HISTOLOGY HISTOLOGY ОДЕСЬКИЙ НАЦІОНАЛЬНИЙ МЕДИЧНИЙ УНІВЕРСИТЕТ THE ODESSA NATIONAL MEDICAL UNIVERSITY Áiáëiîòåêà ñòóäåíòà-ìåäèêà Medical Student’s Library Серія заснована в 1999 р. на честь 100-річчя Одеського державного медичного університету (1900–2000 рр.) The series is initiated in 1999 to mark the Centenary of the Odessa State Medical University (1900–2000) 1 L. V. Arnautova O. A. Ulyantseva HISTÎLÎGY A course of lectures A manual Odessa The Odessa National Medical University 2011 UDC 616-018: 378.16 BBC 28.8я73 Series “Medical Student’s Library” Initiated in 1999 Authors: L. V. Arnautova, O. A. Ulyantseva Reviewers: Professor V. I. Shepitko, MD, the head of the Department of Histology, Cytology and Embryology of the Ukrainian Medical Stomatologic Academy Professor O. Yu. Shapovalova, MD, the head of the Department of Histology, Cytology and Embryology of the Crimean State Medical University named after S. I. Georgiyevsky It is published according to the decision of the Central Coordinational Methodical Committee of the Odessa National Medical University Proceedings N1 from 22.09.2010 Навчальний посібник містить лекції з гістології, цитології та ембріології у відповідності до програми. Викладено матеріали теоретичного курсу по всіх темах загальної та спеціальної гістології та ембріології. Посібник призначений для підготовки студентів до практичних занять та ліцензійного екзамену “Крок-1”. Arnautova L. V. Histology. A course of lectures : a manual / L. V. Arnautova, O. A. Ulyantseva. — Оdessa : The Оdessa National Medical University, 2010. — 336 p. — (Series “Medical Student’s Library”). ISBN 978-966-443-034-7 The manual contains the lecture course on histology, cytology and embryol- ogy in correspondence with the program. The theoretical course on all the top- ics of general and special histology is given. The manual can be used by stu- dents during their preparation to the practical trainings and to the license test examination “Krok-1”. The manual is recommended for the students of medical and dental facul- ties of higher medical universities. UDC 616-018:378.16 BBC 28.8я73 © L. V. Arnautova, O. A. Ulyantseva, 2011 ISBN 978-966-7733-47-6 (series)©The Odessa National ISBN 978-966-443-034-7 Medical University, 2011 4 PREFACE A necessity in the lectures course publication for English-speaking stu- dents has appeared because there is no textbook corresponding to the regu- lations of education issued by the Ministry of Public Health of Ukraine. As a result of conversion to the credit-modular system of education the hour- time of the lectures and practical trainings was changed, that resulted in redistribution of the schedule of the lectures course. The current textbooks on histology published abroad do not meet the requirements to our syllabus. Thus, for example biochemical processes of the cell are overviewed out in detail excessively while the morphology of vital organs (the hypothalamus, the spinal cord and so on) is not described sufficiently. Moreover, there are differences in classifications of the organs. The histology lectures are designed to prepare the student for practical trainings. They are dedicated to the overview of basic and secondary char- acteristics of tissue structure and training student’s skills of histological sample analysis. Some topics of the lectures course contain too much information, so that the students really have no time to scrap-note the data along with the lecturer; this fact and the lack of a textbook makes the preparation to prac- tical classes even more complicated for them. Therefore, we present the lectures course in English that answers the requirements of the Ministry of Public Health of Ukraine and can be used for education of students of medical and dental faculties. 5 Lecture 1 INTRODUCTION TO THE COURSE OF HISTOLOGY, CYTOLOGY AND EMBRYOLOGY. CYTOLOGY The Department of Histology and Embryology of the Novorossiysk Uni- versity was organized in May 1900 and it was one of the first fundamental departments of the Medical university. The founder of the histologists a Kiev school professor V. V. Podvysot- sky was directed to Odessa for organization of the medical faculty of histol- ogists and microbiologists in 1989. In May 1900 he was nominated as a chief of the Histology Department. Scientific activity of professor V. V. Podvysotsky was directed to cancer research and development of some questions of immunology. Professor A. F. Mankovsky had headed the Department of Histology, Cytology and Embryology from 1905 to 1917. The range of his scientific interests covered the questions on embryology and endocrinology. From 1917 to 1920 the Histology Department of the Odessa Medical University had been consecutively managed by professor V. V. Voronin, academician D. K. Tretiyakov and assistant professor V. E. Yanishevsky. Professor F. N. Zhmaylovich had headed the department from 1920 to 1923. The main direction of his scientific studies were the questions of cytology. Professor M. N. Zaevloshin had headed the Histology Department for a long time (1923–1944). He is an author of more than 40 scientific works on histology and patomorphology. Assistant professor B. I. Kardashevich had been at the head of the de- partment twice, from 1944 to 1946, and from 1950 to 1954. During that period the main scientific trend of the department were the questions to age morphology. 6 Professor S. D. Shakhov has managed the Histology Department from 1954 to 1958. He is an author of more than 60 scientific works. The range of his scientific interests was very wide. Professor N. D. Zaytsev had headed the Histology Department from 1958 to 1976. He is an author of 67 scientific works devoted to the ques- tions of embryogenesis and problem of neurohistology. Professor N. D. Zaytsev has changed the professor V. F. Pchelyakov who had managed the department from 1976 to1993. His scientific works were devoted to study of the cornea of the eye in amimals and humans. Professor V. K. Napkhanyuk was elected the head of the department in May 1993 and prepared more than 20 research assistances. A young and perspective leader professor V. A. Ulyanov has headed the department since 2006. CYTOLOGY The cell is a functional unit of all living tissues, having a capacity to perform all the essential life functions. Within the different tissues and or- gans of the body, the constituent cells exhibit a wide range of functional specialization. Despite this extraordinary range of morphological forms and cells conform to a basic structural model which is the subject of this lecture. The process by which cells assume specialized structure and function is known as differentiation. Even with a primitive light microscopy, it was evident that cells are divided into at least two components, the nucleus and the cytoplasm, and as microscopic techniques advanced it became increasingly obvious that both the cytoplasm and nucleus contain a number of subcellular elements called organelles. The resolving power of the light microscope is limited to about 0.2 mm (200 nm) and the study of the ultrastructure of the cell had to await the advent of electron microscopy. The capacity of electron microscopes in current use permits the resolution of structures as small as 1.0 nm, this however, falls far short of many cellular processes which occur at the mo- lecular level. Light and electron microscopy have been successfully com- bined with biochemical and immunological techniques to define the location 7 of many biological processes; these techniques are known as histochemical and immunohistochemical respectively. All cells are bounded by an external limiting membrane called the plasma membrane or plasmolemma, which serves as a dynamic interphase between the internal environment of the cell and various external environments. The nucleus is the largest organelle and its substance, often referred to as the nucleoplasm, is bounded by a membrane system called the nuclear enve- lope. The cytoplasm contains the variety of organelles, most of which are also bounded by membranes. An extensive system of membrane-bound tubules, saccules and flattened cisternae, known as the endoplasmic reticu- lum, is widely distributed throughout the cytoplasm. A more distended sys- tem of membrane-bound saccules, the Golgi apparatus, is typically located close to the nucleus. Scattered free in the cytoplasm are a number of rela- tively large elongated organelles called mitochondria, which have a smooth outer membrane and convoluted inner membrane system. The cytoplasmic organelles are suspended in a fluid medium called the cytosol in which much of the intermediary metabolism of the cell takes place. Within the cytosol, there is a network of minute tubules and filaments collectively known as the cytoskeleton, which provides structural support for the cell and its organelles as well as providing a mechanism for cellular and intracellular movement; elements of the cytoskeleton are only visible with very high magnification. The current concepts of membrane structure derive from the work of Singer and Nicholson in the early 1970 year. In this model, cell membranes consist basically of phospholipid molecules arranged as a bilayer. Phos- pholipid molecules consist of a polar, hydrophilic (water-loving) head and non-polar, hydrophobic (water-hating) tail. The polar heads are mainly de- rived from glycerol conjugated to a nitrogenous compound such as choline, ethanolamine or serine via a phosphate bridge. The phosphate group is negatively charged whereas the nitrogenous group is positively charged. The non-polar tail of the phospholipid molecule consists of two long-chain fatty acids each covalently linked to the glycerol component of the polar head. The weak intermolecular forces which hold the bilayer together allow individual phospholipid molecules to move relatively freely within each lay- er and sometimes to ‘flip” between layers. Cholesterol molecules regulate the fluidity and stabilize the phospholipid bilayer. Associated with the bilay- er are a variety of protein molecules, which make up almost half of the total 8 mass of the membrane.