct of this rattlesnake is thin-walled and is ined by low cuboidal ces with dark staining basal nucle. The colle sections of the dct are separated from each other by connective tissue in which many myoepithelial cells (arrowed) ‘are embedded, The entire ducts surrounded by the ‘temporal and masseter skeletal muscles which, when they contract, augment the myoepithelial cells in ‘0 and through the fangs. H & E.»43,Comparative Veterinary Histology with Clinical Correlates Glands All glands are derived from either ectoderm and mesoderm or endoderm and mesoderm. The ecto- derm and endoderm form the epithelial secretory cell, the parenchyma. The mesoderm forms the sup- porting connective tissue framework, the stroma. Where the demand for the secretion is low, a si gle secretory cell is sufficient. The mucus-secreting goblet cell ofthe small intestine, for example, is ade~ quate. At the other extreme is the liver, the largest gland in che body, which is required to cope with the food absorbed by the gut. Exocrine glands Exocrine glands secrete onto the membrane surface cither singly — the unicellular mucus-secreting gob- let cell - or by a duct system. The simple tubular sland of the endometrium consists of a single layer 224 225 28 of cells secreting into a duct opening into the wer ine lumen (2.24). Compound glands have a branched duct system draining a number of secre tory units. The connective tissue capsule extends into the gland, carrying blood vessels and nerves, and divides it into lobes and lobules. The secretory units may take the form of acini where the height ofthe lining cell greater than the diameter of the lumen, or of alveoli/saccules where the lumen exceeds the height of the lining cel (2.25-2.27). There are wo types of glandular secretion: mero- crine and holocrine. In the meroerine gland, the he seeretion in the cytoplasm fuse with the cell membrane and release the con: tents onto the cell surface. In the holocrine gland the cell builds up the secretion in the eytoplasm, migrates away from the basement membrane and the source of nutrient, and dies. The cell debris itself becomes the secretion. The sebaceous glands are 2.24 Simple tubular glands. Uterus (cat). (1) Lumen of the uterus (2) Simple tubular glands inthe endometrium of the uterus Hae 20, 2.25 Compound acinar gland, Pancreas (dog), (1) The acinus is lined by secretory epithelial cells with 2 basal nucleus. (2) The excretory duct Islined bya stratified cuboidal epithelium. H & E5250,ng into the uter= here the height diameter of the the cytoplasm clease the con olocrine gland the cytoplasm, membrane and cell debris itself ous glands are glands. n of the uterus rland, acinus ined ls with a tory duct ida The Cell 226 Alveolar gland. Carpal skin (Gi) The secretory alveol are cut in osesection; the dlameter ofthe lumen exceeds the height of the lining secretory cells # & E. «100, (Gee also mammary gland, Fig. 2.6.) 227 Compound tubuloacinar gland. ed veromucous salivary gland. The us-secreting cells fre filled with secretion and almost bitrate the lumen, The serous cells thick sand of connective tissue With a blood vesel represents the Supporting framework of the gland Hae «125, 2.28 sebaceous gland. Skin (dog). The gland consists of pale staining tel filles with sebum, fatty Substance. This forms the secretion, fan example of» holoctine gland GGomor’s rich sComparative Veterinary Histology with Clinical Correlates | Endocrine glands Endocrine glands have no ducts: Small groups of distance away (2.29). Detaled discussion of these glands can be found in Chapter 10. Clinical correlates A large variety of benign and malignant epithe lial rumours are recognized in domestic animals. A squamous cell carcinoma, a malignant tumour ‘of squamous epithelium taken from the eyelid of a cow, is shown in 2.30, The neoplastic cells are large with abundant eosinophilic cytoplasm and form nests where cells differentiate and keratinize towards the centres. There is variation in nuclear and cellular size and mitotic figures can be seer Bovine ocular squamous carcinoma is rela- tively common and of some economic impor: the adenohypophyss (pituitary Gland; cat) The secretory cells are closely associated with a rich network of blood vessels. H & E200, tance. Its occurrence is related to ultraviolet light exposure, particularly in white-faced breeds such as Herefords. The tumour devel- ops through premalignant stages before pro- gressing to carcinoma én situ and finally to A pulmonary adenoma, a benign tumour of glandular origin, is shown in 2.31. The cells are ‘uniform in size and appearance and arranged in a recognizable papillary patterr epithe framework, This tumour was an incidental find: ing at necropsy in an aged cat. The columnar cells all rest on a supporting stromal y | 2a 2.30 Squamous cell carcinoma from the eyelid of a ‘cow. H&E, 3200, 2.31 Pulmonary adenoma in an aged cat. H & E.x125. g yey3. CONNECTIVE TISSUE Gonnective tissue can be classified as follows wound. There are few cells, which are usually stl @ Embryonal connective tissues late undifferentiated fibroblasts; the ground sub- Mesenchyme and mucoid connective ti ance is abundant and gelatinous with very few BGonnective tissue proper, including loose fibres. This type of tissue stains poorly with haema (areolar) and dense (regular and irregular) toxylin and eosin (FI & F) (3.2), but stains well with issues, as well as the special types, reticulas, mucin dyes. elastic and adipose © Cartilage and bone a1 # Blood cells and blood-forming tissues. , J i (tutary +200. Embryonal, mesenchymal x and mucoid connective rs ! tissues ‘ Monnective tissue is derived from & 4 p ultraviolet loose embryonic packing tissue of mesodermal ori- a> ite et el cls arc ong sede procenes a mout devel nd are embecided in an amorphous gelatinous sub- 1 before pro: ietance, the extracellular matrix (3.1). Mucoid con- eee id finally to Mective tissue is found in the embryo, and also 3, nbilical cord (foal). (1) Nucleus of the stellate Pes ot Peeurs in limited regions in adult animals, the comb — mesenchymal cell. (2) Long cell processes. (3) Extracellular cect Bnd wattle of the chicken, and around a healing mat (2) Blood vessels H & €.x125, anged in he columnar ing stromal idencal find- 2a1 3.2. Mucoid connective tissue. Comb (chicken. (1) Stratified squamous epithelium. 2) Lamina propria, (2) Mucoid connective tissue. H&E, 50Comparative Veterinary Histology with Clinical Correlates Connective tissue proper Connective tissue proper fills the interstices of tissues and organs, and forms a continuous structure that {G) Nerves. 4) Extracellular matrix {5} Fibrooytes. (6) Collagen fibres. {i} Smooth muscle of the uterine wall Hae. «100, glycans. These sub- sue < collagenous fibres, ns few cells. Fibrous: sommonest cel isthe 3:26 Dense connective tissue (dog) Fibrocyes (arrowed). (1) Collagen flores (2) Blood vessels. Masson's trichrome. 100 327 Dense connective {Gheep). The collagen fibres been. Massons trichrome, sue cat) The petioyte Is fe. (1) Vein.) tymphatle ar mate H & x10.Comparative Veterinary Histology with Clinical Correlates Special types of connective tissue Reticular tissue is composed of numerous reticular fibres and stellate reticular cells, forming a sup- portive network for structures such as the sple« lymph node, kidney and bone marrow. Elastic tis sue, characterized by numerous regularly or irreg. ularly arranged elastic fibres, is exemplified by the ligamentum nuchae and the elastic fascia of the ruminant abdomen, Adipose tissue consists of groups of adipocytes (see above) Cartilage and bone Cartilage Cartilage is a specialized form of connective tissue combining a degree of rigidity with flexibility and strength. There are three types of cartilage: hyaline, clastic and fibrocartilage; differing only in the dis tribution of the main components: the cells, fibres and matrix. Hyaline cartilage This type of cartilage is binish/white in the fresh state and is the most prevalent form, In the embryo, the pre- cursors of the long bones begin as cartilage models, (3.28), As the neonate grows, the cartilaginous tem- plate undergoes progressive mineralization. In post natal life, cartilage is present in the rings ofthe trachea and in plates in the larynx and nose. With ageing and under certain conditions of hypervitaminosis-D, and hypercaleaemia, cartilage may become pathologically mineralized. Cartilage also caps the ends of bones in articulating joints (3.29), At predetermined sites in the embryo, mesenchy mal cells round off and differentiate into chondro- blasts (cartilage-forming cells) and secrete a matrix consisting of proteoglycans and collagen fibrils. The space occupied by each cell isa lacuna and once the matrixis laid down, the cells are called chondrocytes {cartilage cells; 3.30). Chondrocytes are capable of dividing and several cells may come to occupy 2 lacuna; then they are known as an isogenous group or cell nest (3.31). Compared with the bulk: of the tmatrix, which stains poorly with H 8 F, the matrix in the immediate vicinity ofthe cells stains intensely with metachromatic dyes because ofthe presence of elycosaminoglycans. Mesenchymal tissue surrounds the developing cartilage and forms a fibrous cover- ing, the perichondrium. The inner layer of the perichondrium is capable of generating new chon- droblasts. Cartilage is thus able to grove from the pericardium by appositional grow, and by in stitial growth from within by chondrocyte division Iris avascular — the and deposition of new matr cells are nourished by diffusion 3.28 Developing hoot (foal). (1) skin (2) Hyaline cartilage models of the digits. (2) Joint cavity. H & E25,ation. In post- agen fbr: % The & E, the matrix stains intensely le surrounds fibrous cover- r of the ring new chon. ow from the ind by inter rocyte division avascular ~ the Connective Tissue 1329 Scapulohumeral articulation of lous spaces filled with (2) Aricular cartilage 3.30 Hyaline cartilage. Costal gartilage (dog). (1) Hyaline cartilage. (2) chondrocytes. (3) Perichondrium. Hae. xi00. 3.31 hyaline cartilage. Costal Extracellular matrix. H & EComparative Veterinary Histology with Clinical Correlates Elastic cartilage Elastic cartilage is specially adapted to give resilience and withstand zepeated bending. The matrix contains elastic fibres, Examples of elastic cartilage can be found in the epiglottis and the inna (3.32-3.34) 3.33 Elastic cartilage. Pinna (dog). (1) Perichondium, (2) chondrocyte in lacuna. (3) Extracellular matrix with red elastic fibres. Gomor's trichrome, x160. Fibrocartilage Fibrocartilage occurs at the site of tendon insertions and in the intervertebral discs, where firm support and tensile strength are necessary. The chondroblasts lie in rows between parallel bundles of collagen fi and secrete cartilage matrix (3.35-3.37) 3.32 Elastic cartilage. Pinna (dog). (t) Perichondrium. (2) Chondrocyte in lacuna. @) Extracellular n red elastic fibres. Masson's 3.34 Elastic cartilage. Pinna (dog). (1) Chondracyte in lacuna. (2) Extracellular matrix with red elastic fibres. Gomori’ trichrome. 3250re firm suppore Pinna (do), ) Chondroeyte in ar mavixwith sons trichrome. | ss es| rote in tes 3.35 Fibrocartilage (ow). (1) chondrocyte. @) Collagen fibres arranged as dense regular connective fisue, H&E 125, 3.36 fibrocartilage (0) 3.37 Firacartilage (ox (1) Chondrocyte. (2) Collagen fibres Gomons trichrome, »250.338 Comparative Veterinary Histology with Clinical Correlates Bone Bone isa rigid form of comnective tissue composed of cells embedded in an intercellular matrix of col- lagen fibres, glycosaminoglycans and calcium phos- phate deposited as hydroxyapatite erystals. Bone provides the framework of the body’ and serves as a lever for muscle action, as protection for viscera, asa haemopoietic organ, and as a reservoir of body stores of calcium and phosphorus. Small bones are found in soft tissues to provide extra rigidity (8 penis in the dog and ossa cordis in the ox. Bone is a living tissue that is supplied with blood vessels, and nerves, and is constantly changing in response to body stresses and circumstances. There are two forms of bone: cancellous (spongy, medullary) and compact (cortical, dense). Alll bones have both cancellous and compact forms of bone deposition, Cancellous bone consists of ieregular interconnecting bars, the trabeculae, forming a three-dimensional network of lined spaces filled with bone marrow (3.38 and 3.39). Compact bone is solid continuous mass in which the spaces are only visible with the aid of a microscope (3.40). Bones are covered with a specialized connective tissue, the periosteum, the inner layer of which is ‘osteogenic (capable of laying down new bone; 3.40) Spaces in bone, ike the marrow cavity (3.38-3.40) and the canal system (3.40-3.42), are lined with a single layer of osteogenic cells, the endosteum, The characteristic feature ofall bone tissue isthe arrange ment of mineralized bone matrix in layers, the lamellae. Small | ‘occupied by a single bone cell, the osteocyte. Tubular passages, the canaliculi, radiate from each lacuna and link up with canaliculi from adjacent lacunae to create an extensive system of interconnecting canals, This arrangement is clearly defined in compact bone where the lamellae are arranged concentrically around a longitudinal canal to form Systems 3.41-3.43). The central canal of the osteon carries blood vessels, imellae are nae present in the an osteon (Haversi 3.38 Spiculated bone (deg). (1) Periosteur. 2) Bone spicules with osteocytes in the lacunae (arrowed) {@) Bone marrow filled spaces lined by osteoblasts (arrowhead). H & €, 100. 3.39 Spiculated bone (dog) (1) Periosteum. (2) Bone spicules with ‘osteocytes (arrowed). (3) Osteoblasts {rrowhead) on the free surface of the bone, H&E, x160,Compact bone vthe spaces are scope (3.40) ized connective yer of which is ew hone; 3.40} ity (3.38-3.40) ce lined with a ndosteum. The ¢ lamellae are ceyte. Tubular m each lacuna jacent lacunae are arranged canal to form 1-3.43). The Stood vessels, oa) spicules with paces lied by HAE x100, 2) cules with Dsteoblasts urface of the aa 3.40 Compact bone (dog). (1) Compact bone af the daphyss of the femur. (2) Periosteum, (2) Endosteum. (8) Marrow with a hig proportion of fat cells. M & E20, Sree) oie | “ ; J | ae | | sa" 3.42 Compact bone (do9). (1) The osteon fils the field. The central canals ined by endosteum and carries blood vessels and nerves. (2) Osteocytes in lacunae in the al lamellae. (3) Periosteum, H&E, x100, Connective Tissue . a ee . ! 3.41 Compact bone (dog). (1) Compact bone with osteocytes lacunae arranged in lamellae with a central canal carrying blood vestels and nerves. The canal is ined by endostourm. (2) Periostoum, {@) Endosteum of the marrow cavity. H & E. x62. 343 3.43. Compact bone (dog). Ths isa high power view of Fig, 3.42 Ha e9250, 45Comparative Veterinary Histology with Clinical Correlates branches of the perpendicular canals (of Volkmann). ‘These are part of the main blood supply to the bone and link the endosteal and periosteal surfaces. The lamellae may be regular circular rings as in the osteon, surround the shaft of the bone as ciceum- ferential lamellae, or fill in the angular spaces between lamellae as interstitial lamellae. They are often the result of bone remodelling. Bone contains a vast continuous network of canals that are essential for the nutrition of the bone cel in the lacuna, the osteocyte (3.41-3.45). The cll body lcs inthe lacuna and extends in long processes into the canaliculi to contact similar processes from adjacent osteocytes Bone is a living tissue and is constantly being. remodelled; osteoclasts are multiny leate non- dividing cells that are found in resorption bays (Howship’s lacunae) at the site of bone remodelling, (3.46). Osteoclasts are derived from a progenitor 46 cell in bone marrow. The cell migrates to the developing tissue, The ruffled (brush) border is the undulating mobile cell membrane. Osteoblasts are present on the inner surface of the periosteum and endosteum, and cover the surface of bone spicules in an active osteogenic area (3.38, 3.39 and 3.46). The cell has a central or slightly eccentric nucleus with chromatin granules, and the cytoplasm stains deeply with haematoxylin because of the concenteation of organelles. Long cytoplasmic processes extend out from the osteoblast and the matrix is deposited around them. This provides a fine canalicular network and allows nutrients to pass to the bone cells. Unlike cartilage there is no diffusion in bone. Once the bone has been deposited the trapped cells (osteocytes) function to ‘maintain the bone and retain contact withthe blood vessels through the canalicular system, FT] 3.48 Developing periosteal bone (foal. (1) Periosteum, (2) Bone spicules. (3) Osteogenic tse fils the spaces between the spicules, (4) Endosteum. (5) Marrow. # 8 E 2625, 3.45 Developing periosteal bone (toa). (1) periosteum. (2) Bone spicules. (2) Osteogenic tissue fills the spaces between the spicules. (@) Endosteum. (5) Marrow. H & x25,rates to the h) border is the nner surface of and cover the osteogenic area 5 a central natin granules, » haematoxylin ganelles. Long nthe osteoblast nd them. This rk and allows inlke cartilage sone has been es) function to withthe blood jesteal bone 2)Bone jtsue fills ne spicules TOW. BE steal bone 2) Bone ctu fills e spicules nO. H BE. Endochondral ossification Inthe embryo, m sites differentiate into chondroblasts and lay down Eailage models of the long bones (3.47). Later in éstaton, the mesenchyme surrounding the carti- Tage model becomes very vascular and the cartilag +hymal cells at predetermined B46 Compact bone (cat). Osteoclast (arrowed). {)) Osteocytes in lacunae. (2) Osteoblasts on the free furlace ofthe bone, H & . x250, 3.48 Endochondral ossification (cat). (1)Zone of resting cartilage. (2) Zone of proliferating cartilage. {3) Zone of hypertrophied chondrocytes (@) Basophilic calcified cartilage with freshly deposited eosinophilic bone. (5) Osteogenic tissue Hae x625, Connective Tissue matrix is calcified. Nutrients are unable to diffuse through this calcified cartilage and the cells die. Cell death is followed by breakdown of the matrix. Vascular mesenchymal tisstie moves in, differenti tes into osteogenic tissue and deposits bone on the remains of the calcified cartilage (3.48 and 3.49 347 3.47 Whole mount section of the left forelimb of a smal Viviparous yucca night lizard (Xantusa vigils). Note the transition between the blue staining cartilaginous ends and the mineralized diaphyseal compact bone. H & E.»S. aas 3.49. Endochondral ossification (cat). (1) Zone of hypertrophied chondrocytes.) Calcified cartilage with bone deposits. (3) Osteogenic tisue. H & E x160,Comparative Veterinary Histology with Clinical Correlates This is an ossification centre. The cartilage is replaced by bone beginning at the centre of the dia- physis and extending to the epiphysis. The epiph: ysis has a separate centre of ossification and a plate ‘of cartilage persists, the epiphyseal plate; this sep- arates the epiphysis from the diaphysis (3.50). The perichondrium becomes the periosteum. Multinucleated osteoclasts remove mineralized ‘osseous matrix (3.52). In metabolic bone disease (fibrous osteodystrophy, secondary nutritional hyperparathyroidism, ‘rubber jaw’, ‘renal rickets’), which may be caused by an improper dietary calcium : phosphorus ratio, or in some cases of severe chronic renal disease the osteopenic bone is replaced by fibrous connective tissue (3.51). 3.50 Growth of a long bone Bones are able to grow in width and length and do so until the adult size is reached. Thereafter, bone continues co remodel as circumstances demand. Growth in width is appositional from the ‘osteogenic inner layer of the periosteum and new bone is formed locally, the periosteal collar (3.44) This ability to deposit bone is utilized in bone grafts. Growth in length is accomplished at the di: physeal/epiphyscal junction at the persistent layer of the epiphysis. Cartilage grows intersitially on the epiphyseal side and dies on the diaphyseal side of the plate. Osteogenic tissue invades and deposits bone, the epiphyses are pushed apart and the bone grows in length (3.50). 3.50 Epiphyseal pate (at {1) Epiphysis. 2) Spiculated bone (8) Growth area of endochondral ‘ossification. H&E 25, 3.51 Section of femur from a green iguana (iguana iguana) with ‘metabolic bone disease. Note the multinucleated osteoclasts (arrowed) ‘hat have been removing ossfied ‘Yasue and fibrous connective tissue ‘hat has replaced the cancellous bone. H&E. 5250,igth and reafter, ed. T tional from the scum and new collar (3.44) lized in bone shed atthe dia persistent layer rsttally on the physeal side of s and deposits and the bone te (c). uated bone, ndodhondral froma green 2, Note the dass (arrowed) ing osified Intramembranous ossification Small bones and flat bones develop directly from mesenchymal cells; these differentiate into osteo- blasts. Bone is laid down as a network of bony spicules that are gradually enlarged by the deposi- tion of new bone lamellae. The spaces are filled in and compact bone is formed in the out plates c ical correlates Although a variety of benign and malignant tumours of various origins arise inthe skeleton, osteosarcomas, a malignant mesenchymal tumour in which the neoplastic cells produce tumour osteoid or bone, account for approxi- mately 80% of canine and 50% of feline skele- tal neoplasms. Osteosarcomas can be subdivided 3.52 Osteogenic osteosarcoma (og) showing tumour bone formation. H & E520. 3.53 Giant cel osteosarcoma (dog). Nete the numerous ‘muitinucleae cells H&E, x250, Connective Tissue which are continuous with the more central can cellous bone. The surrounding mesenchy denses to become the periosteum; the inner layer is osteogenic (see 3.44). The osteoblasts on the inner surface lining the spicules of cancellous bone also retain their osteogenic ability according to their location within bone or histo logical appearance. An osteogenic, or bone pro ducing, osteosarcoma that has multifocal formation of well mineralized new bone is shown in 3.52, This was taken from an 8-year-old St Beard dog. An example of a giant cell osteosar- ‘coma (3.53) from a German Shepherd dog shows numerous multinucleate syneytia formed by fusion of the neoplastic cells. 49 382Comparative Veterinary Histology with Clinical Correlates Arthritis isa general term for inflammatory dog in which the thickened synovial membrane is | disease of the joints. There are many causes and invaded by large numbers of plasma cels is | many types of arthritis but often a similar pattern shown in 3.54. The predominance of this cell type of reaction and joint damage may be observed. A suggests an immunological basis for the disease synovial biopsy from an adult German Shepherd in this case. 354 3.54 Plasmactic arthritis in the ‘dog. The synovial membrane is infltrated by plasma cells H & € x25,membrane is sma cells is this cell type the disease its inthe moran col H BE. 4. BLOOD Blood isa special rype of connective tissue: the inter- cellular matrix, the plasma, isa fluid, and ane red and white blood corpuscles. Small fragments of cells, che platelets, and large proteins such as fib- finogen, albumin and globulin are non-cellular ele- ments carried in the plasma. Both red and white blood cells are derived from the same primitive cel, the haemocyroblast. The red blood cells, or ery throcytes, ace contained within the blood vessels and carry oxygen and carbon dioxide. The white blood cells, oF leucocytes, are part of the body's defence mechanism and use the circulation as a Ieave the blood vessel and enter the tissues. The cel [ular elements account for 40% of whole blood and the plasma for 6 The preparation of blood for histological exam- ination is remarkably simple. A drop of blood is spread on a glass slide, fixed immediately in air and by immersion in absolute methanol, and stained with one of the Romanowsky dyes, mixtures of methylene blue and the derivatives azure blue and eosin, Cell nuclei stain purple; the haemoglobin containing erytheacytic cytoplasm stains pink or tan; and the eytoplasm of leucocytes assumes a blue to bluc-grey hue. Cytoplasmic granules react either with cosin and aze eosinophilic, or with the blue stains and are basophilic. They often display some actlity, Erythroeytic and leucocytic haemoparastes stain variably, depending upon their iype. Some granules do not stain with either and are regarded as neutral. This method allows for iden tification of all the cell types ina blood smear, and tifferental cell counts are used t0 assess the blood picture (sce Appendix Table 2 for species variation). where they ansport co particular sic degree of 1 Blood cells and platelets Erythrocytes The mature mammalian erythrocyte is a highly dif- ferentiated cell lacking a nucleus, ribosomes and tnitochondria. It isa biconcave dise with the cell membrane enclosing the cytoplasm, and is filed with haemoglobin, the protein carrier of oxygen and carbon dioxide (4.1, 4.2; see also Chapter 1, 1.21). The diameter varies in size according to species. Mammalian erythrocytes are round and they measure from 4.1 jim in the goat to 7 jm in the dog, The erythrocytes of birds, fish, amphib- ians and reptiles are clongated and they can mea. sure nearly 20 x 100 jim in some amphibians and reptiles. Erythrocytes from one animal are the same size, except in the cow and sh in size (anisocytosis) is not unusual. The erythro- cyte is flexible enough to pass through the smallest capillary. The average life span is 3 months in mammals, but in some reptiles it is as long as 3 years, Immature erythrocytes may appear in the ep where variation Th Te eo 4.1 Blood (ox). (1) Enythrocytes. 2) Large lymphocyte. {@) Small lymphocyte. Leishman. «200, ical correlate a2 42 Feline prolymphocytic leukaemia, The nuclei | ‘are large, and the nucleol are retained. Wright’. 7625,« a Comparative Veterinary Histology with Clinical Correlates circulation in response to urgent need; the com: monest of these is che reticulocyte (4.3 and 4.4) Proerythroblasts, basophilic erythroblasts and the metarubricyte are also seen occasionally (4.5). The nuclei of avian, amphibian and reptilian erythrocytes are elongated and retained into maturity. Nucleus- free, but otherwise intact, erythrocytes (erythroplas- tids) are occasionally found in reptilian blood, Similarly, nuclei with a tiny amount of haemoglobin= containing cytoplasm may be seen. These cell-like objects called haematogones, appear to be intact ery throcytie nuclei or nuclear remnants that have been extruded from other red blood cells, Because they are nucleated, the cells of birds, fish, amphibians and. reptiles are capable of mitosis and amitotic division even after they have matured. Although the erythro- cytes of the lower vertebrates retain their nuclei, the presence of their immature phase, the reticulocyte, is easily demonstrated by staining unfixed blood films with the supravital dye new methylene blue, During metamorphosis in amphibians, the ery throcytes change from the larger larval cells to the smaller adult cells. The nucleus is retained in both, but the amount of endoplasmic reticulum is greater im the adult cells. The haemoglobin also changes during the development of larval amphibians into adults. Larval haemoglobin possesses a greater affinity for oxygen; this is consonant with the aquatic habitat. Larval erythrocytes are more able to incorporate the amino acids uridine and thymi- dine than are the adule cells. The site of haemo- poiesis shifts during larval and metamorphic development. As an embryo, the erythracytes are formed in the ventral blood islands. Later, the pronephric and nephric kidneys are major sites for red blood cell production. In late metamorphosis and into adulthood, the liver and then the spleen and bone marrow predominate as the sites where erythropoiesis occurs. 4.3 Reticulocytes (cal). Early stage erythrocytes the rnuceus has just been extruded and a fine network of residual nuclear materials still present. Leishman. x160, 4.4 Mitotic division in a previously mature, haemoglobin Containing erythrocyte of a western diamondback rattlesnake (Crotalus atrox) demonstrates the ability of these cells, under certain conditions of anaemia, to revert 10 a blastic phase and undergo mitosis. Benzidine peroxidase, 625, 52 45 ‘ a nucleated early stage etythracyte seen on rare occasions in circulating blood, Leishman, «200.Blood ibians, the ery Leucocytes numerical equivalent of the PML in mammals, ae Heterophils display the greatest diversity in size and etained in both, There are two main types of lencocytes, granulo- granule shape from species to species of any of the a Bytes and agranulocytes, both of which can leave leucocytes. In most amphibians, lizards, snakes and changes nd enter the circulating blood by crossing the cap-crocodilians, the heterophil granules are small mphibians into lary wall between the endothelial cells. They are round and orange. In chelonians (curtls, tortoises esses a greater artof the body's defence system and involved in and terrapins} they are elongated, needle-shaped pant with the theimmune cesponse. The most numerous of the and often a muddy brown colour. Like their mam s are more able BRM ix docoestic:ehimals;withithe exception) miallan'PMIscoun tetpartsdié Nevesophils ate ved fine and thymi- lfoxen and sheep, are the polymorphonuclear leu- somally active, recruited to sites of bacterial site of haemo. eeytes (PMs), large geanulocytes (10-12 jum in infection and can act as phagocytes. Because oftheir metamorphic iameter) with lobed nuclei, usually five lobes reddish-orange coloured granules, heterophils arc rythrocytes are joined by chin strands of chromatin. They are often mistaken for eosinophil nds. Later, the Mescribed as PMLs (4.6). Staining of the granules ‘The basophilic leucocytes (basophils) are the major sites for Is ery variable and the individual granules are rarest of the granulocytes in mammals, forming metamorphosis BBall and difficult to identify. The eosinophilic leu- only 0.5% of the white cell population. However, hen the spleen Eocytes (eosinophils, acidophils) have large red _as one descends the phylogenetic scale, the num the sites where Branules in the cytoplasm (particularly large in the ber of basophils enumerated in a differential blood Horse), bilobed nuclei and form about 5% of the cell count increases markedly to as many as 10% in White cell population (4.7). They are associated some species. Charac With response to parasitic infections and allergic granules, containing histamine and hepa A Feactions. the bilobed, multilobed, or unlobed nuclei, and the ally similar to mast cells (4.8). .¢ basophilic n, obscure cates network of The heterophil is another granulocyt ishman, 160. Beitin the lower vertebrates. It is the functional and thatis pre- cells are esse: 136 Blood (dos). Polymorphonuclear leucocytes are omy +6 fxtowed. Leshan, x60. | §. last isa 187 Fosinophil. Blood (dog). Large rod-shaped granules 4.8 Basophil Blood (dog). The dense basophile granules Me present inthe cytoplasm. Lelshman, «625, ‘obscure the cell Leishman, x20, 53Comparative Veterinary Histology with Clinical Correlates op 4.9 Blood (dog). (1) Small lymphocyte. {@) Polymorphonuclear leucocytes. Leishman. 200 410 ‘The most common of the agranulocytes isthe | small lymphocyte, which constitutes 30% of the white cell population, except in oxen and sheep where it makes up 70% of the rota. eis 7-8 ymin diameter, with a dark blue nucleus and a thin rim of cytoplasm (4.1, 4.9 and 4.10). Medium and large lymphocytes are up to 10 ym in diameter with a dark blue nucleus and pale blue eytoplasm (4.11 and 4.12). The largest cell of the agranulocytic series is the monocyte, which is 12-20 wm in diameter, with athorseshoe-shaped nucleus lying in abundant cyto plasm (4.13). Iteaves the blood and transforms ino a variety of macrophages at various sites in the body 1s part of the mononuclear phagocyte system. 4.10 Lymphocyte. Blood (dog), A small ymphocyt les in ‘the middie of the field. Leishman. »200. 44.11 Lymphocyte. Blood (or). A large lymphocyte lis ‘near the middle of the field, Leishman, 3200. 4.12 Blood (ox). (1) Large Lymphocyte, (2) Small Iymphocytes {G) Polymorphonuclear leucocytes Leishman, x160,ulocytes is the 5 30% of the cen and sheep [tis 7-8 pm in dla thin rim of jum and ameter with a asm (4.11 and, oeytic series is diameter, with bundant cyto- ransforms into es in the body .¢ : oct les ‘ 98 imphocytes, leucocytes, aay Imonocye isthe largest cel present. 1413 Monocyte. Blood (dog). A Leaman. 200, a a Platelets Platelets are cytoplasmic fragments, about 3 jum Ti diameter, derived from large multinucleated Bills in the bone marrow called megakaryocytes. Mhese cells function with fibrinogen to repair Bamaged blood vessels by forming a thrombus or flor (4.14). In birds, fish, amphibians and reptiles, Thrombocytes are responsible for blood clotting; This ellis not present in mammalian blood. The Hhrombocyte is nucleated, elongated and usually Bomewhat smaller than an erythrocyte. It pos: $esses a pale blue, agranular cytoplasm, and is Eapable of active phagocytosis, amitotic division TAB Platelets. blood (do). A group of platelets are Birowed, Leishman. 625, Blood yas & and, amazingly, pluripotentiality inasmuch as it has the ability under conditions of severe acute and chronic blood loss o be transformed into a haemoglobin-rich, respirationally functional ery throcyte. Whether the thrombocyte, like its mam- malian stem cell cousin, can transform into other cell types is unknowa. In reptiles, thrombocytes bud off from large multinucleated cells with dis tinctly granular cytoplasm that are present in the bone marrow. Under conditions of severe anaemia, they can also be found in extramedullary sites such as the liver and spleen. Because of their delicacy, the megakaryocyt bone marrow touch ike giant cells are best seen in preparations (4.15). 4.15 Megakaryocyticlike multinucleated cell from which eptilian thrombocytes originate. Jenner-Giemsa, 200.Avian blood cells ctophils, eosinophils. and basophils. The het | crophils are 8-10 im in diameter and are so-called i The various corpuscles of mammalian blood are _ because of the variable staining of the cytoplasmic present in birds, but exhibit distinctive features, rods; the nucleus is bilobed or trilobed (4.18). The ] These include retention of the nucleus by the eosinophils are about 7 um in diameter and i mature erythrocyte and the presence of a true rounded granules are present in the cytoplasm ] thrombocyte, in contradistinction to the mam- They are less regularly rounded than are the her | malian blood platelet. The erythrocyte is a nucle- _erophils and most have a bilobed nucleus. The spe- ated oval cell with an eosinophilic cytoplasm, cific granules are round or oval and smaller than ‘measuring approximately 9-12 jum in diameter the granules of the heterophils. The cytoplasm is (4.16). The thrombocyte is analogous to the mam- basophilic (4.20). The basophil is more numerous malian platelet, is oval or round in shape and is than in the mammal; the granules are deep blue smaller than the erythrocyte, The nucleus stains staining. Agranular leucocytes are identical to the deeply basophilic, as does the cytoplasm. Prominent lymphocytes and monocytes of the mammal and red granules are commonly present in vacuoles form 72% of the white cell population (4.18, 4.21 adjacent to the nucleus (4.17). and 4.22). The bone marrow haemopoietic tissue Granular leucocytes account for approximately is basically similar to the mammalian, with the 24% of the white cells and are classified as het- notable absence of megakaryocytes. fay aye Dis 4.17 Blood (bird). A group of thrombocytes ae arrowed Leishman, »250, ae {4.18 Blood (bird). The cytoplasm of the avian heterophil 4.19 Blood (bir). A small lymphocyte. Leishman. +480. {silled with eosinophilic granules, Leishman, »480.ils. The her dare so-called cytoplasmic ed (4.18). Th liameter and ¢ cytoplasm, n are th feus. The spe- cytoplasm is re deep blue poietic tissue an, with the €* aa7 x E » are arrowed. ® 419 Deaee ual pan, 880, Blood Le 6 oO | > it 4.22 Blood (bird). Monocyte. Leishman. 480. Reptilian, amphibian and fish blood cells The formed cellular components of amphibian and reptilian blood are similar to those present in avian blood. Erythrocytes elongated and vary widely beeween species. The largest erythrocytes are found in the ruatara, Sphenodon punctatus (a quadruped reptile that superficially resembles a large lizard), and manders, particularly Amphima In some instances the erythrocytes of animals are proximately 100 jum long and 20 um wide Generally, lizards and snakes possess smaller ery throcytes and leucocytes than do chelonians and odilians, in some aquatic sal However, there are individual variations between species. Although all of the blood cells of fish, amphibians and reptiles retain their nuclei throughout their life- spans, true reticulocytes supravital staining (see 4.4 be demonstrated by Because the blood cells, retain their nuclei, it is nor unusual co find both mitotic and amitotic division in stained films of 4.21 Blood (bird). Monocyte. Leishman, x480, peripheral blood from amphibians and reptiles. These cells in division usually reflect a and, thus, do nor necessarily suggest neoplastic pro yonse to blood loss and reptiles are also similar to those present in birds. Th clude acidophils (amphiphils) comprising het erophils and eosinophils, the intracytoplasmic gran ules of which stain reddish-orange or muddy b with Romanowsky dyes, are ovoid to distinctly spin dle-shaped and possess pale blue lobed or unlobed ccentrially placed. The het jor leucocytes that functions enzymatically in opsinization and chemotaxis in rnuclei that rend to be cerophil is one of the n amphibians and reptiles. Itis also capable of phago- ceytosing particulate matter, such as bacteria and cel detritus. The percentage of heterophils in the differential leucocyte counts of reptiles and amphib- ians normally ranges wi ing on the species, season of animal from which they were obtained. Clinically, they usually range from 20 to are spherical and stain de identical dyes used to stain blood films of mammals y from 2.t0 65%, depend year and sex of the The granules of red with the ceosinophi and birds. In some instances, one or mote giant red granules are found rather than dozens of smaller ones. The nuclei of eosinophils tend to be more centrally located and may be lobed, unlobed or concentric: Eosinophils may account for 0=10% (or more) of the differential leucocyte coune of most reptiles. They ‘increase during parasitism and other antigenic chal- lenges, The basophil granulocytes are probably the most readily identified because of theit uniformly blue or purple staining, Their nucle or unlobed, and ce may be lobed ally of eccente cally located against the inner surface of the cell mem: brane. Basophils account for 010% of the leucocytes, ‘of most normal amphibians and reptiles. 57Comparative Veterinary Histology with Clinical Correlates Another granulocyte observed in amphibians and reptiles i the azurophil. This cell is characterized by its large, usually unlobed, nucleus composed of densely clumped chromatin and finely granular cyto- plasm, which usually contains tiny azurophilic gran ules. Amphibian and reptilian aznrophils are often called PMLs. In instances of bacteraemia, azurophils. may engulf particulate matter. Azurophils account for approximately 2-10% of the differential cell count. The mononuclear leucocytes of amphibians and reptile include small and large lymphocytes, plasma cells and monocytes. Fach of these cell types resem- bles its counterpart in mammals and birds. Unlike mammals, for which the cellular compo: nent of the blood clotting mechanism involves nnucleus-free platelets, fish, amphibians and reptiles possess nucleated thrombocytes. These elongated cells characteristically contain a pale blue, usually non granular, cytoplasm and a single elongated central nucleus. Superfically, they resemble erythrocytes but they lack haemoglobin. Under certain conditions involving acute and chronic blood loss, thrombocyts display pluripotentiality: they can be transformed into erythrocytes. In doing, so, they acquire increasing amounts of haemoglobin and may undergo mitotic or amitotie division. Like their leucocytic counter. parts, the amphibian and reptilian thrombocytes can also serve as phagocytes and engulf bacteria and cx: lular detritus, including senescent erythrocytes. Because they are nucleated, all of the cellular ee- ments of amphibian and reptilian blood can be involved in haemopoietic neoplastic disorders (4.23), in addition to the many Iymphoreticulac and myelo- poietic disorders that occur in mammals and birds 1 4.23 5as0-cosinophilic myelogenous leukaemia in a ‘boa constrictor (Boa C. constrictor). Note the dual population of eosinophilic and basophilic granules that characterize a clonal population of neoplastic ‘granulocytic leucocytes. Wright's. 625. 4.28 Bone marrow (dog). (1) Basophilic erythrablast, Prorubricyte. (2) Polyehromatophil erythroblast rubricyte ) Erythrocytes. (2) Myeloblast. (5) Promyelacyte. (6) Myelocyte. (7) Polymorphonuclear leucocytes. (8) Monocyte Giemsa, «250, : “9 — ee gee @ 4.25 Bone marrow (dog). (4) Basophilic erythrobiast, rorubricyte. 2) Polychromatophil enythroblast,rubricyte ) Erythrocytes. (a) Myeloblast (©) Promyelocyte. (6) Myelocyt. () Polymorphonuclear leucocytes. Giemsa, 2500. 4 ca susually non: gated central throcytes but n conditions hrombocytes isformed into lergo mitotic ytic counter- nbocytes can lood can be ders (4.23), rand myelo. sand birds as, rubrignte (6) | Monooyt. . ‘ mmatophil slat. lot. ucocjtes Bone marrow: production of blood cells Haemopoietic tissue located in the adult bone mar row is responsible for the production of red cells, granular and agranular white cells, and platelets. The precursors of these cells are present in bone marrow (all of which derive originally from the ‘multipotential stem cell, the haemocytoblast), but as they develop the individual characteristics of each cell type predominate and the various stages may be identified (4.24-4.29), Blood a 426 4.26 Bone marrow (doa). (1) Basophilc eryth prorubricyte. 2) Polychromatophil erythrobl (2) Promyelocyte. (4) Myelocyte. Giemsa. «480, 4.27 Bone marrow (dog). (1) Basophilic erythroblast, prorubricyte. 2) Polychromatophil exythroblas, rubrcyte. (3) Promyelocyte (@) Myelocyte. Giemsa, 80, 429 Red bone marrow (dog) (1) Megakaryocytes are separated by hhsemopoietic tissue, (2) Blood vessel Giemsa, x250 Bid ' el 4.28 Megakaryocyte. Bone marrow (dog). Giemsa. 880.Comparative Veterinary Histology with Clinical Correlates The erythrocyte series begins with the rubriblast {proerythroblast), a large cell with a large clear nucleus with two nucleoli in a basophilic cyto- plasm, The prorubricyte (basophilic erythroblast) is smaller, with a dense nucleus and diffuse basophilic cytoplasm. The rubricyte (polychro matophilic erytheoblast) has a dense nucleus in sgrey-pink cytoplasm, which is caused by the syn= thesis of haemoglobin. There are no further cell divisions; the aueleus condenses, the haemoglobin content of the cytoplasm increases and the cell is called a metarubrieyte (normoblast). The nucleus then becomes extruded and the cell becomes a reticulocyte (an immature form of erythrocyte), small numbers of which are present in circulating blood. ‘A similar series forms the gramulocytes, begin- ning with the myeloblast, a large cell with a clear nucleus and a pale rim of agranular cytoplasm. This divides and forms the promyelocyte, a very large cell with a large clear nucleus and some in the basophilic cytoplasm. The myelocyte is a markedly smaller cell. The deeply indented nucleus is eccentrically disposed in the cytoplasm, and spe cific granules indicate the type of leucocyte. Late ‘metamyelocytes have a band nucleus; again, these are found in the circulating blood, The mature lew cocyte has a lobed nuciew ‘Where haemopoietic tissue predominates, with small amounts of adipose tissue, the active marrow is red in appearance (4.24, 4.25 and 4.29). Where adipose tissue predominates the inactive marrow is yellow (4.30). Blood sinusoids lined by reticuloen: dothelial cells are present in marrow (4.30). Lymphopoiesis is complicated by the fact that although lymphocytes arise from stem cells in the bone marrow, they become two separate and func tionally different cell populations: T lymphocytes and B lymphocytes. T lymphocytes from the thy mus are segregated from the blood by the thymic barrier and are responsible for cell-mediated immunity. When they leave the thymus T lym: phocy n diffuse sites in secondary lymphatic tissue, such as the paracortex of a lymph node. The B lymphocytes are present in bone marrow. As a result of antigen stimulus {the humeral response), they become plasma cells and produce immunoglobulin, es are found '30 Yellow bone marrow (dog). The empty spaces indicate the site of the fat cells. Giemsa. »250cocyte. Late in, these he mature lew: pminates, with 14.29). Where by reticuloen 4.30 the fact that emcells in che rate and func Iymphocytes from th bythe thymic in secondary acortex of a re present in stimulus (the sma cells and Clinical correlates Disorders of the blood can be attributed ro abnormalities in production of blood cells from the bone marrow o other central blood-form- ing organs or to abnormal loss or consumption. of these cells, Anaemia, the deficiency of oxy- gen-carrying erythrocytes, can be regenerative (ic. the bone marrow is able to respond by releasing new cells, including imma into the circulation) or non-regenerative (in which this response does not occur). A blood film that illustrates haemolytic anaemia in a dog is shown in 4.31. In this regenerative anaemia there is variation in cell size (anisocytosis) with red cell precursors, such as the large, stippled reticulocyte (1) and nucleated normoblast (2), Blood Visible. Some of the red cells have small, dense dots (Howell-Jolly bodies), which rep remnants of nuclear chromatin and are charac- teristic of regenerative anaemias. Spherocytes, red cells that have lost their biconcave shape and become globular due to immunological attack, are present (3), There is an increased neutrophil count with some immature neu trophils (reactive neutrophilia with a left shift), which typically accompanies haemolysis. These findings are diagnostic of autoimmune haemolytic anaemia, the most common type of anaemia in the dog. In immune-mediated anaemia, erythrocyte destruction follows the attachment of antibody to the cell membrane. In most cases the aetiology is unknown, 4.31 Haemolytic (regenerative) anaemia in @ dog, Note the variation in appearance of the red cells. Reticlocytes, nucleated normoblasts and. Sspherocytes are present. Giemsa, »200, 61Comparative Veterinary Histology with Clinical Correlates ‘The blood film in 4.32 was prepared from a dog with acute lymphoblastic leukaemia. Very hhigh numbers of malignant lymphoblasts, which are larger than normal lymphocytes and have bluer cytoplasm, are seen. In some cases of enkaemia, although the bone marrow is invaded by neoplastic haemopoietic cells, no abnormal cells are released into the peripheral blood. Such cases may be termed aleukaemic leukaemias. 433 Non-regenerative anaemia is a common finding with many leukaemias. The liver of a cat in which the sinusoids (vas ‘cular channels) are heavily infiltrated by large neoplastic megakaryocytes with multilobed nuclei is shown in 4.33. This is megakaryocytic myelosis, a rare, chronic disease of dogs and cats characterized clinically by bleeding and thrombosis of the ear and tai tips 4.32 Acute lymphoblastic leukaemia (dog) showing large numbers of ‘malignant lymphoblasts. Giemsa, >255, 4. with the multilebed nuciel in the celular inflate. H & €. 255.Blood Avian bone The medullary bone is basophilic and decreas on finding during shell deposition and increases at other A onique feature of avian bone in egg-producing times. The basophilia is caused by a change in th sods (vas- birds is the accumulation of spiculated bone in the density of the matrix and in the glycosaminogly d by larg medullary cavity, under the combined influence of cans that are present. During resorption, osteo. nulilobed pestrogens and androgens. This medullary bone clasts are active; during deposition, osteoblasts are aryocytic is particulary labile and the stored calcium is uti- prominent on the surface of the bone spicules dogs and lized in the formation of the calcareous egg shell. (4.34 and 4.35) eding and 4.34 Head of the humerus (do cortical bone. (2) The speul cavity is ined by osteoblasts. H tic her). (1) Outer layer of eosinophilic medullary bone is basophilic, (3) The m: 625, 4.35 Head of the humerus (domestic her). (1) The osteocytes in the tosinophilic cortical bone form an osteon, (2) Osteocytes in the open lacunae I Of the basophilic medullary bone. (3) Marrow cavity lined by osteoblasts ‘ H&E xi28,Reptilian, amphibian and fish bone and bone marrow Reptiles do not possess pneumatized bone such as that found in birds capable of flight. The box-like carapace and plastron of chelonians are composed ‘of specialized bone that must be both strong and relatively lightweight in order for this bone to pro- tect efficiently the delicare internal structures. Compressional stresses applied to the dorsal and ventral surfaces are distributed widely and are then borne upon buttress-like vertical supporting pillars ‘of bone that are at each end of the “bridge” that joins the carapace to the plastron on each side, The strength of their shells is farther enhanced by the curved shape and by additional internal struts that distribute compressive forees so that they are not concentrated onto a single focus. The shell is com posed of parallel layers of inner and outer tables of compact bone with an intervening layer of spongy cancellous bone characterized by spaces rative Veterinary Histology with Clinical Correlates filled with bone marrow. In form and function the bony shell resembles the calvaria that covers the brain case of a mammalian skull. ‘The bone of amphibians and reptiles contains ‘numerous sites of haemopoiesis; long bones (see 3.29), ribs, skull and mandibles are locations in which active blood cell formation normally occurs. During severe blood loss and a few other condi: tions, sites other than bone marrow are recruited for extramedullary haemopoiesis; liver, spleen and kidney are then most often involved. In fish, the major organ of haemopoietic activity is the cranial pole of each kidney (see Chapter 9), with lesser amounts occurring in other extra- medullary sites during times of severe anaemia, infection or stress. The presence of numerous megakaryocytes in the splenic sed pulp is a common finding in the healthy house mouse, Mus musculus, whereas extramedullary haemopoiess is usually considered an abnormal finding in most other animals (4.36). 4.36 Extramedullary haemopoless inthe spleen of @ domestic mouse (Mus ‘muscuts). Note the multinucleated megakaryocytes which are normal in ‘murine splenic tissue. H & E, 125function the les contains s bones (see jacations in ally occurs it condi- re recruited , spleen and Chapter 9), ther extra dis s, whereas i in the ‘considered nals (4.36 5. MUSCLE Contractility, a fundamental property of cytoplasm, is developed to a highly specialized degree in mus. cle tissue. The elongated muscle cell is commonly referred to as a muscle fibre, the plasma membrane as the sarcolemma and the cytoplasm as the sar n, There are basically two types of muscle: smooth, visceral or involuntary muscle; and striated muscle, which is further subdivided into skeletal voluneary muscle and cardiac involuntary muscle. Muscle contraction depends upon the proteins actin and myosin in the sarcoplasm. In skeletal and cardiac muscle the longitudinal arrangement of these proteins is aligned in register to give cross: stciations, which are absent from smooth muscle. Muscle types Smooth muscle Smooth muscle cells are elongated fusiform fibres (2-50 um long) with a single, centrally located nucleus with several nucleoli. These fibres may occur singly, as in the lamina propria of intestinal vill, but are more commonly arranged in sheets or hay tubes of the alimentary, urogenital, respiratory and cardiovas cular systems. The fibres are packed in a staggered shion with the thickest nucleated portion of one fibre juxtaposed to the thin tapered end of an adjoining one. Individual smooth musele fibres are supported by reticular fibres, with collagen and elastic fibres forming a supporting framework of connective tissue carrying blood vessels and nerves. mooth muscle is under involuntary control; it is in the walls of a wide ed by the autonomic nervous system '5. Smooth muscle. Buodenum (dog). (1) Fusiform smooth muscle fibre, H & E200, 5.1 Smooth muscle. Uterus (cat). (1) Nucleus, 2) Sarcoplasm. H & €.x100. 5.2. Smooth muscle. Uterus (at). (1) Nucleus @) sarcoplasm. (3) Longitudinal fibres, () Transverse fibres. H&E, «200, 65eS Comparative Veterinary Histology with Clinical Correlates Striated (skeletal) muscle contractile myofilaments are arranged in alternat- ing isotropic I bands and anisotropic A bands. This imparts the cross-striated effect. Skeletal muscle fibres are bound into large bun- dles by an outer connective tissue investment, the ‘epimysium. This dips into the muscle and invests tundles of muscle fibres (fascicles) in conneetive Skeletal muscle fibres are multinucleated giant cells that range in length from a few millimetres to sev- ceral centimetres. The nuclei lie immediately beneath the sarcolemma, and the myofibrils give both a lon- gitudinal and a cross-striation arrangement. The bi 5.4 Striated muscle. Tongue (0%) (3) Transverse section (TS) muscle fibres, (2) Longitudinal section (5) muscle fibres, Masson's trichrome. 5.5 Striated muscle, Tongue (a. (1) Nucleus. (2) Sarcoplasm {G) Endomysiuim. H&E. »200. 5.6 1S striated muscle. Tongue (eat). (@) Longitudinal striated muscle. {@) Nuclei. @) Endomysium stained freen, Maston's trichrome, x200,ed in alterna: Abands, This nto large bun wvestment, the le and invests in connective longue (0%) (15) muscle section(s) s trichrome page (cat) 3200. Tongue (cath {muscle fissue, the perimysium, This in turn invests each muscle fibre in a vascular loose connective tissue, the endomysium. A fine network of reticular fib Ties against the sarcolemma, The collagen fibres of the tendon extend into the epimysium and allow nent (5.4-5.9) muscular contraction to effect mov 57 TS striated muscle, Tongue (cat) ()ransverse section of muscle Aibres. (2) Connective tissue Betimysiom. Masson's trichrome. 625. 58 striated muscle, Tongue (ox) Muscle fibres showing crose-striations Masson's tricnrome, x625, 5. Tendonmuscle junction (dog). (i) collagen flores of the tendon, (2) striated muscle fibres. H & E 20, Muscle Every skeletal muscle fibre receives an axon tetmi: nal from a motor neuron at the myoneural junction, the motor endplate (see 13.27). Muscle spindl which are attenuated skeletal muscle fibres, act as stretch receptors that are innervated both by motor 67‘Comparative Veterinary Histology with Clinical Correlates Cardiac muscle (myocardium) Cardiac muscle is exclusive to the myocardium, the muscular wall of the heart. The fibres are smaller than skeletal muscle fibres and branch repeatedly (5.10). The nucleus lies in the centre of the fibre (occasionally two nuclei are present) and the fibres, have strong areas of attachment at the intercalated disc. These are visible as a dark cross-striation at the end of one fibre and the beginning of the next, and confer structural integrity on the heart muscle to allow contraction to spread throughout the myocardium (5.11 and 5.12). These are represented at the ultrastructural level by tight junctions and gap junetions. The cardiac conducting system is composed of several specialized muscle fibres in the sinoatrial and atrioventricular nodes. In the sinoa- trial node (5.13, 5.14), small cardiac muscle fibres, which are low in myofilaments, have an intrinsic ability to contract at a species-specific rate and act as the pacemaker for cardiac muscle contraction. The atrial wave of depolarization cor second node, from where specialized large muscle fibres spread throughout the ventricular muscle and initiate contraction, ‘These specialized conducting fibres are binucleate and the nucleus lies in a clear area of sarcoplasm (5. The various types of muscle tissue can be dis cerned, even in invertebrates low on the phyloge- netic scale. For example, the striated muscle fibres of a spider are structurally similar to those found in mammals and the multiple heart-like pumping chambers that circulate the haemolymph and 7 TI] 5:10 Cardiac muscle horse) Z (1) Nucleus of the muscle fibre. (2) Sarcoplasm, H&E, »125, 5.11. Cardiac muscle (horse. (1) Cardiae muscle fibre. (2) Vascular connective tissue of the endomysium, {G) imercalated die. Heidenhain’s ron haematoxylin, 625,usele fibres, ate and act ges on the from where 1e phyloge- uscle fibres hose found e pumping ymph and fire 5, <) rains +2 haemolymphocytes through the coelom of earth. ‘worms are formed from myocardium, As all fish and amphibians and most reptiles (all -ocodilians) possess a the except the: chambered heart with paired atria and a single ventricle, there ferences between the hearts of these animals. For example, a ridge of myocardium helps di ventricle so as to reduce the mixing of oxy genated and deoxygenated blood in non-crocodil ect the flow of blood through an reptiles. Blood flow through twin aortic arches rnd atrioventricular valves is controlled by typical wart valve leaflets composed of myxoid connective tissue that are covered by a thin endothelial lining all vertebrates. 5.13 Cardiac muscle (horse) (1) Cardiac muscle. 2) Purkinje’s flores. H& €, 62.5, 5.15 Cardiac muscle (horse) 1) Endocardium. (2) Impulse conducting fibres (Purkinje’s flees). Muscle fibre, H & £125 5.12 Cardiac muscle (horse). (1) Cardiac muscle fibre, @) Vascular connective tissue ofthe endomysium {@) Intercalated dsc. Heidenhain’ iron haematonyir 200. 5.14, Cardiac muscle (horse). (1) Cardiac muscle. (2) Purkinje’ fibres. (3) Loose vascular connective tissue. H&E 125, 695.16 Comparative Veterinary Histology with Clinical Clinical correlates Myopathies, primacy disorders of muscle struc- ture, may be congenital, metabolic or inflamma: tory, Viral, bacterial, fungal, parasitic, protozoal | and metazoal agents can be implicated in infec- tive myopathies ‘Where there is frank inflammation the disease would be termed a myositis. In eosinophilic myositis, a condition that affcets the masticatory muscles of dogs, in particular in the German Shepherd breed (5.16), the affected muscle is dif- fusely infileraced by numerous. eosinophils accompanied by lesser numbers of lymphocytes and other inflammatory cells. There is muscle | degeneration and atrophy on of abnor- mal antibodies which attack these muscles is believed to initiate the process, which then becomes dominated by a cellular response 5.16 Eosinophilic myositis. Masticatory muscles from a ddog, Phosphotungstic acid haematoxylin. x125, eee Correlates Progressive destruction of these muscles leads to fixation of the jaws. Other non-neoplastic conditions that affect the muscle may be loosely divided into neu- ropathies which result from disturbance of innervation and myasthenic conditions of the motor end plate. Muscle atrophy (reduction in ‘cross-sectional area of the muscle fibres) will tend to occur Under certain circumstances, all types of mus- cle can be vulnerable to pathological deposition ‘of mineral salts. This may be induced by condi- tions characterized by persistent hypercalcaemia (see 6,28) or at sites of previous injury. Primary neoplasms of muscle are quite rare with malignant tumours (rhabdomyosarcoma, 5.17) ournambering benign ones (rhabdomyoma). ‘Muscle can also be affected by neoplasms of a3s0- ciated connective tissue origin. 5.17 Rhabdomyosarcoma in a 10-year-old male cat. Large, rather pleomorphic. elongate to strap~ like cells invade and replace skeletal muscle. These ‘tumours although uncommon, are highly malignant. HAE x100,leads to at affect ce of ction in es) will of mas: position y condi- jgnant. 6. CARDIOVASCULAR SYSTEM The mammalian circulator of tubes with an endothel through the arceries into wun ), unica media (mid- trough the veins ae layer) and tunica outer layer; 6.1 and 6.2). The tunica intima consists of elongated f endothelial cells resting on loose areolar co Arteries tissue. The tunica media of the clastic arteries has high proportion of concentric lamellae 0 rated elastic fibres interspersed with smooth 6.1. Artery (dog). (1) Lumen. (2) Tuniea Intima, (2) Tunica media, (@) Tunica adventitia. Masson's trichrome. 62.5. 6.2 Artery (dog). (1) Lumen. 2) Tunica intima with the internal elastic lamina (arrowed). (3) Tunica media, Masson's trichrome. 125,Comparative Veterinary Histology with Clinical Correlates 63 6.3 Aorta (horse). Elastic artery 5 wavy elastic fibres, Weigert's elastin 100, R 1 to illustrate th muscle fibres (6.3). These allow the vessels to dilate. The recoil sends the blood onwards, creat: ing the pulse in the major elastic artery, the aorta In muscular (distributing) arteries, the content is reduced and the smooth muscle increased. Internal and external elastic lamina are present (6.2, and 6.4-6.6), The arterioles (6.7) reduce the pressure of the blood and supply the capillary bed. The tunica media may consist of only one layer of smooth muscle cells and the luminal diameter is less than the thickness of the wall (6.8). The tunica adventitia is composed of collagen and elastic fibees, and contains vasorum, the small nutrient arteries and veins in the walls of the larger blood vessels. 6.4. Muscular artery (sheep). (1) Lumen. 2) Tunica intima with the internal elastic lamina, (3) Tunica media. (4) Tunica adventitia. #& E 35, 6.5 Muscular artery (sheep) (§) Lumen. (2) Tunica intima, (G) Tunica media. @) Tunica adventitia, Masson's trichrome, »250 66 an (1) sma @ um 67 4 (on). ( ()uyn 6a: Garry the aorta, s the elastic cle increased. » present (6.2 esure of the nia H&E =p) 6.6 Artery and vein, Stomach (dog) (() small artery. 2) Smal vein, 3) Lymphatic vessel. H&E. x62.5. 6.7 Arteriole, vein and a lymphatic ‘esse in connective tissue, Tongue () Arteriole, (2) Vein. Ghlymphatic vessel. H&E x62. 2) Veins 6.8 Arterioles in the artery (horse). (1) a) Artery. H&E.» Cardiovascular SystemComparative Veterinary Histology with Clinical Correlates Capillaries and venules The capillary isthe smallest unit ofthe vascular sys- tem. The diameter of the lumen is no larger than an erythrocyte and permits these cells to passin sin- ale file only (6.9). The capillary wal is two-layered: 2 tunica intima of one or two squamous endothe- lial cells resting on a basal lamina, and a fine tunica adventitia of collagen and elastic fibres. The endothelial cells usually form a continuous layer, but fenestrated capillaries occur in the renal slomerulus, the endocrine glands, intestinal villi and the choroid plexus where gaps are present between adjoining cells closed by a membrane diaphragm The wall also contains pericytes: undifferentiated cells believed capable of becoming fibroblasts or ascle cells. Venules collect the blood from the cap- illaries. Their lumina are wider than those of the arterioles. The tunica intima (there is no tunica media and adventitia) in each venule consists of a continuous layer of endothelial cells and areolar connective tissue. Pericytes are also present. 6.9 Capillaries (sheep) in the connective tissue of the cervix (arrowed). Masson's trichrome, «125, Sinusoids Sinusoids are found in the liver, spleen, bone mar- row and adenohypophysis. Their wide lumina are lined by endothelial cells interspersed with fixed " macrophages of the mononuclear phagocyte scav enging and defence system of the body {6.10-6.12) Similar thin-walled venous sinuses are found in endocrine glands.inal villi and diapheagm ifferentiated those of the consists of a and areolar (6.10-6.12), re found in 6:10 Liver (sheep). (1) Centra 2) Wer sinusoids. H & 6:11 Spleen thors), Sinusoids filled with erythrocytes. H & E125. 6:12 Pars dstlis of the adenohypophyss (cat) (1) Sinusoids (2) Cords of hypophyseal cell (Orange G. 250. Cardiovascular System z iat sy 610 y 6.12Comparative Veterinary Histology with Clinical Correlates Veins Veins are lined by a continuous layer of endothelial Mile and areolar connective tissue. ‘The tunica sedia, which is always narrow contains a few cit ‘cular smooth muscle fibres, some elastic fibres, but saastie lamina, The tunica adventitia consists of Tongitadinal collagen fibres and, in the large veins ame smooth muscle (6.13-6.16). The lumen com valves that are projections of the tunica intima tate allow only unidirectional blood flow (6.17 and 6.18). 6.14 Caudal vena cava (doo). (0) Tunica Intima. 2) Tunice mecia. {G) Tunies adventitia with vasa Vasorum (arrowed). Gomori’ ‘vicheome. 125. 66:15 Caudal vena cava (horse) (1) Tunica intima. Gy Tunica media. Masson's trichrome. x125,sfendothelial The tunica ins afew cir tie fibres, but ia consists of large veins, ¢ lumen con dl flow (6.17 (doa Cardiovascular System 6.16 Cranial ver (9) Tunica intima Hae x25. Intima. (4) Tunica media. (5) Tunica Aventitia, Masson's trichrome. »25, 6.18 Valve inthe brachial vein (at erytirocytes intima. H&E6.19 Comparative Veterinary Histology wil Clinical Correlates Arteriovenous anastomoses Arteriovenous anastomoses are special areas of the skin of the nose, lips and pads where the art riole opens directly into a venule withou ee a Heart The cardiac wall consists of three layers: endo- cardium (inner), myocardium (middle) and epi- The endocardium contains continuous squa- pres, The myocardium is vascular areolar mous endothelial cells tissue and conduetiny composed of cardiac muscle and also contains vas- cular areolar connective tissue, The epicardium is through the capillary bed (6.19). This provides an Iternative channel of blood supply and regula tion of heat loss. 16.19 Arteriovenous anastomosis in loose connective tissue (a. (1) Artery (2) Velo, H & E3625. thicker than the endocardium, and fat deposits in the r ind coronary blood v Fibrous rings support the heart valves (6.22). The dense connective tissue sels are often found (6.20 and 6.21). provide a means of insertion for the cardiac mus. cle and may be re skeleton (6.23 may becor let horses the aortic rinj calcified to form the ossa cordis. Tw the oss bones form in the aortic rings of cattl 6.20 Cardiac muscle (ox). (1) Cardiac muscle. (2) Connective tissue of the fibrous skeleton. (3) Atrioventricular FL node, Masson's trichrome, x25. 6.21 car () cardi (2) Cond H&Ex 622 + atclum artery. ‘chroand regula- stomossin deposits in d coronary and 6.21). ordis, Two e, the ossa 1) Cardiac ue of the ventricular Fas Cardiovascular System 6.21 caraiac muscle (horse). 5 roe el {) Cardiac muscle bres Ce . ! @) Conducting Purkinje) fibres, | Pa \ ’ : HAE. x125. I ¢ ‘ ? 4 4 Use v ; 1 ; 1 2 4 We hwy ; ; : ) ee 4 \ y \ No \ #0 : “Ss ‘ \ ‘ ee , 6.22 Heart (doo). (1) Lumen of the atrium. 2) Lumen of the pulmonary artery. @) Valve cusps. Masson's trichrome. x62. 6.23 Heart (doa). (1) Valve cusps (Q) Dense connective issue part of the fibrous skeleton of the heart. Masson’ trichrome, »62.5. 621 622Comparative Veterinary Histology with Clinical Correlates Lymphatic vessels cess fluid fr made up of a thin Lymphatic vessels drain nthe tis sues and are + of connective ing. Larger lymphatic vessels, such as the thoracic duct, may have a few smooth muscle fibres in the wall Amphibians and reptiles have perilymphatic and endolymphatic systems that are particularly well developed in some species. These Iymph-filled Clinical correlates ‘Accange of cardiovascular diseases are importan in veterinary medicine. The heart itself can be affected by congenital, degenerative, inflamma tory and neoplastic conditions with a varicty of tinderlying causes. Disease caused by congeni |_ tal malformations is naturally recognized most offen in the young animal. Mitral valve dyspla Sia in an TL-week old Bearded Collie is shown in 6.24. The heart is opened to show the left atri- ular (mitral) valve, The valve leaflets are thickened and malformed and the chordae tend- inae ate short, chick and partially fused. This con genital defect leads to mitral valve incomperenc | Shd produces a holosystolic murmur centred round the fifth intercostal space near the left sternal border In ad ost often encountered in the form of cardiomy wve chang oventri ft animals primary cardiac disease is opathy or degen in the muscle of the heart. In 6.25 | cen used ro highlight ore from the same case, a 12-year-old dog with 624 a ah a 6.26 special stains hat atures of interest. Both te steuctures serve several functions, Perilymphatic pathways encitcle the auditory apparatus and mai participate in the transmission of sounds. Th ndolymphatic system consists of receptor organs of the inner ear as well as either bilateral separate or fused thin-walled sacs. These communicate with the skull via narrow ducts and serve as reservoirs for the storage of calcium carbonat amphibians, endolymphatic sacs form of the extension rebral canal around the brain. myocardial degeneration. In 6.25 a Sirius Red stain, which colours collagen red, demonstrates the lege amount of fibrosis replacing muscle | bundles, which are stained yellow. In 6. Masson's trichrome stain highlights muscle fibres undergoing degeneration. Myocardial degener- ation and fibrosis may be found in cases of dilated cardiomyopathy or as a non-specific response of the cardiac muscle to injury or insult ‘One of the most important diseases to affect the blood vessels is haemangiosarcoma, a malignant rumour ¢ line blood v ad skin astasis can be very widespread the endothelial cells thar spleen, right re common primary sites and Other re also affected. A haemangiosarcoma sels, In dogs tl in a 6-year-old thoroughbred gelding is shown | in 6.27. The smooth muscle in the tunica media of od vessels can be affected by the deposition of calcium salts. induced by oversupplementation of th herbivorous reptiles or amphibians with vi min D, either directly or by inclusion of com mercial dog, cat or prima hypervitaminosis D, ma plemented goldfish ai ‘and amphibians. Early arteriosclerotic mineral- This change is fre food. Similarly be induced when sup- fed to fish-eating reptiles | ination of the tunica media in a la pulmonary artery in an iguana is shown in 6.24 Mitral valve dysplasia in an 11-wweek-old Bearded Collie. Note the thickened, distorted chordae tendinae 625 6.25 ll fir wit ec Tehlymphatic sand may unds. The mist | fe, g reptiles ilmonary iaBearded Cardiovascular System = 6.25, 6.26 Myocardial degeneration and fibrosis in a 12-year-old dog with cardiomyopathy. In 6.25, the 10d red, Sirius Red, x45, In 6.26, degenerative he striations of the muscle fibres are also demonstrated emangiosarcoma in 3 my ans 627 tumour cals are spindle a17. RESPIRATORY SYSTEM “The respiratory system has wo functions: conduc tion and respiration. Conduction is carried out via a continuous system of tubes carrying air from the nos trils into the nasal cavity through the nasopharynx and larynx to the trachea and bronchi. The air is warmed, moistened and filtered in these passages before reaching the organs responsible for respira: Conduction of air The skin around the nostrils has long tactile hairs and numerous sebaceous and sweat glands (7.1). II bur the finer Respiratory epithelium lines divisions of the respiratory tract and consists of sd columnar ciliated cells and mu tion: the hung parenchyma, the respiratory bronchi alveolar sacs and alveoli. It is between the alve 1d the capillaries tha ange takes place The paranormal sinu: avities found in skull bones. The mucoendosteum lining these cm nucous membrane of secreting goblet cells, The lamina propria is con- tinuous with the perichondrium or periosteum where appropriate, is very vascular, contains both mn and elastic fibres, and warms the inspired feromucous glands secrete into the tumen colla through the epithelium (7.2-7.4) 7A Skin, Nostril (hors). (1) Sinus hair. (2) Lamina propria (3) sebaceous glands. (4) Sweat glands. H & E,»20. 7.2 Respl (1) Pseude epithelin @) Lamin: glands. 4 2625, 73 Re (1) Psew epithe @ Lam Hae. 7a the ¢ actiory bronchi- cen the alve= takes place es found in g these cav jembrane of spria is con periosteum tains by the inspired the lumen. 7.2 Respiratory epithelium (horse). (1) Pseudostratified columnar ciliated epithelium with goblet cells, (2) Lamina propria, (3) Seromucous lands, (@) Smooth muscle, H & E 1625. 723 Respiratory epithelium (horse), (1) Preusostratitied columnar ciliated ‘epithelium with goblet cells {2) Lamina propria. (3) Blood vessels. Hae «160 74 Respiratory epithelium (horse) The cla project from the surface of the epithelial cel as fine strands active mucus-secreting cals ie boetween the clited cells (arrowed), Scanning electron micrograph. «1500. Respiratory System ] 72 73Comparative Veterinary Histology with Clinical Correlates Cilia beat towards th mon both to the dig stive and to the respiratory to the middle ear and is common to the digestive The gut h of equids isa diverticulum of the audi yystem. The auditory tube connects the espiratory and auditory systems (7.5 tural pou tory tube. The digestive surface is covered by a stratified squamous epithelium that is continuous with the oral cavity, and the respiratory surface by 78 7.5 Auditory tube (hors (G) Seromucous glands. H & €, x100. respiratory epithelium continuous with the nasal cavities (7.6 and 7.7), The epiglotts is a flap-like structure projecting into the pharynx. A plate of elastic cartilage provides internal support. The upper digestive surface mucous membrane is cov~ ered by a non-keratinizing stratified squamous epithelium (7.8) and the lower respiratory surface by respiratory epithelium (7.9). Taste buds may by present on the laryngeal aspect. (1) Respiratory epithelium. (2) Lamina propria. 7.6 Guttural pouch (horse). (1) Respiratory epithelium. @2) Lamina propria. (@) Seromucous glands. H&E. 250 73 Ge Respir ‘ells ar Q)Lam lands 78 & squan propr Hae 79 col Qt lanRespiratory System a flap-like A plate of s ae glands, Alcian BlUe/PAS. »200, squamous nry surface ids may be 28 Epiglotts (horse) (1) Stratified squamous epithelium. (2) Lamina 9) Seromucous glands. HE, »200. DF @itamina propria. (3) Seromucous BI lands. alcian biuerPas. 125,Comparative Veterinary Histology with Clinical Correlates a 242 1 The larynx is lined by respiratory epithelium. The seromucous glands opening into the lumen. epitheli amina propria is continuous with the perichon- Incomplete rings of hyaline cartilage keep the lumen (2) Seror Jrium of the laryngeal cartilages. The vocal cords patent and smooth muscle fibres bridge the gap at fartlage re covered by stratified squamous epithelium (7.10 the dorsal aspect of the trachea (7.12). In the and 7-11), The trachea extends from the larynx to bronchus the hyaline cartilage has a plate-like the bifurcation of the extrapulmonary bronchi. arrangement. The smooth muscle forms a spiral and These tubes have the same structure: a lining of res- appears as discontinuous blocks in transverse sec piratory epithelium rising on a lamina propria of tions (7.13 and 7.14). A fibrous adventitial coat co Jose connective tissue with elastic fibres and mixed ers the trachea and the extrapulmonary bronchi | 723 | (Re (2) tar | muse open pith | (ry ; | | } 7.10 Larynx (vocal cord: dog), (1) Stratified squamous epithelium. (2) Lamina propria. (2) Simple tubular glands. H & E. 62.5. \ 74 } epit pro @: Gor 7A1 Larynx vocal ied squamous epitheliu Covering the vocal cord, (2) Connective tissue core ofthe lamina propria. (G) Parasympathetic ganglion. H & E,»160.he lumen the lumen the gap at 2). In the plate-like spiral and Respiratory System 712 Trachea (sheep). (1) Respiratory epithelium. @) Lamina propria £2) Seromucous glands. () Hyaline Cantlage. Gomor's trichrome. 125. 7.43 Bronchus (sheep) (1) Respiratory epithelium. 2) Lamina propria. (3) Smooth muscle. (2) Simple tubular glands into the lumen through the helium. (5) Perichondrium. 6) Hyaline cartilage. H & E225. 7.8 Bronchus (ox). (1) Respiratory epithelium; the goblet cells are Stained specifically. (2) Lamina propria. (3) Smooth muscle {@) Simple tubular glands. Gomorilaldehyde fuchsin, »200Comparative Veterinary Histology with Clinical Correlates Respiration Z 6 The bronchi, both e pulmonary and intrapul- | >) monary, bring air to che lungs and branch out [ig ‘within the lungs into the bronchioles, which culmi- nate in clusters of minuce sacs: the alveoli. In the feral lung the duct system is developed wheseas th ‘ respiratory part develops slowly. (7.15-7.17) & Expansion begins with the first respiratory move 2 ments after birth. Thereafter, the lung expands : tandem with the growth of the animal. 1 Each lung is covered by elastic connective tissue a + with an outer layer of mesothelium, the visceral 7.45 Bovine fetal (160-day) lung. Blood vessels (arrowed), (1) Pleural @) Duct system. H&E jesothelium. (2) Mesenchy pleura (7.18). Connective tissue septa divide the Tung into lobes and lobules and the intrapulmonary bronchi have the same structure as the extrapul- monary bronchi (7.19-7.23), The epithelium of ovine fetal (160-day) lung. ERE SLRS 7216 2.16 Braj (1) Large duct lined by a simple columnar epithelium. (2) Smooth eee z muscle. (@) Vascular mesenchyme, k * (@) Small ducts. H & €, 25. aol ‘ 2 a 4 77.17 Fetal lung at term (cow). (1) Respiratory bronchiole connective tissue lined by simple columnar epithelium. (2) Alveol H&E of the visceral pleura 525. Bod vas 225, 720 (1) Lume 2) tami (@ Hyali (6) Adve 720 In () Lum Q) tam (4) Hyal lands. @)Alvenme, pune Respiratory System 7.49 Intrapulmonary bronchus. Lung (cow). (1) Lumen lined by respiratory epithelium. {Q) Lamina propria. (3) Smooth muscle. @)Hyaline cartilage, (5) Blood vessel. (6) ANeol. Gomorvaldehyde fuchsin. «52 7.20 Intrapulmonary bronchus. Lung (sheep) 1) Lumen lined by respiratory epithelium. 2) Lamina propria. (3} Smooth muscle. 4) Hyaline cartilage. (5) Simple tubular Glands. (6) Blood vessel (7) Bronchiole. {@} Alveolar duct, (8) Alveoi. H & E. x62. 721 7.21 Intrapulmonary bronchus. Lung (sheep) (1) Respiratory epithelium. (2) Lamina propria. (4) Hyaline cartilage. H & E.x200, 7.22 intrapulmonary bronchus. Lung (cow). A parasympathetic ganglion is present surrounded by vascular connective tissue of the lamina propria. Gomoraldehyde fuchsin. 160. 7.23 Intrapulmonary bronchus (dog). (1) The intrapulmonary bronchus is lined by respiratory epithelium (arrowed). (2) Alveall. Scanning electron ‘micrograph, »500Comparative Veterinary Histology with Clinical Correlates the bronchioles is columnar or even cuboidal and ciliated. In the smaller bronchioles the epithelium is thinner, the lamina propria is elastic, the smooth muscle forms a complete ring and there is no adventitia (7.24 and 7.25) Non-ciliated bronchiolar cells (Clara cells) are tall, dome-shaped and protrude into the bronchio- lar lumen, They replace the mucus-secreting gob- let cells at this level. Both ciliated cells and Clara cells are present in the terminal and respiratory bronchioles (in the dog and cat they are lined by the latter exclusively). Clara cells divide to form other Clara or ciliated cells and have an important role in the repair of damaged epithelium. Their secretion also keeps the small airways parent Respiratory bronchioles are lined with a low columnar or cuboidal epithelium with ciliated and bronchiolar cells, an elastic lamina propria and a smooth muscle layer. This opens into the alveolar ae ahah oe duct lined by squamous epithelium, interrupted by atria and alveoli along its length (7.26). Alveoli are the functional exchange part of the lung. The septa are very thin, with both elastic and collagen fibres, and contain one of the most extensive capillary net works in the body. Cells of the immune system, derived from blood monocytes, are also present and migrate through the alveolar epithelium into the air space, where they phagocytose particulate matter and micro-organisms to become dust cells (alveolar phagocytes). The respicatory membrane where gas exchange takes place consists of capil lary endothelial cells, alveolar epithelial cells and a fused basement membrane (7.27). The squamous alveolar cell, the lining cell responsible for gas exchange, is a type I pneumocyte. The great alve- lar cell secretes surfactant to reduce surf sion. It is a type Il pneumocyte, is cuboidal and projects into the lumen. 7.24 Bronchiole. Lung (sheep). (1) The bronchiole is ined by cuboidaViow Columnar epithelium with ciated and non-ciiated cells arrowed). (2) Smooth muscle. 3) Alveolar duct, (4) Alveol, The free cells in the lumen are macrophages. H&E. 125, 90 arte arRespiratory System RAP hee” * ee 4 aComparative Veterinary Histology with Clinical Correlates [ec The patterns of disease seen in the respiratory system reflect its structure and function. The ical correlates respiratory tract is constantly challenged by potentially injurious agents, by both the aeroge- hous (arrive in inspired aie) and haematogenous {arrive in blood supply) routes. | "These can include micro-organisms such as bacteria, fungi and viruses or toxic substances or particles (see 7.37) in che ai. This is especially important where large numbers of animals are housed in the same airspace. A bronchiole sur rounded by small cells with dark nuclei and scant cytoplasm is shown in 7.28. These cells, which 92 are lymphocytes, also invade the bronchiolar ‘wall, Often termed ‘cuffing’” pneumonia because of the arrangement of the lymphoid cells around. the bronchioles, this is an example of a chronic, non-suppurative pneumonia, commonly seen in calves. Infection with Mycoplasma species is the ‘The very rich blood supply to the lungs (swhich have the largest capillary bed in the body) also ‘makes them a common target for haematog metastasis from tumours at other sites in the body, Primary lung tumours, both beni malignant, are recognized in older animals. A bronchiolar-alveolar carcinoma, a malignant pri- mary tumour of the lung is shown in 7.29. gn and — 7.28 cutting pneumonia (ah. Lymphocytes cluster around a bronchiole. H & €. 125. 7.29 Bronchiolar-alveolar Carcinoma in an 11-year-old entire bitch. Nodules of tumour, composed of convoluted layers of ‘cuboidal epithelium, infitate interstitial tissue and fil alveolar spaces. Several mitoses are seen. H&E 9250, Avi Vest pass In bir atiniz cond nar ci glandchiolar pease around ron ss the (which ly) also. >genous sin the ign and mals. A ant pi 9, (calf. da ayes of veolar Avian respiratory system Vestibule and conducting passages In birds the vestibule is lined with a distinctive ker atinized stratified squamous epithelium, and the conducting passages with pseudostratified colum nar ciliated epithelium with simple alveolar mucous elands (7.30 and 7.31 7.30 Avian vestibule. (1) The vestibule i ined by distinctive stratified squamous keratinized epithelium. 2) Hyaline cartilage. (3) Perichondtium. () Pseudostratfied columnar ciliated epithelium with intraepithelial alveolar mucus-secreting g respiratory epithelium. H & E>. 7.32 Avian syting pessulus. (1) Respiratory e Respiratory System Trachea chelial tracheal glands are lined with respiratory epithelium and rest on a co: The trachea and intraepi neetive tissue lamina propria, Overlapping rings of ossified hyaline cartilage form the wall. The tra: chea is compressed just cranial to the bifurcation into the primary bronchi. Thin vertical bars of car rod in the angle of the bifurcation (7.32). This is (1) Pseudostratified ial mucus-secreting ,ed). (2) Lamina propria and perichondrium, tilage. HaematonjlivPAS. «160, @) Hal ieee) 732 helium. (2) Vascular lamina propria and perichondrium. (3) Hyaline cartilage. Gomori’ trichrome. >.Comparative Veterinary Histology with Clinical Correlates the teacheobronchial syzinx, the avian sound box. secondary bronchi branch into anastomizi S analogous to the parabronchi. Each parabronchus forms the centre The tympaniform membrai of a pulmonary lobule and is lined with simp! mammalian vocal cords are covered with a strat fied squamous epithelium (7. squamous epithelium. Bundles of smooth muse form spiral bands that are encased by a thin lay of connective tissue he parabronchial wall is perf ading to spaces lined with squamous or Bronel ated with open: The primary bronchi are lined with respiratory ings ium thar rests on a connective tissue lam- cuboidal epithelium: the atria. The air capillaries nrtilage embedded arise from the base of the atria via infundibula and towards the periphery of each lobule. Thi epit na propria with hemirings of fn fibrous connective tissue and smooth muscle. radi hi pass into the lung, where air capillaries are lined with type I epithelial cells they give off secondary bronchi. The epithelium forming the respiratory surface and surrounded by of the secondary bronchi contains goblet cells, rather than glands, and cartilage is absent. The tissue (7.34). The primary bror a mass of blood capillaries to form the air exchange chotrachea! laryins S733 Avian br () The tympa covered by a stratified squamous epithelium. (2) Respiratory epithelium, 3) Lamina propria, (a) Pericnoncrium. (5) Kyaline cartilage, H & E200, = 7.34 Lung (bite. (1) Lumen of the porabronchus (tertiary bronchus) (2) Conical ducts, atria.) Alr Capillaries (8) Blood vessels filled ith nucleated erythrocytes. H & E H625.tomizing the centre th simple th muscle thin layer vith open- -apillaries ibula and ule, The lial cells punded by exchange 2a larynx ] pra ine en of the pchus) air Is filled eH, Air sacs Air sacs are thin-walled structures lined by a squa- ‘mous epithelium (they may be ciliated or colurnnar resting on a thin layer of connective tissue. The blood supply is poor and, with the exception of the abdominal air sac (10 branches), they are connected to the secondary bronchi. The humerus and the ster num ate some of the bones penetrated by extensions Reptilian, amphibian and fish respiratory systems Most fish (except lungfish, the bowfin, some cat fish and a few other teleosts) either lack lungs or possess only primitive elongated sac-like lungs. They must rely upon vascularized gills in order to extract oxygen from and excrete carbon dioxide into their aq) nposed of parallel rows of gill filaments, the primary lamel- lae, which are supported by cartilaginous or bony ic environment. Gills are ¢ rays forming semilunar folds: the secondary lamel- Jae. The gill arches contain a fine vascular network of branchial arteries, arterioles and capillaries, across which respiratory gases are exchanged and osmoregulation (in conjunction with the kidneys) is maintained. In salmonids, eels and other fish Respiratory System that alternate between freshwater and marine 4 electrolyte secreting cells of th cyclic roles in osmoregulation. branch gills play major eleost fish also isa moderately ure that is derived dur- 1g embryological development from the first gill arch (7.35). Its function is believed co be the reg possess a pseu compressed gill-like seru ulation of blood oxyg The gills of amphibians are similarly structured and function in a similar manner. Some amphib: ians possess both external gills and internal sac-like lungs, which serve not only as organs of respira tory gas exchange but also have a hydrostatic func tion. When the sac-like lungs are filled with air, the amphibian becomes more buoyant. When these lungs are empty, buoyancy is lost and the animal sinks to the bottom of the water, thereby requiring lietle or no effort to remain submerged, Adult plethodontid salamanders lack lungs entirely their gas respiratory exchange is accomplished solely by diffusion across the well vascularized moist integument. In some amphibians, lungs are much reduced in size; in others, only a single lung is present. Many amphibians augment their pul monary and integumentary respiration by buccal movements that help move gases across their oropharyngeal mucosae where some gas exchange 7.35 Pseudobranch gland of a zebra dani fish This structure is unique to teleost ish and is derived embryologically from the first gil. Its composed of a cartilaginous ‘skeleton’ from which parallel lamellae that resemble compressed gill laments int lersect perpendicularly. This organ fs thought ilation of blood oxygenation, H & E, x20737) Comparative Veterinary Histology with Cli Clinical correlates The hungs of many diurnal amphibians and rep- tiles are heavily pigmented with melanin (see also Chapter 3). This pigment is believed to confer | protection against the effects of solar cadiation. AA section of lung from a pipiens) is shown in 7.36. [As described on page 92, lungs (and similarly 1 Correlates. gills) are vulnerable to pathogens present in both the extemal environment and the blood (7.37). In addition, the delicate capillary bed through which respiratory gases exchange is also predisposed to thromboembolism, because of the small cross-setional area of these vessels. 7.36 Lung of a terrestrial frog Rana pipiens). H & E200. 7.37 Aspiration pneumonia ina lizard, Note the plant fibre within the airspace. H & E125,e blood ary bed seof the fog 0 within 8. DIGESTIVE SYSTEM The digestive or alimentary tra e. The mucous membrane on th nective tis avity and terminates at the undersurface consists of non-keratinized stratified tube lined with a mucous squamous epithelium with a lamina propria (8.1) nd the alimentary canal. tongue, where the epithelium is keratinized, is oath Th bai Wan eal Oral cavity Snake heavily Tongue The bulk of the tongue is made up of interlacing bundles of skeletal muscle fibres and loose con- Tongue (cat) (1) Stratified jamous non-keratinized epithelium, (Q) Vascular lamina propria with small projections. @) Striated muscle fibres. H&E 225, 8.3 Tongue (cow). (1) Flferm papi Lenticular papilla, Masson's trichrome. 21.5‘comp: walls, The circumvallate papilla is surrounded by a moatlike trough oF vallum and is level with the surface of the tongue (8.4). The fungiform papilla is, as the term suggests, mushroom- shaped with a narrow base, is partly or non ative Veterinary Histology with Clinical Correlates keratinized, and projects above the surface of the tongue (8.5). The foliate papilla is large, non-ker- atinized and leaf-like, is crossed by transverse fur rows and appears in section as a row of fungiform papillae (8.6), £84 Circumvallate papilla. Tangue (cow). (1) Stratified squamous epithelium. {@) Vallum. @) Taste buds. (4) Lamina propria. (5) Striated muscle. (6) Mixed salivary gland. Gomor's trichrome, «100, 5 Tongue (cow). (1) Taste buds in the stratified squamous epithelium of the lsteral wall of the papilla, The taste pore i arrowed. (2) Connective tissue lamina propria, Masson's trichrome. >250,Digestive System face of the ¢ epithelial structures associ ibres of the facial and gl sustentaculas (support 1g) cells. Food dissolved in sion passes into the reser- sophar} (8.7) fangiform pore, whic nto the surface of the tongue, The lingual tonsil isa localized mass of lymphoid and a taste chamber lined with a taste receptor and tissue that is often present in the base af che tongue e, non-ker: the salivary glan: sverse fur Within each bud is a taste voir of hai 86 Foliate papila, fungiform papillae. H&E 625, (rabbi) Ths papilla appears as a taste buds are arranged along the la ‘inl 1ud. Tongue (cow). (1) Taste pore. (2) Taste receptor and sustentacular cells H & E5625Comparative Veterinary Histology with Clinical Correlates Sali ary glands nds_are compound tubuloacinar Salivary gh exocrine gl mucous of mixed salivary glands, a mixture of enzymes and mucus. The secretory component of each gland is the parenchyma and the supporting 8.10). In serous and mucous units connective tissue is the stroma ( mixed salivary glands t may be separate or the tinctive cap on one side of am ith caused by the vulum, The demilune (8.11). The serous cell is column a basal nucleus and basal basophil presence of rough endopl: ‘uminal eosinophilia is caused by dl granules accumulating before secretion. The ¢ with a basal flattened mucous cell is trian; nucleus and a pale staining vacuolated cytoplasm. Specialized epithelial cells, the myoepithelial or 100 basket cells, are capable of contracting: these lie between the secretory cells and the basement mem= brane (see Chapter The dilute salivary secretion leaves the acinus and js concentrated in the first part of the duct system: the striated duct. The basal cell membrane is, infolded and the cytoplasm is lined by mitochon dria, This allows water to be removed, passed into the tissue fluid and back to the blood. Palate The hard palate is lined with stratified squamous epithelium with the lamina propria continuous with the underlying periosteum (8.12). The oral surface of the soft palate is also lined with stratified squamous epithelium, but the lamina propria has mucus-sec glands and lymphatic nodules. The nasal surface is covered by respiratory epithelium (8.13). 8.8 Mixed salivary gland (cow). The ‘of seromucous acini. (1) nterlobular connective tisue with (2) blood vessels. (3) Interlobular ducts. Hae o2s, 8.9 Mixed salivary gland (cow), ‘Mixed seromucous acinus with () pale staining mucous cells and (@) darkly stained serous cells ‘Masson's trichrome, 125,Digestive System 8.10 Mixed salivary gland (clog). 0 bya lumnar epithelium. @) ye tissue stroma, ©) ganglion. acinus and @) Seromucous acini H&E. «125, act system mbrane is mitochon- assed into squamous h I surface of squamous sal surface 3 ‘ound fucleus. Gomor's trichrome, x125, els and cl, B12 Hord palate (ox). (1) Stratified squamous 18.13 Soft palate (ox) (1) Respiratory epithelium, ‘epithelium. (2) Lamina propria, (2) Mucus-secreting @) Lymph nodule. (3) Sone. H&E, ¥625, glands inthe lamina propria. H & E125,Comparative Veterinary Histology with Clinical Correlates Teeth secrete cementum in a similar pattern to that of ee pone. Enamel and dentine are involved with the ere- smn al papillae within the enamel organs (8.14). The involved in the soot. The root is formed by an the ou late reticulum (8.15). The mesenchymal cells of the _ brane of collagen fibres em papilla dif oblasts laying down enamel continuous with (8.17 and 8.18). The tooth is held in the develop. enamel epithelium and enclosing the stel- ing mandible and maxilla by the periodontal mem edded in the cementum, dontoblasts, the Temporary teeth develop first. The permanent teeth dentine-forming cells (8.16) and cementoblasts, and are secondary offshoots on the lingual side of th rentiate to become o¢ aaa 8.14 Developing tooth (cat embryo). 1) Oral epithelium. 8.15 Developing tooth (cat embryo). (1) Mesenchymal @) Enamel organ surrounded by mesoderm.H &E. 125. papilla with a layer of odontoblasts (arrowed). (2 In 8.17 enamel epithelium, continuous with (2) the outer ena en epithelium. (4) Stellate reticulum. H & E, x62. lat (arr 102Digestive System ' 1.16 Developing tooth (at embryo) (1) Mesenchymal to that of sertal papa. @) Amelobasts inthe Inner enamel ‘ ith the cre ter enamel epithelium (8) Stelate a ig tion of the foot tubule enchymal Es ) 2) ner 8.18 Developing tooth (al). The ameloblasts are ter enamel tall columnar cells with a basal nucleus (arrowed) 5s. ithe ot ne Hae «125, lum atthe point of root development 103Comparative Veterinary Histology with Clinical Correlates meal temporary teeth. The tooth is divided into a crown and a root (8.19). In the carnivore, teeth cease to Dentine ‘grow after eruption and the ameloblast layer is lost: brachydont teeth. In the horse, ruminant and pig, teeth are much longer and continue to grow for all ‘or part of adult life: hypsodone teeth. In these the dental sac covers the whole of the tooth before — Pulpcavity | eruption and cementum covers the entire tooth, pre venting loss of the ameloblasts and Pulp cavity Oropharynx This is a short junctional area between the oral cav- ity and the alimentary canal with some mucus- secreting glands (8.21). Periodontal membrane Alveolar 77) bone 8.19 lagram of an incor. a20 cementum crown ~ Enamel Dentine Pulp cavity cementum Enamel ah ——— Dentine Pulp cavity Root Maxillary ME attntor | bone 8.20 Diagram of a molar 8.21 Oropharyne. (1) Stratified squamous epithelium. (2) Mucus secreting glands open onto the surface (arrowed). H & E625,re ee ee th cease to at and pig row for all oth before tooth, pre ese species ) Mucus st the 225, Clinical correlates Any by inflammation. Gingivitis, or inflammation of level of the intestinal tract can be affecte: the gums, is common in dogs and cats, of association with dental dis A gingival biopsy from a dog in which the gin gival epithelium is irregularly hyperplastic is shown in 8.22. A dense inflammatory infiltrate ‘occupies the superficial submucosa and extends into the mucosal epithelium, Plasma cells (mature immunoglobulin-secreti Is} predominate in the infiltrate. Their presence indicates a persistent antigenic stimulus, The initiating disease may be Alimentary canal The alimentary canal (digestive tract) is a muscular tube extending from the oropharynx to the anus, ‘comprising the oesophagus, stomach, and small and large intestines. ‘Two large glands, the liver and pan reas, are also derived from the embryonic alimen. tary canal, Each part of the canal has a specific function and the histology reflects this (8.23). The canal wall is derived from endoderm and splanchnic mesoderm and consists of four layers: ‘© Tuna mucosa (mucous membrane), with epithe lial lining, supporting vascular lamina propr 8.23 Diagram TS, Alimentary canal Extratubular glands diver and pancreas) Digestive system £8.22 Plasmacytic gingivitis (dog). H & E125. and lymphatic cells. Mucosal glands, which are derived from the epithelium, ate variably present. The outer muscularis mucosae (absent from the mouth, pharynx, portions of the oesophagus and rumen) is smooth muscle Tela submucosa: a connective tissue layer with mphatic tissue and nerve plexi. Submucosal glands may be present. Tunica muscularis: smooth muscle (except in the ‘oesophagus and the anus where the muscle is vo uuntary, striated muscle 823rr TTT, Comparative Veterinary Histology with Clinical Correlates Oesophagus The oesophagus is lined with stratified squamous epithelium, and both mucosal and submucosal ‘mucus- or seromucus-secreting glands may be pre sent. In ruminants the muscularis externa is skele~ tal muscle; in the pig and cat the distal part is smooth muscle (8.24-8.27) £8.24 esophagus (ca) (1) stratified squamous Keratinized epithelium, (2) Lamina propria. 3) S vein. @) nner layer of circular smooth musce. (5) Outer layer ‘of longitudinal smooth muscle. # & E. LP. orien esas Gs 825 18.25 esophagus (doa) (1 Stratified squamous ‘epithelium. (2) Mucus secreting glands in the lamina propria, (Inner layer of circular skeletal muscle. (4) Outer fayer of longitudinal skeletal muscle. H & E x12. £8.26 esophagus (cat) (1) Stratified squamous ‘epithelium. @) Lamina propria. 3) Muscularis mucosae. {@) Suamucosal mucous lands. (5) Muscularls externa Masson's trichrome. «125,8.27 Oesophagus (dog). (1) Stratified squamous epithelium. (2) Lamina propria, (3) Muscularis mucosae : (@) Submucosal mucous glands (5) Striated muscle of the muscularis externa, H& E,»62.5, 7 Stomach Bea: eck: crates in the sagle nomset cf teed lle cf ences glanlas a SE Gia incseyeuere a daeeopanc tier seop aen gion soda glandular n'8.29) Inth 8.28 Oesophagealstomach junction (horse). The ‘epithelium changes abruptly from stratified squamous to simple columnar at the junction (arrowed). H & €.>62:5 scosae F229 oesophageatstomach junction thors). The cena epithelium changes abruptly from stratified squamous to Simple columnar at he junction (arrowed). H&E. 10, Digestive System 107Comparative Veterinary Histology with Clinical Correlates 830 8.30 Rumen (sheep). (1) Stratified squamous epithelium lines the rumen. @2) Lamina propria. Masson's trichrome, «125, i eegees eae eat The lining epithelium is stratified 8.32 Reticular groove (goat). The mucosa is folded; this, squamous, The lamina propria is loose connective tissue, allows stretching, The lining epithelium is stratified | stained green, Gomor’s trichrome. x125, Squamous. H&E. x12. omasum; the glandular stomach is a separate com: partment: the abomasum (8.30-8.35 The non-glandular stomach is lined with a stra ified squamous ¢} yn, In the ruminant clear vacuolated thelium with some keratiniza cells in the epithelium give ita distinctive appearance and allow the transfer of water, electrolytes and short-chain fatty acids. Muscularis mucosae is present in the from the rumen. The glandular mucosa of the stomach is folded and lined with a simple columnar mucus-secreting epithelium. The gastric pits, the foveoli, are surface depressions co uous with the simple tubular gas tric glands (8.36). Three histological regions are 18.33 Omasum (sheep). The muscularis mucosae is present in the long omasal fold (arrowed). H &E-x125.Digestive System 7 | 835 (Goat). (1) Mucosa cosa, (4) Muscuaris 2s, 1836 Stomach (horse). [ ATT 836 ) Simp pits. H&E. 625. oh gions are 109Comparative Veterinary Histology with Clinical Correlates ining, basophilic opria HAE W128, ng. (3) sculatis mucos 8.38 Cardiac glands. Stomach (horse). (1) Parietal cel ll. (3) Lamina pro TRS £8.39 Fundic glands. Stomach (horse) Simple mucursecreting epitheliun ‘extends into the gastric pits ved, (1) Parietal celDigestive system P recognized: the cardia, the fundus and the pylorus. Glands are sparse E with few cells in the cardia, but are abundant and cellular in the fun dus (8.37-8.42) ynogen 8.40 Fundic glands. Stomach (dog). Simple columnar rmucus-secreting epithelium extends into the gastric pits (errowed). (1) Parietal cll (2) Zymogen cel (@) Muscularis mucosae, (4) Submucosa. H & E125, ‘ 8.41 Fundic glands. stomach (dog). Simple columnar mucus secreting epithelium extends into the gastric pits (arrowed), (1) Parietal cell. 2) Zymogen cell. H&E, 160 hs (hors 8.42 Fundic gland, Sn Stomach (cad (1) Parietal ell.) Zymogen cell H&E, n250, .Comparative Veterinary Histology with Clinical Correlates There Pars ¢ five cell types in the gastric glands: m cells at the neck of the gland divide and lace the surface epithelium. pl © Mucous neck cells atthe neck of the gland secrete © Parietal (oxyntic) cells are large polyhedral cells with a central nck and cosinophilic cytoplasm. They secrete hydrochloric acid into canaliculi and claborate invaginations of the plasma membrane ‘© Chief (rymogenic, peptic) cells secrete the enzyme pepsinogens this ic acid. In common with all enzyme-secret converted into pepsin by cells, the rounded basal nucleus is surrounded with basophilic cytoplasm (rough endoplasmic m). The apical cytoplasm is eosinophilic (stored secretory geanules * Enteroendocrine cells are a diffuse population that are identified with specialized silver stains and are also known as argentaffin and argyrophil cells, Chemical messengers (serotonin, gastrin, somatostatin and enteroglucagon) are secreted locally to control digestion. These cells are 8.43 Pylori glands, Stomach (pia) Simple columnar ‘mucus-Secreting epithelium extends into the gastric pits (arrowed). (1) Simple tubular mucus-secrating glands. (@) Muscular mucosae, (3) Submucosa. H & E.»25. 2 regarded as part of the uuptake-decarboxylation’ (APUD) cell system, which is characterized by the ability to take up and process biogeni ; not all of these cells process amines and the term ‘i fuse neuroendocrine system’ is more ‘amine-precursor mines. Howeve The pyloric glands are mucus secreting (8.43 and 8.44) The lamina propria is loose cellular connective tissue with Iymphatic cells present asa local popu- lation and part of tl t-associated lymphoid tis- sue (GALT). The muscularis mui 's of smi mucosa is aglandular loose connective tissue with parasympathetic nerve plexi (8.37-8.44). The mus: cularis externa consists of three layers of smooth agi coca muscle fibres. The sub: of several lay oblique, circular and longitudinal. The myenteric parasympathetic nerve plexus (Meiss net's) lies between the muscle layers (8.45}. The outer laye erosa, is vascular connective tis- esothelial cells continuous with the visceral peritoneum, sue covered with n 8.44 Pylori glands, Stomach (dog). Simple columnar mucus-secreting epithelium extends into the gastric pits (arrowed), Simple columnar epithelium lines tubule seen here cut in transverse section, H& €. x10.I system, >take up not all erm “dif 8.43 and cal popu phoid tis: omposed The sub- gsue with The mus f smooth nal. The s (Meiss 45). Th tous with e gland ex100 8.45 Myenteric nerve plexus. Stomach (horse) Parasympathetic neuron cell Bodies (arrowed) lie in the connective tse between the smooth muscle layers. Hae, wi25 Clinical correlates Gastric lesions may be associated with many conditions which have signs that also affect other body systems or other levels of the gas: trointestinal tract. The aetiopathogencsis of peptic ulceration (8.46) is incompletely under- stood, but it results from an imbalance between the damaging effects of gastric acid and pepsin and the protective mechanisms of the gastric Digestive System mucosa. Administration of non-steroidal a inflammatory drugs is known to predispose to gastric ulceration by inhibiting prostaglandin metabolism and damaging the gastric epithe- um. Systemic disturbances, such as endotox aemia or uraemia, may produce gastric lesions and complex factors associated with stress can also be implicated. Gastric ulceration can pro- duce abdominal pain, vomition or haemateme- sis, melaena and anaemia, 346 a 8.46 Peptic ulceration (dog). Loss of mucosal epithelium is seen, with eosinophilic necrotic debris within the defect. Granulation tise Is Aeveloping in the base of the ulcer. H & E. x22,Comparative Veterinary Histology with Clinical Correlates | Small intestine f The sn jejunu | 7 | 2 x oe 8.47 D1 m (dog). (1) Mucosa. (2) Submucosa (3) Muscular externa. (4) Serosa. H & E25. 8.48 Duodenal vill (doo) into the lumen of the duodenum. 8fenum, nof the sof the res and che sur villus is | 849 Goblet ined Digestive system | formed by the lamina propri lular and reticular, with loc phoid cells. The tall columna ions of lym: lls that line the intestine have a striated border containi ting goblet cells; these increase in h distance from the stomach. At the bases of villi, the epithelium dips into the lamina pro: > See , of Lieberk cells lining che crypts are colun nnd replacement of the epithelium, Paneth cells, 8.51 Duodenum (horse). (1) Simple columnar epithelium with goblet cell: 2) Lamina propria with smooth muscle fibres (arrowed). (3) Contractile crypts, H & E250, 8.52 Duodenum (do). (1) Mucosal glands. Muscularis 8.53. Ouodenum (do). The lamina propia filed with ‘ucosee. (3) Submcoral glands. H & E9160. lymphatic tisue and imphoeytes ate seen migrating fhrough the epithelium (o Payers pa obular leucocyte 8.85 Enteroendocrine argentatfin cel (cat). Argentaffin inthe epithelium of the intestinal cell tains black in the intestinal gland epithelium, present in the lamina propria Methanamine slversafranin, ~500,EE Comparative Veterinary Histology with Clinical Correlates which contain secretory granules that contain pep- sidase, may also be present in horses and ruminants The muscul smooth muscle, nal, and separates the crypts from the underlying mucosa. A strip of muscle extends into each villus utlaris mucosae; 2 lacteal (lymphatic mucosae consists of two layers of ner circular and outer longitudi- from the m the villi, called contractile crypts, are created by the contraction of the central strip of the muscle. Mucus- and seromucus-seereting submucosal glands are found in the horse in 6-7 m of the intestine, 3-S m in the pig, 4 m in the cow and 60-70 cm in sheep. The muscularis externa consists of two layers of smooth muscle dispersed in a gentle spiral, appearing fas an inner circular and outer longitudinal layer. As in yarasympatheric nerve plexus (Meissner’s) can be found between the layers. The serosa consists of loose connective tissue and the mesothelium is continuous with the visceral Large intestine There are no villi in the large intestine (caecum, colon, rectum and rectal canal). Goblet cells are abundant in the surface epithelium and in the mucosal glands, which are simple regular tubules. Lymphoid tissue is present in the lamina propria, a he epithe- bathes the against luminal are eosinophil leucocytes associated with nfestations, Lymphocytes are present in lium when immunoglobulin, wh epithelial cell surface as a defer antigen, is released. ‘There are no submucosal glands. A muscularis mucosae is present, and the muscu: laris externa consists of an inner circular and outer £8.56 Colon (horse) (1) Mucosa, (2) Muscularis mucosae. (@) Submucosa, H & E97. £8.57 Colon (horse). (1) Simple columnar epithelium with goblet cells (2) Intestinal mucosal glands. H&E. «125. longitudinal layer of smooth muscle. The outer layer, in bands and is charac the horse and pig. In the horse, elastic fibres replace muscle fibres. The serosa is continuous with the peritoneum (8.56 and 8.57) The rectum is lined with simple columnar epithe lium. The mucosal glands decrease in number and as the anus is approached, 1 taenia coli, is istic of the colon o may disappear entirely ‘where there is an abrupt change to a stratified squa. mous epithelium, The muscularis externa is thicker here and becomes striated at the anal sphincter. Part of the rectum is covered by a serosa and the rest b dventitia. Tubuloacinar anal glands are present at the cutancous-rectal junetion where they secrete lipids in carnivores and mucus in the pig. In carn vores, circumanal, sebaceous-secreting glands are nal canals. Anal sacs, opened by small ‘glands and lined with stratified squa- found in the a tubular alveo ‘mons epithelium, open into the perianal region (8.58) GALT is part of the immune system. Both T and B lymphocytes, as well as macrophages and eosinophils, are present. The tissue may be so pro- fuse that the enterocytes mass. Lymphocytes may fare stretched over a bulging rate through the epithe lium (see 8.53). sq ti als Hierlayer, charac In the j 857) ithe rand oached, squat thicker rer Part rest by nds are by smal bulging epithe 357 Digestive system 8.58 Anal sa¢ (dog). (1) Stratified Us Keratinized epithelium lines the sac. 2) Tubuloalveclar lands in the lamina propria, Hae x25 Alimentary system of reptiles and amphibians When amphibians met adults, si e from la take place in fo funetion. Many larval amphi (or oblig ory) herbivores, the alimentary tracts of which are elongated and of tly coiled (par ticularly in frog and toad larvae). During the latter ges of metamorphosis, postlarval amphibians ally cease eating and, therefore, must subsist on r tails and other sources of readily catabolized tissue. As adults, most amphibians are carnivorous. The lingual ap 9 Tongue a potonarow rg Dendrobte sp) 3 dra igual race) scovred by an usual reptiles is modified forthe apprehension of prey: and comple epithelium composed ofa, doening nus (889 and 8.60 cchersarechracteied by Summa es thtnattan aovting of ghee min ‘numerous papillary projections at the lingual tip to. The muscle fibres (2 are primarily arranged in a ‘a ain ociections EOL eet ee er Ma occ Parcs ance and are then brought into the mandible, This facilitates the tongue being rapidly fh, When not being used, the tongue of protruded and retracted in order to catch smal cts into lingual sheath thats ined by fnetebats W €on2 ratus of many amphibians and snakes re £8.60 The tongue of some lizards overlies a sublingual salivary gland (1), a silustrated by this longitudinal section of the tongue of a small sink (Sc ali The dorsal surface is covered by a nor-keratinized stratified squamous epithelium @2) in which cup-shaped taste receptors are embedded. Some lizards (for example, Iguanines) possess tongues with a terminal tip ©f papillary projections that are kept moist and mucus secreted by goblet cells and several salivary glands. H&Esz a1 f Comparative Veterinary Histology with Clinical Correlates mucus-secreting glands. It does not contain glands bur is lubricated when it comes into contact with the luminal surface of the lingual sheath, The congues of other reptiles contain taste receptors that are similar to taste buds found in the tongues of mammals. The dental histology of amphibians and reptiles is similar to that in mammals, although the teeth of these animals are periodically and continually shed. throughout life, Chelonians (turtles, tortoises and. terrapins) lack teeth entirely. Theit premaxillae nd mandibles are covered with hard and ramithecae, with food items. maxillae horny ke which ehese animals cut th The salivary glands of amphibians and reptiles. are similar to those found in mammals. They may be either entirely serous, entirely mucus-secreting, or a mixture of the two. tinous surfaces, ca In venomous snakes and helodermatid lizards (the Gila monster lizard, Heloderma suspectum, and the Mexican beaded lizard, Heloderma horridtem) some salivary glands are greatly modified into structures (see Chapter 2) that secrete extremely toxic secre- tions that help these animals capture their prey and defend themselves. In venomous snakes the secre tions from these glands are conducted to the hollow needle-like fangs through coiled venom ducts. The passage of venom through these ducts is aided by the contraction of the temporal and masseter skele- tal muscles that surround the glands and myoep- ithelial cells that surcound the ducts (see Chapter 2 The fangs are replaced periodically throughout a snake's life, They are formed with a separate hollow channel (8.61). Some nominally non-venomous snakes, especially many colubrids, possess modified salivary (Duvernoy’s) glands (8.62), the secretions 8.61 The fangs of venomous snakes are continually being renewed. Mastrated are several teeth primordia of a juvenile rattlesnake {Crotalus spp), forming modified fangs with a cantral enamet-lined channel theough which venom i Conducted. H & E3425. 8.62 Some non-venomous snakes possess modified (Duvernoys) maxillary and premaxillary salivary Glands connected to short ducts that empty into the oral cavity. Current Studies indicate thatthe secretions from some of these glands manifest venonlike bioactivity on the lower vertebrate prey ofthese snakes. Also, mild clinical envenomation Of sensitive humans bitten by these snakes has been reported. Nlustrated fare two lobules of gland froma ‘watersnake (Natrix cycopion) H&E 625. paand the ehollow cts. The aided by er skele- | myoep. apter 2) wghout a e hollow modified ys snakes 1 Jesnake sified ined 9) salary ducts that Current the lower kes. these ihsrated ‘of which induce a toxic reaction when injected into particularly sensitive pi Generally, the alimentary system of the low tebrates is similar to that found in mammals, but fey and humans. major variations exist in species that are highly adapted to a particular die reptilian herbivores utilize hindgut rather than foregut fermentation to accomplish the processing Folivorous (leaf eating) of cellulose and other complex carbohydrates. Modifications that aid in this process are an expanded sai tion to bits and hares) and some herbivorous rodents, and co the massive caecum and colon of equids. In all hese organs, the surface of the luminal lining is aug. ented by numerous mucosal villous projections, which greatly increase the area available for micro- slated colon, which is similar in func tundus of lagamorphs (rab: bial digestion and nutrient absorption. Thus, the sacculated colon of reptilian folivores serves the same purpose as the large rumen complex of rumi: 8.63 The oesophagea! lumen of many chelonians, such as this green sea turtle (Chelonia mydas), is heavily goblet cells, These characteristics, reflect the scabrous diet of these marine animals. (1) Stratum corneum. (2) Stratum lucidum. (3) Stratum ‘granulosum. (4) Stratum spinosum 5) Stratum basale. (6) Musculars eterna, H&E 125. 8.68 The oesophagus of most snakes and many lizards is characterized by its extensive plating which permis the agus to stretch to accommodate enormous prey llstrated i a cross-section of the oesophagus af a kingsnake (Lampropeltistriangulum). Because of the 1ecessity for abundant lubrication during the swallowing of furry, feathered or scaly prey the oesophageal lume red by a mucous epithelium composed of simple ron-keratinized columnar cells bearing basal nucle! H&E x25 Digestive System nants, even though it is part of the hindgut rather than the foregut. he anterior alimentary tracts of various reptiles are modified. The oropharynx and oesophagus of some sea turtles have a heavily keratinized lining (8.63) that helps to protect the lumen from traum: when scabrous food items such as rocky and silic rich coral are swallowed. The egg-eating snake (Dasypeltis scabra) ingests eggs with calcareous shells. As the egg enters the cranial oesophagus, the nake contracts its thro: the egg against multiple horny ridges that extend from the ventral region of the cervical vertebra: ‘eby compresses After the eggshell is slit, the snake swallows the fluid and/or embryonic contents and regurgitates the she fragments en masse, Most snakes and many lizards possess an oesophagus with walls formed into mul iple longitudinal plaits that permit the swallow. ng of enormous meals (8.64), many times the diameter of their necks, Other reptiles, such as mostComparative Veterinary Histology with Clinical Correlates crocodilians, have thick-walled muscular stomachs in which their prey are macerated with the aid of ingested stones, The gastric mucosa of reptiles is similar to that found in mammals, except that only chief and clear cells are present; parietal cells are lacking (8.65 and 8,66). The small intestine lacks Briinner’s glands (see Chapter 3). The serosa covering most or all of the coelomic viscera of many diurnal lizards ish pigmented (8,67). Lymphoid patches or aggregates sughout the length of the alimen tary tract. Discrete lymph nodes are not present. 865 8.66 Gastric mucosa of a boa constrictor (Boa constrictor. The lumen Is covered by tall columnar epithelium. The gastric glands consist of only granular, cuboidal, pink staining chief cells (1) with large vesicular rnudel, and pale staining clear cells (2) whose nuclei are ddatk and basa Some gastric pits are lined by both cell types. H&E, 5250, Many secreting glands through which hyperosmolar solu tions containing sodium, potassium and chloride ions are secreted. In many lizards these glands are izards and some snakes possess salt situated in the nasal cavity. Some sea snakes possess sublingual salt glands. In some crocodilians, par ticularly crocodiles that inhabit salt marshes and travel between oceanic islands, salt-secreting glands are located on the dorsal surface of the tongue. All Of these aforementioned glands permit the non renal secretion of electrolytes without the appre ciable loss of water 18.65 Whole mount cross-section ofthe fundic stomach of ‘a small skink (Scincella laterals). A very thin serosa covers, the outermost visceral surface, The gastric wal is ‘composed of an outer external longitudinal muscularis ‘externa (1), a circular muscularis externa, the muscularis mucosa), and immediately beneath isthe glandular mucosa (2) whichis composed of pink staining granular chief cals and clear cells. The lumen is ined by tall mucus: secreting columnar cells Parietal cells, present in mammalian gastric mucosae, are lacking in amphibians ‘and reptiles. The outermost surface of the stomach is ‘covered by a delicate serosa (2) formed of non-keratinized squamous cls. H&E «125, 867 37 The colon of some lizards, particularly folivorous specie, js highly modified sacculated organ divided into ‘multiple chambers that are funetionally analogous tothe hindgut of lagamorphs and some (herbivorousfolivorous) rodents, and the forestomachs of ruminants. Digestion enhanced because the villous surface of the colon i covered by a highly absorptive columnar mucosa across ‘wich nutrients processed from cellulose-digesting micro ‘organisms are assimilated. The elongated vil that cover the surface are supported, and stiffened, by thin cores of smooth muscle. lustrated is the sacculated colon of a ‘reen iguana iguana iguana). H & Exess sal pole sol chloride Binds er ans, par ee Al he appre stomach of, phibians keratinized vided into Falivorous) | | | cl Alimentary system of reptiles and amphibians al correlates Squamous metaplasia of the nasal and pharyn- geal mucosa (8.68 and 8.69) is a frequent clin- ical condition in reptiles fed diets deficient in B-carotene, Once this alteration ‘occurs, the lubricative mucoid glandular secre- more susceptible to respiratory and oropha: ryngeal diso: vitamin A 0} ses and the affected animal becomes 8.68 Massive pharyngeal hyperkeratosis in a desert The pharyngeal glands are replaced by pearlike masses of desquamated keratin. The luminal epithelial surface is thickened and covered by dense keratin debris. A similar alteration is seen in birds and ‘mammals suffering from vitamin A deficiency Hae x25 8.69 Cross-section of the pharyns of a ed-eared sider turtle that was fed a diet setiously deficient in earotene (or preformed vitamin A. The pharyngeal glands display Squamous metaplasia and, asa ‘sult, have lost their mucus secreting, gabletcal-rich lands, which have been replaced by masses of ddesquamated keratin debris (i). The stratified squamous epithelium lining the pharyngeal lumen is thickened and hyperplastic. H & E. 32.5 Digestive System Ulcerative stomatitis is one of the most com. mon conditions found in the cranial alimentary tract of captive snakes. This infectious matory di ase is caused by a va genic Gram-negative and so bacteria. Depending upon the the inflammatory response may be sup or non-suppurative, In suppurative lesions, het erophil granulocytes peedominate; in non-sup- purative inflammations, heterophils may be entirely absent. Glossitis, pharyngitis, oesophagitis and gast tis also occur in captive amphibians and reptiles, ‘ety of patho: Gram-p ‘ologic ase 269Comparative Veterinary Histology with Clinical Correlates These inflammatory conditions are often caused he delicate by items in the diet that injure ‘mucous membranes that cover the tongue or line these cavities, In snakes, and to a lesser extent in lizards, gastric eryptosporidiosis is a serious clin ical problem. Although typically termed ‘hyper trophic’ gastritis in the literature, the anatomical and histological features of gastric cryp rosporidiosis in snakes are a marked hypertro. phy of the muscular tuniea comprising the wall Of the stomach, together with an atrophy of dl gastric mucosa. Gastric biopsy (or gastric specimens of infected snakes revea bers of protozoan organisms attached to the brush border of the epit tric lumen and gastric pits (8.70). itis is accompanied by an over- ‘myriad num- clial cells lining the gas Usually, production of protective mucus by the goblet cells. The inflammation may be suppurative, in which heterophils are easily identified, or now: suppurative, in which the predominant leuco- cytes are mononuclear (8.71). The actiologic agent may or may not be immediately apparent Intussusception (the telescoping of one seg: ‘ment of intestine into another, or into th ach) occurs relatively frequently in some reptiles, particularly in iguanas and Old World 122 chameleons (8.72 and 8.73). The reasons for thi igh incidence are unknown, but endopa and dietary problems, especially hypocalcaemia, are suspected as predisposing factors, Benign and malignant neoplasia of the stom- ach and small intestine are relatively common in captive reptiles, particularly snakes and lizards. This may b onsiderably consequence of living, longer while in captivity than under natural (wild) conditions. Adenomata, carcinomata, leiomyom and leiomyosarcoma have been recorded in many snakes and in fewer lizards Prencoplastic leukoplakia and inva ‘mous cell carcinoma have lonians. These those observed in mammals Adult green iguanas (which are folivorous herbivores) have a simple stomach and a shor small intestine that transports the partially processed leafy ingesta into the sacculated and much expand oliferative lesions are similar to -olon. Villous projections (8.67), covered with pseudostratified, non-ciliated columnar epithelium overlying a thin lamina pro- ptia and a core of smooth muscle and blood ves- sels, extend into the colonic lumen and create a arger surface area for the processing of cellulose and absorption of nutrients. 8.70 Gastric cryptosporidiosis in an Australian tiger snake Notechis scutatus). A myriad number of round organisms arrowed) are attached to the brush border of the mucosal cells ining the gastric lumen and gastric pits. H & E250for this rasitism eaemia, lizards. iderably al wild) fe squa- lin che- milar to a shore artially ted and (8.67), ciliated a the a ells 8.71 Non-suppurative enteritis ina desert tortoise, Most of the leucocytes are iymphoplasmacytic 8.72, 8.73 Duodenal jejunal intussusception ina Fisher's chameleon 8.72) and a A segment telescoped into the jejunum causing the two serosa layers tole adjacent to each other. Hae xe Digestive SystemComparative Veterinary Histology with Clinical Correlates Liver The liver is the largest gland in the body, Blo drains to it from the intestines in the hepatic por tal vein, and the products of digestion are m ized, harmful material detoxified, senesce erythrocytes removed from the circullation and bile secreted. The liver is surrounded by mesothelium. The connective tissue capsule extends into the gland and divides it into lobes and lobules, The structure of the classic lobule is most clearly visualized in the pig. because of its plentiful array of connective tis: sue dividing the liver into discrete hexagonal lob- ales with a portal area at the corners of each hexagon. This is not the case in the other domesti animals, except under pathological conditions such, three or more lobules and each contains one or more branches of a hepatic artery, hepatic po ‘cin, a lymphatic vessel and a parenchyma consists of p of endodermal origin, the hepat verging on the central vein. The sinusoids are lin with fenestrated Sinusoid Liver cord cells hepatocytes 8.74 Hepatic lobule. The classi lobule Is the hexagon, clearly seen in the figure by the outline (green) of connective tissue), Portal areas (P) occur between the lobules, A porta lobule is defined as the central ‘unetional ans of the lobule (the black dotted triangle). 124 bile duct. The iyhedral epithelial cells eytes, arranged in rows separated by sinusoids con- ndothelial cells and macrophages, part of the mononuclear phagocyte system. Blood flows through the sinusoids to the central vein. This in turn leaves the liver lobale to travel separate branches of the hepatic vein, Bile is secreted by each hepatocyte into the bil canalicu| branes of the hepatocytes, channels that are lined with the plasma lows from there to a small bile duet in the portal area. Where the bile duct is the central functional axis of the lobule instead of the central vein, the term ‘portal lobule’ is used. The liver aci rus i the smallest functional unit of the liver. It con -d by a terminal branch o the portal vein and the hepatic by two central veins and termi bile duct. It has functional and pathological sigr icance. Bile duets are lined with cuboidal in the portal areas; the lar Jobul lined with columnar epith Reptiles, amphibians and fish The livers of fish, amphibia ns and reptiles are super ficially similar to those of ma are some differences. Many of the lower vertebrates have abundant melanin pigment scattered through a Bile Zone 1 Zone2 zone canaliculi Liver cord Central Branch of the hepatic Branch of the hepatic artery 8.75 Liver acinus (functio is served by t nal unit). The liver parenchyma ferminal branch of the portal vein and icartery. The acinus is divided into three h indicate the relative position of the cells in realtion to the oxygen gradient. Hepatocytes in Zone 1 are closest to the fresh, oxygen-rich hepatic arterial blood, and those in Zone 3 are further away. Equally, Cells in Zone 1 are first inline for toxins, ete, carried in the portal blood, with Zone 3 the cells the least affected 877 inrem. Blood vein, This eparately as nto the bile the plasma n adjoining liver: It branch of dis drained ches of the gical signif lepithelium ar duets are 5.80) wever, there vertebrates ed through: 75: 63 liver card parenchyma al vein and three {the ces s inZone | arterial LEqualy carried in gst affected Digestive system ‘out che hepatocellular parenchyma Usually, that are a see Chapter 3 pigment is contained in melanophages regaced together in packet-like groups of cells beariny ine dark-brown granules. In som: species the liver is ed in narrow cords radi ating outward from a thin-walled centeal vein, It has one or more portal triads consisting of an arte- olar branch of the hepatic artery and one or more small bile ducts {as in mammals). In other species the central veins are scattered randomly through: cout the liver and more than one portal triad or ti ads with multiple arterioles and bile canaliculi or ducts are p An admixture of hepatocellular and pancreatic tissues, thus forming a hepatopancreas, is 8.76 Liver (pig). The heaxagonal iver lobule is delineated by the green strands of the interlobular esent connective tissue. (1) Central vein, Masson's trichrome in many fish and in some amphibians and reptiles. _»125 - ane 8.77 Liver (00. (1) Hepatocytes arranged in cords. £8.78 Portal canal (trad) (sheep). (1) Branch of the bile (2) Sinusoids lined by endothelial cells and macrophages duct. (2) Hepatic artery. (3) lymphatic vessel.) Hepatic ‘pen into (3) the central vein, Safranivvhaematonylin portal vein, (5) Liver cords. (6) Sinusoids. H & E. «125, a7 2.20 £8.80 Liver (dog). The hepatocytes lie in anastomesing cords separated by sinusoids lined by macraphages ds of hepatocytes. Carbor-injected with H & E (arrowed), Safraninvhaematoxylin after carbon injection, erstain. 125. 250,Comparative Veterinary Histology Clinical correlates With its pivotal role in processing material c from the intestine via the portal system, th is exposed to toxic factors and potentially ful micro-organisms passing from the gut. Metabolic or nutritional disease (8.81), infectious sia (8.82), both local also affect the liver. ned hepacitis in excess fat storage in disease (see 8.84) and net and metastatic, can Inflammation of the liver is te (8.83). Hepatic lipidosis, 18.82 Cholangiocarcinoma in an Asiatic leopard (Panthera pardus). The cubokdal to columnar neoplastic cells form frondsike ducts and tubules with ‘hin walls composed of two cel layers separated by a scant fibrovascular connective tise soma, H&E 125, I Correlates | | the live, is seen as a clinical problem in obese ani- mals under physiological stress: often pregnant pony mares, dairy cattle after parturition and ewes arrying twins in late pregnancy (pregnancy tox: aemia). Mobilization of large amounts of triglye erides causes fatty acid to be presented tothe liver n excess ofits capacity to handle them. This prob- fatal and cases of sud may be quite rap den death due to liver rupture are not uncommon. The liver has a great capacity for regeneration of hepatocyte mass, but fibrosis is also a charac: 18.81 Hepatic lipidosis (horse) This micrograph of horse liver shows a ‘central vein surrounded by radiating cords of hepatocytes ‘that contain large, smoothly ound vacuoles whieh occupy most Of the cell and displace the rhucleus to the periphery. These are fat vacuoles, The horse was hyperlipidaemic with hepatic ipidoss. H & E. 250. 8.83 Giant cell hepatitis ina Syear-old neutered female Siamese cat, Note that many of the hepatocytes are much larger than normal and frequently have ‘multiple nuciel. A mixed mononuclear and ppolymorphonuclearleucocytic response is present. H&E, Haemosiderin is present. x125.regnant nd ewes the liver is prob- of sud- peration charac- rse)-This shows 2 4 tes hy Upy most These atic esa patocytes teristic reaction of the liver to chronic injury. Any hepatic injury severe enough to result in hepatic necrosis results in some fibrogenesis, but pro: ive fibrosis can develop when the insult per sists or when the initial damage is severe and provokes an extensive reaction. A canine liver with micronodular cirrhosis (8.84) shows pro- aressive fibrosis in which the normal architecture of the liver is lost and cels are divided into small ‘proups surrounded by fibrous tissue. 8.84 Micronodular ctrhosis in ‘8 T-year-old Cocker Spaniel. The Yellowestained hepatocytes vary in Size and shape and the developing fibrous tisue i stained red, With this technique, red blood cells stain yellow. Sirius Red, x250. 28s ies Zi et 8.85 Viral hepatitis in a Colombian boa constrictor (Boa C.constricton, Most of the hepatocytes contain ‘eosinophilic, intracytoplasmic val inclusion bodies (arrowed), most of which are surrounded by narrow clear haloes H & E.>250, Digestive System Amphibians and reptiles As with domestic animals, numerous chemical and viral agents induce severe liver disease (8.85 The hepatic parenchyma is sensitive to changes fn calcium and other minerals in the blood and, under conditions of hypervitaminosis Ds, may undergo severe mineralization and even ossifica tion (8.86), 884 286 8.86 Nutrition related, massive hepatic mineralization secondary to hypervitaminosis D, in an African leopard tortoise (Geochelone pardalis). Most of the hepatocellular parenchyma has undergone gross alteration and s replaced by bone. As a consequence, few normal hepatocytes remain. Several osteocytes surrounded fy concentric lameliae of compact bone fare present. H&E. x125,Gall bladder The gall bladder (absent in the horse, dolphins, chi nos and hippos) is a reservoir for bile and is attached to the visceral surface or between the lobe: of th ne is folded in the cid state, and the epithelial lining consists of tal columnar cells with a striated border {8.87}. Goblet cells and mucus- and serous-secreting glands may be present in ruminants. The muscularis externa is a circular layer of smooth muscle and the other tinuous with the p The gall bladder of lizard embedded in or surrounded by the liver, as itis in mammals and birds (8.88). The g: ‘itoneum, a Me 128 Comparative Veterinary Histology with Clinical Correlates snakes is located a a variable distance from the liver and is contiguous wit creas. A ong bile bile duct(s) to the tual release into Cli ical correlates The gall bladder can be a site of inflammation (cholangitis), holeliths), neoplasia and foreign bodies such as parasites (Fasciola hepatica). ST 8.87 Gal bladder (cow), (1) Tal columnar epithelium lining the lumen. @2) Mucosal folds Muscular. (4) Serosa. H & E 8.88 The liver and pancreas of some fish, amphibians and rept fused or admixed with one and form a hepatopancreas. lustrated is a section of such a mixed organ in an axolotl, a neotenic form of the aquatic salamander (Ambystoma maculetum), The ‘he bottom Fight; an islet isin the upper Ha E9250, (arcoplasi Fasciol: (ntl ithe ae uch a mixed ten fo The the bottom ere Pancreas A fine connective tissue capsule extends into the gland and divides it into lobules. The parenchyma is compos idocrine tissue; both of exocrine and are derived from the endoderm of the foregut, The wand xocrine portion of the pancreas is a comp ubuloacinar gland thar secretes enzymes into the Pancreas (dog). (1) Serous acini 3) Interiobular duct. (@) Pancreat let, the endocrine pancreas, H&E. 8.90 Pancreas (dog). (1) Serous acinus with a centroacinar cel rowed), (2) Interlobular connective ue, @) Interlobular duct. H & E 8.91 Pancreas (dog) (1) Nucleus ies in the basal basophilic cytoplasm o the serous cel, 2) Eosinophil granules (the secretion le in the luminal cytoplasm. H & E.»250. Digestive System lumnar with 1. Where the duodenum. The acinar cells are tall a basal nucleus in basophilic c is eosinophilic. Projections of duct cells are com monly seen in the acinus; these are the centroacinai cells chat ate typical of the pancreas. Smaller duct are lined with cuboidal epithelium and larger ducts with columnar epithelium (8.89-8.91). EE 289 q 3 3 : 1 4 i 391 ° 129Comparative Veterinary Histology with Clinical Correlates The endocrine pancreas is responsible for the and B, or beta, cells secreting insulin (a peptide hor control of blood sugar concentrations, and isolated mone released into the blood in response to a groups of pale staining islet cells (pancreatic cells or in concentration of blood glucose or amino acids the islets of Langerhans) are found scattered among Rate D, or delta, cells secrete somatostatin and the secretory units (8.92). These have two main cell cells secrete pancreatic polypeptide, These cell ypes: A, or alpha, cells secreting glucagon ( ongto the APUD cell group {see enteroendocrine 'ypeptide hormone secreted in response to hypo- cells of the stomach) tlyeaemia or to stimulation by growth hormone 32 % ake a 8.92 Pancreatic eet. Pancreas (ca) 5 (1) Pale staining endocrine eels form ords associated with cpilaies, | A ay Clinical correlates 8.93). Polycystic deformities, diabetes mellitus, pancreatic amyloidosis (8.94), acute and Essentially all of the various pancreatic disorders chronic pancreatitis, intraductal calculosis, and | that occur in humans also occur in domestic mam~ both benign and malignant ne oplasms are rec mals and in the so-called ‘lower’ vertebrates ognized in diverse species 393 Sean | ast reatc islet amyloidosis in a neutered female 2). Essentially, this cat's islets are 2 I2year-old dog. Part 8.94 9 nodular mass wit iy sold pattem af poorly ocelot (Fells pat Gifferentiated or cuboidal-to-low columnar cellsis_hyalinized and replaced w shown. H&E. »625, amyloid. H&E. 125, 8.93 Pancreatic car orphous, eosinophilic M 1ide hor nino acids). atin and F These cells endocrine a0. lars 25. nophilic Digestive system Reptilian, amphibian and fish pancreas Morphologically, the pancreas of teleost fish sue amphibians and mammals (8.9! o be conventional s similar to that found in arranged and buted, ‘giant’ islets of bur two major differences are Langerhans are characteristic in the pan observed in some species. Many fish, and soi mphibians and reptiles, possess a pancreas th sand boas); rather thar ind admixture of cells, being scattered in a more or less random manner spleen or liver (8.96). This combined throughout the pancreatic exocrine tis some snakes, particularly members of the family Boidae (py these lized in ;ermed a ‘splenopancreas’ or ‘hepato- huge islets of endocrine cells tend te respectively, and the cells and tissues of specific areas of pancreatic tisst arge aggregate anda portion of the on the left The islet displays 28 of some snakes Ww rython, is a4 Characterized by possessing formed into oiant throughout the parenchyma from a regal (bal) python (Python regius). He, 125,Comparative Veterinary Histology with Clinical Correlates Clinical correlates Rep’ jan pancreas Certain species of reptiles appear to have a her than expected incidence of some tumours. In captivity, some lizards, especially savanna monitors (Varanus exai seem to show a high incidence of adult-onset diabetes mellitus and exocrine deficiency, N evidence suggests that diabetes mellitus or exocrine deficiency are as prevalent in wild savanna monitors. Therefore, these disorders seem to be artefacts of captive husbandry rly by overfeeding and lack exercise) resulting in spontaneous (caused part of adequate acute and chronic pane quent autodigestion of the parenchyma, The migration of helminth larvae can als titis with secondary fibrosis and induce pan loss of secretory function both of exocrine and of endocrine components Avian digestive system The horny beak replaces functionally the lips and ;eeth of mammals. Oral cavity and oesophagus The oral cavity is lined with stratified squamous The tongue is also lined with this type of epithelium, epithelium with some keratinized areas, The main mass of the tongue consists of striated muscle and a small bar of cartilage 0 There are no teeth. The glands in the lam ina propria of the oral cavity, tongue and pharynx are simple-branch and mucus secreting. oesophagus is lined with stratified squamous non-keratinized epithelium, wich simple mucou glands in the connective tissue lamina propria 8.98). The muscularis externa consists of a thick innet layer of circular and a thin outer layer of lon. gitudinal smooth muscle. Lymphoid tissue accu. caudal ‘oesophageal tonsil. The crop is an aglandular eau: mulates in the esophagus as the dal diverticulum situated two-thirds of the way down the oesophagus (8.99). In the pigeon two la eral glomerular sacs secrete crop milk. 132 18 esophagus (bie). (1) Stratified squamous non. keratinizing epithelium. (2) Simple tubular mucosal glands. @) Lamina propria H & E. 625. 8.99 Crop (bir). (1) Stratified squamous non keratinized epithelium. 2) Lamina propria. (3) Musc @) Muscularis externa, H & E. 12.5Stomach ‘The stomach consists of the glandular proventricu lus and a muscular ventriculus. The gastric epithe- lium of the proventriculus is simple columnar and. mucus secreting, A thin lamina propria separates. it from the lobules of the submucosal glands. These glands form an almos with adjacent lobules separated by fine strands of connective tissue. Each gland lobule contains a cen. teal cavity with straight secretory tubules radiating, continuous mass of tissue, to the interlobular connective tissue. An excretory 3.99 | 8.101 Proventriculus bir). (1) Simple columnar mucus- secreting epithelium, (2) Lamina propria, (3) Submucosal lands. (4) Muscularis externa, Masson’ trkhyome. »25, Digestive system duct drains onto the gastric mucosal surface. The lands contain only one type of cell, which secretes acid and pepsinogen, thus combining the functions of both the chief and The muscularis externa is arranged as inner circu lar and outer longitudinal layers of smooth muscle (8.100-8.102), The ventriculus is the aglandular stomach or giz: zard. The luminal st product of the mucosal surface to form a hard cuticle of koilin, The epithe ce is lined with secretory ands, which solidifies ar the lium is low columnar and continues within the | 8.100 Proventriculus bird). (1) Simple columnar mucus secreting epithelium. (@) Lamina propria. @) Submucosal glands. (@) Musculris externa, H & E,x62:5 18.102 Proventriculus bird). The submucosal gland lobules are separated by thin strands of connective tissue ‘Maszon's trichrome. 3250,Comparative Veterinary Histology with Clinical Correlates 8.103 * 8.103 Ventrculus, gizzard (bir). (1) Corified lining, 2) Epithelium lining the gizzard, (3) Mucosal glands (4) Lamina propria, W& E.»2 simple straight tubular mucosal glands inthe lamin propria. A submucosa is present, and the muscularis extema isa thick layer of smooth muscle (8.103 ani 8.104). T Intestine The small intestine is similar to that of mammals at is more uniform throughout its length, Diffuse lymphatic tissue is present in the lamina propria and 8,104 Ventriculus, glzard (bird). (1) Comitied lining. Wacosal glands. H & E125, submucosa, and a third layer of circular smooth (2) Lining epithelium muscle may be present in the muscularis externa (8.105-8.10: The caeca are two blind sacs at the junction of the nar with mucous cells. Lymphatic tissue is particu ‘caecal tonsil in the in domestic birds. The epithelium is simple colum- row proximal part of the caecum (8.108 and 8.109) small and large intestine and are of considerable size larly abundant, forming, 8.105 18.105 Duodenum (bird). (1) Simple fruezal gland (3) Connwctiee Hive core of the villus. (4) Muscularis 8.106 |Lining e128 nthe nat ind 8.109) Digestive system {duodenum has a striated bord bsorption and goblet cel strchrome. «125. Piece ectretum reconeComparative Veterinary Histology with Clinical Correlates the same histological the connective tissne capsule extends ance as the caeca (8.110). The cloaca is lined and divides it into lobes and lobules, with tall columnar epithelium with a variable num- cytes are arranged in row per of mucus-secreting cells. it from che muscularis mucosae a The avian exocrine pancreas is similar to the mam externa See Chapter 15 for the cloacal _mallian, but has less interlobular connective nasa, (8.112). The endocrine pancreatic islets are of three ight (beta) islets, dark (alpha) islets and mixed a0 8.110 Rectum (bird). The lining epithelium is simple imnar with goblet cells extending into the simple red). H&E tubular glands. ‘tubular mucosal glanas 8.112 Pancreas (bird). (1) Serous 2) Pancreatic islet, the endoctine land. H&E. x125, 136 . often two cells thick,| 9. URINARY SYSTEM ls thick ee s ureters, bladder and urethra, _etin, which stimulates erythe ane mixed Kidneys connective tissue capsule extending into. the The kidneys are highly vascularized organs that fl: the medulla (9:1-9/4). Smioerh macle may be pee water and electrolytes via the ure dder medial border of the kidney and contains the renal as.urine (see Appendix artery, the renal vein and the ureter, j 92 9.1 Kidney horse. (1) Capsule.) Outer area ofthe cortex. 6) Renal corpuscle. 8) Uriniferous tubules H&E, x6255, oe ‘medulla (dog) (1) Blood vessel 125 3} Ascending thin limb, (4) Descending thin limb H&E, x25, \ 9.3 Kidney cortex (dog) (1) Renal (2)Uriniferous tubules. (3) Medullary ray. H & E 137Comparative Veterinary Histology with Clinical Correlates Nephron The nephron is the functional unit of the kidney. The major subdivisions are the renal corpuscle and the uriniferous tubule, Renal corpuscle The blind end of the proximal tubule is indented with a network of capillaries and supporting cells to form a filtering system: the renal corpuscle. Each renal corpuscle consists of a glomerulus and a glomerular (Bowman's) capsule. The outer layer of the glomerular capsule is the capsular (parietal wall, which is separated from the glomerula (vis ceral) layer by the capsular (urinary) space (9.5 ‘The capillaries are lined with a fenestrated endothe- lium resting on a basal lamina, The visceral epithe: lial cells, or podocytes, closely invest the capillary endothelium of the glomerulus and develop pri mary processes wrapped around each capillary. These processes develop secondary foot processes called pedicles The foor processes of adjacent podocytes in of small gaps called slit pores (9.6). The podocyte basal lamina is fused with the endothelial basal lamina and blood passing through the capillary is filtered through this coms lamina into the capsular space (9.6 and 9.7). Mesangial perivas- cular cells are present between the endothelium and the basal lamina

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