t^T^^G VA'^? Columbia ^nttosittp ISitUxmt^ SItbrarg ^ AJyC / fru (i.uU^ ^ ^-CU..^ -7 , 1^/. - SIi[ps for Librarians to paste on Catalogue Cards. N.B. — Take out carefully, leaving about quarter of an inch, at the back. To do otherwise would, in some cases, release other leaves. MARTIN, H. NEWELL.— The Human Body. An Account of its Structure and Activities, and the conditions of its healthy working, by H. New- ell Martin, D. Sc, M. A., M.B., Professor of Biol- ogy in the Johns Hopkins University ; Fellow of University College, London ; late Fellow of Christ's College, Cambridge. New York : Henry Holt & Co., 1881. Large i2mo, pp. xvi. 606, Appendix 33. (American Science Series.) HUMAN BODY, THE.— An Account of its Structure and Activities, and the conditions of its healthy working, b}^ H. Newell Martin, D. Sc, M. A., M. B., Professor of Biology in the Johns Hop- kins University; Fellow of University College, London; late Fellow of Christ's College, Cam- bridge. New York: Henry Holt & Co., iSSi. Large lamo, pp. xvi. 606, Appendix t^2>- (Ameri- can Science Series.) ANATOMY.— The Human Body. An Account of its Structure and Activities, and the conditions of its healthy working, by H. Newell IMartin, D. Sc, M. A., M. B., Professor of Biology in the Johns Hopkins University; Fellow of University College, London ; late Fellow of Christ's College, Cambridge. New York: Henry Holt & Co., 1S81. Large i2mo, pp. xvi. 606, Appendix 33. (Ameri- can Science Series.) PHYSIOLOGY.— The Human Body. An Ac- count of its Structure and Activities, and the con- ditions of its healthy working, by H. Newell Mar- tin, D. Sc, M. A., M. B., Professor of Biology in the Johns Hopkins University ; Fellow of Univer- sity College, London ; iate Fellow of Christ's Col- lege, Cambridge. New York : Henry Holt & Co., 1881. Large i2mo, pp. xvi. 606, Appendix 33. (American Science Series.) Digitized by the Internet Archive in 2010 with funding from Open Knowledge Commons http://www.archive.org/details/humanbodyaccount1881mart THE AMERICAN SCIE^'CE SERIES FOR HIGH SCHOOLS AND COLLEGES. The principal objects of this series are to supply the lack — in some subjects very great — of authoritative books whose principles are, so I'ar as practicable, illustrated by familiar American facts, and also to supply the other lack that the advance of Science perennially creates, of text-books which at least do not contradict the latest generaliza- lions. Tlie volumes are large 12mo. The books arranged for are as follows. They systematically out- line the field of Science, as the term is usually employed with refer- ence to general education. Those marked * had been published up to February 1, 1881. I. Physics. By Arthur W. "Wright, Pro- fessor in Yale College. II. Chetnisfry. By Samuel W. Johkson and William G. Mixter, Professors in Yale College. III. Asffononit/.* By Simon Newcomb, Supt. American Nautical Almanac, and Edward S. Holdek, Professor in the United States Naval Ob- servatory. $2.50. ir. Geology. By Raphael Pumpeluy, late Professor in Harvard University. I'. Bofaiii/.* By C. E. Bessey, Professor in the Iowa Agricultural College and Lite Lecturer in the Uni- versity of California. !j;2.75. VI. Zoolo(/i/.* By A. S. Packard, Jr., Pro- fessor of Zoology and Geology in Brown University, Editor of the A merican Naturalist. $3. VII. The Ilunian liodij.* By H. Newell ]\L\rtix, Pro- fessor in the Johns Hopkins Uinversity. $2.75. Copies with- out the Appendix on Reproduc- tion will be sent when specially ordered VIII. Psyehologif. By ^Villiam James, Piofessor in Harvard University. IX. Political Economy. By Francis A. Walker, Pro- fessor in Yale College. A". Government. By Edwin L. Godkin, Editor of the Nation. HENRY HOLT & CO., Publishers, NEW YORK. AMERICAN SCIENCE SERIES THE HUMAN BODY AN ACCOUNT OF ITS STRUCTURE AND ACTIVITIES AND THE CONDITIONS OF ITS HEALTHY WORKING H. NEWELL MAPiTIA. D.,S< .,, :\l.A., M.B. Professor of Biology in- the Johns Hopkins University ; Fellow of University College, London; Late Fellow of Christ's College, Camhndge NEW YORK HEKEY HOLT AXD COMPAXY 1881 \ O /7 Copyright, 1881, BY Henry Holt & Co. A -50. J. Campbeli-, PRINTER, 15 VaudewatiT St., N. Y. PEEFACE. In the following pages I have endeavored to give an account of the structure and activities of the Human Body, which, while intelligible to the general reader, shall be accurate, and sufficiently minute in details to meet the requirements of students who are not making Human Anatomy and Physiology subjects of special advanced study. Wherever it seemed to me really profitable, hygienic topics have also been discussed, though at first glance they may seem less fully treated of than in many School or College Text-books of Physiology. Whoever will take the trouble, however, to examine critically wdiat passes for Hygiene in the majority of such cases, will I think find that, when correct, much of it is platitude or truism: since there is so much that is of importance and interest to be said it seems hardly worth while to occupy space with insisting on the commonplace or obvious. It is hard to write a book, not designed for specialists, without running the risk of being accused of dogmatism, and some readers will, no doubt, be inclined to think that, in several instances, I have treated as established facts matters which are still open to discussion. General readers and students are, however, only bewildered by the production of an array of observations and arguments on each side of every question, and, in the majority of cases, the chief responsi- bility under which the author of a text-book lies is to select what seem to him the best supported views, and then to state them simply and concisely: how wise the choice of a side has been in each case can only be determined by the discoveries of the future. Others will, I am inclined to think, raise the contrary iv PBEFACE. ol)jcction, thfit too many (lisi»ute(l matters luive been dis- cussed: tliis was deliberately done as the result of an experi- ence ill teacliing Physiology which now extends over more than ten years. It would have been comparatively easy to slip over things still uncertain and subjects as yet unin- vestigated, and to represent our knowledge of the workings of the animal body as neatly rounded off at all its contours and complete in all its details — iotus, tei^es, et rotundus. Ikit by so doing no adequate idea of the present state of ])hysiological science would have been conveyed; in many directions it is much farther travelled and more completely known than in others; and, as ever, exactly the most in- teresting points are those Avhich lie on the boundary Ijetween what we know and what we hope to know. In gross Anatomy there are now but few points calling for a suspension of judgment; with respect to Micro- scopic Anatomy there are more; but a treatise on Physiology which would pass by, unnientioned, all things not known but sought, would convey an utterly unfaithful and untrue idea. Physiology has not finished its course; it is not cut and dried, and ready to be laid aside for reference like a specimen in an Herbarium, but is comparable rather to a living, growing plant, with some stout and useful branches well raised into the light, others but jiart grown, and many still represented by unfolded buds. To the teacher, moreover, no jDupil is more discouraging than the one who thinks there is nothing to learn; and the boy who has "finished" Latin and " done" Geometry finds sometimes his counterpart in the lad who has " gone through" Physiology. For this unfortunate state of mind many Text-books are, I believe, much to blame: difficulties are too often ignored, or opening vistas of knowledge resolutely kept out of view: the forbidden regions may be, it is true, too rough for the young student to be guided through, or as yet jiathless for the pioneers of thought; but the opportunity to arouse the re- ceptive mental attitude apt to be produced by the recogni- tion of the fact that much more still remains to be learnt — to excite the exercise of the reasoning faculties upon dis- puted matters — and, in some of the better minds, to arouse PREFACE. V the longing to assist in adding to knowledge, is an inesti- mable advantage, not to be lightly thrown aside through the desire to make an elegantly symmetrical book. " While I trust, therefore, that this volume contains all the more imjDortant facts at present known about the working of our Bodies, I as earnestly hope that it makes plain that very much is yet to be discovered. A work of the scope of the present volume is, of course, not the proper medium for the publication of novel facts; but, while the "Human Body," accordingly, professes to be merely a compilation, the introduction of constant ref- erences to authorities would have been out of place. I trust, however, that it will be found throughout imbued with the influence of my beloved master, Michael Foster; and on various hygienic topics I have to acknowledge a special indebtedness to the excellent series entitled Health Primers. The majority of the anatomical illustrations are from Henle's Anatomie des Mensclien, and a few from Arendfs Schulatlas, the publishers of each furnishing electrotypes. A considerable number, mainly histological, are from Qiiaiji's Anatomy, and a few figures are after Bernstein, Carpenter, Frey, Haeckel, Helmholtz, Huxley, McKen- drick, and Wundt. About thirty, chiefly diagrammatic, were drawn specially for the work. Quantities are throughout expressed first on the metric system, their approximate equivalents in American weights and measures being added in brackets. H. Newell Martin. Baltimore, October, 1880. coi^rTEisrTs. CHAPTER I. THE GENERAL STRUCTURE AND COMPOSITION OF THE IHIMAN BODY. PAGE Definitions. Tissues and organs. Histology. Zoological posi lion of man. The vertebrate plan of structure. The mam- malia. Chemical composition of the Body 1 CHAPTER n. THE FUNDAMENTAL PHYSIOLOGICAL ACTIONS. The properties of the living Body. Physiological properties. Ceils. Cell growth. Cell division. Assimilation and repro- duction. Contractility. Irritability. Conductivity, Spon- taneity. Protoplasm. The fundamental physiological proper- ties 15 CHAPTER HI. THE DIFFERENTIATION OF THE TISSUES, AND THE PHYSI- OLOGICAL DIVISION OF EMPLOYMENTS. Development. The physiological division of labor. Classifica- tion of the tissues. Undifferentiated tissues. Supporting tissues. Nutritive tissues. Storage tissues. Irritable tissues. Co-ordinating and automatic tissues. 3Iotor tissues. Conduc- tive tissues. Protective tissues. Reproductive tissues. Or- gans. Physiological mechanisms. Anatomical systems. The Body as a working whole 26 Vlll CONTENTS. CHAPTER IV. TITK INTEKNAIi MEDIUM. PA(!K The external medium. Tlie internal medium. The blood. The lymph. Histology of l)lood. Blood crystals. Histology of lymph r 39 CHAPTER V. THE CLOTTING OF THE BLOOD. The coagulation of blood. Causes of coagulation. Whipped blood. The Ijuffy coat. Uses of coagulation. The fibrin factors. Artificial clot. The fibrin ferment. E.\citing causes of coagulation. Relation of blood-vessels to coagulation. Composition of the blood. Quantity of blood. Origin and fate of the blood corpuscles. Chemistry of lymph 50 CHAPTER VI. THE SKELETON. Exoskeleton and endoskeleton. The bony skeleton. Segmenta- tion of the skeleton. Peculiarities of the human bony skele- ton 62 CHAPTER VII. THE STRUCTURE AND COMPOSITION OF BONE. .JOINTS. Gross structure of the bones. Microscopic structure of bone. Chemical composition of bone. Hygiene of the bony skeleton. Articulations. Joints. Hygiene of the joints 86 CHAPTER VIII. CARTILAGE AND CONNECTIVE TISSUE. Temporary and permanent cartilages. Varieties of cartilage. The connective ti-ssues. Elastic cartilage and fibro-cartilage. CONTENTS. ix PAGE Homologies of the supportiiiLr tissues. Hygiene of tbe develop- ing skeleton. Adipose tissue 100 CHAPTER IX. TIEE STRCCTUKE OF THE JIOTOll ORGANS. Motion in animals and plants. Ama?boid cells. Ciliated cells. The muscles. Histology of striped muscle. Uustriped mus- cles. Cardiac muscular tissue. The chemistry of muscular . tissue. Beef -tea and Liebig's extract 11; CHAPTER X. THE PROPERTIES OP MTSCTTLAR TISSUE. Contractility. Irritability of muscle. A simple muscular con- traction. Tetanus. Causes influencing the degree of muscu- lar contraction. The measurement of muscular work. Mus- cular elasticity. Physiology of plain muscular tissue. Hy- giene of the muscles. Exercise 128 CHAPTER XI. MOTION AND LOCOMOTION. The special physiology of muscles. Levers in the Body. The erect posture. "Walking. Running. Leaping 143 CHAPTER XII. THE ANATOMT OF THE NERVOPS STSTEil. Nerve-trunks. Plexuses. Xerve-centres. The cerebro-spinal centre and its membranes. The spinal cord. The spinal nerves. The brain. The cranial nerves. Ganglia and com- munications of the cranial nerves. The sympathetic system. The sporadic ganglia. The histology of nerve-fibres. The histology of nerve-cells. The structure of the spinal cord 1.54 CONTENTS. CHAPTER XIII. THE GENERAL PHYSIOLOGY OF TIIE NERVOUS SYSTEM PAGK The properties of nerve tissues. The functions of nerve-centres and nerve-trunks. Excitant and inhibitory nerves. The clas sification of nerve libres. Intercentral nerve-fibres. The stimuli of nerves. General and special nerve stimuli. Specific nerve energies. All nerve-fibres are fundamentally alike. The nature of a nervous impulse. The rate of transniis.sion of a nervous impulse. Functions of the spinal nerve-roots. Tlie intercommunication of nerve-centres ISO CHAPTER XIV. THE ANATOMY OP TIIE HEART AND BLOOD VESSELS. General statement. Position of the heart. The membranes of the heart. The cavities of the heart. The heart as viewed from the outside. The interior of the heart. The valves of the heart. The arterial system. The capillaries. The veins. The pulmonary circulation. The course of tlie blood. The portal circulation. Arterial and venous blood. The structure of arteries, capillaries, and veins 201 CHAPTER XV. THE WORKING OF THE HEART AND BLOOD-VESSELS. The beat of the heart. The cardiac impulse. Events in a cardiac period. Use of the papillary muscles. Sounds of the heart. Function of the auricles. The work done by the heart. The blood flow outside the heart. The circulation as seen with the microscope. Internal friction in the vessels. The conver- sion of an intermittent into a continuous flow 219 CHAPTER XVI. ARTERIAL PRESSDRE AND THE PULSE. Weber's schema. Arterial pressure. Modifications of arterial pressure, and how they may be produced. The pulse. The CONTENTS. xi PAGE rate of the blood-flow. Secondary causes of the circulation Proofs of the circulation of the blood 233 CHAPTER XVII. THE REGULATION OF THE HEART AND BLOOD \T:SSELS BY THE NERVOUS SYSTEM. The need of co-ordination in the vascular sj'stem. The intrinsic nerves of the heart. Nerves showing the heart's beat. The cardio-inhibitory mechanism. The accelerator nerves of the heart. The nerves of the blood-vessels. The vasomotor centre. Taking cold. Vaso-dilator nerves 247 CHAPTER XVIII. THJ<; SECRETORY TISSI^S AND ORGANS. Definition. Organs of secretion. Glands. Physical processes in secretion. Chemical processes in secretion. The mode of activity of secretory cells. Influence of the nervous system upon secretion. Summary 259 CHAPTER XIX. THE INCOME AND EXPENDITURE OP THE BODY. The material daily losses of the Body. The daily losses of the Body in energy. The conservation of energy. Potential and kinetic energy. The energy of chemical afiinity. The relation between matter lost by the Body daily and the energy spent by it. The conditions of oxidation in the living Body. The fuel of the Body. Oxidation by successive steps. The utilization of energy in the living Body. Summary 277 CHAPTER XX. Foods as tissue-formers. The food of plants. Non-oxidizable foods. Definition of foods. Conditions which a food must XI 1 CONTENTS. PAGE fuUill Alimentary principles. 'J'lie couijjusition of the more important foods. Cooking. Alcohol. The advantage of a mixed diet 2*j;i CHAPTER XXI. Tnii ALIMKNTAKY ( AN.\fi AND ITS AITENDAOEK. General arrangement. Tlic teeth. The tongue. The salivary glands. The fauces. The pharynx. The gullet. The stomach. The histology of the gastric glands. The small intestine The large intestine. The liver. The pancreas. . . . 308 CHAPTER XXII. THE LYMPHATIC SYSTEM AND THE Ul'CTLESS GLjVNDS. Distribution and structure of lymph vessels. The thoracic duct. The serous cavities. The lymphatic glands. The movement of the lymph. The spleen. The thymus. The thyroid body. The supra-renal capsules 32!) CHAPTER XXIII. DIGESTION. The object of digestion. U.-?es of saliva. Deglutition. The gastric juice. Gastric digestion. The chyle. The pancreatic secretion. The bile. The intestinal secretior-s. Intestinal di geslion. Ab.sorption from the intestines. The digestion of an ordinary meal. Dyspepsia 334. CHAPTER XXIV. THE EESPrRATORY MECHANISM. Definitions. Respiratory organs. The air-passages and lungs. The pleura. The respiratory movements. The structure of the thorax. The mechanism of inspiration. Expiration. Forced respiration. The respiratory sounds. Tlie capacit}- of the lungs. Hygiene of respiration. The aspiration of the thorax. Influence of respiratory movements upon the flow of blood and lymph 353 CONTENIS. xiii CHAPTER XXV. THE CHEMISTEY OF RESPIRATION. PAGE Nature of the problems. Changes prockiced m air by being once breathed. Ventilation. Changes undergone by blood in the kings. The blood gases. Causes of color of arterial and venous blood. Laws governing the absorption of gases by liquids. The absorption of oxygen by the blood. Conse quences of the way in which oxygen is held in the blood. The general oxygen interchanges of the blood. The carbon dioxide of the blood. Internal respiration 3T2 CHAPTER XXVT. THE NERVOUS FACTORS OF THE RESPIRATORY MECHANISM. ASPHYXIA. The respiratory centre. Is the respiratory centre reflex? The stimulus of the respiratory centre. The cause of the respira- tory rhythm. The influence of the pneumogastric nerves upon the respiratory centre. The expiratory centre. Asphyxia. Carbon - monoxide htnemoglobin. Modified respiratory move- ments 390 CHAPTER XXVII. THE KIDNEYS AND THE SKIN. The general arrangement of the urinary organs. The structure of the kidneys. The renal secretion. The mechanism of the renal secretion. The function of the renal epithelium. The skin. Epidermis and cutis vera. Hairs. Nails. Glands of the skin. Relation of nerves to sweat secretion. Hygiene of the skin. Bathing 404 CHAPTER XXVIII. NUTRITION. The problems of animal nutrition. The seat of the oxidations of the Body. Tissue-building and energy-yielding foods. The source of the energy spent in muscular work. Luxus con- sumption. The antecedents of urea. Proteid starvation and XIV COI^TEI^TS. PAOE over-feedinc;. The storage tissues. Glycogen. Diabetes. The history of fats. Dietetics 428 CHAPTER XXIX. THE PnODUCTION AND REGULATION OF THE HEAT OF THE BODY. Cold- and warm-blooded animals. The temperature of the Body. The sources of animal heat. Energy lost l)y the Body in twenty-four hours. The superiority of the Body as a machine for executing mechanical work. The maintenance of an average temperature. Local temperatures. Thermic nerves. Clothing 449 CHAPTER XXX. SENSATION AND SENSE-ORGANS, The subjective functions of the nervous system. Common sen- sation and organs of special sense. The peripheral reference of sensations. Differences between sensations. The essential structure of a sense organ. The cause of the modality of sensa- tions. The psycho-physical law. Perceptions. Sensory illu- sions 461 CHAPTER XXXI. THE EYE AS AN Ol'TICAL INSTUMENT. The essential structure of an eye. The appendages of the eye. The lachrymal apparatus. The muscles of the eyeball. The anatomy of the eyeball. The structure of the retma. The refracting media of the eye. The properties of light. Accom- modation. Short sight and long sight. Hygiene of sight. Optical defects of the eye 479 CHAPTER XXXII. THE EYE AS A SENSORY APPARATUS. The excitation of the visual apparatus. Idio-retinal light. The parts of the retina on which light directly acts. The vision purple. Intensity of visual sensations. Duration of luminous sensations. The localizing power of the retina. Color vision. CONTENTS. XV PAGE Colorblindness. Fatigue of the retina and after-images. Con tvasts. Heriug's theory of vision. Visual perceptions. Single vision with two eyes 50(! CHAPTER XXXIII. THE EAR AND HEARING. The external ear. The tympanum. The auditory ossicles. The internal ear. The organ of Corti. The loudness, pitch, and timbre of sounds. Sympathetic resonance. Functions of tlie tympanic membrane, of the auditory ossicles, of the cochlea, and of the vestibule. Auditory perceptions 535 CHAPTER XXXIV. TOUCH, THE TEMPER.VTURE SENSE, THE MUSCULAR SENSE, COrMON SENSATION, SMELL, AND TASTE. Nerve-endings in the skin. The pressure sense. The localizing power of the skin. The temperature sense. Modality of skin sensations. The muscular sense. Pain. Hunger and thirst. Smell. Taste 556 CHAPTER XXXV THE FUNCTIONS OF THE BRAIN AND SPINAI, CORD. The special physiology of nerve-centres. The spinal cord as a centre. Reflex actions. The least-resistance hypothesis. The educafiou of the spinal cord. The inhibition of reflexes. Psy- chical activities of the cord. The cord as a conductor of nervous impulses. Functions of the brain in general. Func tions of medulla oblongata. Functions of cerebellum, pons Varolii, and mid-brain. Sensations of equilibrium and func- tions of semicircular canals. Functions of the fore-bram Hygiene of the brain 578 CHAPTER XXX^a. VOICE .\ N D SPEECH. Anatomy of larynx. The vocal cords. Causes of the varying pitch of the voice. Ran^e of the human voice. The produc- tion of vowels. Consonants 595 xvi VoyTE^TS. APPENDIX. REPRODUCTION AND DEVELOPMENT. Ri^production in general. Sexual and asexual reproduction. Male reproductive organs. Female reproductive organs. Puberty, Ovulation. Menstruation. Hygiene of menstruation. Impregna- tion, Pregnancy. The fatal ai)peiidages. The intra-utcrine nutrition of the embryo. Parturition. Lactation. Feeding of infants. The stages of life. Death. THE HUMAN BODY. CHAPTER I. THE GENERAL STRUCTURE AND COMPOSITION OF THE HUMAN BODY. Definitions. The living human Body may be considered from either of two aspects. Its structure may be especial- ly examined, and the forms, connections and mode of growtli of its parts be studied, as also the resemblances or differences in such respects, which appear when it is com- pared with other animal bodies. Or the living Body may be more especially studied as an organism presenting defi- nite ]n-operties and performing certain actions; and then its parts will be investigated with a view to discovering what duty, if any, each fulfills. The former group of studies constitutes the science of Anatomy, and in so far as it deals with the human Body alone, of Human Anatomy; while the latter, the science concerned with the uses — or in tech- nical language the functions — of each part is known as Physiology. Closely connected with physiology is the science of Hygiene, which is concerned with the conditions which are favorable to the healthy action of the various parts of the Body ; while the activities and structure of the diseased body form the subject-matters of the sciences of Pathology and Pathological Anatomy. Tissues and Organs. Histology. Examined merely from the outside, our Bodies present a considerable com- plexity of structure. We easily recognize distinct parts as head, neck, trunk and limbs ; and in these again smaller '^ THE HUMAN BODY. constituent purts, us eyes, nose, ears, mouth; arm, fore- arm, hand; thigh, leg and foot. We can, with such an external examination, go even farther and recognize different materials as entering into the formation of the larger parts. Skin, hair, nails and teeth are obviously different substances ; simple examination by pressure l)roves that internally there are haixler and softer solid parts; vrliile the blood that flows from a cut finger shows that liquid constituents also exist in the Body. The con- ception of complexity, which may be thus arrived at from external observation of the living, is greatly extended by dissection of the dead Body, which makes manifest that it consists of a great number of diverse parts or organs, which in turn are built up of a limited number of materials, the same material often entering into the comjiosition of many different organs. These primary building materials are known as the tissues, and that branch of anatomy which deals with the characters of the tissues and their arrange- ment in various organs is known as Histology; or, since it is mainly carried on with the aid of the microscope, as Microscopic Anato7ny. If, with the poet, we compare the Body to a house, we may go on to liken the tissues to the bricks, stone, mortar, wood, iron, glass and so on used in building; and then walls and floors, stairs and windows, formed by the combination of these, would answer to ana- tomical organs. Zoological Position of Man. External examination of the human Body shows also that it presents certain re- semblances to the bodies of many other animals: head and neck, trunk and limbs, and various minor parts enter- ing into them, are not at all peculiar to it. Closer study and the investigation of internal structure demonstrates further that these resemblances are in many cases not su- perficial only, but that our Bodies may be regarded as built upon a plan common to them and the bodies of many other creatures: and it soon becomes further apparent that this resemblance is greater between the human Body and the bodies of ordinary four-footed beasts, than between it and the bodies of birds, reptiles or fishes. Hence, from THE PRIMATES. 3 a zoological point of view, man's Body marks liim ont as belonging to the group of Mammalia (see Zoology), wliicli includes all animals mwhicli the female suckles the young; and among mammals the anatomical resemblances are closer and the differences less between man and certain apes than between man and the other mammals; so that zoologists still, with Linnaeus, include man with the mon- keys and apes in one subdivision of the Mammalia, known as the Primates. That civilized man is mentally far superior to any other animal is no valid objection to such a classifi- cation, for zoological groups are defined by anatomical and not by physiological characters ; and mental traits, since we know that their manifestation depends upon the structural integrity of certain organs, are essentially phenomena of function and therefore not available for purposes of zo- ological arrangement. Man however walks erect v.'ith the head upward, while the great majority of Mammals go on all fours with the head forward and the back upward, and various apes adopt intermediate positions, so in considering corre- sponding parts in such cases confusion is apt to arise unless a precise meaning be given to such terms as " an- terior" and ''posterior." To avoid this difficulty anato- mists give these words definite arbitrary significations in all cases and these we shall use in future. The head end is always antei'ior whatever the natural position of the animal, and the opposite end posteriorj the belly side is spoken of as ventral, and the opposite side as dorsal; rigid and left of course present no difficulty. Moreover, that end of a limb nearer the trunk is spoken of as proximal Avith reference to the other or distal end. The words upper and loiver may be conveniently used for the relative position of parts in the natural standing position of the animal. The Vertebrate Plan of Structure. Neglecting such merely apparent differences as arise from the differences of normal posture above pointed out, we find that man's own zoological class, the Mammals, differs very widely in its broad structural plan from the groups including sea anem- 4 THE HUMAN BODY. ones, insects, or oysters, but agrees in many points with the groups of fishes, amphibians, reptiles, and birds. These four are therefore placed with man and all other Mammals in one great division of the animal kingdom known as the Vertehrata. The main anatomical character of all vertebrate animals is the presence in the trunk of the body of two cavities, a dorsal and a ventral, separated by a solid partition, and in the adults of nearly all vertebrate animals a hard axis, the vertebral column {hachhone or spine), develops in this partition and forms a central support for the rest of the body (Fig. 2, e e). The dorsal cavity is con- tinued through the neck, when there is one, into the head, and there widens out. The bony axis is also continued through the neck and extends into the head in a modified form. The ventral cavity, on the other hand, is confined to the trunk. It contains the main organs connected with the blood-flow and is thus often called the limmal cavity. Upon the ventral side of the head is the mouth o^jen- inff leading into a tube, the alimentary canal, f, which passes back through tlie neck and trunk and opens again on the outside at the posterior part of the latter. In its passage through the trunk region this canal lies in the' ventral cavity. The Mammalia. In many vertebrate animals the ven- tral cavity is not subdivided, but in the Mammalia it is; a membranous transverse partition, the midriff or diaphragm (Fig. 1, z), separating it into an anterior chest or thoracic cavity, and a posterior or ahdominal cavity. The alimen- tary canal and whatever else passes from one of these cavi- ties to the other must therefore perforate the diaphragm. In the chest, besides part of the alimentary canal, lie important organs, the heart, h, and lungs, In, the heart being on the ventral side of the alimentary canal. The abdominal cavity is mainly occupied by the alimentary canal and organs connected with it and concerned in the digestion of food, as the stomach, ma, the liver, le, the pancreas and the intestines. Among the other more prom- inent organs in it are the 'kid7ieys and the spleen. In the dorsal cavity lie soft white organs, the hrain and THE MAMMALIAN TYPE. spinal cord, the former occupying its anterior enlargement in the head. Brain and spinal cord together form the cerebi'o-spinal nervous centre, but in addition to this there are found in the ventral cavity a number of small nerve cen- FiG. 1.— The Body opened from the front to show the contents of its ventral cavity. tres united together by connecting cords, and. with their offshoots forming the sympathetic nervous system. The walls of the three main cavities are lined by smooth, THE HUMAN BODY. moist serous membranes. That lining the dorsal cavity is the arachnoid ; that lining the chest the pleura ; that lining the abdomen the perito- netwi ; the abdominal cavity is in consequence often called the per- itoneal cavity. Externally the walls of these cavities are covered by the sM7i, which consists of two layers : an outer horny layer called the epidermis, which is constantly being shed on the surface and re- newed from below ; and a deeper layer, called the dermis and con- taining blood, which the ej^ider- mis does not. Between the skin and the lining serous membranes are bones, muscles (the lean of meat), and a great number of other structure's which we shall have to consider hereafter. All cavities inside the body, as the alimentary canal and the air j^as- sages, which open directly or indi- rectly on the surface are lined by soft and moist prolongations of the skin known as mucous mem- branes. In these the same two layers are found as in the skin, but the superficial bloodless one is called epitheUum and the deep- er one the cerium. Diagrammatically we may rep- I'esent the human Body in longi- , • , 1,- , ■ * *T, tudinal section as in Fig. 2, where ber, opening behind into the ° ' pharj'nx, from which one tube aa' is the dorsal or luural cavitii, leads to the lungs, I. and another • i i to the stomach. /; /i, the heart; and b and c, respectively, the k. a kidney; s, the sympathetic ,, . ..-, ., ''.-.. nei-\-ous chain. From the stom- tlioracic and abdommal subdivi- ach, /, the intestinal tube leads . « ,-, , i • , ^ through the abdominal cavity to sious 01 the vcntral cavity: d rep- mliJtaS'cana?''''"^''' ''"" ""'" rescuts the diaphragm separating Fig. 2.— Diagrammatic longi- tudinal section of the body, a, the neural tube, with its upper enlargement in the skull cavity at a'; N, the spinal cord: N', the brain; ee, vertebrae form- ing the solid partition between the dorsal and ventral cavities; 6, the pleural, and c, the abdom- inal divisions of the ventral cav- ity, separated from one another by the diaphragm, d; i, the nasal, and o, the mouth cham- CROSS-SECTION OF THE BODY. 7 them; ee is the vertebral cohimn with its modified prolon- gation into the head beneath the anterior enlargement of the dorsal cavity; / is the alimentary canal opening in front through the nose, i, and mouth, o; h is the heart, / a lung, s the sympathetic nervous system, and k a kidney. A transverse section through the chest is represented diagrammatically in Fig. 3, where x is the neural canal containing the spinal cord. In the thoracic cavity are seen the heart, h, the lungs, //, part of the alimentary canal, a, and the sympathetic nerve centres, sy; the dotted line on each side covering the inside of the chest wall and the outside of the lung represents i\i.Q pleura. Fig. 3.— a diagrammatic section across the Body in the chest region, x. the dorsal tube, which contains the spinal cord : the black mass surrounding it is a vertebra; a. the gullet, a part of the alimentary canal; h, the heart; sy, sym- pathetic nervous system: //, lungs; the dotted lines around them are the pleurae; rr, ribs; st, the breastbone. Sections through corresponding parts of any other Mam- mal would agree in all essential points with those repre- sented in Figs. 2 and 3. Tlie Limbs. The limbs present no such arrangement of cavities on each side of a bony axis as is seen in the trunk. They have an axis formed at different jiarts of one or more bones (as seen at U and R in Fig. 4, which represents a cross-section of the forearm near the elbow joint), but around this are closely packed soft parts, chiefly muscles, and the whole is enveloj^ed in skin. The only cavities in the limbs are branching tubes which are filled with liquids du^^ing life, either hloocl or a watery-looking fluid known as ly^ph. These tubes, the blood and lymph vessels respec- 8 THE HUMAN BODY. tively, are not however characteristic of the limbs, for they are present in abundance in the dorsal and ventral cavities and in their walls. Fig. 4.— a section across the forearm a short distance below the elbow-joint. R and U, its two supporting bones, the radius and ulna; e, the epidermis, and d the dermis of the skin ; the latter is continuous below with bands of connective tissue, s, which penetrate between and invest the muscles, which are indicated by numbers; n, n, nerves and vessels. Chemical Composition of the Body. In addition to the study of the Body as composed of tissues and prgans which are optically recognizable, we may consider it as composed of a number of different chemical substances. This branch of knowledge, which is still very incomplete, really j)resents two classes of joroblems. On the one hand we may limit ourselves to the examination of the chemical substances which exist in or may be derived from the dead Body, or, if such a thing were possible, from the living Body entirely at rest; such a study is essentially one of structure and may be called Chemical Anatotny. But as long as the Body is alive it is the seat of constant chemical trans- formations in its material, and these are inseparably con- nected with its functions, the great majority of which are in the long-run dependent upon chemical changes. From this point of view, then, the chemical study of the Body presents physiological problems, and it is usual to include all the known facts as to the chemical composition and metamorphoses of living matter under the name of Physio- logical Chemistry. For the present we may confine our- selves to the more important substances derived from or known to exist in the Body, leaving questions concerning the chemical changes taking place Avithin it for consideration along with those functions which are performed in connec- tion with them. CHEMISTRY OF THE BODY. 9 Elements Composing the Body. Of the elements known to chemists only sixteen have been found to take part in the formation of tlie human Body. These are carbon, hy- drogen, nitrogen, oxygen, sulphur, iDliosphorus, chlorine, fluorine, silicon, sodium, potassium, lithium, calcium, magnesium, iron, and manganese. Copper and lead have sometimes been found in small quantities but are probably accidental and occasional. Uneombined Elements. Only a very small number of the above elements exist in the body uneombined. Oxygen is found in small quantity dissolved in the blood; but even there most of it is in a state of loose chemical combination. It is also found in the cavities of the lungs and alimentary canal, being derived from the inspired air or swallowed with food and saliva; but while contained in these spaces it can hardly be said to form a part of the Body. Nitro- gen, also exists uneombined in the lungs and alimentary canal, and in small quantity in solution in the blood. Free hydrogen has also been found in the alimentary canal, be- ing there evolved by the fermentation of certain foods. Chemical Compounds. The number of these which may be obtained from the Body is very great; but with re- gard to very many of them we do not know that the form in which we extract them is really that in which .the ele- ments they contain were united while in the living Body; since the methods of chemical analysis are such as always break down the more complex forms of living matter and leave us only its dehris for examination. We know in fact, tolerably accurately, what compounds enter the Body as food and what finally leave it as waste; but the inter- mediate conditions of the elements contained in these com- pounds during their sojourn inside the Body we know very little about; more especially their state of combination dur- ing that part of their stay when they do not exist dissolved in the bodily liquids, but form part of a solid living tissue. For present purposes the chemical com])ounds existing in or derived from the Body may be classified as organic 10 THE HVMAN BODY. and inorganic, and the former be subdivided into those which contam nitrogen and those which do not. Nitrogenous or Azotized Organic Compounds. These fall into several main groups: jjroieids, 2^e2)iones, albu- 7ni)ioixh, cnjstaUiae substances, and coloring 'matters. Proteids are by far the most characteristic substances ob- tained from the Body, since they are only known as exist- ing in or derived from living things, either animals or plants. The type of this class of bodies may be found in the Avhite of an egg, where it is stored uj) as food for the developing chick ; from this typical form, which is called egg albumin, the jsroteids in general are often called albuminous bodies. Each of them contains carbon, hydro- gen, oxygen, sulj)hur, and nitrogen united to form a very complex molecule, and although different members of the family differ from one another in minor points they all agree in their broad features and have a similar percentage composition. The latter in different examples ajoj^ears to vary within the following limits, but it is almost imj)ossible to get any one of them pure for analysis: Carbon 53 to 54 per ceut. , Hydrogen 7 to 7.5 " ! Oxj^gen 21 to 24.0 " 1 Nitrogen 15 to 17.0 " j Sulphur 0.8 to 2.0 " | Proteids are recognized by the following characters: 1. Boiled, either in the solid state or in solution, with strong nitric acid they give a yellow liquid which becomes orange on neutralization Avitli ammonia. This is the xa)itho-proteic test. 2. Boiled with a solution containing subnitratc and per- nitrate of mercury they give a pink precipitate, or, if in very small quantity, a pink-colored solution. This is known as MiUon^s test. 3. If a solution contidning a proteid be acidulated Avitli strong acetic acid and be boiled after the addition of an equal bulk of a saturated w^atery solution of sodium sul- phate, the proteid wall be precipitated. PR0TEID8. PEPTONES. 11 Among the more important proteids obtained from tlie human Body are the following: Serum albumin. This exists in solution in the blood and is very like egg albumin in its proj^erties. It is coagulated (like the white of an egg) when boiled, and then passes into the state of coagulated proteid which is, unlike the original serum albumin, insoluble in dilute acids or alka- lies or in water containing neutral salts in solution. All other proteids can by appropriate treatment be turned into coagulated proteid. Fibrin. This forms in blood when it " clots," either in- side or outside of the body. It is made by the interaction of two other proteids known SiS fibrinogen and fibrinojjlastin. It is insoluble in water. Myosin. This is derived from the muscles, in which it develops and solidifies after death, causing the ''death- stift'ening." Globulin exists in the red globules of the blood and dis- solved in some other liquids of the body. In the blood corpuscles it is combined with a colored substance to form hmmoglobin, which is crystallizable. Casein is found in milk. It is insoluble in water but soluble in dilute acids and alkalies. Its solutions, ^^nlike those of fibrin or myosin, do not coagulate spontaneously, or like that of serum albumin on boiling. In the milk it IS held in solution by the free alkali jiresent; when milk be- comes sour this is neutralized and the casein is precipitated as the '"curd." Cheese consists mainly of casein. Peptones. These are formed in the alimentary canal by the action of some of the digestive liquids upon the proteids swallowed as food. They contain the same elements as the proteids and give the xantho-proteic and Millon's reactions, but are not preci]iitated hj boiling with acetic acid and sodium sulphate. Their great distinctive charac- ter is however their difEusibility. The proteids proper will not dialyze (see Physics), but the peptones in solution pass readily through moist animal membranes. Albuminoids. Tliesc contain carbon, hydrogen, oxy- gen and nitrogen, but rarely any sulphur. Like the 12 THE HUMAN BODY. j)rotcids, the nearest chemical allies of which they seem to be, they are only known in or derived from living beings. Gelatin, obtained from bones and ligaments by boiling, is a typical albuminoid; as is chondrin, which is obtained similarly from gristle. 3fucin, which gives their glairy tenacious character to the secretions of the mouth and nose, is another albuminoid. Crystalline Nitrogenous Substances. These are a heterogeneous group, the great majority of them being materials which have done their work in the Body and are about to be got rid of. Nitrogen enters the Body in foods for tlie most part in the chemically complex form of some j)rotcid. In the vital processes these proteids are broken down into simpler substances; their carbon being partly combined with oxygen and passed out through the lungs as carbon dioxide; tlieir hydrogen is similarly in large part combined with oxygen and passed out as water; while their nitrogen, with some carbon and hydrogen and oxygen, is usually passed out in the form of a crystalline compound, containing what chemists call an ''ammonium residue." COl Of these the most important is urea (Carbamide H^ >■ W), which is eliminated through the kidneys. U)'ic acid is an- other nitrogenous waste product, and many others, such as krcaiin and Tcreatinin, seem to be intermediate stages be- tween the proteids which enter the body and the urea and uric acid which leave it. In the bile or gall, two crystallizable nitrogen-contain- ing bodies, glycocliolic and taurocliolic acids, are found com- bined with soda. Nitrogenous Coloring Matters. These form an arti- ficial group whose constitution and origin is ill known. xVmong the most important are the following: HcBinatin, derived from the red corpuscles of the blood in which a residue of it is combined with a proteid residue to form licBinoglohin. Biliriihin and hiliverdin, which exist in the bile ; the former predominating in the bile of niau and of carnivo- FATS. CARBOHYDRATES. 13 rous animals and giving it a reddish yellow color, while biliverdin predominates in the bile of Herbivora which is green. Non-Nitrogenous Organic Compounds. These may be conveniently grouped as liydrocarbons or fatty bodies; carbohydrates or amyloids ; and certain non-azotized acids. Fats. T\\efats all contain carbon, hydrogen and oxygen, the oxygen being present in small proportion as compared with the hydrogen. Three fats occur in the body in large quantities, viz. : palmatin (CsiHgsOe), stearin (CsTHuoOe), and oleifi (C57H104O6). The two former when pure are solid at the temperature of the Body, but in it are mixed with olein (which is liquid) in such proportions as to be kept fluid. The total quantity of fats in the Body is sub- ject to great variations, but its average quantity in a man weighing 75 kilograms (165 pounds) is about 2.75 kilo- grams (6 pounds). Each of these fats when heated with a caustic alkali, in the presence of water, breaks up into a fatty acid {stearic, palmitic, or oleic as the case may be) and glycerine. The fatty acid unites with the alkali present to form a soap. Carbohydrates. These also contain carbon, hydrogen and oxygen, but there is one atom of oxygen present for every two of hydrogen in the molecule of each of them. Chemically they are related to starch. The more impor- tant of them found in the Body are the following: Glycogen (CeHioOs) found in large quantities in the liver, where it seems to be a reserve of material answering to the starch stored up by many plants. It exists in smaller quantities in the muscles. Glucose, or grape sugar (CbHisOb), which exists in the liver in small quantities ; also in the blood and lymph. It is largely derived from glycogen which is very readily converted into it. Iiiosit, or muscle sugar (CsHiaOs + 2HoO), found in muscles, liver, spleen, kidneys, etc. Lactose, or sugar of milh (OiaHooOn + H^O), found in considerable quantity in milk. Organic Non-Nitrogenous Acids. Of these the most 14 THE HUMAN BODY. important is carbonic dioxide (CO*), which is the form in which by far the greater part of the carbon taken into the Body ultimately leaves it. United with calcium it is found in the bones and teeth in large proportion. Formic, Acetic, and Butyric acids also are found in the Body; stearic, palmitic, and oleic have been above men- tioned as obtainable from fats. Lactic acid is found in the stomach and develops in milk when it turns sour. A body of the same percentage composition, CaHeOa {sarcolac- tic acid), is formed in muscles when they work or die. GlycerO'iihosplioric acid (CsHoPOe) is obtained on the decomposition of lecithin, a complex nitrogenous fat found in nervous tissue. Inorganic Constituents. Of the simpler substances en- tering into the structure of the body the following are the most important: Water ; in all the tissues in greater or less proportion and forming about two thirds of the weight of the whole Body. A man weighing 75 kilos (165 lbs.), if completely dried would therefore lose about 50 kilos (110 lbs.) from the evaporation of water. Of the constituents of the Body the enamel of the teeth contains least Avater (about two per cent) and the saliva most (about 99.5 per cent); between these extremes are all intermediate steps — bones containing about 23 per cent, muscles 75, blood 79. Common salt — Sodium chloride — (NaCl) ; found in all the tissues and liquids, and in many cases playing an important part in keeping other substances in solution in water. Potassium chloride (KCl); in the blood, muscles, nerves, and most liquids. Calcium phosphate (Ca32P04); in the bones and teeth in large quantity. In less proportion in all the other tissues. Besides the above, ammonium chloride, sodium and potassium jDhosphates, magnesium phosphate, sodium sul- phate, potassium sulphate and calcium fluoride have been obtained from the body. Uncombined Hydrochloric acid (HOI) is found in the stomach. CHAPTER II. THE FUNDAMENTAL PHYSIOLOGICAL ACTIONS. The Properties of the Living Body. Wheu we turn from the structure aud compositiou of the living Body to consider its powers and properties we meet with the same variety and complexity, the most superficial examination being sufficient to show that its parts are endowed with very different faculties. Light falling on the eye arouses in us a sensation of sight but falling on the skin has no such effect; pinching the skin causes pain, but pinching a hair or a nail does not: when the ears are stopped, sounds arouse in us no sensation; we readily recognize, too, hard parts formed for support, joints to admit of movements, ajjertures to receive food and others to get rid of wastes. We thus perceive that different organs of our Bodies have Tery different endowments and serve for very distinct pur- poses; and here again the study of internal organs shows us that the varieties of quality observed on the exterior are but slight indications of differences of property which per- vade the whole, being sometimes dependent on the specific characters of the tissues concerned and sometimes u]3on the manner in which these are combined to form various organs. Some tissues are solid, rigid and of constant shape, as those composing the bones and teeth; others, as the muscles, are soft and capable of changing their form; and still others are capable of working chemical changes by which such peculiar fluids as the bile or the saliva are produced. "We find elsewhere a number of tissues com- bined to form a tube adapted to receive food and carry it through the Bodv for digestion, and again similar tissues 16 THE HUMAN BODY. differently arranged to receive tlie air which we breathe-in, and expel after abstracting from it part of its oxygen and adding to it certain other things; and in the heart and blood-YCSsels we find almost the same tissues arranged to propel and carry the blood over the whole Body. The working of the Body oiiers clearly even a more comjilex subject of study than its structure. Physiological Properties. In common with inanimate objects the Body possesses many merely physical properties, as weight, rigidity, elasticity, color, and so on; but in ad- dition to these we find in it while alive many others which it ceases to manifest at death. Of these perhaps the power of executing spontaneous movements and of maintaining a high bodily temperature are the most marked. As long as the Body is alive it is warm and, since the surrounding air is nearly always cooler, must be losing heat all day loug to neighboring objects; neverthe- less we are at the end of the day as warm as at the begin- ning, the temperature of the Body in health not varying much from 37.5° 0. (99° F.), so that clearly our Bodies must be making heat somehow all the time. After death this production of heat ceases and the Body cools down to the temperature in its neighborhood; but so closely do we associate with it the idea of warmth that the sensation experienced in touching a corpse produces so powerful an impression as commonly to be described as icy cold. The other great characteristic of the living Body is its l^ower of executing movements; so long as life lasts it is never at rest; even in the deepest slumber the regular breathing, the tap of the heart against the chest-wall, and the beat of the pulse tell us that we are watching sleep and not death. If to this we add the possession of conscious- ness by the living Body, Avhether aroused by forces im- mediately acting upon sense-organs or not, we might de- scribe it as a heat-producing, moving, conscious organism. The production of heat in the Body needs fuel of some kind as much as its jiroduction in a fire; and every time we move ourselves or external objects some of the Body is used up to supply the necessary working power, just CELLS. 17 as some coals are burnt in the furnace of an engine for every bit of work it does; in the same way every thought that arises in us is accompanied with the destruction of some part of the Body. Hence these primary actions of keeping warm, moving, and being conscious necessitate many otliers for the supply of new materials to the tis- sues concerned and for the removal of their wastes; still others are necessary to regulate the production and loss of heat in accordance with changes in the exterior tempera- ture, to bring the moving tissues into relation with the thinking, and so on. By such subsidiary arrangements the working of the whole Body becomes so complex that it would fill many pages merely to enumerate what is known of the duties of its various parts. However, all the proper physiological properties de^Dend in ultimate analysis on a small number of faculties which are possessed by all living things, their great variety in the human Body depending upon siDccial development and combination in different tissues and organs; and before attempting to study them in their most complex forms it is advantageous to examine them in their simplest and most " produced. a mass consisting of a number of similar units and called the mulberry mass or the morula. At this jDcriod then, long before birth, there are no distinguishable tissues en- tering into the structure of the Body, nor are any organs recognizable. For a short time the morula increases in size by the growth and division of its cells, but very soon new pro- cesses occur which ultimately give rise to the complex DIVISION OF LABOB. 37 adult body with its many tissues and organs. G-roups of cells ceasing to grow and multiply like their parents begin to grow in ways peculiar to themselves, and so come to differ both from the original cells of the morula and from the cells of other groups, and this unlikeness becoming- more and more marked, a varied whole is finally built up from one originally alike in all its parts. Peculiar growth of this kind, forming a complex from a simple whole, is called developinent; and the process itself in this case is known as the differentiation of the tissues, since by it they are, so to speak, separated or specialized from the general mass of mother-cells forming the morula. As the difference:^ in the form and structure of the con- stituent cells of the morula become marked, differences in property arise, and it becomes obvious that the whole cell aggregate is not destined to give rise to a collection of in- dependent living things, but to form a single human being, in whom each part, while maintaining its own life, shall have duties to perform for the good of the whole. In other words, a single compound individual is to be built up by the union and co-operation of a number of simple ones represented by the various cells, each of which thenceforth, while primarily looking after its own interests and having its own peculiar faculties, has at the same time its activi- ties subordinated to tlie good of the entire community. The Physiological Division of Labor. The fundamen- tal physiological properties, originally exhibited by all the cells, become ultimately distributed between the different modified cells which form the tissues of the fully developed Body much in the same way as different employments are distributed in a civilized state; for the difference between the fully developed human Body and the collection of amoeboid cells from which it started is essentially the same as that between a number of wandering savages and a civi- lized nation. In the former, apart from differences de- pendent on sex, each individual has no one special occu- pation different from that of the rest, but has all his own needs to look after: he must collect his own food and prepare it for eating, make his own clothes if he wears 'Z8 THE HUMAN BODY. any, provide liis own shelter, and defend himself from Avild beasts or his fellow-men. In the civilized countr}', on the other hand, we find agriculturists to raise food and cooks to prepare it, tailors to make clothes, and policemen and soldiers to provide i)rotection. And jnst as we find that when distribution of employments in it is more minute the more advanced a nation is in civilization, so is an ani- mal higher or lower in the scale according to the degree in Avhich it exhibits a division of physiological duties between its different tissues. From the subdivision of labor in advanced communities several important consequences arise. In the first place, each man devoting himself to one kind of work mainly and relying upon others for the supply of his other needs, every sort of work gets better done. The man who is constantly making boots becomes more expert than one whose atten- tion is constantly distracted by other duties, and he will not only make more boots in a given time, but better ones; and so with the performance of all other kinds of work. In the second place, a necessity arises for a new sort of indus- try, in order to convey the produce of one individual in excess of the needs of himself and his family to those at a distance who may want it, and to convey back in return the excess of their produce which he needs. The carriage of food from the country to cities, and of city produce to country districts, and the occupation of shopkeeping, are instances of these new kinds of labor which arise in civilized communities. In addition there is developed a need for arrangements by which the work of individuals shall be regulated in j^roportion to the wants of the whole community, such as is in part effected by the agency of large employers of labor who regulate the activities of a number of individuals for the production of various articles in the different quantities required at different times. Exactly similar phenomena result from the subdivision of labor in the human Body. By the distribution of em- ployments between its different tissues, each one specially doing one work for the general community and relying on CLASSIFICATION OF TISSUES. 29 the others for their aid in turn, each necessary work is better performed. And a need arises for a distributive mechanism by which the excess products, if any, of various tissues shall be carried to others which require them, and for a regulative mechanism by which the activities of the various tissues shall be rendered proportionate to the needs of the whole Body at different times and under different circumstances. Accordingly, as we may classify the in- habitants of the United States into lawyers, doctors, clergy- men, merchants, farmers, and so on, we may Classify the Tissues, by selecting the most distinctive properties of each of those entering into the construc- tion of the adult Body and arranging them into physio- logical groups; those of each group being characterized by some one prominent employment. No such classification, however, can be more than approximately accurate, since the same tissue has often more than one well-marked physiological property. The following arrangement, how- ever, is practically convenient. 1. Undifferentiated Tissues. These arc composed of cells which have developed along no one special line, but retain very much the form and properties of the cells form- ing the very young Body before different tissues were re- cognizable in it. The lymph corpuscles and the colorless corpuscles of the blood belong to this class. 2. Supporting Tissues. Including cartilage (gristle), hone, and contiective tissue. Of the latter there are several subsidiary varieties, the two more imj)ortant being wliite fibrous connective tissue, composed mainly of colorless in- extensible fibres, and yelloiv fibrons tissue, composed mainly of yellow elastic fibres. All the supi^orting tissues are used in the Body for mechanical purposes: the bones and carti- lages form the hard framework by which softer tissues are supported and protected; and the connective tissues unite the various bones and cartilages, form investing mem- branes around different organs, and in the form of fine networks penetrate their substance and support their con- stituent cells. The functions of these tissues being for the most part to passively resist strain or pressure, none of 30 THE HUMAN BODY. them litis any very marked j^liysiological property; they are not, for example, irritable or contractile, and their mass is chiefly made up of an intercellular substance which has been formed by the actively living cells sparsely scattered through them, as for instance in cartilage. Fig. 42,* where the cells are seen imbedded in cavities in a matrix which they have formed around them; and which matrix by its firmness and elasticity forms the functionally im})ortant part of the tissue. 3. Nutritive Tissues. This is a large group, the mem- bers of which fall into three main divisions, viz. : Assimilative tissues, concerned in receiving and prepar- ing food materials, and including — {a) Secretory tissues, com])osed of cells Avhich make the digestive liquids poured into alimentary canal, and bringing about chemical or other changes in the food, {h) Receptive tissues, represented by cells which line parts of the alimentary canal and take up the digested food. Eliminative or excretory tissues, represented by cells in the kidneys, skin, and elsewhere, whose main business it is to get rid of the waste products of the various parts of the Body. Respiratory tissues. These are concerned in the gas- eous interchanges between the Body and the surrounding air. They arc constituted by the cells lining the lungs and by the colored corpuscles of the blood. As regards the nutritive tissues it requires especially to be borne in mind that although such a classification as is here given is useful, as helping to show the method i)i!rsued in the domestic economy of the Body, it is only imperfect and largely artificial. Every cell of the Body is in itself assimilative, respiratory, and excretor}^ and the tissues in this class are only those concerned in the first and last interchanges of material between it and the external world. They provide or get rid of substances for the whole Body, leaving the feeding and breathing and excre- tion of its individual tissues to be ultimately louiced after by themselves, just as even the mandarin described by Robin- sou Crusoe who found his dignity promoted by having *P. lui. STORAGE TISSUES. 31 servants to put the food into his month, had finally to swallow and digest it for himself. Moreover, there is no logical distinction between a secretory and an excretory cell: each of them is characterized by the formation of cer- tain substances which are ponred out on a free surface on the exterior or interior of the Body. Many secretory cells too have no concern with the digestion of food, as for example those which form the tears and sweat. 4. Storage Tissues. The Body does not live from hand to mouth: it has always in health a supply of food materials accumulated in it beyond its immediate needs. This lies in part in the individual cells themselves, just as in a pros- perous community nearly every one will have some little pocket-money. But apart from this reserve there are cer- tain cells, a sort of capitalists, which store up considerable quantities of material and constitute what we will call the storage tissues. These are especially represented by the liver-cells and fat-cells, which contain in health a reserve fund for the rest of the Body. Since both of these, to- gether with secretory and excretory cells, are the seats of great chemical activity, they are all often called nietaholic tissues. 5. Irritable Tissues. The maintenance, or at any rate the best prosperity, of a nation is not fully secured when a division of labor has taken place in food-supjjly and food- distribution employments. It is extremely desirable that means shall be i^rovided by which it may receive infornui- tion of external changes which may affect it as a whole, such as the policy of foreign countries ; or which shall en- able the inhabitants of one jiart to know the needs of an- other, and direct their activity accordingly. Foreign min- isters and consuls and newspaper correspondents are em- ployed to place it in communication with other states and keep it informed as to its interests ; and we find also orga- nizations, such as the meteorological department, to warn distant parts of approaching storms or other climatic changes which may seriously affect the pursuits carried on in tliem. In tlie human Body we have a comparable class of ' ktelligence-gaining tissues lying in the sense organs. 32 THE HUMAN BODY. "whose business it is to ascerttiin and communicate to the whole, external changes which occur around it. Since the usefulness of these tissues depends upon the readiness with which slight causes excite them to activity, we may call them the irritahJe tissvef<. 6. Co-ordinating and Automatic Tissues. Such infoi-- mation as that collected by ministers in foreign parts or by meteorological observers, is usually sent direct to some cen- tral office from which it is redistributed; this mere redis- tribution is, however, in many cases but a small j^art of the work carried on in such offices. Let us suppose informa- tion to be obtained that an Indian chief is collecting his men for an attack on some point. The news is probably first transmitted to Washington, and it becomes the duty of the executive officers there to employ certain of the con- stituent units of the society in such definite Avork as is needed for its protection. Troops have to be sent to the place threatened; perhaps recruits enlisted; food and clothes, weapons and ammunition must be provided for the army; and so on. In other words, the work of the various classes composing the society has to be organized for the common good; the mere spreading the news of the danger would alone be of little avail. So in the Body: the information forwarded to certain centres from the irritable tissues is used in such a way as to arouse to orderly activity other tissues whose services are required; we find thus in these centres a group of co-ordinating tissues, represented by nerve-cells and possibly by certain other con- stituents of the nerve centres. Certain nerve-cells are also automatic in the physiological sense already pointed out. The highest manifestation of this latter faculty, shown objectively by muscular movements, is subjectively known as the ''will," a state of consciousness; and other mental phenomena, as sensations and emotions, are also associated with the activity of nerve-cells lying in the brain. How it is that any one state of a material cell should give rise to a particular state of consciousness is a matter quite beyond our powers of conception; but not really more so than how it is that every portion of matter attracts every other por- MOTOR TISSUES. 33 tion according to the law of inverse squares. In the living Body, as elsewhere in the universe, we can study phenomena and make out their relations of sequence or co-existence; but why one phenomenon is accom])anied by another, why in fact any cause produces an effect, is a matter quite beyond our reach in every case; whether it be a sensation accompanying a molecular change in a nerve- cell, or the fall of a stone to the ground in obedience to the law of gravitation. 7. Motor Tissiies. These have the contractility of the original protoplasmic masses highly developed. The more important are ciliated cells and muscular tissue. The for- mer line certain surfaces of the body, and possess on their free surfaces fine threads which are in constant movement. One finds such cells, for example (Fig. 47*) lining the in- side of the windpipe, where their threads or cilia serve, by their motion, to sweep any fluid formed there towards the throat, where it can be coughed up and got rid of. Mus- cular tissue occurs in two main varieties. One kind is found in the muscles attached to the bones, and which are used in the ordinary voluntary movements of the body. It is composed of fibres which present cross-stripes when viewed under the microscope (Fig, 53f), and is hence known as striped or striated muscular tissue. The other kind of muscular tissue is found in the walls of the alimentary canal and some other hollow organs, and con- sists of elongated cells (Fig. ,55 J) which present no cross striation. It is known as plain or unstriated muscular tissue. The cells enumerated under the heading of ''undiffer- entiated tissues'' might also be included among the motor tissues, since they are capable of changing their form. 8. The Conductive Tissues. These are represented by the nerve fibres, slender threads formed by modification and fusion of cells, and having the conductivity of the amoeboid cells of the morula highly developed; that is to say, they readily transmit molecular disturbances. When its equilibrium is upset at one end, a nerve-fibre will transmit to its other a molecular movement known as a *P. 115. t P^m JP. 134 " 34 TEE HUMAN BODY. ^'nervous impulse,'' and so can excite in turn parts distant from the original exciting force. Nerve-fibres place, on the one hand, the irritable tissues in connection with the automatic, co-ordinating, and sensory; and on the other put the three latter in communication with the muscular, secretory, and other tissues. 9. Protective Tissues. These consist of certain cells lining ca^itics inside the body and called epithelial cells, and cells covering the whole exterior of the Body and forming epidermis, hairs, and nails. The enamel which covers the teeth belongs also to this group. The class of protective tissues is, however, even more artificial than that of the nutritive tissues, and cannot be defined by positive characters. Many epithelial cells are secretory, excretory, or receptive; and ciliated cells have already been included among the motor tissues, although from the fact that the movements of their cilia go on in separated cells and independently of recognizable exter- nal stimuli, they might well have been put among the au- tomatic. The protective tissues may be best defined as including cells which line free surfaces, and whose func- tions are mainly mechanical or physical. 10. The Reproductive Tissues. These are concerned in the production of new individuals, and in the human Body are of two kinds, located in different sexes. The conjunction of the products of each sex is necessary for the origination of offspring, since the ovum, or female pro- duct which directly develops into the new human being, lies dormant until it has been fertilized or acted ui)on by the product of the male. The Combination of Tissues to Form Organs. The va- rious tissues above enumerated forming the building mate- rials of the Body, anatomy is primarily concerned with their structure, and physiology with their properties. If tliis, however, were the whole matter, the problems of anatomy and physiology would l)e much simpler than they actually are. The knowledge about the living Body obtained by studying only the forms and functions ORGANS. 35 of the individual tissues would be comparable to that at- tained about a great factory by studying separately the boilers, pistons, levers, wheels, etc., found in it, and leav- ing out of account altogether the way in which these are combined to form various machines; for in the Body the various tissues are for the most part associated to form organs, each organ answering to a complex machine like a steam-engine with its numerous constituent parts. And just as in different machines a cogged wheel may perform very different duties, dependent upon the way in which it is connected with other parts, so in the Body any one tis- sue, although its essential properties are everywhere the same, may by its activity subserve very various uses accord- ing to the manner in which it is combined with others. For example: A nerve-fibre uniting the eye Avith one part of the brain will, by means of its conductivity, when its end in the eye is excited by the irritable tissue attached to it on which light acts, cause changes in the sensory nerve- cells connected with its other end and so arouse a sight sensation; but an exactly similar nerve-fibre running from the brain to the muscles will, also by virtue of its conduc- tivity, when its ending in the brain is excited by a change in a nerve-cell connected with it, stir up the muscle to con- tract under the control of the will. The different results depend on the different parts connected with the ends of the nerve-fibres in each case, and not on any difference in the properties of the nerve-fibres themselves. It becomes necessary then to study the arrangement and uses of the tissues as combined to form various organs, and this is frequently far more difficult than to make out the structure and properties of the individual tissues. A\\ or- dinary muscle, such as one sees in the lean of meat, is a very complex organ, containing not only contractile mus- cular tissue, but supporting and uniting connective tissue and conductive nerve-fibres, and in addition a complex commissariat arrangement, composed in its turn of several tissues, concerned in the food supply and waste removal of the whole muscle. The anatomical study of a muscle has 36 THE HUMAN BODY. to take into account the arrangement of all these parts within it, and also its connections with other organs of the Body. The physiology of any muscle must take into ac- count the actions of all these parts working together and not merely the functions of the muscular fibres themselves, and has also to make out under what conditions the muscle is excited to activity by changes in other organs, and what changes in these it brings about when it works. Physiological Mechanisms. Even the study of organs added to that of the separate tissues does not exhaust the whole matter. In a factory avc frequently find machines arranged so that two or more shall work together for the performance of some one work : a steam-engine and a loom may, for example, be connected and used together to weave carpets. Similarly in the Body several organs are often arranged to work together so as to attain some one end by their united actions. Such combinations are known as physiological cqjparatuses. The circulatory apparatus, for example, consists of various organs (each in turn composed of several tissues) known as heart, arteries, capillaries, and veins. The heart forms a force-pump by which the blood is kept flowing through the whole mechanism, and the rest, known together as the blood-vessels, distribute the blood to the various organs and regulate the supply accord- ing to their needs. Again, in the visual apparatus we find the co-operation of (a) a set of optical instruments which bring the light proceeding from external objects to a focus upon (5) the reifi/^a, which contains highly irritable parts; these, changed by the light, stimulate (^•) the optic nerve, which is conductive and transmits a disturbance Avhich arouses finally (d) sensory parts in the brai7i. In the pro- duction of ordinary sight sensations all these parts are con- cerned and work together as a visual apparatus. So, too, we find a respiratory apparatus, consisting primarily of two hollow organs, the lungs, which lie in the chest and com- municate by the ivindpipe with the back of the throat, from Avhicli air enters them. But to complete the respi- ratory apparatus are many other organs, bones, muscles, nerves, and nerve-centres, which work together to renew ANATOMICAL SYSTEMS. 37 the air in the lungs from time to time; and the act of breathing is the final result of the activity of the whole apparatus. Many similar instances, as the alimentary apparatus, the auditory apparatus, and so on, will readily be thought of. The studv of the working of such complicated mechanisms forms a very important part of physiology. Anatomical Systems. From the anatomical side the whole collection of bodily organs agreeing in structure with one another is often spoken of as a system; all the muscles, for example, are grouped together as the muscular system, and all the bones as the osseous system, and so on, without any reference to the different uses of different muscles or bones. The term system is, however, often used as equivalent to " apparatus:" one reads indifferently of the "circulatory system" or the '^ circulatory apparatus." It is better, however, to reserve the term system for a collec- tion of organs classed together on account of similarity of structure; and "'apparatus" for a collection of organs con- sidered together on account of their co-operation to execute one function. The former term will then have an anatomi- cal, the latter a physiological, significance. The Body as a Working Whole. Finally it must all through be borne in mind that not even the most complex system or apparatus can be considered altogether alone as an independently living part. All are united to make one living Body, in which there is throughout a mutual inter- dependence, so that the whole forms one human being, in whom the circulatory, respiratory, digestive, sensory, and other apparatuses are constantly influencing one another, each modifying the activities of the rest. This interaction is mainly brought about through the conductive and co- ordinating tissues of the nervous system, which place all parts of the Body in communication. But in addition to this another bond of union is formed by the blood, which by the circulatory apparatus is carried from tissue to tissue and organ to organ, and so, bringing materials derived in one region to distant parts, enables each organ to influence all the rest for good or ill. 38 THE HUMAN BODY. Besides the blood another liquid, called lymph, exists in the Body. It is contained in vessels distinct from those which carry the blood, but emptying into the blood-vessels at certain points. This liquid being also in constant move- ment forms another agency by which products are carried from part to part, and the welfare or ill-fare of one member enabled to influence all. CHAPTER IV. THE INTERNAL MEDIUM. The External Medium. During the whole of life inter- changes of material go on between every living being and the external world; b}^ these exchanges material particles that one time constitute parts of inanimate objects come at another to form part of a living being; and later on these same atoms, after having been a part of a living cell, are passed out from the Body in the form of lifeless com- pounds. As the foods and wastes of various living things differ more or less, so are more or less different environ- ments suited for their existence; and there is accordingly a relationship between the plants and animals living in any one place and the conditions of air, earth, and water prevailing there. Even such simple unicellular animals as the amceba live only in water or mud containing in solu- tion certain gases and, in suspension, solid food particles; and they soon die if the water be changed eitlier by essen- tially altering its gases or by taking out of it the solid food. So in yeast we find a unicellular plant which thrives and multiplies only in liquids of certain composition, and which in the absence of organic compounds of carbon in solution will not grow at all. Each of these simple living things, which corresponds to one only of the innumerable cells composing the full-grown human Body, thus requires for the manifestation of its vital properties the presence of a surrounding medium suited to itself: the yeast would die, or at the best lie dormant, in a liipiid containing only the solid organic particles on which the amoeba lives; and the amoeba would die in such solutions as those in which veast thrives best. 40 TEE HUMAN BODY. The Internal Medium. The same close relationship between the living being and its environment, and the same cyclical interchange between the two which we find in the amoeba and the yeast-cell, occur also in even the most complex living, beings. When, however, an animal comes to be composed of many cells, some of which are placed far away from the surface of its body and so from immediate contact with the environment, there arises a new need — a necessity for an internal medium or jjhiftma which shall play the same part toward the individual cells as the surrounding air, water, and food to the whole animal. This internal medium kept in movement, and receiving at some regions of the bodily surfaces materials from tlie exterior, while losing other substances to the exterior at other sur- faces, thus forms a sort of middleman between the in- dividual tissues and the surrounding world, and stands in the same relationship to each of the cells of the Body as the water in which an amoeba lives does to that animal or beer-wort does to a yeast-cell. We find accordingly the human Body pervaded by a liquid plasma, containing gases and food material in solution, and the presence of which is necessary for the maintenance of the life of the tissues. Any great change in this medium will affect injuriously few or many of the groups of cells in the Body, or may even cause their death; just as altering the media in which they live will kill an amoeba or a yeast-cell. The Blood. In the human Body the internal medium is primarily furnished by the Mood, which, as every one knows, is a red liquid, very widely distributed over the frame, since it flows from any part when the skin is cut through. There are in fact very few portions of the Body into which the blood is not carried. One of the exceptions is the epidermis, or outer layer of the skin : if a cut be made through it only, leaving the deeper skin-layers in- tact, no blood will flow from the wound. Hairs and nails also contain no blood. In the interior of the Body the epithelial cells lining free surfaces, such as the inside of the alimentary canal, contain no blood, nor do the hard parrs of the teeth, the cartilages, and the refracting media THE INTERNAL MEDIUM. 41 of the eye (see Chap. XXXI.), but these interior parts are moistened with liquid of some kind, and unlike the epi- dermis are protected from rapid evaporation. All these bloodless parts together form a group of non-vascular tis- sues; they alone excej^ted, wounding any part of the Body will be followed by bleeding. In many of the lower animals there is no need that the liquid representing their blood should l)e renewed very rapidly in different parts. Their cells live slowly, and so require but little food and produce but little waste. In a sea anemone, for example, there is no special arrangement to keep the blood moving ; it is just pushed about from part to part by the general moyements of the body of the animal. But in higher animals, especially those with an elevated temperature, such an arrangement, or rather ab- sence of arrangement, as this would not suffice. In them the constituent cells live very fast, making much waste and using much food, and so alter the blood in their neigh- borhood very raj^idly. Besides, we have seen that in com- plex animals certain cells are set apart to get food for the whole organism, and certain others to finally remove its wastes, and there must be a sure and rapid interchange of material between the feeding and excreting tissues and all the others. This can only be brought about by a rapid movement of the blood in a definite course, and this is ac- complished by shutting it up in a closed set of tubes, and placing somewhere a pump, which constantly takes in blood from one end of the system of tubes and forces it out again into the other. Sent by this pump, the heart, through all parts of the Body and back to the heart again, the blood gets food from the receptive cells, takes it to the working cells, carries off the waste of these latter to the excreting cells ; and so the round goes on. The Lymph. The blood, however, lies everywhere in closed tubes formed by the vascular system, and does not come into direct contact with any cells of the Body except those which float in it and those which line the interior of the blood-vessels. At one part of its course, however, the vessels through which it passes have extremely thin A. Jb_ic_ 42 THE HUMAN BODY. coats, and through the walls of these capillaries liquid transudes from the blood and bathes the various tissues. The transuded liquid is the lymph, and it is this which forms the immediate nutrient plasma of the tissues except the few which the blood moistens directly. Dialysis. When two liquids containing different mat- ters in solution are separated from one another by a moist animal membrane, an interchange of material will take place under certain conditions. If A be a vessel (Fig. 9) completely divided vertically by such a membrane, and a solution of com- mon salt in water be placed on the side h, and a solution of sugar in water on the side c, it will be found after a time that some gram of a diaiy- Salt has got into c and some sugar into J, al- contammg\wo though there are no visible j^ores in the parti- c,*separated'^by ^ioi^' Sucli an interchange is said to be due membrane!'™^^ ^^ £?mZys*s or osmosis, and if the process were allowed to go on for some hours the same })roportions of salt and sugar would be found in the solu- tions on each side of the dividing membrane. The Renewal of the Lymph. Osmotic processes play a great part in the nutritive i)rocesses of the Body. The lymph jjresent in any organ gives up things to the cells there and gets things from them; and so, although it may have originally been tolerably like the liquid part of the blood, it soon acquires a different chemical composition. Diffusion or dialysis then commences between tlie lymph outside and the blood inside the capillaries, and the latter gives up to the lymph new materials in place of those which it has lo^t and takes from it the waste products it has received from the tissues. When this blood thus altered by exchanges with the lymi^h gets again to the neighborhood of the re- ceptive cells, having lost some food materials it is poorer in these than the richly sujjplied lymph around those cells, and takes up a supply by dialysis from it. When it reaches the excretory organs it has previously picked up a quantity of waste matters and loses these by dialysis to the lymph there present, which is specially j^oor in such matters,. LYMPHATICS. 43 since the excretory cells constantly deprive it of them. In consequence of the different wants and wastes of various cells, and of the same cells at different times, the lym])h must vary considerably in composition in various organs of the Body, and the blood flowing through them will get or lose different things in different places. But renewing during its circuit in one what it loses in another, its aver- age composition is kept pretty constant, and, through in- terchange with it, the average composition of the lymph also. The Lymphatic Vessels. The blood, on the whole, loses more li(juid to the lymph through the capillary walls than it receives back the same way. This depends mainly on the fact that the pressure on the blood inside the ves- sels is greater than that on the lymph outside, and so a certain amount of filtration of liquid from Avithin out occurs through the vascular wall in addition to the dialysis proper. The excess is collected from the various organs of the Body into a set of hjmpliatic vessels Avhich carry it directly back into some of the larger blood-vessels near where these empty into the heart; and by this flow of lymph, under pressure from behind, it is renewed in various or- gans, fresh liquid filtering through the capillaries to take its place as fast as the old is carried off. The Lacteals. In the walls of the alimentary canal cer- tain food materials after passing through the receptive cells into the lympli are not transferred locally, like the rest, by dialysis into the blood, but are carrried off' bodilv in the lymph-vessels and poured into the veins of a distant part of the Body. The lymphatic vessels concerned in this work, being frequently filled with a white li(piid during di- gestion, are called the milky or lacteal vessels. Summary. To sum up: the blood and lym})]i form the internal medium in which the tissues of the Body live; the lymph is primarily derived from the blood and forms the immediate jdasma for the great majority of the living cells of the Body; and the excess of it is finally returned to the blood. The lympli moves but slowly, but is constantly renovated by the blood, which is kept in rai)id movement, 44 THE nUMAN BODY. and which, besides contiiining a store of new food matters for the lymph, carries off the wastes which the various cells have poured into the latter, and thus is also a sort of sewage stream into Avhich the wastes of the whole liody are pri- marily collected. Microscopic Characters of Blood. If a finger be pricked, and the drop of blood flowing out be received on a glass slide, covered, protected from evaporation, and ex- amined with a microscope magnifying about 400 diameters, it Avill be seen to consist of innumerable solid bodies float- ing in a liquid. The solid bodies are the hlood corpuscles, and the liquid is the Uood pilasma or liquor sanguinis. The corpuscles are not all alike. While currents still exist in the freshly spread drop of blood, the great majority of them are readily carried to and fro; but a certain num- ber more commonly stick to the glass and remain in one place. The former are the red, the latter the pale or color- less hlood corpuscles. Red Corpuscles. Form and Size. The red corpuscles as they float about frequently seem to vary in form, but by a little attention it can be made out that this appearance is due to their turning round as they float, and so presenting different aspects to view; just as a silver dollar presents a different outline according as it is looked at from the front or edgewise or in three-quarter i)rofile. Sometimes the corpuscle (Fig. 10, B) appears circular; then it is seen in full face; sometimes linear (C), and slightly narrowed in the middle; sometimes oval, as the dollar when half-way between a full and a side view. These appearances show that each red cori)uscle is a circu- lar disk, slightly hollowed in the middle (or biconcave) and about four times as wide as it is thick. The average trans- verse diameter is 0.008 millimeter (-g^o-jp inch). — Color. Seen singly each red corpuscle is of a pale yellow color; it is only when collected in masses that they appear red. The blood owes its red color to the great numbers of these bodies in it; if it be spread out in a very thin layer it, too, is yellow. The layer must, however, be very thin or the drop will still look red on account of the immense number BLOOD. 45 of these corpuscles present; in a cubic millimeter (o^ inch) of blood there are about five millions of them. — Structure. Seen from the front the central part of each red corpuscle in a certain focus of the microscope appears dimmer or darker than the rest (Fig. 10, B), except a narrow band near the outer rim. If the lens of the microscope be raised, however, this previously dimmer central part becomes brighter, and the previously brighter part obscure {E). Fig. 10.— Blood corpuscles. A, magnified about 400 diameters. The red cor- puscles have arranged themselves in rouleaux ; a, a, colorless corpuscles ; B, red corpuscles more magnified and seen in focus ; E, a red corpuscle slightly out of focus. At the right-hand top coi'ner is a red corpuscle seen in three- quarter face, and at C one seen edgewise. F, O, H, I, white corpuscles highly magnified. This difference in appearance does not indicate the presence of a central part or nucleus different from the rest, but is an optical phenomenon due to the shape of the corpuscle, in consequence of which it acts like a little biconcave lens (see Physics). Eays of light passing through near the centre of the corpuscle are refracted differently from those passing through elsewhere; and when the microscope is so focused that the latter reach the eye, the former do not^ 46 THE HUMAN BODY. and vice versa; iliiis when the central parts look bright, those around them look obscure, and the contrary. There is no .satisfactory evidence that these corpuscles have any enveloping sac or cell-wall. All the methods used to bring one into view under the microscope are such as Avould coagulate the outer layers of the substance com- posing the corpuscle and so make an artificial envelope. ISo far as optical analysis goes, then, each corpuscle is ho- mogeneous throughout. By other means we can, however, show that at least two materials enter into the structure of each red corpuscle. If the blood be diluted with several times its own bulk of water and bo then examined with the microscope, it will be found that the red corpuscles are col- orless and the plasma colored. The dilution has caujjed tlie coloring matter to pass out of the corpuscles and dis- solve in the liquid. This coloring constituent of the cor- puscle is licemoglohin, and the colorless residue which it leaves behind and which swells up into a sphere in the di- luted plasma is the stroma. In the living corpuscle the two are intimately mingled throughout it, and so long as this is the case the blood is opaque; but when the coloring matter dissolves in the plasma, then the blood becomes transparent, or, as it is called, lahy. The difference may be very well seen b}' comparing a thin layer of fresh blood diluted with ten times its volume of tcn-per-cent salt so- lution with a similar Liver of blood diluted with ten vol- umes of Avater. The watery mixture is a dark transparent red; the other, in which the coloring matter still lies in the corpuscles, is a brighter opaque red. — Consistency. Each red corpuscle is a soft jelly-like mass which can be readily crushed out of shape. Unless the pressure be such as to rupture it, the corpuscle immediately reassumes its pro])er form when the external force is removed. The cor- puscles are, then, highly elastic; they fretpiently can be seen much dragged out of shape inside the vessels when the circulation of the blood is watched in a living animal (Chap. XA'.), but immediately springing back to their nor- mal form when they get a chance. Blood-Crystals. Hcemoglobin is, as above shown, readily BL00D-CRTSTAL8. 47 soluble in water. In this it soon decomposes if kept in a warm room, breaking up into a proteid substance called globulin and a red-colored body, luematin. By keeping the hemoglobin solution very cold, however, this decompo- sition can be greatly retarded, and at the same time the solubility of the haemoglobin in the water much diminished. In dilute alcohol haemoglobin is still less soluble, and so if its ice-cold watery solution have one fourth of its volume of cold alcohol added to it and the mixture be put in a re- frigerator for twenty-four hours, a part of the hc^moglobin will often crystallize out and sink to the bottom of the vessel, where it can be collected for examination. Tlie haemoglobin of the rat is less soluble than that of man, and therefore crys- tallizes out especially easily; but these haemo- globin crvstals, or. as they are often called, blood-crystals, can be obtained from human blood. In 100 parts of ^^^ „_b1ouc1 -crystals, or hemoglobin dry human red blood- crystals. corpuscles there are 90 of haemoglobin. The ha?moglobin is the essential constituent of the red blood corpuscles, enabling them to pick up large quantities of oxygen in the lungs and carry it to all parts of the Body, (See Ees- piration, ) Haemoglobin contains a considerable quantity of iron, much more than auv other proximate constituent of the Body. The Colorless Blood Corpuscles (Fig, 10, F, H, G). The colorless, pale, or wJiife corpuscles of the blood are far less numerous than the red; in health there is on the ave- rage about one white to three hundred red, but the pro- portion may vary considerably. Each is finely granular and consists of a soft mass of protoplasm enveloped in no definite cell-wall, but containing a nucleus. The granules in the protoplasm commonly hide the nucleus in a fresh 48 THE HUMAN BODY. corpuscle, but dilute acetic acid dissolves most of them and brings the nucleus into view. Tliese ])ale corpuscles belong to the group of undifferentiated tissues and differ in no important recognizable character from the cells which make up the whole very young human Body, nor indeed from such an unicellular animal as an Amaba. Like the latter, they have the power of sloAvly changing their form spontaneously, and so have not the definiteness of outline which belongs to the red corpuscles. At one moment (Fig. 12) a pale corpuscle will be seen as a spheroidal mass; a few seconds later processes will be seen radiating from this, and soon after these pro- cesses may be retracted and others thrust out; and so tlie corpuscle goes on changing its shape. These slow amcehoid moi^ements are greatly pro- FiG. 12.-A white blood nioted by keeping the specimen of cori^uscle sketched at sue- j i o i cessive intervals of a few ijlood at the temperature of the Body seconds to illustrate the ^. . -n t changes of form due to its wllllc Under examination. By thrust- am ceboid movements. . , ■ t , i mg out a process on one side, then drawing the rest of its body up to it, and then sending out a process again on the same side, the corpuscle can slowly change its place and creep across the field of the micro- scope. Inside the blood-vessels these corpuscles execute quite similar movements; and they sometimes bore right through the capillary walls and, getting out into the lymph spaces, creep about among the other tissues. This emigra- tion is especially frequent in inflamed parts, and the ptis or " matter''' which collects in abscesses is largely made up of white blood corpuscles which have in this way got out of the blood-vessels. The size of the white corpuscles is not so constant as that of the red; on the whole, however, they are larger, their average diameter being about 0.0127 milli- meter (-gifiro i"ch). The general properties of those cor- puscles have already been described in Chap. II. Blood of Other Animals. In all animals with blood the pale corpuscles are pretty much alike, but the red corpus- cles, which with rare exceptions are found only in Verte- LYMPH. 49 brates, vary considerably. In all the class of the mammalia they are circular biconcave disks Avith the exception of the camel tribe, in which they are oval. They vary in diam- eter from .002 mm. (musk deer) to .011 mm. (elephant). In the dog they are nearly the same size as those of man. In no mammals do the fully developed red corpuscles possess a nucleus. In all other vertebrate classes the red corpus- cles possess a central nucleus, and are oval slightly biconvex disks except in a few fishes in which they are circular. They are largest of all in the amphibia. Those of the frog are 0.02 ram. { -^-^^ inch ) long and .007 mm. broad. Histology of Lymph. Pure lymph is a colorless watery- looking liquid; examined with a microscope it is seen to contain numerous pale corpuscles exactly like those of the blood, and no doubt largely consisting of pale blood cor- puscles which have emigrated. It contains none of the red corpuscles. The lymph flowing from the intestines durmg digestion is, as already mentioned, not colorless but white and milky. It is known as chyle and will be considered with the process of digestion. During fasting the lymph from the intestines is colorless like that from other parts of the Body. CHAPTER V. THE CLOTTING OF BLOOD. The Coagulation of the Blood. When blood is first drawn from the living Body it is perfectly liquid, flowing in any direction as readily as water. This condition is, however, only temporary ; in a few minutes the blood be- comes viscid and sticky, and the viscidity becomes more and more marked until, after the lapse of five or six min- utes, the Avhole mass sets into a jelly Avhich adheres to the vessel containing it so that this may be inverted without any blood whatever being spilled. This stage is known as that of gehdinization and is also not permanent. In a few minutes the top of the jelly-like mass will be seen to be hollowed or ''cupped" and in the concavity will be seen a small quantity of nearly colorless liquid, the hlood serum. The jelly next shrinks so as to pull itself loose from the sides and bottom of the vessel containing it, and as it shrinks, squeezes out more and more serum. Ulti- mately we get a solid clot, colored red, and smaller in size than the vessel in which the blood coagulated but retain- ing its form, floating in a quantity of pale yellow serum. If, however, the blood be not allowed to coagulate in per- fect rest, a certain number of red corpuscles Avill be rubbed out of the clot into the serum and the latter will be more or less reddish. Tlie longer the clot is kept the more serum will be obtained: if the first quantity exuded be decanted off and the clot put aside and protected from evaporation, it will in a short time be foiind to have shrunk to a smaller size and to have pressed out more serum; and this goes on as long as it is kept, until putrefactive changes commence. CAUSES OF COAGULATION. 51 Cause of Coagulation. If a drop of fresh-drawn blood be spread out and watched with a microscope magnifying 600 or 700 diameters, it will be seen that the coagulation is due to the separation of very fine solid threads which run in every dii-ection through the plasma and form a close network entangling all the corpuscles. These threads are composed of a proteid substance known as fibrin. When they tirst form, the whole drop is much like a sponge soaked full of water (represented by the serum) and having solid bodies (the corpuscles) in its cavities. After the fibrin tlireads have been formed they tend to shorten ; hence when blood clots in mass in a vessel, the fibrinous network tends to shrink in every direction just as a network formed of stretched india-rubber bands would, and this shrinkage is greater the longer the clotted blood is kept. At first the threads stick too firmly to the bottom and sides of the vessel to be pulled away, and thus the first sign of the contraction of the fibrin is seen in the cupping of the surface of the gelatinized lilood where the threads have no solid attachment, and there the contracting mass presses out from its meshes the first drops of serum. Finally the contraction of the fibrin overcomes its adhesion to the vessel and the clot pulls itself loose on all sides, j^yressing out more and more serum, in which it ultimately floats. The great majority of the red corpuscles are held back in the meshes of the fibrin, but a good many pale corpuscles, by their amoeboid movements, work their way out and get into the serum. Whipped Blood. The essential point in coagulation being the formation of fibrin in the plasma, and blood only forming a certain amount of fibrin, if this be removed as fast as it forms the remaining blood will not clot. The fibrin may be separated by what is known as "whipping" the blood. For this purpose fresh-drawn blood is stirred up vig- orously with a bunch of twigs, and the sticky fibrin threads as they form adhere to these. If tlie twigs be withdrawn after a few minutes a quantity of stringy material will be found attached to them. This is at first colored red by adhering blood corpuscles: but l)y washing in water these 52 THE HUMAN BODY. may be removed, and the piu-e librin thus obtained is per- fectly white and in the form uf highly elastic tlireads. It is insoluble in water and in dilute acids, but swells up to a transparent jelly in the latter. The ''whipped" or "defi- brinated blood" from which the fibrin has been in this way removed, looks just like ordinary blood, but has lost its power of coagulating spontaneously. The Buffy Coat. That the red corpuscles are not an essential part of the clot, but are merely mechanically caught up in it, seems clear from the microscopic ob- servation of the process of coagulation; and from the fact that perfectly formed fibrin can be obtained free from cor- puscles by whipping the blood and washing the threads which adhere to the twigs. Under certain conditions, moreover, one gets a naturally formed clot containing no red corpuscles in one part of it. The corpuscles of human blood are a little heavier, bulk for bulk, than the plasma in which they float; hence, when the blood is drawn and left at rest they sink slowly in it; and if for any reason the clotting takes place more slowly or the corpuscles sink more rapidly than usual, a colorless top stratum of plasma, with no red corpuscles in it, will be left before gelatiniza- tion occurs and stops the farther sinking of the corpuscles. The uppermost part of the clot formed under these cir- cumstances is colorless or pale yellow, and is known as the huffy coat; it is especially apt to be formed in the blood drawn from febrile j^atients, and was therefore a point to which physicians paid much attention in the olden times when bloodletting was thought a panacea for all ills. In horse's blood the difference between the specific gravity of the corpuscles and that of the plasma is greater than in hu- man blood, and horse's blood also coagulates more slowly, so that its clot has nearly always a buffy coat. The color- less buiiy coat seen sometimes on the top of the clot must, however, not be confounded with another phenomenon. When a blood-clot is left floating exposed to the air its top becomes bright scarlet, while the part immersed in the serum assumes a dark purple-red color. The brightness of the top layer is due to the action of the oxygen of the air. USES OF COAGULATION. 53 which forms a bright reel compound with the coloring mat- ter of the red corpuscles. If the clot be turned upside down and left for a short time, the previously dark bottom layer, now exposed to the air, will become bright; and the previ- ously bright top layer, now immersed in the serum, will become dark. Uses of Coagulation. The clotting of the blood is so important a process that its cause has been frequently in- vestigated; but as yet it is not perfectly understood. The living circulating blood in the healthy blood-vessels does not clot; it contains no solid fibrin, but this forms in it, sooner or later, when the blood gets by any means out of the vessels or if the lining of these is injured. In this way the mouths of the small vessels opened in a cut are clogged up, and the bleeding, Avhich would otherwise go on indefinitely, is stopped. So, too, when a surgeon ties up an artery be- fore dividing it, and the tight ligature crushes or tears its delicate inner surface, the blood clots where this is injured, and from there a coagulum is formed reaching up to the next highest branch of the vessel. This becomes more and more solid, and by the time the ligature is removed has formed a firm plug in the cut end of the artery, which gi-eatly diminishes the risk of bleeding. The Fibrin Factors. As regards the formation of fibrin the folloAving points seem to be made out with toler- able certainty. Fresh-drawn blood contains or develops two substances, fihrinoplastin and fihrinogen, which by their interaction form fibrin, under the influence of a third body called the fihrin ferment; moreover, fibrin is only formed if a certain proportion of neutral mineral salts, such as are found dissolved in the blood plasma, is present. Blood serum does not clot of itself at ordinary tempera- tures: it contains fihrinoplastin and fibrin ferment and the requisite quantity of salts, but not the fibrinogen; that which originally existed in the plasma having apparently been used up with the proper proportion of fihrinoplastin to form fibrin, leaving over an excess of fibriaoplastin in solution in the serum. On the other hand, the liquids found in the cavities of 54 THE HUMAN BODY. the Body which are lined by serous membraneSj commonly contain iibrinogen and the salts but no fibrinoplastin, and therefore they do not coagulate spontaneously. But if a little blood serum be added to one of these liquids, coagula- tion takes place. Artificial Clot. If serum l)e slightly diluted with water and kept ice-cold while a stream of carbon dioxide gas is passed through it for some hours, a white precipitate is thrown down which contains fibrinoplastin and the fibrin ferment. This precipitate after washing may be dissolved in cold water containing the merest trace of caustic potash. If the liquid moistening a serous cavity be treated in a similar way a precipitate is formed, containing fibrinogen instead of the fibrinoplastin, and but little of the ferment. If this precipitate be washed and dissolved and the solution be added to the solution of the blood-serum precipitate, no clot is formed; but if about one per cent of sodic car- bonate or other neutral salt be added to the mixture, then it clots. This shows the necessity of the salts, which is perhaps better proved in another way. If serum be put in a dialyzer (see Physics) with distilled water on the other side of the membrane, all the salts will gradually pass out from the serum into the water: as the last portions of them pass out, the fibrinoplastin and ferment, which are "colloids''' (that is, bodies which will not dialyze), are pre- cipitated; they may be redissolved by the addition of a trace of caustic potash. Similarly the salts may be re- moved from the liqiiid obtained from a serous cavity, and the precipitated fibrinogen redissolved. If these solutions be now mixed no clot is formed; but if the salts which have been dialyzed ottt, or an equivalent portion of other neu- tral salts, be added to the mixture, it will clot. The Fibrin Ferment. Tlie activity of the ferment i.- proved as follows: If scrum be diluted with a large bulk of water and then caribou dioxide gas be passed through it, fibrinoplastin will be precipitated, with little or none of the ferment. If this fibrinoplastin be dissolved and added to the liquid from a serous cavity it will not cause it to clot, or only very slowly, according as no fibrin ferment or FIBRIN FERMENT. 56 but a little is present. But if some of the ferment be uclcled, then the mixture co-igulutes nipidl}-. The ferment may be obtained b}^ adding a large quantity of strong al- cohol to some fresh blood serum. The alcohol precipi- tates albumen, fibrinoplastin, and the ferment. The pre- cipitate is let stay under alcohol for some months, during which time the albumen and fibrinojilastin are altered so as to become insoluble in water. The alcohol is then de- canted off and the residue treated with water which dis- solves the ferment. This solution added to the above mixture containing fibrinoplastin, fibi'inogeu, and salts, will make it clot. Of these four bodies which play a part in the coagula- tion of the blood, the fibrinoplastin and fibrinogen jiri- marily determine the quantity of fibrin formed. The fer- ment seems to act on them in some way so as to make them interact, but it does not enter into the fibrin; it is not used up in the process, and the quantity of fibrin formed is thus independent of the quantity of the ferment joresent; but the more of it there is, the more quickly does the coagula- tion occur. The part the salts play is obscure: probably part of them are necessary constituents of the fibrin, since it leaves a large proportion of ash when burnt. But they seem to act in some other way when present in certain proportions, since too large a percentage of them stops coagulation as completely as their total absence. If fresh blood be mixed with an equal bulk of a saturated solution of magnesium sulphate (Epsom salts) or of common salt, it will not clot; but if this mixture be largely diluted with water, then clotting Avill take place. Exciting Causes of Coagulation. The above facts show clearly enough that the coagulation of the blood is a physico-chemical process, but still leave unexplained why it does not occur in circulating blood inside healthy blood- vessels. It is, in fact, much easier to point out what arc not the proximate reasons of the coagulation of drawn blood than what are. Blood when removed from the Body and received in a vessel comes to rest, cools, and is exposed to the air. from 56 THE HUMAN BODY. which it may receive or to which it may give off gaseous bodies. But it is easy to prove that none of these three things is the cause of coaguhition. Stirring the drawn blood and so keeping it in movement does not prevent but hastens its coaguhition; and blood carefully imprisoned in a living blood-vessel, and so kept at rest, will not clot for a long time: not until the inner coat of the vessel begins to change from the "want of fresh blood. Secondly, keeping the blood at the temperature of the Body hastens coagulation, and cooling retards it; blood received into an ice-cold vessel and kept surrounded with ice will clot more slowly than blood drawn and left exposed to ordinary temperatures. Finally, if the blood be collected over mercury from a blood-vessel, without having been exposed to the air even for an instant, it will still clot perfectly well. The formation of fibrin is then due to changes taking place in the blood itself when it is removed from the blood-vessels; clotting depends upon some rearrangement of the blood constituents. There is a good deal of reason to believe that what occurs is a breaking up of a number of the colorless corpuscles; that these then form fibrino- plastin and fibrin ferment, and, the fibrinogen and salts already existing in solution in the blood plasma, fibrin is formed. \Yhen fluids which contain no red corpuscles clot, as for instance vaccine lymph, the first threads of fibrin developed can be seen under the microscope to radiate from the pale corpuscles present. Relation of the Blood- Vessels to Coagulation. As to the role of the vessels with respect to coagulation when the blood is flowing in them two views are held, between which the facts at present known do not permit a decisive judg- ment to be made. One theory is that the vessels actively prevent coagulation by constantly absorbing from the blood some substance, as for example the fibrin ferment, which may be supposed constantly to develop, and the presence of which is a necessary condition for the formation of fibrin. The other view is that the blood-vessels are passive and completely neutral. They simply do not excite those changes in the blood constituents which give rise to the COMPOSITION OF THE BLOOD. 57 formation of fibrinoplastin or the ferment, while foreign bodies in contact with the blood do excite these changes and so cause coagulation. Whatever the part which the blood-yessels play, it is only exhibited when their inner surfaces are healthy and unin- jured. If this lining be ruptured or diseased the blood clots. Accordingly, after death, Avhen post-mortem changes have affected the blood-vessels, the blood clots in them; but often very slowly, since the vessels only gradually alter. If the Body be left in one position after death, the clots furmedinthe heart have often a marked buffy coat, because the corpuscles have had a long time to sink in the plasma before coagulation occurred. In medico-legal cases it is thus sometimes possible to say what was the position of a corpse for some hours after death, although it has been subsequently moved. The lymph clots like the blood, but not so firmly; since it contains no red corpuscles, the clot formed is of course colorless. Composition of the Blood. The average specific gravity of human blood is 1055. It has an alkaline reaction, which becomes less marked as coagulation occurs. About one half of its mass consists of moist corpuscles and the remainder of plasma. Exposed in a vacuum, 100 volumes of blood yield about 60 of gas consisting of a mixture of oxygen, carbon dioxide, and nitrogen. Chemistry of the Serum. The blood plasma cannot well be examined as to its elieniical constituents, since it clots under manipulation. The scrum is, however, essentially blood plasma minus fibrin, and from an analysis of it we can draw conclusions as to the plasma. In 100 parts of serum there are about 90 parts of water, 8.5 of proteids, and 1.5 of fats, salts, and other less-known solid bodies. Of the proteids present the most abtindant is serum albumin, which agrees with egg albtimin in coagulating when heated: so that serum when boiled sets into an opaque white mass, just as the white of an Qgg does. Chemically, serum albu- min differs from egg albumin in being coagulated by ether; and physiologically, in the fact that although present in such large quantities in the blood, it does not pass through 68 THE HUMAN BODY. tlie kidneys, whereas t-^^g albiiiniu when injected into the blood-vessels of an animal is rapidly excreted by those organs. In health the fats are only present in the serum in small (piantity except after a meal at which fatty sub- stances have been eaten ; serum obtained from the blood of an animal soon after such a meal is often milky in appear- ance from the fats present, instead of being perfectly color- less or i)alc yellow, and transparent as it is after fasting. The salts dissolved in the serum are mainly sodium chloride and carbonate; but small quantities of sodium, calcium, and magnesium ])h<)sp]i;itos arc also ]iresent. Chemistry of the Red Corpuscles. In these in the fresh moist state there are in 100 parts, b% of water and 44 of solids. Of the solids about one per cent is salts, chiefly potassium phosphate and chloride. The remaining organic solids contain, in IdO parts, 90 of haemoglobin and about 8 of other proteids; the residue consists of less well-known bodies. Chemistry of the White Corpuscles. These yield be- sides much water, several proteids, some fats, glycogen (see Chap. XXVIIL), and salts; and smaller quantities of other bodies. The predominant salts, like those of the red corpuscles, are j^otassium phosphates. Variations in the Composition of the Blood. Hygienic Jh'Duirl-s. The above statements refer only to the average comjoosition of the healthy blood, and to its better known constituents. From what was said in the last chapter it is clear that the blood flowing from any organ will have lost or gained, or gained some things and lost others, when compared with the blood which entered it. But the losses and gains in particular parts of the Body are in such small amoimt as, with the exception of the blood gases, to elude analysis fi)r the most part: and the blood from all parts being mixed up in the heart, they balance one another and produce a tolerably constant average. In health, however, the specific gravity of the blood may vary from 1045 to 1075; the red eorjniscles also are present in greater propor- tion to the plasma after a meal than before it. Ilealthy sleep in projjer amount also increases the proportion of red BLOOD CORPUSCLES. 59 corpuscles, and want of it diminishes tlieir number as may be recognized in the pallid aspect of a person ^vho has lost several nights' rest. Fresh air and plenty of it has the same effect. The proportion of these corpuscles has a great import- ance since, as we shall subsequently see, they serve to carry oxygen, which is necessary for the performance of its func- tions, all over the Body. Aiicemia is a diseased condition characterized by pallor due to deficiency of red blood cor- puscles, and accompanied by languor and listlessness. It is not unfrequent in young girls on the verge of womanhood, and in persons overworked and confined within doors. In such cases the best remedies are oj^en-air exercise and good food. Summary. Practically the composition of the blood may be thus stated: It consists of (1) plasma, consisting mainly of water containing in solution serum albumin, sodium salts, smaller amounts of those of other metals, and extractives of which the most constant are urea, kreatin, and grape sugar; (2) red corpuscles, containing rather more than half their weight of water, the remainder being main- ly haemoglobin, other proteids, and potash salts; (3) white corpuscles, consisting of water, various proteids, glycogen, and potash salts; (4) gases, partly dissolved in the plasma or combined with its sodium salts, and (oxygen) partly combined with the haemoglobin of the red corpuscles. Quantity of Blood. The total amount of blood in tlie Body is difficult of accurate determination. It is, how- ever, about ^ of the whole weight of the Body, so tlie (|uantity in a man weighing 75 kilos (165 lbs.) is about 5.8 kilos (12.7 lbs.). Of this at any given moment about one fourth would be found in the heart and big blood-vessels; and equal quantities in the capillaries of the liver, and in those of the muscles which move the skeleton; while the remaining fourth is distributed among the remaining parts of the Body. The Origin and Fate of the Blood Corpuscles. Tlie white blood corpuscles vary so rapidly and frequently in number in the blood that they must be constantly in pro- 60 TEE HUMAN BOD T. cess of alteration or removal, and formation; their number is largely increased by taking food, even more than that of the red, so that their i)roportion to tlic red rises, from 1 io 1000 during fasting, to 1 to 250 or 300 after a meal. They no doubt multiply to a certain extent by division while circulating in the blood, but the majority come from liie lymi)hatic glands and similar structures (see Chap. XXII.) found in many parts of the Body, which con- tain many cells like pale blood corpuscles, and often in l)rocess of division. From tliese organs the corpuscles en- ter the lymph-vessels and are carried on into the blood. From the capillary blood-vessels many again migrate, and it is probable that these emigi'ahts take part frequently in the repair or regeneration of injured tissues. Being un- differentiated and specialized to no line of work they are ready to take up any that comes to hand, and may be com- pared to the young men in a community who have not yet selected an occupation and are on the lookout for an open- ing. On the other hand there seems little doubt that a great many Avhite corpuscles give rise to red ones, and this is perhaps to be regarded as their special function. The corpuscles of nearly all invertebrate animals are colorless only, although the blood plasma of some contains haemo- globin in solution. Amphioxus, the lowest undoubted vertebrate animal (see Zoology), also possesses only colorless corpuscles in its blood. But higher and more complex ani- mals need more oxygen, and as blood plasma dissolves very little of that gas, they develop in addition the hgemo- globin-containing corpuscles which pick it up in the gills or lungs and carry it to all parts of the Body, leaving it where wanted (see Chap. XXV. ). In cold-blooded vertebrates the red corpuscles are not nearly so many in proportion as in the warm-blooded, Avhich use far more oxygen. The older view was that the mammalian red corpuscle repre- sented the nucleus of one of the white, in which hemoglo- bin had l>cen formed and from about which the rest of the corpuscle had disappeared. This, however, does not seem to be the case; but the pale corpuscle develops or forms I haemoglobin in its cell protoplasm, and flattens and as- LYMPH. 61 sumes the form of a red corpuscle, while its nucleus disap- pears. Occasional transitional forms between the pale and the red corpuscle are seen in blood when examined with the microscope; and if blood be put fresh on a cold slide and examined in a cold room these transitional forms are more numerous, since at ordinary temperatures they very rapidly break down and fall to pieces when blood is drawn. How long an individual red corpuscle lasts is not known, nor with certainty how or when it disappears. There is, however, some reason to believe that a great many are de- stroyed in the spleen (see Chap. XXII.). Chemistry of Lymph. Lymph is a colorless fluid when pure, feebly alkaline, and with a specific gravity of about 1045. It maybe described as blood minus its red corpuscles and considera])ly diluted, but of course in various parts of the Body it will contain minute quantities of substances derived from neighboring tissues. It contains a considera- ble quantity of carbon dioxide gas which it gives up in a vacuum, but no oxygen, since any of that gas which passes into it by diffusion from the blood is immediately picked up by the living tissues among which it flows. CHAPTER V I. THE SKELETON. Exoskeleton and Endoskeleton. The skeleton of an aniiiKil includes all its liard protecting or supporting parts, and is met with in two main forms in the animal kingdom. First as an exoskeleton develo23ed in connection with either the superficial or deeper layer of the skin, and represented by the shell of a clam, the scales of fishes, the horny plates of a turtle, the bony plates of an armadillo, and the feathers of birds. In man the exoskeleton is but slightly developed, but it is represented by the hairs, nails, and teeth; for al- though the latter lie within the mouth, the study of devel- opment shows that they are developed from an offshoot of the skin which grows in and lines the mouth long before birth. Hard parts formed from structures deeper than the skin constitute the endoskeleton, which in man is highly developed and consists of a great many bones and cartilages or gristles, the bones forming the mass of the hard frame- work of the Body, Avhile the cartilages finish it off at vari- ous parts. This framework is what is commonly meant by the skeleton, and it primarily supports the softer parts and is also arranged so as to surround cavities in which delicate organs, as the brain, heart, or spinal cord, may lie with safety. The skeleton thus formed, however, is completed and supplemented by another made of the connective tissue, which not only, in the shape of tough bands or ligaments, ties the bones and cartilages together, but also in various forms pervades the whole Body as a sort of subsidiary skeleton running through all the soft organs, forming net- works of fibres around their other constituents; so that it nuikes, as it were, a microscopic skeleton for the individu.tl modified cells of which the Body is so largely composed. AXIAL SKELETON. 63 and also forms partitions between tlie muscles, cases for sncli organs as the liver and kidneys, and sheaths around the blood-vessels. The bony and cartilaginous framework with its ligaments might be called the skeleton of the organs of the Body, and- this finer supporting mcshwork tlie skeleton of the tissues. Besides forming a support in the substance of various organs, the connective tissue is also often laid down as a sort of packing material in the crevices between them; and so widely is it distributed everywhere from the skin outside to the lining of the alimentary canal inside, that if some solvent could be employed which would corrode away all the rest and leave only this tissue, a very perfect model of the whole Body would be left; something like a " skeleton leaf," but far more minute in its tracery. The Bony Skeleton (Fig. 13). If the hard framework of the Body were joined together like the joints and beams of a house, tlie whole mass would be rigid; its parts could not move with relation to one another, and we would be un- able to raise a hand to the mouth or put one foot before another. To allow of mobility the bony skeleton is made of many separate pieces which are joined together, the points of union being called articulations, and at many places the bones entering into an articulation are movably hinged together, forming what are known as joints. The total number of bones in the Body is more than two hundred in the adult; and the number in children is still greater, for various bones which are distinct in the child (and remain distinct throughout life in many lower animals) grow together so as to form one bone in the full-grown man. The adult bony skeleton may be described as con- sisting of an axial skeleton found in the head, neck, and trunk; and an appendicular sheleton, consisting of the bones in the limbs and in the arches (w and s, Fig. 13) by which these are carried and attached to the trunk. Axial Skeleton. The axial skeleton consists primarily of the vertettral column or spine, a side view of which is represented in Fig. 14. The up^ier part of this column is composed of twenty-four se])arate bones, each of which is a vertebra. At the posterior part of the trunk, beneath the 64 THE HUMAN BODY. '%^ Fig. 13.- The bony and cartilaginous skeleton. Fig. 14.— Side view of the spinal column. VERTEBRA. 65 movable vertebrae, comes the sacrum (;S' 1), made up of five vertebrae, which in the adult grow together to form one bone, and below the sacrum is the coccyx {Co 1-4), consisting of four very small tail vertebrae, Avhich in ad- vanced life also unite to form one bone. On the top of the vertebral column is borne the skull, made up of two parts, viz., a great box above which in- closes the brain and is called the cranium, and a large number of bones on the ventral side of this which form the skeleton of the face. Attached by ligaments to the under side of the cranium is the liyoid bone, to which the root of the tongue is fixed. Of the twenty -four separate vertebrae of the adult the seven nearest the skull (Fig. 14, C 1-7) lie in the neck and are known as the cervical vertebrce. These are fol- lowed by twelve others which have ribs attached to them (see Fig. 13) and lie at the back of the chest; they are the dorsal vertebrce (2)1-12). The ribs (Fig. 35*) are slender curved bones attached by their dorsal ends, called their heads, to the dorsal vertebra? and running thence round the sides of the chest. In the ventral median line of the lat- ter is the breast-bone or siernum {d, Fig 13), Each rib near its sternal end ceases to be bony and is composed of cartilage. These parts — skull, hyoid bone, vertebral column, ribs, and sternum — constitute the axial skeleton, and Ave have now to consider its parts more in detail. The Dorsal Vertebrae. If a single vertebra, say the eleventh from the skull, be examined carefully it will be found to consist of the following parts (Figs. 15 and 16): First a bony mass, C, rounded on the sides and flattened on each end where it is turned towards the vertebrjB above aud below it. This stout bony cylinder is the "body" or centrum of the vertebra, and the series of vertebral bodies (Fig. 14) forms in the trunk that bony partition betAveen the dorsal and ventral cavities of the body spoken of in Chapter I. To the dorsal side of the body is attached an tirch — the neural arcli. A, which Avith the back of the body incloses a space. Fv, the neural ring. In the tube formed *p773. 66 THE nUMAN BODY. by the rings of the successive vertebras lies the spinal cord. Projecting from the dorsal side of the neural arch is a long bony bar, Fs, i\\Q sjnnous process: and the projections of these processes from the various vertebras can be felt through the skin all down the middle of the back. Hence the name of spinal coliDiin often given to the whole back-bone. Six other i)rocesses arise from the arcli of the vertebra: two l)roject forwards, i.e. towards the head; these, Pas, are the anterior articular processes and have a smooth surface, covered with cartilage on their dorsal sides. A pair of sim- ^t Paa Fig. 15. Fig. 16. Fig. 13.— a dorsal vertebra seen from behind, i.e. the end turned from the head. Fig. 16.— Two dorsal vertebrsB viewed from the left side, and in their natural relative positions. C, the body; A. neural arch; Fi\ the neural ring; Ps. spi- nous process; Pas, anterior articular process ; Pai, posterior articular process; Pi, transverse process; Ft, facet for articulation with the tubercle of a rib; Pes, Fci, articular surfaces on the centrum for articulation with a rib. ilav posterior a?'ticular processes, Pai, runs back from the neural arch, and these have smooth surfaces on their ven- tral aspects. In the natural position of the vertebra, the smooth surfaces of its anterior' articular processes fit upon the posterior articular processes of the vertebra next in front, forming a joint, and the two processes are united by ligaments. Similarly its posterior articular processes form joints (Fig. 10) Avith the anterior articular processes of the vertebra next behind. SEGMENTATION OF SKELETON. 67 The remaining processes are the transverse, Pt, which rnn outwards and a little dorsally. Eueli of these has a smooth articular surface, Ft, near its outer end. On the •"body" are seen two articular surfaces on each side: one, Fes, at its anterior, the other, Fri, at its poste- rior end, and both close to the attachment of the neural arch. Each of these surfaces formed with corresponding areas on the vertebrse in front and behind a pit into which the end of a rib fitted and the rib attached in this way to the anterior part of the " body" also fitted on, a little way from its dorsal end, to the articular surface at the end of the transverse process. The Segments of the Axial Skeleton. If a dorsal verte- bra, say the first (Fig. 17), be detached with the pair of ribs, Cv, belonging to it and the bit of the sternum, S, to which these ribs are fixed ventrally, we would find a bony partition formed by the body of the vertebra, lying betweeij tv/o arches which surround cavities. The dorsal cavity inclosed by the " body" and " neural ^ ,„ ^. •^ ^ . . ,, Fig. 17.— Diagrammatic representation arch contained originally of a segment of the axial skeleton. F, a J, . , . , ? .„, vertebra; C, C'l', ribs articulating above part Ot the SJJlUal cord, i lie with the body and transverse process of ,1 J i_ J.1 the vertebra; S, the breast-bone. The other ring, made up by the lighter-shaded part between 5f and C is body of the vertebra dor- '^"^ "'^ •^^'•"^^^"• sally, the sternum ventrally, and the ribs on the sides, sur- rounds the chest cavity with its contents. All of these parts together form a typical segment of the axial skeleton, which, however, only attains this completeness in the thoracic region of the trunk. In the skull it is greatly modified; and in the neck and the lower part of the trunk the ribs are either absent or very small, appearing only as processes of the vertebrae; and the sternal portion is wanting altogether. Nevertheless we may regard the whole axial skeleton as made up of a series of such segments placed one in front of 68 THE HUMAN BODY. the other, but having ditlerent portions of the complete segment much niodiliod or rudiuieutarv, or even altogether wanting in some regions. Parts which in this sort of way reuUy correspond to one another though they differ in de- tail, wliich are so to speak different varieties of one thing, are said in anatomical language to he hoinolocjnus to one an- other; and when they succeed one another in a row, as the trunk segments do, the homology is spoken of as serial. The Cervical Vertebrae. In the cervical region of tlie vertebral column tlie bodies of the vertebra? are smaller than in the dorsal, but the arches are larger; the spinous processes are short and often bifid and the transverse pro- cesses api)ear perforated by a canal, the vertebral foramen. The bony bar bounding this aperture on the ventral side, however, is in reality a very small rib which has grown into continuity with the body and true transverse process of the vertebra, al- though separate in very early life: the transverse process proper bounds the vertebral foramen dorsally. In this latter during life runs an artery, which ultimately enters the skull cavity. The Atlas and Axis. The first and second cervical ver- tebra differ considerably from the rest. The first, or atla>! (Fig. 19), which carries the head, has a very small body. Aa, and a large neural ring. This ring is subdivided by a cord, or the transverse ligament, L, into a dorsal moiety in which the spinal cord lies and a ventral into which the bony process D projects. This is the odontoid process, and arises from the front of the axis or second cervical vertebra (Fig. 20). Around titis peg the atlas rotates when the head is turned from side to side, carrying the skull (which ar- ticulates with the large hollow surfaces Fas) with it. The odontoid process really re])resents a large })iece of the body of the atlas which in early life separates from its own vertebra and grows on the axis. Fig. 18.— a cervical vertebra. Frt, vertebrai foramen ; Pai, anterior artic- ular process. SACRUM. 69 The Lumbar Vertebrae (Fig. 21) are the largest of all the movable vertebrae and have no ribs attached to them. Their spines are short and stout and lie in a more horizontal A a Fas Fig. 19. Fio. 20. Fig. 19.— The atlas. Fig. 20.— The axis. Aa, body of atlas; D, odontoid pro- cess; Fas, facet on front of atlas with which the skull articulates; and in Fig. 20, anterior articular surface of axis; £, transverse ligament; Frt, vertebral fora- men. plane than those ot the vertebrae in front. The articular and transverse processes are also short and stout. The Sacrum, which is represented along with the last lumbar vertebra in Fig. 22, consists in the adult of a single bone; but cross-ridges on its ventral surface indicate the Fig. 21.— a lumbar vertebra seen from the left side. Ps, spinous process; Pat. anterior articular process: Pai, posterior articular process. limits of the five separate vertebrae of which it is composed in childhood. It is somewhat triangular in form, its base being directed upv^ards and articulating with the under 70 THE HUMAN BODY surfiice of the body of lla- lit'th luinbai' vertebra. On its sides are large surfaces to which the arrli benring the lower Pas Fsa Fig. 22. — The last lumbar vertebra and the sacrum seen from the ventral side. Fsa. anterior sacral foramina. limbs is attached (see Fig. 13). Its yentral surface is con- cave and smooth and presents four pairs of anterior sacral foramina, Fsa, which communicate "with the neural canal. Its dorsal surface, convex and roughened, has fonr similar pairs of pos- terior sacral foramina. The coccyx (Fig, 23) calls for no special remark. The four bones which grow togeth- er, or anhylose, to form it represent only the bodies of vertebrae, and even that imperfectly. It is in reality a short tail, although not visi- ble as such from the exterior. The Spinal Column as a Whole. The vertebral column Fig. 23.— The coccyx. SPIJ^^E AS A WHOLE. 71 is ill a man of average height about twenty-eight inches long. Viewed from one side (Fig. 14) it presents four cur- vatures ; one with the convexity forwards in the cervical region is followed, in the dorsal, by a curve with its concav- ity towards the chest. In the lumbar region the curve has again its convexity turned ventrally, while in the sacral and coccygeal regions the reverse is the case. These curvatures give the whole column a good deal of springiness- such as would be absent were it a straight rod, and this is farther secured by the presence of compressible elastic pads, the intervertebral dii^ks, made up of cartilage and connective tissue, which lie between the bodies of those vertebra? which are not ankylosed together, and fill up completely the empty spaces left between the bodies of the vertebriB in Fig. 14. By means of these pads, moreover, a certain amount of movement is allowed between each pair of ver- tebra* ; and so the spinal column can be bent to consider- able extent in any direction ; while the movement between any two vertebrre is so limited that no sharp bend can take place at any one point, such as might tear or injure other- wise the spinal cord contained in the neural canal. The amount of movement permitted is greatest in the cervical region. In the case of the movable vertebras, where the arch Joins the body on each side, it is somewhat narrowed; this narrowed stalk being known as the pedicle [li, Fig. 16), while the broader remaining portion of the arch is its lam- ina. Between the pedicles of two contiguous vertebrae there are in this way left apertures, the intervertebral holes which form a series on each side of the vertebral column, and one of which, Fi, is shown between the two dorsal vcrtebrtB in Fig. 16. Through these foramina nerves run out from the spinal cord to various regions of the Body. The sacral foramina, anterior and posterior, are the repre- sentatives of these apertures, but modified in arrangement, on account of the fusion of the arches and bodies of the vertebrjB between which they lie. Sternum. The sternum or hreast-hone (Fig. 24 and d, Fig. 13) is wider from side to side than dorso-ventrally. It 72 THE HUMAN BODY. Icl Is y/^'yr'jf^^ consists in the adult of three pieces, and seen from the ven tral side has somewhat the form of a dagger. The piece M nearest the head is called the handle or manuhrivyn, and presents anteriorily a notch, Icl, on each side, with which the collar-bone articulates {u, Fig. 13); on each side are two other notches, Ic 1 and Ic 2, to wiiich the sternal ends of the first and second ribs are attached. The middle piece, (7, of the sternum is called the body; it completes the notch for the second rib and has on its sides others, Ic 3-7, for the third, fourth, fifth, sixth, and seventh ribs. The last piece of the sternum, P, is called the ensiform or xiphoid process; it is composed of cartilage, and has no ribs attached to it. The Ribs (Fig. 25). There are twelve pairs of ribs, each being a slender curved bone attached dorsally to the body and transverse process of a vertebra in the manner already mentioned, and con- tinued ventrally by a costal cartilage. In the case of the anterior seven pairs, the costal cartilages are attached direct- ly to the sides of the breast-bone; the next three cartilages are each attached to the cartilage of the preceding rib, Fig. 24.-The sternum while the Cartilages of the tenth and Kanubrium'^^^^^^^ twelfth rlbs are quite unattached ven- ijF^^^i^: trally, so these are called the free or \ i-;j-.notches for the -floaUnq ribs. The convexity of each nb cartilages. . . • curved rib is turned outwards so as to give roundness to the sides of the chest and increase its cavity, and each slopes downwards from its vertebral at- tachment, so that its sternal end is considerably lower than its dorsal. The Skull (Fig. 26) consists of twenty-two bones in the adult, of which eight, forming the cranium, are ar- ranged so as to inclose the brain-case and protect the auditory organ, while the remaining fourteen support THE SKULL. 73 the face and surround the mouth, the nose, and the eye- sockets. Fig. 25.— The ribs of the left side, with the dorsal and two lumbar vertebrae, the rib cartilages and the sternum. Cranium. The cranium is a box with a thick floor and thinner walls and roof. Its floor or lase represents in the head (as is depicted diagrammatically in Fig. 2) that par- 74 TUE HUMAN BODY. tition between the dorsal and ventral cavities which in the trunk is made np of the bodies of the vertebrae. In very early life it presents in the middle line a series of four bones, the hasi-occipital, basi-sphenoid, presjjhenoid, and Tsp Fio. 26 — \ side MOAN of flu skull O, occipital bone; T, temporal; Pr, parie- tal; F, fitjiital, ubic symphysis in the ventral median Ime. the liumerus. At the elbow the humerus is succeeded by two bones, the radius and tilna, c and h, which lie side by side, the radius being on the thumb side. At the distal ends of these bones come eight small ones, closely packed PELVIC GIRDLE. 79 and forming the wrist, or carpus. Then come five cylindri- cal bones which can be felt through the soft parts in the palm of the hand; one for the thumb, and one for each of the fingers. These are the metacarpal hones, and are dis- tinguished as first, second, third, and so on, the first being that of the thumb. In the thumb itself arc two bones, and in each finger three, arranged in rows one after the other; these bones are all called plialanges. The Pelvic Girdle (Fig. 29). This consists of a large bone, the on innaminatum, Oc, on each side, which is firml}^ fixed dorsally to the sacrum and meets its fellow in the middle ventral line. In the child each os innominatum consists of three bones, viz., the ilium, the ischium, and pubis. Where these three bones meet and finally ankylose there is a deep socket, the acetabulum, into which the head of the thigh-bone fits (see Fig. 13). Between the pubic and ischial bones is the largest foramen in the whole skeleton, known as the doorlike or thyroid foramen. The pubic bone lies above and the ischial below it. The ilium forms the upper expanded portion of the os innominatum to which the line drawn from Oc in Fig. 29 points. The Hind Limb. In this there are thirty bones, as in the fore limb, but not quite similarly arranged; there being one less at the ankle than in the wrist, and one at the knee not present at the elbow-joint. The thigh-bone or femur [a, Fig. 31) is the largest bone in the body and extends from the hip to the knee-joint. It presents above a large rounded Jiead. which fits into the acetabulum and, below, it is also enlarged and presents smooth surfaces which meet the bones of the leg. These latter are two in number, known as the tibia, c, or shin-bone, a^nd fibula, d; the tibia being on the great-toe side. In front of the knee-joint is the knee-cap, or patella, b. At the distal end of the leg-bones comes the foot, con- sisting of tarsus, metatarsus, and phalanges. The tarsus, which answers to the carpus of the fore limb, is made up of seven irregular bones, the largest being the heel-bone, or calcaneum, h. The metatarsus consists of five bones lying &-*de by side, and each carries a toe at its distal end. In 80 THE HUMAN BODY. the great toe (or hallux) there are two phalanges, in each of the others three, arranged as in the fingers, but smaller. Comparison of the Anterior and Posterior Limbs. It Fig. 30. Fig. 31. Fig. 30.— The bones of the arm. a, humerus; 6. ulna; c. radius; d, the carpus: e, tlie fifth metacarpal; /. the three phalanges of the fifth digit (little finger), g, the phalanges of the pollex (thumb). Fig. 31.— Bones of the leg. a, femur; b. patella; c. tibia; d, fibula; 7i, calca- neum; e, remaining tarsal bones: /, metatarsal bones; g, phalanges. is clear that the skeletons of the arm and leg correspond pretty closely to one another. Both are in fact quite alike in very early life, and their differences at birth depend upon HOMOLOGIES OF THE LIMBS. 81 their diverging in different ways as tliey develop from their primitive simplicity; as both may be regarded as modifications of the same original structure, they are ho- o P Fio. 3-3. —The skeleton of the arm and leg. H. the humerus: Cd, its articular head which fits into the glenoid fossa of the scapula; £/. the ulna; iJ, the ra- dius; O, the olecranon; Fe, the femur; P, the patella; Fi, the fibula: T. the tibia. mologous. The pelvic girdle clearly corresponds generally to the pectoral arch, tlie tibia and fibula to the radius and S2 THE HUMAN BODY. ulna ; the five metatarsal bones to the five metacarpal, and the phalanges of the toes to those of the thumb and fingers. On the other hand, there is in the arm no separate bone at the elbow-joint corresponding to the patella at the knee, but the ulna bears above a bony process, the olecranon {0, Fig. 32), which at first is a separate bone and is the rej)- resentative of the patella. There are in the carpus eight bones and in the tarsus but seven. The astragalus of the tarsus {Ta, Fig. 35) represents two bones Avhich, however, />**. Fig. 33.— Diagram showing the relation of the pectoral arch to the axial skel- eton. have grown together. The elbow-joint bends forwards and the knee-joint backwards. Comparing the limbs as a whole, greater differences come to light, differences which are mainly correlated with the different uses of the two limbs. The arms, serving as ( prehensile organs, have all their parts as movable as is consistent with the requisite Fig. »i.-Diagram showing the attach- strength, while the lower Ston ^^^ '''''''''' ^"""^ **' ^^^ ^^'^^ limbs, having to bear the whole weight of the Body, require to liave their parts much more firmly knit together. Accordingly we find the shoulder girdle, represented red in the diagram (Fig. 33), only directly attached to the axial skeleton by the union of the inner ends of the clavicles with the sternum, and capable of considerable independent move- ment, as seen, for instance, in " shrugging the shoulders." The pelvic arch, on the contrary, is firmly and immovably HAND AND FOOT COMPARED. 83 fixed to the sides of the sacrum. The socket of the scapula, into which the head of the humerus fits, is very shallow and allows a far greater range of movement than is per- mitted by the deeper socket on the pelvis, into which the head of the femur fits. Further, if we hold the right humerus tightly in the left hand and do not allow it to move, we can still move the forearm bones so as to turn the palm of the hand either up or down: no such move- ment is possible between the tibia and fibula. Finally, in the foot the bones are much less movable than in the hand, and are arranged so as make a springy arch (Fig. 35) which bears behind on the calcaneum, Ca, and in front on the distal ends of the tarsal bones, Os] and over the crown of the arch, at Ta, is the surface with which the leg-bones \ SfW 3 Fig. 35 —The bones of the foot. Co, calcaneum, ov os calcis; Ta, articular surface for tibia on the astragalus; Cb. tlie cuboid bone. articulate and on which the weight of the Body bears in standing. The toes, too, are far less movable than the fingers, and this diiference is especially well marked between the great toe and the thumb. The latter can be made to meet each of the finger-tips and so the hand can seize and manipulate very small objects, while this power of opposing the first digit to the rest is nearly absent in the foot of civilized man. In children, however, who have never worn boots, and in savages, the great toe is far more movable, though it never forms as complete a thumb as in many apes, which use their feet, as well as their hands, for prehension. By practice, however, our own toes can be made much more 84 THE HUMAN BODY. mobile than they usually are, so that the foot can to a certain extent replace the hand; as has been illustrated in the case of persons born without hands who have learned to write and paint Avith their toes. Peculiarities of the Human Skeleton. These are largely connected with the division of labor between the fore and hind limbs referred to above, which is carried farther in man than in any other creature. Even the highest apes frequently use their fore limbs in locomotion and their hind limbs in prehension, and we find accordingly that anatomically they present less differentiation of hand and foot. The other more important characteristics of the human skeleton are correlated for the most part with the maintenance of the erect posture, which is more complete and habitual in man than in the animals most closely allied to him anatomically. These peculiarities, however, only appear fully in the adult. In the infant the head is pro- portionately larger, the curves of the vertebral column are nearly absent, and the posterior limbs are relatively very short. In all these points the infant approaches more closely to the ape, and they all combine to give the centre of gravity of the Body a comparatively very high |)osition and to render the maintenance of the erect posture difficult and insecure. The subsequent great relative length of the posterior limbs, which grow disproportionately fast in childhood as compared with the anterior, makes progression on them more rapid by giving a longer stride and at the same time makes it almost impossible to go on "all fours" except by crawling on the hands and knees. In other Pri- mates this disproportion between the anterior and posterior limbs does not occur to nearly the same extent. In man the skull is nearly balanced on the top of the vertebral column, the occipital condyles which articulate with the atlas being about its middle (Fig. 27*), so that but little effort is needed to keep the head erect. In four-foot- ed beasts, on the contrary, the skull is carried on the front end of the horizontal vertebral column and needs special ligaments to sustain it. For instance, in the ox and sheep there is a great elastic cord running from the cervical ver- CHARACTERISTICS OF HUMAN SKELETON. 85 tebrai to tlie back of the skull and helping to hold up the head. Even in the highest apes the skull does not balance on the top of the spinal column; the face part is much heavier than the back, while in man the face parts are rela- tively smaller and the cranium larger, so that the two nearly equipoise. To keep the head erect and look things straight in the face, "like a man," is for the apes far more fatiguing, and so they cannot long maintain that position. The human spinal column, gradually widening from the neck to the sacrum, is well fitted to sustain the weight of tlie head, upper limbs, etc., carried by it, and its curvatures, which are peculiarly human, give it considerable elasticity combined with strength. The pelvis, to the sides of which the lower limbs are attached, is proportionately very broad in man, so that the balance can be more readily maintained during lateral bending of the trunk. The arched instep and broad sole of the human foot are also very character- istic. The majority of four-footed beasts, as horses, walk on the tips of their toes and fingers, and those animals, as bears and apes, which like man place the tarsus also on the ground, or are plcuitigrade in technical language, have a much less marked arch there. The vaulted human tar- sus, composed of a number of small bones, each of which can glide a little over its neighbors, but none of which can move much, is admirally calculated to break any jar which might be transmitted to the spinal column by the contact of the sole with the ground at each step. A well-arched instep is therefore justifiably considered a beauty ; it makes progression easier, and by its springiness gives elasticity to the step. In London flat-footed candidates for appoint- ment as policemen are rejected, as they cannot stand the fatigue of walking the daily "beat." CHAPTER VII. THE STRUCTURE AND COMPOSITION OF BONE. JOINTS. Gross Structure of the Bones. The bones of the Body have all a similar structure and composition, but on ac- count of differences in shape they are divided by anato- mists into the following groups: (1) Long tones, more or less cylindrical in form, like the bones of the thigh and arm, leg and forearm, metacarpus, metatarsus, fingers and toes. (2) Tabular hones, in the form of exjjanded plates, like the bones on the roof and sides of the skull, and the shoulder-blades. (3) Short hones ; rounded or angular in form and not much greater in one diameter than in another, like the bones of the tarsus and carpus. (4) Ir- regular hones, including all which do not get well into any of the preceding groups,and commonly lying in the middle line of the Body and divisible into two similar halves, as the vertebriE. Living bones have a bluish-white color and possess considerable elasticity, which is best seen in long slender bones such as the ribs. To get a general idea of the structure of a bone, we may se- lect the humerus. Externally in the fresh state it is covered by a dense white fibrous membrane very closely adherent to it and containing a good many small blood-vessels. This membrane is called the periosteum; on its under side new osseous tissue is formed while the bone is still growing, and all through life it is concerned in maintaining the nu- trition of the bone, which dies if it is stripped off. The periosteum covers the whole surface of the bone except its ends in the elbow and shoulder joints; the surfaces there which come in contact with other bones and glide over them STRUCTURE OF A BONE. 87 in the movements of the joint have no periosteum, but are covered by a thin layer of gristle, known as the articu- lar cartilage. Very early in the development of the Body the bone in fact was repre- sented entirely by cartilage; but afterwards nearly all this was replaced by osseous tis- sue, leaving only a thin carti- laginous layer at the ends. The bone itself, Fig. 36, consists of a central nearly cylindrical portion or shaft, extending between the dot- ted lines X and z in the fig- ure, and two enlarged artic- idar extremities. On the upper articular ex- tremity is the rounded sur- face, Cp, which enters into the shoulder -joint, fitting against the glenoid cavity of the scapula; and on the low- er are the similar surfaces, Cpl and Tr, which articulate with the radius and ulna re- spectively. Besides carry- ing the articular sui'faces, each extremity presents sev- eral j)rominences. On the upper are those marked Tmj and Tm (the greater and smaller trochanters), which give attachment to muscles; and similar eminences, the external and internal con- dyles, El and Em, are seen on the lower end. Besides Fig, from the front, text. •igi] For description, seo these, several bony ridges 88 THE HUMAN BODY. Hiid rough patches on tlie shiiit indicate places to which muscles of the arm were fixed. Internal Structure. If the Ijone he divided longitudi- nally, it will he seen that its shaft is hollow, the space being known as the medullary rarity, and in the fresh bone filled with marrow. Fig. 37 represents a longitudi- nal section of the femur, which in this re- spect is quite similar to the humerus. The marrow cavity does not reach into the articular extremities, but there the bone has a loose spongy texture, except a thiii layer on the surface. In the shaft, on the other hand, the outer compact layer is much the thickest, the spongy or cancel- lated hone forming only a thin stratum im- mediately around the medullary cavity. To the naked eye the cancellated bone appears made up of a trellis-work of thin bony 2)lates whicli intersect in all direc- tions and surround cavities rather larger than the head of an ordinary pin; the compact bone, on the contrary, ajoiieaj's to have no cavities in it until it is exam- ined with a magnifying glass. In the spaces of the spongy portion lies, during life, a substance known as the red marroic, which is quite different from the yellow fatty marrow lying in the central cavity of tile shaft. Microscopic Structure of Bone. Tlic microscope shows that the compact bone contains cavities and only differs from the spongy portion in the fact that these are much smaller and the hard true bony plates surrounding them much more nu- merous in })roportiou than in the spongy parts. If a thin transverse section of the shaft of the hu- merus be examined (Fig. 38) with a microscope magnifying Fig. 37— a longi- l • idinal section or the femur, d, dense bone ; a. cancellated bone; b, medullary cavity. HISTOLOGY OF BONE. 89 twenty diameters, it will be seen that numerous openings exist III! over the compact parts of the section and gradu- ally become larger as this passes into the cancellated part, next the medullary cavity. These openings are the cross- sections of tubes known as the Haversian canals, which ramify all through the bone, running mainly in the direc- FiG. 38. — A, a tranverse section of the ulna, natural size ; showing the medul- lary cavity. B, the more deeply shaded part of A magnified twenty diameters. tion of its long axis, but united by numerous cross or ob- lique branches as seen in the longitudinal section (Fig. 39). The outermost ones open on the surface of the bone be- neath the periosteum, and in the living bone blood-vessels run from this through' the Haversian canals and convey 90 THE HUMAN BODY. materials for its growth and nourishment. Tlie average diameter of the Haversian canals is 0.05 mm. {-^^ of an inch). Around each Haversian canal lies a set of plates, or lameU(v, of hard bony substance (see the transverse section Fig. 38), each canal with its lamellae forming an Haver- sian system: and the whole bone is made up of a number of such systems, with the addition of a few lamellse lying in the corners between them, and a certain number which run around the whole bone on its outer surface. In the spongy parts of the bone the Haversian canals are very Inrge and the intervening lamellse few in number. Between the lamella? lie small cavities, the lamnm, each of which is lenticular in form, somewhat like the space which would be inclosed by two ^^T^l^^'lli^'*^^^^!^ ^.j^|-g|-^_g|j^gggg joined by their $P r^tii^^ (^^9 ^^'Ses- From the lacuna? many €^i^^^ ViFI^^^^M ^^^^'^'^^^y ^^^® branching ca- S/MM ^^ l|^^^Ki~^\^S ^^''^^^' ^^■'^ ccmaliculi, radiate f T/^V?/ 'll^^ilsV'^ i^^'^^ penetrate the bony la- ' ^ i> ^if^ ^f,PAik\% melL^ in all directions. The ^ L ^^ ^ it J^ i^jt i} i' innermost canaliculi of each system oiDcn mto the central Fig. 39— a thin longitudinal sec tV • i i jt c tion of bone, magnified about 350 Haversian Can al; and those oi diameters, aa. Haversian canals. • i - , various lacunge intercommu- nicating, these fine tubes form a set of passages through Avhich liquid which has transuded from the blood-vessels in the Haversian canals can ooze all through the bone. The lacunae and canaliculi are well seen in Fig. 39. In the living bone a granular nucleated cell lies in each lacuna. These cells, or hone coiyusdes, are the remnants of those which built up the bone, the hard parts of the lat- ter being really an intercellular substance or skeleton formed around and by these cells, much in the same way as a calcareous skeleton is formed around each Foraminifer (see Zoology) by the activity of its protoplasm. By the co-operation of all the bone corpuscles, and the union of their skeletons, the whole bone is built up. In other bones we find the same general arrangement of COMPOSITION OF BONE. 91 the parts, ;in outer dense layer and an inner spongy por- tion. In the flat and irreguhir bones there is no medullary cavity, and the whole centre is filled up with cancellated tissue with red marrow in its spaces. For example, in the thin bones roofing in the skull we find an outer and inner hard layer of compact bone known as the outer and iimer tables respectively, the inner especially being very dense. Between the two tables lies the spongy bone, red in color to. the naked eye from the marrow within it, and called the diploe. The interior of the vertebra also is entirely occupied by spongy bone. Everywhere, except where a bone joins some other part of the skeleton, it is covered with the periosteum Chemical Composition of Bone. Apart from the bone corpuscles and the soft contents of the Haversian canals and of the spaces of the cancellated bone, the bony sub- stance i)roper. as found in the lamellae, is composed of earthy and organic portions intimately combined, so that the smallest distinguishable portion of bone contains both. The earthy matters form about two thirds of the total weight of a dried bone, and may be removed by soaking the bone in dilute hydrochloric acid. The organic portion left after this treatment constitutes a flexible mass, retain- ing perfectly the form of the original bone. By long boil- ing, especially under pressure at a higher temperature than that at whieli water boils when exposed freely to the air, the oi'ganic portion of the bone is nearly entirely con- verted into gelatine which dissolves in the hot water. Much of the gelatine of commerce is prepared in this manner by boiling the bones of slaughtered animals, and even well- picked bones may be used to form a good thick soup if boiled under pressure in a Papin's digester; much nutri- tious matter being, in the common modes of domestic cooking, thrown away in the bones. The earthy salts of bone may be obtained free from or- ganic matter by calcining a bone in a clear fire, which burns away the organic matter. The residue forms a white very brittle mass, retaining perfectly the shape and structural details of the original bone. It consists mainly of normal 92 THE HUMAN BODY. calcium phos])liate, or boiif earth (Ctia, 2PO4); but there is also present a considerable proportion of calcium carbonate (CaCOa) and smaller ([uantitics of other salts. Hygiene of the Bony Skeleton. In early life the bones are less rigid, from the fact that the earthy matters then present in them bear a less proportion to the softer organic parts. Hence the bones of an aged person are more brittle and easily broken than those of a child. The bones of a young child are in fact tolerably flexible and will be dis- torted by any continued strain; therefore children should never be kept sitting for hours, in school or elsewhere, on a bench which is so high that the feet are not supported. If this be insisted u])on (for no child will continue it volunta- rily) the thigh-bones will almost certainly be bent over the edge of the seat by the weight of the legs and feet, and a permanent distortion may be produced. For the same reason it is important that a child be made to sit straight in writing, to avoid the risk of producing a lateral curvature of the spinal column. The facility with which the bones may be moulded by prolonged pressure in early life is well seen in the distortion of the feet of Chinese ladies, pro- duced by kee23ing them in tight shoes; and in the extraor- dinary forms which some races of man produce in their skulls, by tying boaixls on the heads of the children. Throughout the whole of life, moreover, the bones re- main among the most easily modified parts of the Body; although Judging from the fact that dead bones are the most permanent parts of fossil animals we might be in- clined to think otherwise. The living bone, however, is constantly undergoing changes under the influence of the protoplasmic cells imbedded in it. and in the living Body is constantly being absorbed and r(>constructed. The expe- rience of physicians shows that any continued pressure, such as that of a tumor, will cause the absorption and dis- appearance of bone almost quicker than that of any other tissue; and the same is true of any other continued pres- sure. Moreover, during life the bones ;irc eminently plas- tic; under abnormal pressures they are found to quickly assume abnormal shapes, being absorbed and disappearing ARTICULATION'S. 93 at points where the pressure is most powerful, and increas- ing at other points; tight hxcingmay in this way produce a permanent distortion of the ribs. When a bone is fractured a surgeon should be called in as soon as possible, for once inflammation has been sot up and the parts have become swollen it is much more diffi- cult to place the broken ends of the bone togetlier in their proper position than before this has occurred. Once the bones are replaced they must be held in position by splints or bandages, or the muscles attached to them will soon dis- place them again. With rest, in young and healtliy per- sons complete union will commonly occur in three or four weeks; but in old persons the process of cure is slower and is apt to be imperfect. Articulations. The bones of the skeleton are joined to- gether in very various ways; sometimes so as to admit of no movement at all between them; in other cases so as to per- mit only a limited range or variety of movement; and else- where so as to allow of very free movement in many direc- tions. All kinds of unions between bones are called articu- lations. Of articulations permitting no movements, those which unite the majority of the cranial bones afford a good exam- ple. Except the lower jaw, and certain tiny bones inside the temporal bone belonging to the organ of hearing, all the skull-bones are immovably joined together. This unioji in the case of most occurs by means of toothed edges which lit into one another and form jagged lines of union known as sutures. Some of these can be well seen in Fig. 2G* between 1 he frontal and parietal bones {coronal suture) and between the parietal and occipital bones {lamMoidal suture); while another lies along the middle line in the top of the crown between the two parietal bones, and is known as the sagit- tal suture. In new-born children where the sagittal meets the coronal and lambdoidal sutures there are large spaces not yet covered in by the neighboring bones, which subse- quently extend over them. These oj^enings are known as fontanelles. At them a pulsation can often ])e felt syn- chronous with each beat of the heart, which, driving more 94 THE HUMAN BOBT. blood into the braiu, distends it and causes it to push out the skin where bone is absent. Another good example of an articulation admitting of no movement, is that between the rough surface on the sides of the sacrum and the in- nominate bone. We find good examples of the second class of articula- tions— those admitting of a slight amount of movement — in the vertebral column. Between every pair of vertebra^ from the second cervical to the sacrum is an elastic pad, the intervertebral dish, which adheres by its surfaces to the bodies of the vertebrae between which it lies, and only per- mits so much movement between them as can be brought about by its own compression or stretching. When the back-bone is curved to the right, for instance, each of the intervertebral disks is compressed on its right side and stretched a little on its left, and this combination of move- ments, each individually but slight, gives considerable flexibility to the spinal column as a whole. Joints. Articulations permitting of movement by the gliding of one bone over another, are known as joints and all have the same fundamental structure, although the amount of movement permitted in different joints is very different. Hip-Joint. We may take this as a good example of a true joint permitting a great amount and variety of move- ment. On the OS innominatum is the cavity of the aceta- hnhim (Fig. 40), which is lined inside by a thin layer of articular curtilage which has an extremely smooth surface. The bony cup is also deepened a little by a cartilaginous rim. The proximal end of the femur consists of a nearly spherical smooth head, borne on a somewhat narrower neck, and fitting into the acetabulum. This head also is covered witli articular cartilage; and it rolls in the acetabulum like a ball in a socket. To keep the bones together and limit the amount of movement, ligaments pass from one to the other. These are com])osed of white fibrous connective tissue (Chap. VIII.) and are extremely jDliable but quite inextensible and very strong and tough. One is the cap- sular ligament, which forms a sort of loose bag all round SYJS'^OrTAL JOINTS. 95 the Joint, and another is the rouiid ligament, which passes from the acetabnhim to tlie head of the femur. Should the latter rotate above a certain extent in its socket, the round ligament and one side of the capsular ligament are put on the stretch, and any further movement which might dislocate the femur (that is remove the head from its socket) is checked. Covering the inside of the capsular ligament and the outside of the round ligament is a layer of flat cells, which are continued in a modified form over Fia. 40.— Section thi-ough the hip- joint. the articular cartilages and form the synovial memlrane. This, which thus forms the lining of the joint, is always moistened in health by a small quantity of glairy synovial fluid, something like the white of a raw cg^ in consis- tency, and playing the part of the oil with which the con- tiguous moving surfaces in a machine are moistened; it makes all run smoothly with very little friction. In the natural state of the parts, the head of the femur and the bottom and sides of the acetabulum lie in close contact, the two synovial membranes rubbing together. 96 THE HUMAN BODY. This contact is not nuiintiiiued by the ligaments, which are too loose and serve only to check excessive movement, bnt by the numerous stout muscles which pass from the thigh to the trunk and bind the two firmly togethei-. Moreover, the atmospheric pressure exerted on the surface of the Body and transmitted through the soft parts to the outside of the air-tight joint helps also to keep the parts in contact. If all the muscles and ligaments around the joint be cut away it is still found in the dead Body that the head of the femur will be kept in its socket by this pressure, and so firmly as to bear the weight of the whole limb without dislocation, just as the pressure of the air will enable a boy's " sucker" to lift a tolerably heavy stone. Ball-and-socket Joints. Such a joint as Ifhat at the hip is called a ball-and-socket joint and allows of more free movement than any other. Through movements occurring in it the thigh can be jiexed, or bent so that the knee ap- proaches the chest; or extended, that is moved in the oppo- site direction. It can be abducted, so that the knee moves outwards; and adducted, or moved back towards the other knee again. The limb can also by movements at the hip- joint be circumducted, that is made to describe a cone of which the base is at the foot and the apex at the hip. Fi- nally rotation can occur in the joint, so that with knee and foot joints held rigid the toes can be turned in or out, to a certain extent, by a rolling around of the femur in its socket. At the junction of the humerus with the scapula is another ball-and-socket joint permitting all the above movements to even a greater extent. This greater range of motion at the shoulder-joint depends mainly on tlie shallowness of the glenoid cavity as compared with the acetabulum and upon the absence of any ligament answer- ing to the round ligament of the hip-joint. Another ball- and-socket joint exists between the carpus and the meta- carpal bono of the thumb; and others with the same variety, but a much less range, of movement between each of the remaining metacarpal bones and the proximal phalanx of the finger which articulates with it. Hinge-Joints. Another form of synovial joint is known FORMS OF JOINTS. 97 as a hinge-joint. In it the articulating bony surfaces are of such shape as to permit of movement, to and fro, in one plane only, like a door on its hinges. The joints between the phalanges of the fingers are good exami)]es of hinge- joints. If no movement be allowed where the finger joins the palm of the hand it will be found that each can be bent and straightened at its own two joints, but not moved in any other way. The knee is also a hinge-joint, as is the articulation between the lower jaw and the base of the skull which allows us to open and close our mouths. The latter is, however, not a perfect hinge-joint, since it per- mits of a small amount of lateral movement such as occurs in chewing, and also of a gliding movement by which the lower jaw can be thrust forward so as to protrude the chin and bring the lower row of teeth outside the upper. Pivot- Joints. In this form one bone rotates around an- other which remains stationary. We have a good example of it between the first and second cervical vertebrge. The first cervical vertebra or atlas (Fig. 19*) has a very small body and a very large arch, and its neural canal is subdi- vided by a transverse ligament {L, Fig. 19) into a dorsal and a ventral portion; in the latter the spinal cord lies. The second vertebra or axis (Fig. 20) has arising from its body the stout bony peg, D, called the odontoid process. This projects into the ventral portion of the space sur- rounded by the atlas, and, kept in place there by the trans- verse ligament, forms a pivot around which the atlas, car- rying the skull with it, rotates when we turn the head from side to side. The joints on each side between the atlas and the skull are hinge-joints and permit only the movements of nodding and raising the head. When the head is leaned over to one side, the cervical part of the spinal column is bent. Another kind of pivot-joint is seen in the forearm. If the limb be held straight out, with the palm up and the elbow resting on the table, so that the slioulder-joint be kept steady while the hand is rotated until its back is turned upwards, it will be found that the radius has partly rolled round the ulna. When the palm is upwards and '■ *P.69. 98 TEE HUMAN BODY. the thumb outwards, the lower end of the radius can be felt on the outer side of the forearm just above the wrist, and if this be done while the hand is turned over, it will l)e easily discerned that during the movement this end of the radius, carrying the hand with it, travels around the lower end of the nlna so as to get to its inner side. The relative position of the bones when the palm is upwards is shown at A in Fig. 41, and when the palm is down at B. The former position is known as supination; the lat- ter as pronation. The elbow end of the humerus (Fig, 36*) bears a large articular surface: on the inner two thirds, of this, Tr, the ulna fits, and the ridges R- ■V and grooves of both bones inter- locking form a hinge-joint, al- lowing only of bending or straightening the forearm on the arm. The radius fits on the rounded outer third, Cpl, and forms there a ball-and-socket joint at which the movement takes place when the hand is turned from the supine to the prone position; the ulna forming a fixed bar around which the lower end of the radius is moved. Gliding Joints. These per- mit as a rule but little move- ment: examples are found be- tween the closely packed bones of the tarsus (Fig. 35 f) and carpus, which slide a little over one another when subjected to pressure. Hygiene of the Joints. AYlien a bone is displaced or dislocated the ligaments around the joint are more or less torn and other soft parts injured. This soon leads to in- flammation and swelling which make not only the recogni- tion of the injury but, after diaguo.sis, the replacement of the bone, or the reduction of the dislocation, difficult. Fig. 41. — A, arm in supination; B, arm in pronation; if, humerus; B, radius; 17, ulna. » P. 87. fP. 83. DISLOCATIONS AND SPRAINS. 99 Moreover the muscles attached to it constantly pull on the displaced bone and drag it still farther out of place; so that it is of great importance that a dislocation be reduced as soon as possible. In most cases this can only be attempted with safety by one who knows the form of the bones, and possesses sufficient anatomical knowledge to recognize the direction of the displacement. No injury to a joint should be neglected. Inflammation once started there is often diffi- cult to check and runs on, in a chronic way, until the syno- vial surfaces are destroyed, and the two bones perhaps grow together, rendering the joint permanently stiff. A sprained joint should get immediate and complete rest, for weeks if necessary, and if there be much swelling, or con- tinued pain, medical advice should be obtained. An im- properly cared-for sprain is the cause of many a useless ankle or knee. CHAPTER VIII. CARTILAGE AND CONNECTIVE TISSUE. Temporary and Permanent Cartilages. In earl}^ life a great m;iiiy parts of the supporting framework of the Body, which afterwards become boue, consist of cartilage. Such for example is the case with all the vertebrae, and with the bones of the limbs. In these cartilages subse- quently the process known as ossification takes place, by which a great portion of the original cartilaginous model is removed and replaced by true osseous tissue. Often, how- ever, some of the primitive cartilage is left throughout the whole of life at the ends of the bones in joints where it forms the articular cartilages; and in various other places still larger masses remain, such as the costal cartilages, those in the external ears forming their framework, others finishing the skeleton of the nose which is only incom- pletely bony, and many in internal parts of the Body, as the cartilage of "Adam's apple," which can be felt in the front of the neck, and a number of rings around the windpipe serving to keep it open. These persistent masses are known as the permanent, the others as the temporary cartilages. In old age many so-called permanent cartilages become calcified — that is, hardened and made unyielding l^y deposits of lime salts in them — without assuming the histological character of bone, and this calcification of the permanent cartilages is one chief cause of the want of pliability and suppleness of the frame in advanced life. Hyaline Cartilage. In its purest form cartilage is flexi- ble and elastic, of a pale bluish-white color when alive and seen in large masses, and cuts readily with a knife. In thin pieces it is quite transparent. Everywhere except in the CARTILAGE. 101 Joints it is invested by a tongh adherent membrane, the perichondrium, which resembles in structure and function the periosteum of the bones. When boiled for a long time in water such cartilages yield a solution of chondrin, which differs from gelatin in minor points but agrees with it in the fact that its hot watery solutions " set" or gelatinize on cooling. When a thin slice of hyaline cartilage is examined with a microscope it is found (Fig. 42) to consist of gran- ular nucleated cells, often collected into groups of two, four, or more, scattered through a homogeneous or faintly granular ground substance or mrt/rhr. Essentially, cartilage resembles bone, being made up of protoplasmic cells and a proportionately large amount of non-protoplasmic mtercel- FiG. 43. -Hyaline cartilage, c, a cell with several nuclei, and about to divide ; h. a cell which has divided into two; a, a group of four cells such as would re- sult from a repetition of the division of b The granules of the matrix are represented much too coarse and conspicuous. lular substance, the cells being the more actively living part and the matrix their product. Examples of this hya- line variety (so called from its glassy transj)arent appear- ance) are found in all the temporary cartilages, and in the costal and articular among the permanent. They rarely contain blood-vessels except at those points where a temporary cartilage is being removed and replaced by bone; then blood-vessels run in from the loerichondrium and form loops in the matrix, around which it is absorbed and bony tissue deposited. In consequence of the usual absence of blood-vessels the nutritive processes and ex- changes of material must be small and slow in cartilage, as 102 TEE ETJMAN BODY. might indeed be expected from the passive and merely mechanical rdle which this tissue plays. Hyaline cartilage is the type, or most characteristically developed form, of a tissue found with modifications else- where in the Body. One of its other modifications is the so-called cellular cartilage, which consists of the cells with hardly any matrix, only just enough to form a thin capsule around each. This form is that with which all the car- tilages commence, the hyaline variety being built up by the increase of the cell capsules and their fusion to form the matrix. It persists throughout life in the thin cartilaginous plate of a mouse's external ear. Other varieties of cartilage are really mixtures of true cartilage and connective tissues, and will be considered after the latter. The Connective Tissues. These complete the skeleton, marked out in its coarser features by the bones and car- tilages, and constitute the final group of the supporting tissues. They occur in all forms from broad membranes and stout cords to the finest threads, forming networks around the other ultimate histological elements of various organs. In addition to subsidiary forms, three main varie- ties of this tissue are readily distinguishable, viz., areolar, white fibrous, and yelloiv elastic. Each consists of fibres and cells, the fibres being of two kinds, mixed in nearly equal proportions in the areolar variety, while one kind predominates in one and another in the second of the re- maining chief forms. Areolar Connective Tissue. This exists abundantly be- neath the skin, where it forms a loose flocculent layer, somewhat like raw cotton in appearance but not so white. It is on account of its loose texture that the skin can every- where be moved, more or less, to and fro over the subja- cent parts to which it is united by this tissue. Areolar tissue consists of innumerable bands and cords interlacing, in all directions, and can be greatly distended by blowing air in at any point, from whence it travels widely through the intercommunicating meshes. In dropsy of the legs or feet the cavities of this tissue are distended with lymph. From beneath the skin the areolar tissue extends all through the CONNECTIVE TISSUES. 103 Body between the muscles and around the blood-vessels and nerves; and still finer layers of it enter into these and other organs and unite their various parts together. It consti- tutes in fact a soft packing material which fills up the holes and corners of the Body, as for instance around the blood-vessels and between the muscles in Fig. 4. Microscopic Structure of Areolar Tissue. When exam- ined wdth the microscope areolar tissue is seen to consist of nucleated cells imbedded in a ground substance which is permeated by fibres. The fibres everywhere form the i)re- dominant feature of the tissue (the homogeneous matrix and the cells being inconspicuous) and are of two very dif- ferent kinds. In a strict sense indeed the areolar tissue ought to be considered as a mixture of two tissues, one corresj)onding to each variety of fibres in it. It is charac- terized as a distinct individual by its loose texture and by the fact that the two forms of fibres are present in tolera- bly equal quantities. In many places a tissue containing the same histological elements as the areolar tissue is found in the form of dense membranes, as for example periosteum and perichondrium. White Fibrous Tissue. One of the varieties of fibres pervading the matrix of areolar tissue exists almost un- mixed with the other kind in the cords or tendons which unite muscles to the bones. This form, known as the white fibrous connective tissue, also exists fairly pure in the ligaments around most joints. Physically it is very flexi- ble but extremely tough and inextensible, so that it w^ill readily bend in any direction but is very hard to break r when fresh it has an ojiaijue white color. White fibrous tissue (Fig, 43) consists of a matrix, con- taining cavities in which cells lie and pervaded by bundles of extremely fine fibres. These fibres lie in each bundle tolerably parallel to one another in a wavy course (Fig, 43) and never branch or unite. Their diameter varies from 0.0005 to 0.001 millimeter (go Jo¥ to 2yJ-„T o^ an inch). Chemically this tissue is characterized by the fact that its fibres swell up and become indistinguishable when t'Qated with dilute acetic acid, and by the fact that it 104 THE HUMAN BODY. yields gelatin when boiled in water. The substance in it, and in tlie bones, which is turned into gelatin by such treatment is known as collagen. Glue is impure gelatin obtained from tendons and ligaments, and calf's-foot jelly, so often recommended to invalids, is a purer form of the same substance obtained by boiling the feet of calves, which contain the tendons of many muscles passing from the k'o; to the foot. Fig. 43. Fig. 43a. Fig. 43.— White fibrous connective tissue, highly magnified. The nucleated corpuscles, seen edgewise and appearing spindle-shaped, are seen here and there on the surface of the bundles of fibres. Fig. 43a.— Yellow elastic tissue, magnified after its fibres have been torn apart. Elastic Tissue. This is almost invariably mixed in some proportion in all specimens of white fibrous tissue, even the purest, such as the tendons of muscles; but in certain places it exists almost alone, as for example in the liga- ments {ligamenta suhjiava) between the arches of the vertebra?, and in the coats of the larger arteries. In quad- rupeds it forms the great ligament already referred to (p. 84), which helps to sustain the head. This tissue, in CONNECTIVE-TISSUE CORPUSCLES. 105 mass, is of a dull yellow color and extremely extensible and elastic; when purest nearly as much so as a piece of india- rubber. Sometimes it appears under the microscope to be made up of delicate membranes, but most often it is in the form of fibres (Fig. •13rt) which are coarser than those of white fibrous tissue and frequently branch and unite. It is unaffected by acetic acid and does not yield gelatin when boiled. Connective-Tissue Corpuscles. The fibres of white fi- brous tissue, wherever it is found, are united into bundles by a structureless ground material known as the cement sub- stance, which also invests each bundle, or skein as we may call it, with a delicate coating. In this ground sub- stance are numerous cavities, branched and flattened in Fig. 44.— Connective-tissue corpuscles. one diameter, and often intercommunicating by their branches. In these cavities lie nucleated masses of proto- plasm (Fig. 41), frequently also branched, known as the connective-tissue coiyuscles. These it is which btiild up the tissue, each cell in the course of development forming around it a quantity of intercellular substance, which sub- sequently becomes fibrillated in great part, the remainder forming the cement. The cells do not quite fill the cavi- ties in which they lie, and these opening into others by their offsets there is formed a set of mintite tl^bes ramify- ing through the connective tissues ; and (since these in tttrn permeate nearly all the Body) pervading all the organs. In these cell cavities and their branches the lymph flows before it enters definite lymphatic vessels, and they are ac- 106 THE HUMAN BODY. cordingly known as lymph canaliculi. In addition to the fixed branched connective-tissue corpuscles lying in the cavi- ties of the ground substance there are often found other cells, when living connective tissue is examined by the micro- scope. These cells much resemble white blood corpuscles, and probably are such which have bored through the walls of the finer blood-vessels. They creep about along the canaliculi by means of their faculty of amoeboid movement, and are known as the ''wandering cells." During in- flammation at any point their number in that region is greatly increased. Subsidiary Varieties of Connective Tissue. In various parts of the Body are connective-tissue structures which have not undergone the typical development, but have de- parted from it in one way or another. The cells having formed a non-fibrillated intercellular substance around them, develoj^ment may go no farther and the mass remain permanently as the jelly-Uhe connective tissue; or, as in the vitreous humor of the eye (Chap. XXXI.), the cells having formed the soft matrix, may disajjpear and leave the latter only. In other cases the intercellular substance disappears and the cells branching, and joining by the ends of their brandies, form a network themselves, nucleated or not at the points answering to the centre of each originally separate cell. This adenoid connective tissue is found widely distributed in the Body especially in connection with the lymphatic system, and forming a supporting- framework for the proper nervous elements in the brain and sj)inal cord. In other cases the cells almost alone con- stitute the tissue, becoming flattened, closely fitted at their edges, and united by a very small amount of cement sub- stance. ]\Iembranes formed in this way lie beneath layers of epithelium in many places and are known as dasement memhrcmes, and the flattened cells which line joints and the serous cavities seem really to be closely apposed flattened connective- tissue corpuscles. Elastic Cartilage, and Fibro-Cartilage. We may now return to cartilages and consider those forms which are made up of more or less true cartilage mixed with more or INTEBABTICULAB CARTILAGES. lor less connective tissue of one kind or another. The carti- lages of the ear and nose and some others have their matrix pervaded by fine branching fibres of yellow elastic tissue, which form networks around the groups of cartilage cells. Such cartilages are pliable and tough and possess also con- siderable extensibility and elasticity. They are known as elastic or, from their color, as yellow cartilages. Elsewhere, especially in the cartilages which lie between the bones in some joints, we find forms which have the matrix pervaded by white fibrous tissue and known as fibro-vartilages. For example the articular cartilage on the end of the lower jaw Fig. 4o. — Section through the joint of the lower jaw showing its interarticular fibro-cartUage, x, with tlie synovial cavity on each side of it. does not C(jme into direct contact with that covering its socket on the skull, but lying between the two in the joint (Fig. 45) is an interarticular fibro-cartilage : similar cartilages exist in the knee-joint; and the intervertebral disks are also made up of this tissue. Both elastic cartilage and fibro-cartihige often shade off insensibly into pure elastic or pure white fibrous connective tissue. Homologies of the Supporting Tissues. Bone, cartilage, and connective tissue all agree in broad structural charac- ters, and in the iises to which they are applied in the Body. In each of them the cells which have built up the tissue, 108 THE HUMAN BODY. ■with rare exceptions, form an inconspicuous part of it in its fully developed state, tlie chief mass of it consisting of intercellular substance. In hyaline cartilages this latter is not fibrillated; but these cartilages pass insensibly in va- rious regions of the Body into elastic or fibro-cartilages, and these latter in turn into elastic or fibrous connective tissue. The lamellse of bone, too, when peeled off a bone softened in acid and examined with a very high magnifying power, are seen to be pervaded by fine fibres. Structurally, there- fore, one can draw no hard and fast line between these tis- sues. The same is true of their chemical composition; bone and white fibrous tissue contain a substance (collagen) which is converted into gelatin when boiled in water; and in old people many cartilages become hardened by the de- posit in their matrix of the same lime salts which give its hardness to bone. Further, the developmental history of all of them is much alike. In very early life each is repre- sented by cells only : these form an intermediate substance, and this subsequently may become fibrillated, or calcified, or both. Finally they all agree in manifesting in health no great physiological activity, their use in the Body depend- ing upon the mechanical properties of their intercellular substance. The close alliance of all three is further shown by the frequency with which they replace one another. All the bones and cartilages of the adult are at first represented only by collections of connective tissue. Before or after birth this is in some cases substituted by bone directly (as in the case of the collar-bone and the bones on the roof of the skull), while in other cases cartilage supplants the con- nective tissue, to be afterwards in many places replaced by bone, while elsewhere it remains throughout life. Moreover in different adult animals we often find the same part bony in one, cartilaginous in a second, and com- posed of connective tissue in a third: so that these tissues not only represent one another at different stages in the life of the same animal Ijut permanently throughout the whole life of different animals. Low in the animal scale HYGIENE OF GBOWING SKELETON. 109 we find tliem all represented merely by cells with struc- tureless intercellular substance: a little higher in the scale the latter becomes fibrillated and forms distinct connective tissue. In the highest MoUusks (see Zoology), as the cuttle- fishes, this is partly replaced by cartilage, and the same is true of the lowest fishes; while in some other fishes and the remaining Vertebrates, we find more or less bone ap- pearing in place of the original connective tissue or carti- lage. From the similarity of their modes of development and fundamental structure, the transitional forms which exist between them, and the frequency with which they rej^lace one another, histologists class all three (bone, cartilage, and connective tissue) together as homologous tissues and re- gard them as differentiations of the same original struc- ture. Hygienic Remarks. Since in the new-born infant many parts which will ultimately become bone, consist only of car- tilage, the young child requires food which shall contain a large proportion of the lime salts which are used in building up bone. JSTature provides this in the milk, which is rich in such salts (see Chap. XX. ), and no other food can thoroughly replace it. If the mother's health be such as to render it unwise for her to nurse her infant, the best substitute, apart from a wet-nurse, will be cow's milk diluted with one fourth its volume of water. Arrowroot, corn-flour, and other starchy foods will not do alone, since they are all defi- cient in the required salts, and many infants though given food abundant in quantity are really starved, since their food does not contain the substances requisite for their healthy development. At birth even those bones of a child which are most ossi- fied are often not continuous masses of osseous tissue. In the humerus for example the shaft of the bone is well ossified and so is each end, but between the shafts and eacli of the articular extremities there still remains a cartilad- nous layer, and at those points the bone increases in length, new cartilage being formed and replaced by it. The bone 110 THE HUMAN BODY. increases in thickness by new osseous tissue formed beneath the periosteum. Tlie same tiling is true of tlie bones of the leg. On account of the largely cartilaginous and imperfectly knit state of its bones, it is cruel to encourage a young child to walk beyond its strength, and may lead to "bow-legs" or other j^ermanent distortions. Nevertheless here as else- where in the animal body, moderate exercise promotes the growth of the tissues concerned, and it is nearly as bad to wheel a child about forever in a baby-carriage as to force it to walk beyond its strength. The best rule is to let a healthy child use its limbs when it feels inclined, but not by praise or blame to incite it to efforts which are beyond its age, and so sacrifice its healthy growth to the vanity of parent or nurse. The final knitting together of the bony articular ends with the shaft of many bones takes place only compara- tively late in life, and the age at which it occurs varies miich in different bones. Generally speaking, a layer of cartilage remains between the shaft and the ends of the bone, until the latter has attained its full adult length. To take a few examjjles: the lower articular extremity of the humerus only becomes continuous with the shaft by bony tissue in the sixteenth or seventeenth year of life. The upper articular extremity only joins the shaft by bony con- tiuuity in the twentieth year. The u]iper end of the femur joins the shaft by bone from the seventeeu th to the nine- teenth year, and the lower end during the twentieth. In the tibia the upper extremity and the shaft unite in the twenty-first year, and the lower end and the shaft in the eigiiteenth or nineteenth: while in the fibula the upper end joins the shaft in the twenty-fourth year, and the lower end in the twenty-first. The separate vertebrfe of the sacrum are only united to form one bone in the twenty-fifth year of life; and the ilium, ischium, and pubis unite to form the os iniiominatum about the same period. Up to about twenty-five then the skeleton is not firmly "knit," and is incapable, without risk of injury, of bearing strains which it might afterwards meet with impunity. To let FAT-CELLS. Ill lads of sixteen or seventeen row and take other exercise in plenty is one thing, and a good one; but to allow them to undergo the severe and prolonged strain of training for and rowing a race is quite another, and not devoid of risk. Adipose Tissue. Fatty substances of several kinds ex- ist in considerable quantity in tbe human Body in health, some as minute droplets floating in the bodily liquids or imbedded in various cells, but most in special cells, nearly filled with fat, and collected into masses with supporting and nutritive parts, to form adipose tissue. In fact al- most in every spot where the widely distributed areo- lar tissue is found, there is adipose tissue in greater or less proportion along with it. Considerable quantities ex- ist for example in the subcutaneous areolar tissue, espe- cially in the female sex, giving the figure of the woman its general graceful roundness of contour when compared with that of the male. Large quantities commonly lie in the abdominal cavity around the kidneys; in the eye-sockets, forming a pad for the eyeballs; in the marrow of bones ; around the joints, and so on. Examined with the microscope (Fig. 46) adipose tissue is found to consist of small vesicles from 0.2 mm. to 0.09 mm. (jj^ to jJq- inch) in diameter, clustered together into little masses and bound to one another by connective tissue and blood-ves- ^^^ ^^ _^^^ ^^,j^ ^^^ sels which intertwine around them ; suppoi-tmg connective tis- in this way the little angular masses which are seen in beef suet are formed, each mass be- ing separated by a somewhat coarser partition of areo- lar from its neighbors. The individual fat-cells are round or oval except when closely packed, when they become polygoiud. Each consists of a delicate enve- lope containing oily matter, which in life is liquid at the temperature of the Body. Besides the oily mat- ter, a nucleus is commonly present in each fat-cell: 112 THE HUMAN BODY. and sometimes a thin layer of protoplasm forms a lin- ing to the cell-wall. The oily matter consists of a mixture of palmatin, olein and stearin, which are compounds of palmitic, stearic and oleic acids with glycerine, three molecules of the acid being combined with one of glycerine, with the elimination of water; as for example: o /CisHssO ) r\\ 1 C3H5 ) ^ SfCisHssO) ) /^3 1 QTir\ ^[ Hf^j+ h4^^= C3H5 f^ +'^^^^- stearic acid. Glycerine. Stearin. Water. CHAPTER IX. THE STEUOTURE OF THE MOTOR ORGANS. Motion in Animals and Plants. If one were asked to point out the most distinctive property of liyiiig animals, the answer would probably be, their power of executing sijontane- ous movements. Animals as we commonly know them are rarely at rest, while trees and stones move only when acted upon by external forces, which are in most cases readily re- cognizable. Even at their quietest times some kind of mo- tion is observable in the higher animals. In our own Bod- ies during the deepest sleep the breathing movements and the beat of the heart continue; their cessation is to an on- looker most obvious sign of death. Here however, as else- where in Biology, we find that precise boundaries do not exist; at any rate so far as animals and plants are concerned we cannot draw a hard and fast line between them with reference to the presence or absence of apparently sponta- neous motility. Many a flower closes in the evening to ex- pand again in the morning sun; and in many plants compara- tively rapid and extensive movements can be called forth by a slight touch, which in itself is quite insufficient to produce mechanically that amount of motion in the mass. The Venus's flytrap (Dionaea muscipula) for example has fine hairs on its leaves, and when these are touched by an insect the leaf closes up so as to imprison the animal, which is subsequently digested and absorbed by the leaf. The higher plants it is true have not the power of locomotion, they cannot change their place as the higher animals can; but on the other hand some of the lower animals are perma- nently fixed to one spot; and among the lowest plants many are known which swim about actively through the water in 114 THE HUMAN BODY. which they live. The lowest animals and plants are in fact those which have nndcrgone least- differentiation in their development, and wliich therefore resemble each other in possessing, in a more or less manifest degree, all the funda- mental physiological properties of that simple mass of pro- toplasm which formed the starting point of each individual. With the johysiological division of labor which takes place in the higher forms we find that, speaking broadly, plants especially develop nutritive tissues, while animals are char- acterized by the high development of tissues with motor and irritable properties; so that the jDrej^onderance of these latter is very marked when a comi)lex animal, like a dog or a man, is compared with a complex plant, like a pine or a hickory. The higher animal possesses in addition to greatly developed nutritive tissues (which differ only in detail from those of the plant, and constitute what are therefore often called orfjans of vegetative life), well-devel- oped spontaneous, irritable and contractile tissues, found mainly in the nervous and muscular systems, and forming what have been called the organs of animal life. Since these place the animal in close relationship with the sur- rounding universe, enabling slight external forces to excite it, and it in turn to act upon external objects, they are also often spoken of as orgaiis of relatioti. In man they have a higher develoi)ment on the whole than in any other animal, and give him liis leading place in the animate world, and his power of so largely controlling and directing natural forces for his own good, while the plant can only passively strive to endure and make the best of what happens to it ; it has little or no influence m controlling the haj)- pening. AmcBboid Cells. The simplest motor tissues in the adult Human Body are the amoeboid cells (Fig. 12) already de- scribed, which may be regarded as the slightly modified descendants of the undifferentiated cells which at one time made up the whole Body. In the adult they are not attached to other parts, so that their changes of form only affect themselves and produce no movements in the rest of the Body. Hence with regard to the whole frame they CILIATED CELLS. 115 can hardly be called motoi- tissues, and so are placed in a group by themselves under the name of undifferentiated tissues. Ciliated Cells. As the growing Body develops from its primitive simplicity we find that the cells lining some of the tubes and c:ivities in its interior undergo a very re- markable change, by which each cell differentiates itself into a nutritive, and a highly motile and spontaneous portion. Such cells are found for example lining the ^\ indpipe, and a number are represented in Fig. 47. Each has a conical form, the base of the cone being turned to the cavity of the air-tube, and contains an oval nucleus, with a nucleolus. On the broader free end are a number (about thirty on the average) of extremely fine j)rocesses called cilia. During life these are in constant rapid move- ment, lashing to and fro in the liquid which moistens the interior of the passage; and as the cells are very closely packed, a bit of the inner sur- face of the windpipe examined with a microscope, looks like a field of wheat or barley when the wind blows over it. Each cilium strikes with more force in one direction than in the opposite, and as this di- rection of more powerful stroke is the same for all the cilia on any one surface, the general result is that the liquid in which they move is driven one way. In the case of the windpipe for example it is driven up towards the throat, and the tenacious liquid or mucus which is thus swept along is finally coughed or "hawked" up and got rid of, instead of accumulating in the deej^er air-passages away down in the chest. These cells afford an extremely interesting example of the division of physiological emijloyments. Each proceeds from a cell which was primitively equally motile, automatic, and nutritive in all its parts. But in the fully developed state the nutritive duties have been especially assumed by the conical cell-body, while the automatic and contractile proj)- erties have been condensed, so to speak, in that modified Fig. 47.— Ciliated ceU.s. 116 THE HUMAN BODY. portion of the primitive protoplasmic mass, wliicli forms tlie cilia. These, being supplied with food by the rest of the cell, are raised above the vulgar cares of life and have the opportunity to devote their whole attention to the perform- ance of automatic movements; which are accordingly far more rapid and precise than those executed by the whole cell before any division of kiborhad occurred in it. That the movements depend upon the structure and com- position of the cells themselves, and not upon influences reaching them from the nervous or other tissues, is proved by the fact that they continue for a long time in isolated cells, removed and placed in a liquid, as blood serum, which does not alter their physical constitution. In cold-blooded animals, as turtles, whose constituent tissues frequently retain their individual vitality long after that bond of union has been destroyed which constitutes the life of the whole animal as distinct from the lives of its different tissues, the ciliated cells in the windpipe have been found still at work three weeks after the general death of the animal. The Muscles. These are the main motor organs ; their general appearance is well known to every one in the lean of butcher's meat. While amceboid cells can only move themselves, and (at least in the Human Body) ciliated cells the layer of liquid with which they may happen to be in contact, the majority of the muscles, being fixed to the skeleton, can, by alterations in their form, bring about changes in the form and i:)Osition of nearly all parts of the Body. With the skeleton and joints, they constitute pre- eminently the organs of motion and locomotion, and are governed by the nervous system which regulates their activ- ity. In fact skeleton, muscles, and nervous system are correlated parts: the degree of usefulness of any one of them largely depends upon the more or less complete de- velopment of the others. Man's highly endowed senses and his powers of reflection and reason would be of little use to him, were his muscles less fitted to carry out the dictates of his will or his joints less numerous or mobile. All the muscles are under the control of the nervous system, but all are not governed by it with the co-02)eration of will or VARIETIES OF MUSCLE. 117 consciousness; some moving without our having any direct knowledge of the fact. This is especially the case with cer- tain muscles which are not fixed to the skeleton but sur- round cavities or tubes in the Body, as the blood-vessels and the alimentary canal, and by their movements control the passage of substances through them. The former gi'oup, or skeletal muscles, are also from their microscopic characters known as striped muscles, while the latter, or visceral mus- cles^ are called unstriped or plain muscles. The skeletal muscles being generally more or less subject to the control of the will (as for example those moving the limbs) are frequently spoken of as voluntarij, and the visceral muscles, which change their form independently of the will, as invol- untary. The heart-muscle forms a sort of intermediate link ; it is not directly attached to the skeleton, but forms a hollow bag which drives on the blood contained in it and that quite involuntarily; but in its microscopic structure it resembles the skeletal voluntary muscles. The muscles of respiration might perhaps be cited as another intermediate group. They are striped skeletal muscles and, as we all know, are to a certain extent subject to the will; any one can draw a deep breath when he chooses. But in ordinary quiet breathing we are quite unconscious of their working, and even when attention is turned to them the power of control is limited; no one can voluntarily hold his breath long enough to suffocate himself. As we shall see hereafter, moreover, any one or all of the striped muscles of the Body may be thrown into activity independently of or even against the will, as, to cite no other instances, is seen in the '" fidgets" of nervousness and the irrepressible trembling of extreme terror; so that the names voluntary and involun- tary are not good ones. The functional differences be- tween the two groups depend really more on the nervous connections of each, than ujion any essential difference in the properties of the so-called voluntary or involuntary muscular tissues themselves. The Skeletal Muscles. In its simplest form a skeletal consists of a red soft central part, called the belly, which tapers at each end and there passes into one or more dense 118 THE HUMAN BODY. white cords wliich consist nciirly entirely of white fibrous connective tissue. These terminal cords are called the tendons of the muscle and serve to attach it to parts of the hunj ur cartilaginous skeleton. In Fig. 48 is shuAvn the bice2}s muscle of the arm, which lies in front of the humerus. Its fleshy belly, Bh, is seen to divide above and end there in two tendons, one of which, Bl, is fixed to the scapula, wliile the other joins the tendon of a neighboring muscle (Ihe coraco-hrachial) and is also fixed above to the shoulder- blade. Near the elbow-joint the muscle is continued into a single tendon, B', which is fixed to the radius, but gives an offshoot, B", to the connective-tissue membranes lying around the elbow- joint. The belly of every muscle possesses the power of shorten- ing forcibly under certain conditions. In so doing it pulls upon the tendons, which being composed of inextensible white fibrous tissue transmit the movement to the hard parts to which they are attached, just as a pull at one end of rope may be made to act upon distant objects to which the other end is tied. The tendons are merely passive cords and are sometimes very long, as for instance in the case of the muscles of the fingers, the bellies of many of which lie away in the forearm. If the tendons at each end of a muscle were fixed to the same bone the muscle would clearly be able to produce no movement, unless by bending or breaking the bone; the probable result in such a case would be that the muscle would be torn by its own efforts. In the Body, however, the two ends of a muscle are always attached to two differ- ent pieces of the skeleton, between which more or less movement is permitted, and so when the muscle pulls it alters the relative positions of the parts to which its ten- dons are fixed. In the great majority of cases a true joint lies between the bones on which the muscle can pull, and when the latter contrarfs it produces movement at the joint. Many muscles even pass over two joints and can produce movement at either, as the biceps of the arm which, fixed at one end to the scapula and at the other to the radius, can move the bones at either the shoulder or MUSQLE.s OF ARM. 119 ^-o>i 130 THE HUMAJf BOJjr. elbow joints. Where a muscle passes over an articulation it is nearly always reduced to a narrow tendon; otherwise the bulky bellies lying around the joints would make them extremely clumsy and limit their mobility. Origin and Insertion of Muscles, Almost invariably that part of the skeleton to which one end of a muscle ia fixed is more easily moved than the part on which it pulls by its other tendon. The less movable attachment of a muscle is called its origin, the more movable its insertion. Taking for example the biceps of the arm, we find that when the belly of the muscle contracts and pulls on its upper and lower tendons, it commonly moves only the fore- arm, bending the elbow-joint as shown in Fig. 49. The iia. 40. — The biceps muscle and the arm -bones, to illustrate ho\v, imder ordi- nary circumstances, the elbow joint is flexed when the muscle contracts. shoulder is so much more firm that it serves as a fixed point, and so that end is the origin of the muscle, and the forearm attachment, P, the insertion. It is clear, how- ever, that this distinction in the mobility of the points of fixation of the muscle is only relative for, by changing the conditions, the insertion may become tlie stationary and origin the moved point ; as for instance in going up a rope "hand over hand." In that case the radial end of the muscle is fixed and the shoulder is moved through space by its contraction. Different Forms of Muscles. Many muscles of the Body have the simple typical form of a belly tapering to a FORMS OF MUSCLE. 121 single tendon at each end as a, Fig. 50; but others divide at one end and are called two-headed or bleeps muscles; while some are even three-headed or triceps muscles. On the other hand some muscles have no tendon at all at one end, the belly running right up to the point of attachment; and some have no tendon at either end. In many muscles a tendon runs along one side and the fibres of the belly are attached obliquely to it: such muscles {b, Fig. 50) are coXieA ^^enniform or feath- er-like; or a tendon runs obliquely down the middle of the muscle and has the fibres of the belly fixed obliquely on each side of it (c, Fig. 50), forming a hipennifonn muscle: or even two ten- dons may run down the belly and so form a tripe im if orm muscle. In a few cases a tendon is found in the middle of the terminal teudons; belly as well as at each end of it; such S^^'-.^oukf not/™n muscles are called digastric. A muscle uninterruptedly along '' . Its whole edge as repre- Of this form (Fig. 51) is found in con- seuted in the figure); c, . , T I . T • ^ bipennirorm muscle. nection with the lower jaw. It arises by a tendon attached to the base of the skull; from there its first belly runs downwards and forwards to the neck by the side of the hyoid bone, where it ends in a tendon which passes through a loop serving as a pulley. This is succeeded by a second belly directed upwards towards the chin, where it ends in a tendon inserted into the lower jaw. Eunning along the front of the belly from the pelvis to the chest is a long muscle on each side of the middle line called the rectus ahdominis : it is polygastric, consisting of four bellies sep- arated by short tendons. Many muscles moreover are not rounded but form wide flat masses, as for example the muscle Ss seen on the ventral side of the shoulder-blade in Fig. 48. Gross Structure of a Muscle. However the form of the skeletal muscles and the arrangement of their tendons may vary, the essential structure of all is the same. Each c Fig. 51. — a di- gastric muscle. 123 THE HUMAN BODY. consists of a proper striped muscular tissue. Avliich is its es- sential part, but which is supported by connective tissue, nourished by blood-vessels and lymphatics, and lias its ac- tivity governed by nerves; so that a great variety of things go to form the complete organ. A loose sheath of areolar connective tissue, called the penmysium, envelops each muscle, and from this parti- tions run in and subdivide the belly into hnndhs or fasci- culi which run from tendon to tendon, or for the whole length of the muscle when it has no tendons. Tlie coarse- ness or fineness of butcher's moat depends upon the size of these primary fasciculi, which differs in different muscles of the same animal. These larger fasciculi are subdivided by finer connective-tissue membranes into smaller ones (as shown in Fig. 52, which represents a few pri- mary fasciculi of a muscle and the secondary fasciculi into which these are di- vided), each of which cou- ^ sists of a certain number of Fig. 52.— a small bit ol muscle com- posed of three pnmaiT fasciculi. A, mUSCUkir JlOreS DOUnd to- natural size; B. the same magnified, ii t r. showing the secondary fasciculi of which gether by TCry fine COn- the primary are composed. nectivB tissue and envel- oped in a close network of microscopic blood - vessels. Where a muscle tapers the fibres in the fasciculi become less numerous, and when a tendon is formed disappear altogether, leaving little but the connective tissue. Histology of Muscle. For the present we need only concern ourselves with the muscular fibres. Each of these is from eight to thirty-five millimeters (^ to 1\ inch) long, but only from 0.034 to 0.055 mm. (y|o to j\^ inch) in* diameter in its widest part, and tapering to a point at each end. Hence in long muscles witli terminal tendons, no fibre runs the whole length of a fasciculus, which may be a foot or more long, but the fasciculus is made up of many successive fibres, the narrow end of each fitting in between the ends of those which follow it. In short or penniform PLAIN MUSCULAR TISSUE. 123 muscles, where the fasciculi are short, the fibres may run the whole length of each of the latter. Examined carefully with a good microscope each fibre is seen to possess a very thin homogeneous sheath or envel- ope, called the sarcolermna, within which lies the contrac- tile portion of the fibre, b, which presents a striped appear- ance as if composed of darker and dimmer alternating bands (Fig. 53). During life this substance is very soft or semifluid, but after death it rapidly solidi- fies and death-stiffening, or rigo?- mortis, is produced. Besides the contractile sub- stance a number of oval nuclei, each sur- rounded by a little unmodified protoplasm, lie inside the sarcolemma. The latter is imperforate except at one point where the central portion (or axis cylinder, see Chap. XII.) of a nerve-fibre penetrates it, and ends in an expansion or end plate which is in immediate contact with the striated sub- stance. The larger blood-vessels of a muscle run in the coarser partitions of the connective tissue, and the finer ones lie close around each fibre but entirely outside its sarcolem- ma. Structure of the Unstriped Muscles. Of these the muscular coat of the stomach (Fig. 54) is a good example. They have no definite tendons but form expanded mem- branes surrounding cavities, so that they have no definite origin or insertion. Like the skeletal muscles they consist of proper contractile element.-^, with accessory connective tissue, blood-vessels, and nerves. Their fibres, however, have a very different microscopic structure. They present no striation but are made up of elongated cells (Fig. 55), bound together by a small quan- tity of cementing material. Each cell is flattened in one plane and tapers off at each end; in its interior lies an elongated nucleus with one or two nucleoli. These cells FiG.53.— Asmall piece of muscular fibre highly mag- nified. At a the fibre has been crushedandtwist- ed so as to tear its contents while the tougher sar- colemma, e 1 s e- where so closely applied to the rest as to be invisible, remains iintorn and conspicuous. 124 THE HUMAN BODY. Fig. 54.— The muscular coat of the stomach. have the power of shortening in the direction of their long axis, and so of diminishing the capacity of the cavities in the walls of which they lie. Cardiac Muscular Tissue. This con- sists of flattened branched cells which unite to form a network, in the interstices of which blood capillaries and nerve-fibres run. The cells present transverse striations, but not so distinct as those of the skeletal mus- cles, and are said to have no sarcolemma. The Chemistry of Muscular Tissue. The chemical structure of the living muscular fibre is unknown, since all the methods of chemical analysis at present discovered de- compose and kill it. It contains 75 per cent of water; and, among other inorganic constituents, phosphates and chlorides of potassium, sodium, and magnesium. When at rest a living muscle is feebly alkaline, but after hard work, or when dying, this reaction is reversed through the formation of sarco- lactic acid (CaHeO^). Muscles contain small quantities of grape sugar and glycogen, and of organic Fig. 55.-Unstriped muscle-cells. GHEMISTRT OF MUSCLE. 125 nitrogenous crystalline compounds, especially kreatin (C4H9N3OJ). As in the case of all other physiologically ac- tive tissues, however, the main post mortem constituents of the muscular fibres are proteid substances, and it is proba- ble that like protoplasm itself (p. 34) the essential con- tractile part of the tissue consists of a complex body con- taining proteid, carbohydrate and fatty residues; and that during muscular work this is broken up yielding proteids, carbon dioxide, sarcolactic acid, with probably other things; for this hypothetical substance, which has never yet been isolated, the name inogeii has been proposed. The main proteid substance obtained from muscles is that known as myosin, which is prepared rs follows. Perfectly fresh and still living muscles are cut out from a frog which has just been killed by destruction of its brain and spinal cord, a proceeding which entirely deprives the animal of consciousness and the power of using its muscles, but leaves these latter unaltered and alive for some time. The excised muscles are thrown into a vessel cooled below 0° C. by a freezing mixture and so are frozen hard before any great chemical change has had time to occur in them. The solidified muscles are then cut up into thin slices, the bits thrown on a cooled filter and let gradually warm up to tlie freezing point of water, with the addition of some ice-cold 0.5 per cent solution of common salt. Gradually a small quantity of a tenacious liquid filters through which is at first alkaline to test-paper but soon sets into a jelly and becomes acid. The coagulation and the acidity are due to the breaking up of the muscle substance into the myosin and other bodies referred to above. At first the jelly is transparent, but soon the myosin becomes opaque and shrinks just like blood fibrin, squeezing out a quantity of muscle serum, and remaining itself as the muscle clot. Myosin thus prepared is insoluble in water and in saturated solution of common salt, but soluble in five or ten per cent watery solutions of the latter substance. When boiled it IS turned into coagulated proteid (p. 11) and becomes in- soluble in dilute acids, in which the original ijiyosin was soluble, being at the same time, however, converted into 1 2 6 THE E UMAN BOD T. another proteid, called synto7iin, wliicli "vvas formerly con- sidered to be tlie original proteid yielded by the muscles. Syntonin is insoluble in water but soluble in dilute acids and alkalies and its solutions are not coagulated by boiliug. Beef Tea and Liebig's Extract. From the above stated facts it is clear that when a muscle is boiled in water its myosin is coagulated and left behind in the meat: even if cooking be commenced by soaking in cold water, the myosin still remains as it is insoluble in cold water. Beef tea as ordinarily made, then, contains little but the flavoring matters and salts of the meat and some gelatin, the former making it deceptively taste as if it were a strong solution of the whole meat, whereas it contains but little of the most nutritious proteid portions, which in an insipid shrunken form are left when the liquid is strained off. Various pro- posals have, been made with the object of avoiding this and getting a really nutritive beef tea; as for example chopping the raw meat fine and soaking it in strong brine for some hours to dissolve out the myosin; or extracting it with dilute acids which turn the myosin into syntonin and dissolve it, at the same time rendering it non-coagulable by heat when subsequently boiled. Such methods, however, make unpalatable compounds which invalids, as a rule, will not take. Beef tea is a slight stimulant but hardly a food and cannot be relied upon to keep up a jDatient's strength for any length of time. Liebig's extract of meat is essentially a very strong beef tea; containing much of the flavoring substances of the meat, nearly all its salts and the crystal- line nitrogenous bodies, such as kreatin, which exist in muscle, but hardly any of its really nutritive joarts. From its stimulating eflects it is often useful to persons in feeble health, but other food should be given with it. It may also be used on account of its flavor to add to the " stock" of soup and for similar purposes; but the erroneousness of the common belief that it is a highly nutritious food can- not be too strongly insisted upon. Under the name of liqiiid extracts of meat other substances have been prej^ared LIQUID EXTRACT OF MEAT. 137 by subjecting meat to chemical processes in which it un- dergoes cliauges smiilar to those experienced in digestion: the myosin is thus rendered sohible in water and uncoagu- lable by heat, and such extracts if properly prepared are highly nutritious. The flavor may be imi^roved by adding a little ot Liebio-'s extract if desired. CHAPTER X. THE PROPERTIES OF MUSCULAR TISSUE. Contractility. The characteristic physiological property of muscular tissue, and that for which it is employed in the Body, is the faculty possessed by its fibres of shortening forcibly under certain circumstances. The direction in which this shortenmg occurs is always that of the long axis of the fibre, m both plain and striped muscles, and it is accompanied by an almost equivalent thickening in other diameters, so that when a muscle contracts it does not shrivel up or diminish its bulk in any appreciable way; it simply changes its form. When a muscle contracts it also becomes harder and more rigid, especially if it has to over- come any resistance. This and the change of form can be well felt by placing the fingers of one hand over the biceps muscle lying in front of the humerus of the other arm. When the muscle is contracted so as to bend the elbow it can be felt to swell out and harden as it shortens. Every schoolboy knows that when he appeals to another to " feel his muscle" he contracts the latter so as to make it thicker and apparently more massive as well as harder. In statues the prominences on the surface, indicating the muscles be- neath the skin, are made very conspicuous when violent effort is represented, so as to indicate that they are in vigor- ous action. In a muscular fibre we find no longer the slow, irregular, and indefinite changes of form seen m the undif- ferentiated cells of early development; this is replaced by a precise, rapid, and definite change of form m one di- rection only. Muscular tissue represents a group of cells in the bodily community, which have taken up the one spe- cial duty of executing changes of form, and m proportion MUSCULAR IRRITABILITY. 129 as they have fewer other things to do, they do that better. This contractility of the muscular fibres may be best con- ceived by considering each to possess two natural shapes; one, the state of rest, in which the fibres are long and nar- row; and the other the state of activity, in which they are shorter and thicker: under certain conditions the fibres tend to pass, with considerable force, from their resting to their active form, and in so doing they move parts attached to their tendons. When the state of activity passes off the fibres suffer themselves to be passively extended again by any force pulling upon them, and they so regain their rest- ing shape; and since in the living Body other parts are nearly invariably put upon the stretch when any given muscle contracts, these by their elasticity serve to pull the latter back again to its primitive shape. No muscular fibre is known to have the power of actively expanding after it has contracted: in the active state it forcibly resists ex- tension, but once the contraction is over it suffers itself readily to be pulled out to its resting form. Irritability. With that modification of the primitive protoplasm of an amoeboid cell which gives rise to a mus- cular fibre, with its great contractility, there goes a loss of other properties. All trace of spontaneity seems to disap- pear; muscles are not automatic like native protoplasm or ciliated cells; they remain at rest unless directly excited from without. The amount of external change required to excite the living muscular fibre at any moment is, how- ever, very small; in other words, it is highly irritable, a very little thing being sufficient to call forth a powerful contraction. In the living Human Body the exciting force, or stimulus, acting upon a muscle is almost invariably a nervous impulse, a molecular movement transmitted along the nerve-fibres attached to it, and upsetting the equili- brium of the muscle. It is through the nerves that the will acts upon the muscles, and accordingly injury to the nerve of a part, as the face or a limb, will cause paralysis of its muscles. They may still be there, intact and quite ready to work, but there are no means of sending commands to them, and so they remain permanently idle. Although a 130 THE HUMAN BODY. nervous impulse is the natural physiological muscular stimulus it is not the only one known. If a muscle be exposed in a living animal and a slight but sudden tap be given to it, or a hot bar be suddenly brought near it, or an electric shock be sent tlirough it, or a drop of glycerine or of solution of ammonia be placed on it, it will contract; so that in addition to the natural nervous stimulus, muscles are irritable under the influence of mechanical, thermal, electrical, and chemical stimuli. One condition of the ef- ficacy of all of them is that they shall act with some sud- denness; a very slowly increased pressure, even if ultimately very great, or a very slowly raised temperature, or a slowly increased electrical current passed tlirough it, will not ex- cite the muscle; although far less pressure, warmth, or electricity, more rapidly applied would stimulate it power- fully. It may perhaps still be objected that it is not proved that any of these stimuli excite the muscular fibres, and that in all these cases it is possible that the muscle is only excited through its nerves. For the various stimuli named above also excite nerves (see Chap. XIII), and when we ap- ply them to the muscle we may really be acting first upon the fine nerve-endings there, and only indirectly and through the mediation of these upon the muscular fibres. That the muscular fibres have a piu]ier irritability of their own, independently of their nerves, is, however, shown ]>y the action of certain drugs — for example curari, a South American Indian arrow poison. When this substance is introduced into a wound, all the striped muscles are apparently poisoned, and the animal dies of snfl'ocation because of the cessation of the breathing movements. But the poison does not really act on the muscles themselves: it kills the muscle nerves, but leaves the muscle intact; and it kills the very endings of the muscle-nerves right down \\\ the muscle fibres themselves. Now after its administra- tion we still find that the various non-physiological stimuli referred to above make the muscles contract just as powerfully as before the poisoning, so we must conclude that the muscles themselves are irritable in the absence of all nerve stimuli — or, what amounts to the same thing, when MUSCULAR IBBIT ABILITY, 131 all their nerve-fibres have been i^oisoned. The experiment also shows that the contractility of a muscle is a property belonging to itself, and that its contracting force is not something derived from the nerves attached to it. The nerve stimulus simply acts like the electric shock or sudden l^low and arouses the muscle to manifest a property which it already possesses. The older physiologists seeing that muscular paralysis followed when the nervous connection l^etween a muscle and the brain was interrupted, concluded that the nerves gave the muscles the power of contracting. They believed that in the brain there was a great store of a mysterious thing called vital sjnrUs, and that some of this was ejected from the brain along the nerve to the muscle, when the latter was to be set at work, and gave it its working power. We now know that such is not the case, but that a muscle fibre is a collection of highly irrita- ble material which can have its equilibrium upset in a definite way, causing it to change its shape, under the influ- ence of slight disturbing forces, one of which is a ner- vous impulse ; and since • in the Body no other kind of stimulus usually reaches the muscles, they remain at rest when their nervous connections arc severed. But the muscles paralyzed in this way can still, in the living Body, be made to contract by sending electrical shocks through them. Physiologically, then, muscle is a contractile and irritable, but not automatic tissue. A Simple Muscular Contraction. Most of the details concerning the physiological properties of muscles have been studied on those of cold-blooded animals. A frog's muscle will retain all its living properties for some time after removal from the body of the animal, and so can be experimented on with ease, while the muscles of a rabbit or cat soon die under those circumstances. Enough has, however, been observed on the muscles of the higher ani- mals to show that in all essentials they agree with those of the frog or terrapin. When a single electric shock is sent through a muscle it rapidly shortens and then, if a weight be hanging on it. rapidly lengthens again. The whole series of phenomena 132 THE HUMAN BODY. from the moment of stimulation until the muscle regains its resting form is known as a ' • simple muscular contractioji' or a " tivitch.'' It occupies in the frog about one tenth of a second and is separable into three portions. First, there elaspes a time, after the stimulation and before the com- mencement of the shortening, which is known as the "lost time" or the jieriod of latent excitement. This lasts about one hundredth of a second, and represents the time during whicii molecular changes preparatory to the contraction are taking place in the muscle fibres. Then follows the short- ening, at first slow, then rapid, then slower again up to a maximum, and occupying rather more than half of the re- maining time; the elongation occupies the remainder of the time taken up in the contraction. In warm-blooded ani- mals, the duration of a simple muscular contraction is even less than one tenth of a second and all its stages are quick- ened. In any given animal, cold increases the time taken in a muscular contraction and also impairs the contractile power, as we find in our own limbs when " numbed" with cold. Moderate warmth on the other hand (up to near the point at which heat-rigor is produced) diminishes the duration of the contraction; so that the molecular changes in a muscular fibre are clearly eminently susceptible to slight changes in its environment. The contractility of a muscle does not depend upon a vital force, entirely distinct from ordinary inanimate forces, but upon an arrangement of its material elements, which is only maintained under cer- tain conditions and is eminently modifiable by the sur- roundings. Physiological Tetanus. It is obvious that the ordinary movements of the Body are not brought about by such tran- sient muscular contractions as those described in the last paragraph. Even a wink lasts longer than one tenth of a second. Our movements are, m fact, due to more prolonged contractions which may be described as consisting of several simple twitches fused together, and known as " tetanic co7itr actions"; it might be better to call them "compound contractions," since the word tetanus has long been used by pathologists to signify a diseased state, such as occurs MUSCULAR CONTRACTION. 133 in strychnine jjoisoning and hydrophobia, in which most of the muscles of the Body are thrown into prolonged and powerful involuntary contractions. If, while a frog's muscle is still shortening under the in- fluence of one electric shock, another stimulus be given it, it will contract again and the new contraction will be added on to that already existing, without any period of elongation occurring between them. While the muscle is still con- tracting under the influence of the second stimulus a third electric shock will make it contract more, and so on, until the muscle is shortened as much as is possible to it for that strength of stimulus. If now the stimuli be repeated at the proper intervals, each new one will hot produce any further shortening, but, each acting on the muscle before the effect of the last has begun to pass off, the muscle will be kept in a state of permanent or " tetanic" contraction; and this can be maintained, by continuing the stimuli, until the organ begins to get exhausted or *'' fatigued'' and it then commences to elongate in spite of the stimulation. When our muscles are stimulated in the Body, from the nerve-centres through the nerves, they receive from the lat- ter about 20 stimuli in a second, and so are thrown into tetanic contractions. In other words, not even in tlie most rapid movements of the Body is a muscle made to execute a simple muscular contraction; it is always a longer or a shorter tetanus. AVhen very quick movements are executed, as in performing rapid passages on the piano, the result is obtained by contracting two opposing muscles and alternately strengthening and weakening a little the tetanus of each. Causes affecting the Degree of Muscular Contraction. The extent of shortening which can be called forth in a muscle varies with the stimulus. In the first place, a single stimulus can never cause a muscle to contract so much us rapidly repeated stimuli of the same strength — since in the latter case we get, as already explained, scA^eral simple contractions such as a single stimulus would call forth, piled on the top of one another. With very ]iower- ful repeated electrical stimuli a muscle can be made to 134 THE HUMAN BODY. shorten to one third of its resting length, but in the Body the strongest effort of the will never produces a contrac- tion of that extent. Apart from the rate of stimulation, the strength of the stimulus has some influence, a greater stimulus causing a greater contraction, but very soon a point is reached after which increasing the stimulus pro- duces no increased contraction; the muscle has reached its limit. The amount of load carried by the muscle (or the resistance opposed to its shortening) has also an influ- ence, and that in a very remarkable way. Suppose we have a frog's calf-muscle, carrying no weight, and find that with a stimulus of a certain strength it shortens two milli- meters {^j inch). Then if we hang one gram (15.5 grains) on it and give it the same stimulus, it will be found to con- tract more, say four or five millimeters, and so on, up to the point when it carries eight or ten grams. After that an increased weight will, with the same stimulus, cause a less contraction. So that up to a certain limit, resistance to the shortening of the muscle makes it more able to shorten: the mere greater extension of the muscle due to the greater resistance opposed to its sliortening, puts it into a state in which it is able to contract more powerfully. Fatigue diminishes the working power of a muscle and rest restores it, especially if the circulation of tlie blood be going on in it at the same time. A frog's muscle cut out of the body will, however, be considerably restored by a period of rest, even although no blood flows through it. The Measure of Muscular Work. The work done by a muscle m a given contraction, when it lifts a weight verti- cally against gravity, is measured by the weight moved, mul- tiplied by the distance through which it is moved. In the above case when the muscle contracted carrying no load it did very little work, lifting only its own weight; when loaded with one gram and lifting it five millimeters it did five gram-millimeters of work, just as an engineer would say an engine had done so many kilogrammeters or foot- pounds. If loaded with ten grams and lifting it six millimeters it would do sixty gram-millimeters of work. Jlven after the weight becomes so great that it is lifted MUSCULAR WORE. 135 through a less distance, the work done by the muscle goes on increasing, for the bigger weight lifted more than com- pensates for the less distance through which it is raised. For example, if the above muscle were loaded with fifty grams it would maybe lift that weight only l.o millime- ters, but it would then do seventy-five gram-millimeters of work, which is more than when it lifted ten grams six millimeters. A load is, however, at last reached with which the muscle docs less work, the lift becoming very little indeed, until at last the weight becomes so great that the muscle cannot lift it all and so does no work when stimulated. Starting then from the time when the muscle carried no load and did no work, we pass with increasing weights, through phases in which it does more and more work, until with one particular load it does the greatest amount possible to it with that stimulus: after Ihat, with increasing loads less work is done until finally a load is reached with which the muscle again does no work. What IS true of one muscle is of course true of all, and what is true of work done against gravity is true of all muscular work, so that there is one precise load with wdiich a beast of burden or a man can do the greatest possible amount of work in a day. With a lighter or heavier load the distance through which it can be moved will be more or less, but the actual work done always less. In the living Body, how- ever, the working of the muscles depends so much on other things, as the due action of the circulatory and respiratory systems and the nervous energy or "grit" (upon which the stimulation of the muscles depends) of the individual man or beast, that the greatest amount of work obtainable IS not a simple mechanical problem as it is with the excised muscle. Influence of the Form of the Muscle on its Working Power. The amount of work that any muscle can do de- pends of course largely u])on its physiological state; a healthy well-nourished muscle can do more than a dis- eased or starved one; but allowing for such variations the work which can be done by a muscle varies with its form. The thicker the muscles, th:it is the greater the number of 136 THE HUMAN BODY fibres present in a section made across the long axes of the fasciculi, the greater the load that can be lifted or the other resistance that can be overcome. On the other hand, the extent through Avliich a muscle can move a weight in- creases with the length of its fasciculi. A muscle a foot m length can contract more than a muscle six inches long, and so would move a bone through a greater distance, jirovided the resistance were not too great for its strength. But if the shorter muscle had double the thickness, then it could lift twice the weight that the longer muscle could. We find in the Body muscles constructed on both plans; some to have a great range of movement, others to overcome great resistance, besides numerous intermediate forms which cannot be called either long and slender or short and thick ; many short muscles for example are not specially thick, but are short merely because the parts on which they act lie near together. It must be borne in mind, too, that many apparently long muscles are really short stout ones — those namely in which a tendon runs down the side or middle of the muscle, and has the fibres inserted obliquely into it. The muscle {gastrocnemius) in the calf of the leg for instance (Fig. 50, b) is really a short stout muscle, for its working length depends on the length of its fasciculi and these are shor.t and oblique, while its true cross-section is that at right angles to the fasciculi and is very large. The force with which a muscle can shorten is very great. A frog's muscle of 1 square centimeter (0.39 inch) m section can just lift 2800 grams (98.5 ounces), and a human muscle of the same area more than twice as much. Muscular Elasticity. A clear distinction must be made between elasticity and contractility. Elasticity is a physi- cal property of matter in virtue of which various bodies tend to assume or retain a certain shape, and when re- moved from it forcibly, to return to it. When a spiral steel spring is stretched it will, if let go, "contract" in a certain sense in virtue of its elasticity, but such a contrac- tion is clearly quite different from a muscular contraction. The spring will only contract as a result of previous distor- tion; it cannot originate a chiinge of form, while the mus- MUSCULAR ELASTICITY. 137 cle can actively contract or change its shape when a stimnlns acts npon it, and that without being previously stretched. It does not merely tend to return to a natural shape from which it has been removed, but it assumes a quite new natural shape, so that physiological contractility is a differ- ent thing from mere physical elasticity; the essential differ- ence being that the coiled spring or a stretched band only gives back mechanical work which has already been spent on it, while the muscle originates Avork independ- ently of any previous mechanical stretching. In addition to their contractility, however, muscles are highly elastic. If a fresh muscle be hung up and its length measured, and then a weight be hung upon it, it will stretch just like a piece of indian-rubber, and when the weight is removed, provided it has not been so great as to injure the muscle, the latter will return passively, without any stimulus oi active contraction, to its original form. In the Body all the muscles are so attached that they are usually a little stretched beyond their natural resting length; so that when a limb is amputated the muscles divided in the stump shrink away considerably. By this stretched state of the resting elastic muscles two things are gained. In the first place when the muscle contracts it is already taut, there is no " slack" to be hauled in before it pulls on the parts attached to its tendons: and, secondly, as we have already seen the working power of a muscle is increased by the presence of some resistance to its contraction, and this is always provided for from the first, by having the origin and insertion of the muscle so far apart as to be pulling on it Avhen it begins to shorten. Physiology of Plain Muscular Tissue. What has hith- erto been said aj^plies especially to the skeletal muscles; but m the main it is true of the unstriped muscles. These also are irritable and contractile, but their changes of form are much more slow than those of the striated fibres. Upon stimulation, a longer period of latent excitement elapses before the contraction commences, and when finally this takes place it is extremely slow, very gradually attain- ing a maximum and then gi-adually dying away again. 138 THE HU3IAN BODY. There seems in fact to be some connection 1)etween that ar- ranffement of tlie contractile substance Avliich sliows itself under the microscope as striation and the power of rapid contraction, since we find that the heart, which is not a skeletal or voluntary muscle but yet one that contracts rap- idly, agrees with these in having its fibres striated. Hygiene of the Muscles, The healthy working of the muscles needs of course a healthy state of the Body gener- ally, so that they shall be supplied with proper materials for growth and repair and have their wastes rajndly and efiiciently removed. In other words, good food and pure air are necessary for a vigorous muscular system, a fact which trainers recognize in insisting upon a strict dietary, and in supervising generally the mode of life of those who are to engage in athletic contests. The muscles should also not be exj)osed to any considerable continued j^ressure since this interferes with the flow of blood and lymph through them. As far as the muscles themselves are directly concerned, exercise is the necessary condition of their best develop- ment. A muscle Avhich is permanently unused degenerates and is absorbed, little finally being left but the connective tissue of the organ and a few muscle fibres filled with oil- drops. This is well seen in cases of paralysis dependent on injury to the nerves. In such cases the muscles at first may themselves be perfectly healthy, but lying unused for weeks they rapidly alter and,finally, when the nervous in- jury has been healed, the muscles may be found incapa- ble of functional activity. The physician therefore is often careful to avoid this by exercising the paralyzed muscles daily by means of electrical shocks sent through the part. The same fact is illustrated by the feeble and wasted condi- 1 ion of the muscles of a limb which has been kept for some time in splints. After the latter have been removed it is only slowly, l)y judicious and persistent exercise, that the long idle muscles regain their former size and power. The great muscles of the '' brawny arm" of tlie blacksmith or wrestler illustrate the reverse fact, the growth of the mus- cles by exercise. Exercise, however, must be judicious: re^ MUSCULAR EXERCISE. 139 peatedly continued until exhaustion it does harm; the period of repair is not suthcient to allow replacement of the parts used in work, and the muscles thus waste under too violent exercise as with too little. Rest should alternate with work, and that regularly, if benefit is to be obtained. Moreover violent exercise should never be suddenly undertaken by one unused to it, not only lest the muscles suffer but be- cause muscular work greatly increases the work of the heart, not only because more blood has to be sent to the muscles themselves, but they produce great quantities of carbon dioxide which must be carried off in the blood to the lunsfs for removal from the Body, and the heart must work harder to send the blood faster through the lungs and at the same time the breathing be ha.stened so as to renew the air in those organs faster. The least evil result of throwing too violent work on the heart and lungs in this way, is repre- sented by being '*' out of breath," which is advantageous insomuch as it may lead to a cessation of the exertion. But much more serious, and sometimes permanent, injuries of either the circulatory or respiratory organs may be caused by violent and prolonged efforts without any previous train- ing. No general rule can be laid down as to the amount of exercise to be taken; for a healthy man in business the minimum would perhaps be represented by a daily walk of five miles. Varieties of Exercise. In walking and running the muscles chiefly employed are those of the lower limbs and trunk. This is true also of rowing, which when good is performed much more by the legs than the arms: esjoecially since the introduction of sliding seats. "Hence any of these exercises alone is apt to leave the muscles of the chest and arms imjierfectly exercised. Indeed no one exercise em- ploys equally or proportionately all the muscles: and hence gymnnasia in which various feats of agility are practiced, so as to call different parts into play, have attained a great popularity. It should be borne in mind howe\'er, that the legs especially need strength; while the upper limbs, in which delicacy of movement, as a rule, is more desirable than power, do not require such constant exertion; and the 140 THE HUM AX BODY. fact that gymnastic exercises are commonly carried on in- doors is a great drawback to tlieir value. AVlien the weather permits, out-of-door exercise is far better than that carried on in even the best ventilated and lighted gymnasium. For those who are so fortunate as to possess a garden there is no better exercise, at suitable seasons, than an hour's daily digging in it; since this calls into play nearly all the muscles of the Body: while of games, the modern one of lawn tennis is perhaps the best from a hygienic view that has ever been introduced, since it not only demands gi'eat muscular agdity in every part of the Body, but trains the hand to work with the eye in away that walking, running, rowing and similar pursuits do not. For the same reasons baseball, cricket, and boxing are excellent. Exercise in Infancy and Childhood. Young children have not only to strengthen their muscles by exercise but also to learn to use them. Watch an infant trying to con- vey something to its mouth, and you will see how little control it has over its muscles. On the other hand the healthy infant is never at rest when awake; it constantly throws its limbs around, grasps at all objects within its reach, coils itself about, and so gradually learns to exer- cise its powers. It is a good plan to leave every healthy child, more than a few months old, several times daily on a large bed or even on a rug or carpeted floor, with as little covering as is safe and that as loose as possible, and let it wriggle about as it pleases. In this way it will not only enjoy itself thoroughly, but gain strength and a knowledge of how to use its limbs. To keep a healthy child swathed all day in tight and heavy clothes is cruelty. When a little later the infant commences to crawl, it is safe to permit it as much as it wishes; but unwise to tempt it when disinclined. The bones and muscles are still fee- ble and may be injured by too much work. The same is true of commencing walking. From four or five to twelve years of age almost any form of exercise should be permitted, or even encouraged. At this time, however, the epiphyses of many bones are not firmly united to their shafts and so anything tending to MUSCULAR EXERCISE. 141 throw too great a strain on the joints should be avoided. After that up to commencing manhood or maidenhood any kind of outdoor exercise for healthy persons is good, and ofirls are all the better for being allowed to ioin in their brothers' sports. Half of the debility and general ill-health of so many of our women is dependent upon deficient ex- ercise during childhood, and the day, which fortunately seems approaching, which will see dolls as unknown to, or as despised by, healthy girls as healthy boys, will see the be- ginning of a great improvement in the stamina of the female portion of our population. Exercise in Youth should be regulated largely by sex; not that women arc to be shut up and made pale, delicate, and unfit to share the duties or participate fully in the pleasures of life; but the other calls on the strength of the adult woman render vigorous muscular work often unad- visable, especially under conditions where it is apt to b( followed by a chill. A healthy boy or young man may do nearly anythin,^, but until twenty-two or twenty-three very prolonged effort is unadvisable. The frame is still not firmly knit or as capable of endurance as it will subsequently become. Girls should be allowed to ride or play out-door games m moderation, and in any case should not be cribbed in tight stays or tight boots. A flannel dress and pro])er lawn-tennis shoes are as necessary for the healthy and safe enjoyment of an afternoon at that game by a girl as they are for her brother in the base-ball field. Rowing is excel- lent for girls if there be any one to teach them to do it prop- erly, with the legs and back and not with the arms only, as women are so apt to row. Properly practiced it strengthens the back and improves the carriage. Exercise in Adult Life. Up to forty a man may carry on safely the exercises of youth, but after that sudden ef- forts should be avoided. A lad of twenty-one or so may, if trained, safely run a quarter-mile race, but to a man of forty-five it would be dangerous, for with the rigidity of the cartilages and blood-vessels which begins to show itself about that time appears a diminished power of meeting a 142 THE IIl'MAK BODY. sudden violent demand. On the other hand tlie man of thirty would more safely than the lad of nineuen or twenty undertake one of the long-distance walking matches which have lately been in vogue; the jirolonged effort Avould be less dangerous to him, though a six days' match with its attendant loss of sleep cannot fail to be more or less dan- gerous to any one. Probably for one engaged in active business a Avalk of a couple of miles to it in the morning and back again in the afternoon is the best and most avail- able exercise. The habit which Americans have everywhere acquired, of never walking when they can take a horse-car, is certainly detrimental to the general health: though the extremes of heat and cold to which we are subject often render it unavoidable. For women during middle life the same rules apply: there should be some regular but not violent daily ex- ercise. In Old Age the needful amount of exercise is less, and it i> still moi'e important to avoid sudden or violent effort. Exercise for Invalids. This should be regulated under medical advice. For feeble persons gymnastic exercises are especially valuable, since from their variety they permit of selection according to the condition of the individual: and their amount can be conveniently controlled. Training. If any person attempts some unusual exer- cise he soon finds that he loses breath, gets perhaps a " stitch in the side," and feels his heart beating with un- wonted violence. If he perseveres he will probably faint — or vomit.as is frequently seen in imperfectly trained men at the end of a hard boat-race. These phenomena are avoided by careful gradual preparation known as '"train- ing." The immediate cause of them lies in disturbances of the circulatory and respiratory organs, on which excessive work is thrown, and the further discussion of training must be postponed for the present. CHAPTER XI. MOTION AND LOCOMOTION. The Special Physiology of the Muscles. Having now considered separately the structure and properties in gen- eral of the skeleton, the joints, and the muscles, we may go on to consider how they all work togetlier in the Body. The properties of a muscle for example are everywhere the same, but the uses of different muscles are very varied, by reason of the different parts with which they are connected. Some are muscles of respiration, others of deglutition; many are known as flexors because they bend joints, others as extensors because they straighten them, and so on. The determination of tiie exact use of any particular muscle is known as its special pliijsiologij, as distinguished from its general fliysiology, or pro^aerties as a muscle without refer- ence to its use as a muscle in a particular place. The uses of those muscles forming parts of the physiological mechan- isms concerned in breathing and swallowing will be studied hereafter; for the i^resent we may consider the muscles which co-operate in maintaining postures of the Body; in ])roducing movements of its parts with reference to one another; and in producing locomotion or movement of the whole Bod}^ with reference to its environment. In nearly all case the striped muscles carry out their functions with the co-operation of the skeleton, since nearly all are fixed to bones at each end and when they contract primarily move these, and only secondarily the soft parts attached to them. To this general rule there are, however, exceptions. The muscle for example which lifts the upper eyelid and opens the eye arises from ])one at the back of the orbit, but is inserted, not into bone, but into the evelid 144 THE HUMAN BODY. directly ; and similarly other muscles arising at the back of the orbit are directly fixed to the eyeball in front and serve to rotate it on the pad of fat on which it lies. Many facial muscles again have no direct attachment whatever to boiies, as for example the muscle [orhicularis oris) which surrounds the mouth-opening and by its contraction nar- rows it and purses out the lips; or the orbicularis palpe- hrarum which similarly surrounds the eyes and when it contracts closes them. Levers in the Body. When the muscles serve to move bones the latter are in nearly all cases to be regarded as levers whose fulcra lie at the joint where the movement takes place. Examples of all the three forms of levers recognized in mechanics are found in the Human Body. Levers of the First Order. In this form (Fig. 56) the fulcrum or fixed point of support lies between the " weight" Sli 1 _J^ Fig. 56.— a lever of the first order. 7=^, fulcrum; P. power; W, resistance or weight. or resistance to be overcome, and the ''power"' or moving force, as shown in the diagram. The distance PF, from the power to the fulcrum, is called the " puwer-arm;" the distance FW is the weight-arm. When power-arm and weight-arm are equal, as is the case in the beam of an ordi- nary pair of scales, no mechanical advantage is gained, nor is there any loss or gain in the distance through which the Aveight is moved. For every inch through Avhich P is de- pressed, W Avill be raised an equal distance. When the power-arm is longer than the other, then a smaller force at P will raise a larger weight at W, the gain being propor- tionate to the difference in the lengths of the arms. For example if PF is twice as long as FW, then half a kilo- gram applied at F will balance a whole kilogram at W, and motio:n and locomotion. 145 just more than half a kilogram would lift it; but for every centimeter through which P descended, W would only be lifted half a centimeter. On the other hand when the weight-arm in a lever is longer than the power-arm, there is loss in force but a gain in the distance through which the weight is moved. Examples of the first form of lever are not numerous in the Human Body. One is afforded in the nodding move- ments of the head, the fulcrum being the articulations between the skull and the atlas. When the chin is elevated the power is applied to the skull, behind the fulcrum, by small muscles passing from the vertebral column to the occiput; the resistance is the excess in the weight of the part of the head m front of the fulcrum over that behind it, and is not great. To depress the chin as in nodding does not necessarily call for any muscular effort, as the head will fall forward of itself if the muscles keeping it erect cease to work, as those of us who have fallen asleep during a dull discourse on a hot day have learnt. If the chin however be depressed forcibly, as in the athletic feat of suspending one's self by the chin, the muscles passing from the chest to the skull in front of the atlanto-occipital artic- ulation are called into play. Another example of the em- ployment of the first form of lever in the Body is afforded by the curtsey with which a lady salutes another. In curtseying the trunk is bent forward at the hip-joints, which form the fulcrum; the weight is that of the trunk acting as if all concentrated at its centre of gravity, which lies a little above the sacrum and behind the hip-joints; and the power is afforded by muscles passing from the thighs to the front of the pelvis. Levers of the Second Order. In this form the weiglit or resistance is lietween the power and the fulcrum. The power-arm PFh always longer than the weight-arm WF, and so a comparatively weak force can overcome a consid- erable resistance. But it is disadvantageous so far as re- gards rapidity and extent of movement, for it is obvious that when P is raised a certain distance IFwill be moved a less distance in the same time. As an exam23le of the employ- 146 THE HUMAN BODY. ment of such levers (Fig. 57) in tlie Body, we may take the act of standing on the toes. Here the foot rejj resents the lever, the fulcrum is at the contact of its fore part with P fT -^ W Fig. 57.— a lever of the second order. F. fulerum; P, power; The arrows indicate the direction in which the forces act. PF, weight. the ground ; the weight is that of the Body acting down through the ankle-joints at Ta, Fig. 58; and the power is the great muscle of the calf acting by its tendon inserted into the heel-bone {Ca, Fig. 58). Another example is afforded by holding up the thigh when one foot is kept raised from the ground, as in hopping on the other. Here the fulcrum is at the hip-joint, the power is aj^plied at the em en Fif}. .'iS.— The skeleton of the foot from the outer side. Ta, surface with which the leer-bones articulate ; Ca, the calcaneum into which the tendon itendo Achillis) of the calf muscle is inserted ; Mb, the metatarsal bone of the fifth diRit. knee-cap by a great muscle {redus femoris) inserted there and which arises from the pelvis; and the weight is that of the whole lower limb acting at its centre of gravity, which will lie somewhere in the thigh between the hip and LEVERS IN THE BODY. 147 knee joints, that is between the fulcrum and the point of application of the power. Levers of the Third Order. In these (Fig. 59) the power is between the fulcrum and the weight. In such levers the weight-arm is always longer than the power- arm, so the power works at a mechanical disadvantage, but swiftness and range of movement are gained. It is the lever most commonly used in the Human Body. For example, when the forearm is bent up towards the arm, the fulcrum is the elbow-joint, the power is applied at the insertion of the biceps muscle (Fig. 49*) into the radius (and of another muscle not represented in the figure, the hracliialis anticus, into the ulna), and the weight is that ']L yr 'VfT Fig. 58.— a lever of the third order. F, fulcrum ; P, power ; W, weight. of the forearm and hand, with whatever may be contained in the latter, acting at the centre of gravity of the whole somewhere on the distal side of the point of application of the power. In the Body the power-arm is usually very short so as to gain speed and range of movement, the mus- cles being powerful enough to still do their work in spite of the mechanical disadvantage at which they are thus placed. The limbs are thus made much more shapely than would be the case were the jjower applied near or beyond the weight. It is of course only rarely that simple movements as ,those described above take place. In the great majority of those executed several or many muscles co-operate. The Loss to the Muscles from the Direction of their Pull. It is worthy of note that, owing to the oblique direc- — — * p. 120. ~ 148 THE HUMAN BODY. tion in which the muscles are coniniunly inserted into the bones, much of their force is lost so far as producing move- ment is concerned. Suppose the log of wood in the dia- gram (Fig. 60) to be raised by pulling on the rope in the direction a; it is clear at first that the rope will act at a great disadvantage; most of the pull transmitted by it will be exerted against the pivot on which the log hinges, and only a small fraction be available for elevating the latter. But the more the log is lifted, as for example into the position indicated by the dotted line, the more useful will be the direction of the pull, and the more of it will be spent on the log and the less lost unavailingly in merely increas- ing the pressure at the hinge. If we now consider the ac- tion of the biceps (Fig. 49) in flexing the elbow-joint, we see similarly that the straighter the joint is, the more of Fig. 60.— Diagram illustrating the disadvantage of an oblique pull. the pull of the muscle is wasted. Beginning with the arm straight, it works at a great disadvantage, but as the fore- arm is raised the conditions become more and more favor- able to the muscle. Those who have practiced the gym- nastic feat of raising one's self by bending the elbows when hanging by the hands from a horizontal bar, know practi- cally that if the elbow-joints are quite straight it is very hard to start; and that, on the other hand, if they are kept a little flexed at the beginning the effort needed is much POSTURES. 149 less; the reason being of course the more adA-untageous direction of traction by the biceps in the latter case. Experiment proves that the power with which a muscle can contract is greatest at the commencement of its short- ening, the very time at which, we have just seen, it works at most mechanical disadvantage; in proportion as its force becomes less the conditions become more favorable to it. There is however, it is clear, nearly always a considerable loss of power in the working of the skeletal muscles, strength being sacrificed for variety, ease, rapidity, extent, and elegance of movement. Postures. The term posture is applied to those posi- tions of equilibrium of the Body which can be maintained for some time> such as standing, sitting, or lying, com- pared with leaping, running, or falling. In all postures the condition of stability is that the vertical hne drav\'n through the centre of gravity of the Body shall fall within the basis of support afforded by objects with which it is m contact; and the security of the postitre is proportionate to the extent of this base, for the wider it is, the less is the risk of the perpendicular through the centre of gravity fall- ing outside of it on slight disj^lacement. The Erect Posture. This is pre-eminently character- istic of man, his whole skeleton being modified with refer- ence to it. Nevertheless the power of maintaining it is only slowly learnt in the first years after birth, and for a long while it is unsafe. And though finally we learn to stand erect without conscious attention, the maintenance of that posture always requires the co-operation of many muscles, co-ordinated by the nervous system. The influ- ence of the latter is shown by the fall which follows a severe blow on the head, which may nevertheless have frac- tured no bone and injured no muscle: the "concussion" of the brain, as Ave say," stuns" the man, and until its effects have passed off he cannot stand upright. In standing with the arms straight by the sides and the feet together the centre of gravity of the whole adult Body hes in the articulation between the sacrum and the last lumbar verte- bra, and the perpendicular drawn from it will reach the 150 THE HUMAN BODY. I, 7 I ground between the two feet, within the basis of support afforded by them. With the feet close together, however, the i)osture is not very stable, and in standing we com- monly make it more so by slightly separating them so as to increase the base. The more one foot is in front of the other the more sway- ing back and forward will be com pat i ble with safety, and the greater the lateral distance separating them, the greater the lateral sway which is possi- ble without falling. Consequently we see that a man about to make great movements with the upper part of his Body, as in fencing or boxing, or a sol- dier preparing for the bayonet exercise, always commences by thrusting one foot forwards obliquely, so as to increase his basis of support in both directions. The ease with which we can stand is largely dependent upon the way in which the head is nearly balanced on the top of the vertebral column, so that but little muscular effort is needed to keep it upright. In the same way the trunk is almost balanced on the hip- joints: but not quite, its centre of grav- ity falling rather behind them; so that just as some muscular effort is needed to keep the head from falling forwards, FiGf. 61.— Diagram ii- some is needed to keep the trunk from lustratiiig the muscles , ■>• ■> •> -i xxii- t (drawn in thick black topplmg backwards at the hips. In a ^rb^'ehind ^thf YointI similar manner other muscles are called ^cHvi'Jy keep the^S into play at other joints: as between the rigid and the Body erect, ygrtebral column and the pelvis, and at the knees and ankles; and thus a certain rigidity, due to muscular effort, extends all along the erect Body: which on account of the flexibility of its joints could not other- wise be balanced on its feet as a statue can. Beginning (Fig. 61) at the ankle-joint, we find it kept stiff in standing W WALKI^'0. 151 by the combined and balanced contraction of the muscles passing from the heel to the thigh, and from the dorsum of the foot to the shin-bone {tibia). Others passing before and behind the knee-joint keep it from yielding; and so at the hip-joints: and others again lying in the walls of the abdomen and along the vertebral column, keep the latter rigid, and erect on the pelvis; and finally the skull is kept in position by muscles passing from the sternum and ver- tebral column to it, in front of and behind the occipital condyles. Locomotion includes all movements of the whole Body in space, dependent on its own muscular efforts: such as walking, running, leaping, and swimming. Walking. In walking the Body never entirely quits the ground, the heel of the advanced foot touching the ground in each step before the toe of the rear foot leaves it. The advanced limb supports the Body, and the foot in the rear at the commencement of each step, propels it. Suppose a man standing with his heels together to com- mence to walk, stepping out with the left foot; the whole Body is at first inclined forwards; the movement taking place mainly at the ankle-joints. By this means the cen- tre of gravity would be thrown in front of the base formed by the feet and a fall on the face result, were not simulta- neously the left foot slightly raised by bending the knee and then swung forwards, the toes just clear of the ground and, in good walking, the sole nearly parallel to it. When the step is comj)leted the left knee is straightened and the sole placed on the ground, the heel touching it first and, the base of support being thus widened from before back, a fall is prevented. MeaiiAvhile the right leg is kept straight, but inclines forwards above with the trunk when the latter advances, and as this occurs the sole gradually leaves the ground, commencing with the heel. When the step of the left leg is completed the great toe of the right alone is in contact with the support. With this a push is given which sends the trunk on over the left leg which is now kept rigid, except at the ankle-joint; and the right knee being bent that limb swings forwards, its foot just U-i THE HUMAN BODY. cleariug the groimd as the left did before. The Body is meanwhile sni)ported on the left foot alone, hut wiien the right completes its steptlie knee of that leg is straightened and the foot thus placed, heel first, on the ground. Mean- Avhile the left foot has been gradually leaving the gi'ound, and its toes alone are at that moment upon it: from these a push is given, as before Avith the right foot, and the knee being bent so as to raise the foot, the left leg swings for- wards at the hij)-joint to make a fresh step. During each step the Avhole Body sways up and down and also from side to side. It is highest at the moment Avhen the advancing trunk is yertically over the foot sup- porting it, and then sinks until the moment when the ad- Tancing foot touches the ground, when it is loAvest. From this moment it rises as it swings forward on this foot, until it is vertically over it, and then sinks again until the other touches the ground; and so on. At the same time, as its weight is alternately transferred from the right to the left foot and vice versa, there is a slight lateral SAvay, commonly more marked in women than in men, and which when ex- cessive produces an ugly "waddling" gait. The length of each step is primarily dependent on the length of the legs; but can be controlled within wide lim- its by special muscular effort. In easy walking, little mus- cular work is employed to carry the rear leg forwards after it has given its push. When its foot is raised from the ground it swings on like a pendulum; but in fast walking the muscles jjassing in front of the hijj-joint, from the pel- vis to the limb, by their contraction forcibly carry the leg forwards. The easiest step, that in which there is most economy of labor, is that in which the limb is let SAving freely, and since a short pendulum SAvings faster than a longer, the natural step of short-legged people is quicker than that of long-legged ones. In fast Avalking the advanced or supporting leg also aids in projjulsion; the muscles passing in front of the ankle-joint contracting so as to pull the Body forwards OA'er that foot and aid the push from the rear foot. Hence the fatigue and pain in front of the shin Avhich is felt in RUNNING AND LEAPING. 153 prolonged very fast walking. From the fact that each foot reaches the ground heel first, but leaves it toe last, the length of each stride is increased by the length of the foot. Running. In this mode of progression there is a mo- ment in each step when both feet are off the ground, the Body being unsupported in the air. The toes alone come in contact with the ground in each step, and the knee-joint is not straight when the foot reaches the ground. When the rear foot is to leave the support, the knee is suddenly straightened, and at the same time the ankle-joint is ex- tended so as to push the toes forcibly on the ground and give the whole Body a powerful push forwards and upwards. Immediately after this the knee is greatly flexed and the foot raised from the ground, and this occurs before the toes of the forward foot reach the latter. The swinging lag m each step is violently pulled forwards and not suf- fered to swing naturally as in walking. By this the rapid- ity of the succession of steps is increased, and at the same time the stride is made greater by the sort of one-legged leap that occurs through the jerk given by the straighten- ing of the knee of the rear leg just before it leaves the ground. Leaping. In this mode of progression the Body is i-aised completely from the ground for a considerable period. In a powerful leap the ankles, knees, and hip- joints are all flexed as a preparatory measure, so that the Body assumes a crouching attitude. The heels, next, are raised from the ground and the Body balanced on the toes. The centre of gravity of the Body is then thrown forwards, and simultaneously the flexed joints are straightened, and by the resistance of the ground, the Body receives a propul- sion forwards; much m the same way as a ball rebounds from a wall. The arms are at the same time swung for- wards. In leaping back, the Body and arms are Inclined in that direction; and in jumping vertically there is no lean- ing either way and the arms are kept by the sides. CHAPTER XII. ANATOMY OF THE NERVOUS SYSTEM. Nerve-Trunks. In dissecting the Human Body nnmer- ous white cords are found which at tirst sight might be taken for tendons. That they are something else however soon becomes clear, since a great many of them have no connection with muscles at all, and those which have usually enter somewhere into the belly of the muscle, instead of be- mg fixed to its ends as most tendons are. These cords are nerve-trunks: followed in one direction each (Fig. 62) will be found to break up into finer and finer branches, until the subdivisions become too small to be followed without the aid of a microscope. Traced the other way the trunk will in most cases be found to increase by the union of others with it, and ultimately to join a much larger mass of different structure, and from which other trunks also spring. This mass is a nerve-centre. That end of a nerve attached to the centre is naturally its central, and the other its distal or 'peripheral end. Nerve-centres, then, give origin to nerve-trunks; these latter radiate all over the Body, usually branching and becoming smaller and smaller as they proceed from the centre; they finally become very small, and hoAv they iiltimately end is not in all cases cer- tain, but it is known that some have sense-organs at their terminations and others muscular fibres. Tlie general ar- rangement of the larger nerve-trunks of the Body is shown in Fig. 62. Physically a nerve is not so tough or strong as a tendon of the same size; it may readily be split up into longitudinal strands, each of whicli consists of a number of microscopic threads, the nerve-fibres, bound together by connective tissue. AJVA TOM Y OF NEB VO US S TSTEM. 1 5 o Fig. 62.— Diagrram illustrating the g'eneral arrangement of the nervous system. 156 THE HUMAN BODY. Plexuses. Very fn'(nu'ntly several ncigliboring nerve- trunks send off comimiiiicating branches to one another, each brunch carrying libres from one trunk to the olhei-. 8uch networks are called j^jfen/.ses (Fig. 05*), and through the interchanges taking place in them it often happens that the distal branches of a nerve-trunk contain fibres which it does not possess as it leaves the centre to which it is connected. Nerve-Centres. The great majority of the nerves take their origin from the hrain and sjnnal cord, which together form the great cerebro-sjnnal centre. Some, however, com- mence in rounded or oval masses which vary in size from that of the kernel of an almond down to microscopic dimensions, and which are widely distributed in the Body. Each of these smaller scattered centres is called a ganglion, and the whole of them are arranged in three sets. A considerable number of the largest are united directly to one another by nerve-trunks, and also give off nerves to various organs, especially to the blood-vessels and the viscera in the thoracic and abdominal cavities. These ganglia and their branches form the sympathetic nervous system, as distinguished from the cerebro-si^inal nervous system consisting of the brain and spinal cord and the nerves springing from them. Of the remaining ganglia some are connected with various cerebro-s2nnal trunks near their origin, while the rest, for the most part very small and connected with the peripheral branches of sympathetic or other nerves, are known as the sporadic ganglia. The Cerebro-Spinal Centre and its Membranes. Ly- ing in the skull is the hrain and in the neural canal of the vertebral column the sjnnal cord or spinal viarrow, the two being continuous through i\\Q forameii magnum of the occipital bone and forming the great cerebro-spinal nerve- centre. This centre is bilaterally symmetrical throughout except for slight differences on the surfaces of parts of the brain, which are often found in the higher races of mankind. Both brain and spinal cord are very soft and easily crushed; the connective tissue which pervades them being of the deli- cate retiform variety; accordingly both are placed in nearly *P. 162. MEMBRANES OF THE XERVE-CENTRES. 15? completely closed bony cavities and are also enveloped by membranes wliicli give tliem consistency and support. These membranes are three in number. Externally is the dura mater, very tough and strong and composed of white fibrous and elastic connective tissues. In the cranium this dura mater adheres by its outer surface to the inside of the skull, serving as the periosteum of its bones; this is not the case in the vertebral column, where the dura mater forms a loose sheath around the spinal cord and is only attached here and there to the sur- rounding bones, which have a sep- arate ]3eriosteum of their own. The innermost membrane of the cerebro- spinal centre, lying in immediate contact with the proper nervous parts, is the j*;ia mater, also made up of white fibrous tissue inter- woven with elastic fibres, but less closely than in the dura mater, so as to form a less dense and tough membrane. The pia mater con- tains many blood-vessels which break up in it into small branches before entering the nervous mass beneath. Covering the outside of the pia mater is a layer of flat closely fitting cells, a similar layer lines the inside of the dura mater, and these two layers are described as the third membrane of the cere- bro-spinal centre, called the arach- . , T j_i 1 ; ,1 FiR- 63.— The spinal cord noia. in the space between the an.i meduiia oblongata, a. (.,„„ 1 „ J? J.1 1 -1 ■ from the ventral, and 5, from two layers of the arachnoid is con- tue dor.sai aspect ; c to h. tained a small quantity of watery fevlfs.'^""'"'^ ^^ ^^'''''^ 158 THE HUMAN BODY. cerebro-spi7ial liquid. Part of the surface of the brain is folded and the pia mater does not dip down and line the furrows between tlie folds but stretches across them: in the spaces thus left there is also contained some of the cerebro- si)inal liquid. The Spinal Cord (Fig. 03) is nearly cylindrical in form, being however a little Avider from side to side than dorso- Tentrally, and tapering off at its posterior end. Its aver- age diameter is about 19 millimeters (f inch) and its length 0.43 meter (17 inches). It weighs 42.5 grams (1^ ounces). There is no marked limit between the sijinal cord and the brain, the one passing gradually into the other (Fig. 70*), but the cord is arbitrarily said to com- mence opposite the outer margin of the foramen magnum : from there it extends to the articulation between the first and second lumbar vertebraB, where it narrows off to a slender filament, the filum terminaU (cut off and rej)re- sented separately at B' in Fig. 63), which runs back to the end of the neural canal behind the sacrum. In its course the cord presents two expansions, an upper, 10, the cer- vical enlargement, reiichiugiTom. the third cervical to the first dorsal vertebrae, and a lower or lumhar enlargement, 9, opposite the last dorsal vertebra. Eunning along the middle line on both the ventral and the dorsal aspects of the cord is a groove, and a cross-sec- tion shows tl\at these grooves are the surface indications of fissures which extend deeply into the cord {C, Fig. 64) and nearly divide it into right and left halves. The anterior fissure (1, Fig. 64) is wider and shallower than the posterior, 2. The transverse section, C, shows also that the substance of the cord is not alike throughout, but that its tvJnte superficial layers envelop a central gray substance arranged somewhat in the form of a capital H. Each half of the gray matter is crescent-shaped, and the crescents are turned back to back and united across the middle line by the grag commissure. The tips of each crescent are called its horns or cornua, and the ventral, or anterior cornu, on each side is thicker and larger than the posterior. In the cervical and lumbar enlargements the *P. 169. SPINAL CORD. 159 proportion of white to gray matter is greater than else- where ; and as the coi-d approaches the meduUa oblongata its central gray mass becomes irregular in form and begins to break up into smaller portions. If lines be drawn on the transverse section of the cord from the tip of each horn of the gray matter to the nearest point of the surface, the white substance in each half will be divided into three por- FiG. 64.— The spinal cord and nerve-roots. A. ti small portion of the cord seen from the ventral side ; B, tlie same seen laterally ; C, a cross-section of the cord ; D, the two roots of a spinal nerve ; 1, anterior (ventral) fissure ; 2. poste- rior (dorsal) fissure ; 3, surface groove along the line of attachment of the ante- rior nerve-roots ; 4, line of origin of the posterior roots : 5, anterior root fila- ments of a spinal nerve ; 6, posterior root filaments : 6'. ganglion of the poste- rior root ; 7,7,' the two primary divisions of the nerve-trunic formed by the miion of the two roots. tions: one between the anterior fissure and the anterior cornu, and called the anterior white column ; one between the posterior fissure and the posterior cornu, and called the posterior iclrite column; while the remaining one lying in the hollow of the crescent and between the two horns is the lateral column. In addition to this a certain amount of white substance crosses the middle line at the bottom of 160 THE HUMAN BODY. the iinterior fissure; tliis forms tlie (inferior tvliite commis- sure. There is no jiosterior white commissure, the bottom of the posterior fissure being the only portion of the cord where tlie gray substance is uncovered by white. Running along the middle of the gray commissure, for the whole length of the cord, is a tiny channel, just visible to the unaided eye; it is known as the central canal {canalis cen- tralis). The Spinal Nerves. Thirty-one pairs of spinal nerve- trunks enter the neural canal of the vertebral column through tlie intervertebral foramiiia (p. 71). Each di- vides in the foramen into a dorsal and ventral portion known respectively as the posterior and anterior roots of the nerve (6 and 5, Fig. 64), and these again subdivide into finer branches which are attached to the sides of the cord, the posterior root at the point where the posterior and late- ral Avhite columns meet, and the anterior root at the junc- tion of the lateral and anterior columns. At the lines on which the roots are attached there are superficial furrows on the surface of the cord. On each posterior root is a spinal ganglion (6', Fig. 64), placed just before it joins the an- terior root to make up the common nerve-trunk. Imme- diately after its formation by the mixture of fibres from both roots, the trunk divides into a small posterior prima rp and a larger anterior ptrimary hrancli (7' 7 D, Fig. 64). The former branches of the spinal nerves go for the most part to the skin and muscles on the back, while the anterior primary branches form a series of plexuses from which the nerves for the sides and ventral region of the neck and trunk, and for the limbs, arise. The various spinal nerves are named from the jiortions of the vertebral column through the intervertebral foramina of which they pass out; and as a general rule each nerve is named from the vertebra in front of it. For example the nerve passing out between the fifth and sixtli dorsal verte- brae is the "fifth dorsal" nerve, and that between the last dorsal and first lumbar vertebrfe, the "twelfth dorsal." In the cervical region, however, this rule is not adhered to. The nerve passing out between the occipital bone and the THE SPINAL XERVES. lOl atlas is called the " first cervical" nerve, that between the atlas and axis the second, and so on; that between seventh cervical and first dorsal vertebra? being the "eighth cervi- cal " nerve. The thirty-one pairs of spinal nerves are then thus distributed: 8 cervical, 13 dorsal, 5 lumbar, 5 sacral, and 1 coccygeal; the latter passing out between the sacrum and coccyx. Since the spinal cord ends opposite the upper lumbar vertebrae while the sacral and coccygeal nerves pass out from the neural canal much farther back, it is clear that the roots of those nerves, on their way to unite in the foramina of exit and form nerve-trunks, must run obliquely backwards in the spinal canal for a considerable distance. One finds in fact the neural canal in the lumbar and sacral regions, behind the point where the spinal cord has tapered oif, occupied by a great bunch of nerve-roots forming the so-called " horse's tail" or cauda equina. Distribution of the Spinal Nerves. It would be out of place here to go into detail as to the exact portions of the Body supplied hj each spinal nerve, but the following general statements may be made. The anterior prinuiry branches of the first four cervical nerves form on each side the cervical plexus (Fig. 65) from which branches are sup- plied to the muscles and integument of the neck: also to the outer ear and the back part of the scalp. The anterior primary branches of the remaining cervical nerves and the first dorsal form the bi'achial plexus, from which the upper limb is supplied. The roots of the trunks which form this plexus arise from the cervical enlargement of the spinal cord. From the fourth and fifth cervical nerves on each side, small branches arise and unite to make the phrenic nerve (4, Fig. 65) which runs down through the chest and ends in the diaphragm. Tlie anterior primary branches of the dorsal nerves, ex- cept part of the first which enters the brachial plexus, form no plexus, but each runs along the posterior border of a rib and supplies branches to the chest-walls, and the lower ones to those of the abdomen also. The anterior primary branches of the four anterior lum- bar nerves are united by branches to form the lumbar 162 TEE HUMAN BODY. plexus. It snpi)lies the lower part of the trunk, the but- tocks, the front of llie thigh, and medial side of the leg. The sacral plexus is formed by the anterior primary branches of the fifth lumbar and the first four sacral nerves, which unite into one 'great cord and so form the Fig. 65.— The cervical and brachial plexuses of one side of the Body. xeiatic nerve, which is the largest in the Body and, running down to the back of the thigh, ends in branches for the lower limb. The roots of the trunks which form the sacral plexus arise from the lumbar enlargement of the cord. THE BRAIN. 163 The Brain (Fig. 66) is far larger tlian the spinal cord and more complex in structure. It weighs on the average about 1415 grams (50 ounces) in the adult male, and about 155 grams (5.5 ounces) less in the female. In its simpler forms the vertebrate bram consists of three masses, each with subsidiary parts, following one another in series from before back, and known as the fore-brain, mid- brain, and hind-brain respectively. In man the fore-brain, A, weighing Fig. 66.— Diagram iUustrating the general relationships of the parts of the brain. A, fore-braiu ; b, mid-brain ; B. cerebellum : C, pons Varolii ; D, me- dulla oblongata ; B, C, and D together constitute the hind-brain. about 1215 grams (44: ounces), is much larger than all the rest put together and laps over them behind. It consists mainly of two huge convoluted masses, separated from one "another by a deep median fissure, and known as the cerebral I/eniisjilieres. The immense proportionate size of these is very characteristic of the human brain. Beneath each cerebral hemisphere is an olfactory lobe, inconspicuous in man but often larger than the cerebral liemisiDheres, as in most fishes. Buried in the fore-brain on each side are two large gray masses, the corpora striata and optic thalami. The mid-brain forms a connecting isthmus between the two other divisions and presents on its dorsal side four 164 THE IIUMA^ BODY. hemispherical eminences, the corpora qnadrigemioia. On its ventral side it exhibits two semicylindrical pillars (seen iinder the nerve IV in Fig. 70* and known as the crura cerebri. The hind-brain consists of three main parts: on its dorsal side is the cerebellum, B, Fig. 66, consisting of a right, a /(?//, and a median lobe ; on the ventral side is the pons Varolii, C, Fig. 66, and behind the medulla oblongata, D, Fig. 66, which is continuous with the spinal cord. In nature the main divisions of the brain are not sepa- rated so much as has been represented in the diagram for Cb Fig. 67.— The brain from the left side. Cb, the cerebral hemispheres forming the main bulk of the fore-brain ; Chi, the cerebellum ; Mo, the medulla oblon- gata ; P, the pons Varolii ; *, the fissure of Sylvius. the sake of clearness, but lie close together as represented in Fig. 67, only some folds of the membranes extending between them; and the mid-brain is entirely covered in on its dorsal aspect. Xearly everywhere the surface of the brain is folded, the folds, known as gyri or convolutions, being deeper and more numerous in the brain of man than in that of lower animals; and in the human species more marked in the higher than in the lower races. The brain like the spinal cord consists of gray and white nervous matter but somewhat differently arranged, for while the brain, like the cord, contains gray matter in * P. 169. CEREBRAL VENTRICLES. 1G5 its interior, a great part of its surface is also covei'ed with it. By tlie external convolutions of the cerebellum and the cerebral hemispheres the surface over which this gray substance is spread is very much increased (see Fig. 68). The Ventricles of the Brain. The minute central canal of the spinal cord is continued into the brain and exjaands there at several points into chambers known as the ventricles. Entering the medulla oblongata it approaches its upper surface and dilates into the fourtli ventricle, which has a very thin roof, lapped over by the cerebellum. From the front of the fourth ventricle runs a narrow pas- Ccr Fig. 68.— a vertical section across the cerebral hemispheres. Ccl^. the corpus callosum ; IT, the anterior end of the riisrht lateral ventricle: the gray mass on its exterior is the corpus striatum. On the left side the superficial gray matter covei'iiig the convolutions is shaded. sage (ifer a tciiio ad qiiartum vent ricul urn) Avhicli enters another dilatation lying in the middle line near the under side of the fore-brain (just above the two small rounded masses seen between the nerves // and III in Fig. 70) and known as the tliird ventricle. From the third ventricle two apertures (the foramens of Monro) lead into the first and second, or lateral ventricles, one of which lies in each 166 THE HUMAN BODY. of the cerebral hemispheres. The front ends of these two ventricles are seen in the vertical transverse section of the brain reprcscnied in Fig. 68. The ventricles contain a small amount of cerebrospinal liquid and are lined by epithelium which is ciliated in early life. Note. A frequent cause of apoiilexy is a hemorrhage into one of the lateral ventricles; the outpoured blood accu- mulating and pressing upon the cerebral hemispheres their functions are suppressed and unconsciousness produced. When a person is found in an apoplectic fit therefore the best thing to do is to leave him perfectly quiet until medi- cal aid is obtained: for any movement may start afresh, bleeding into the ventricle which had been stopped by clots formed in the mouths of the torn blood-vessels. Sections of the Brain. Having got a general idea of the parts composing the brain, the best way to complete a knowledge of its anatomy is to study sections taken in various directions. Two such are given in Figs. 08 and GO. Fig. 69 represents tlie right half of a vertical section of the brain, taken from before back in the middle line and viewed from the inner side. Above, the knife has passed between the two cerebral hemispheres, in the longitudinal fissure, without cutting either, and the convoluted inner surface of the right one is seen. The sickle-shaped mass lower down, CcV to Ccf represents the cut surface of a connecting band of white nervous tissue called the corinis callosum, which runs across the middle line from one cerebral hemisjsliere to the other and puts them in communication. SI, the septum lucidum, is a thin membrane which forms the inner wall of the lateral ventricle of the hemisphere. Between the two septa lucida on the sides (in the natural position of the parts) and the corpus callosum above is inclosed a narrow space known as the fifth ventricle. It is, however, quite different from the remaining cerebral ventricles, not being a continuation of the canalis centralis of the spinal cord. Tlie space beneath the septum lucidum and the back part of the corpus callosum is the third ventricle, which, lying in the middle line, har^ been laid open in the MEDIAX SURFACE OF TEE BRAIN. 167 section. It is deep from above down but narrow from side to side. From its under side a prolongation runs down to H, the pituitary lodij ; behind, the aqueduct of Sylvius, A, is seen passing back from the third to the fourth ven- tricle, Vq. At F3I is the aperture {foramen of Monro) leading into the right lateral ventricle. Crossing the third ventricle and putting the two halves of the fore-brain in direct communication are three small commissures, Coa, C,l8 Cam sM Fig. 69.— The i-ight half of the brain as seen on its median side after a section made througli the organ in the middle line. Vq, fourth ventricle ; Mo, medulla oblongata ; P. pons Varolii ; II, optic nerve ; H, pituitary body ; Coa. ante- rior commissure ; FM, foramen of 3Ionro leading from the third ventricle, in the cavity of which the lower end of the line SM lies, to the right lateral ven- tricle ; Com, soft commissure, running from side to side of the third ventricle, divided ; Cop, posterior commissure ; Lq, corpora quadrigemina ; .-1, aque- duct of Sylvius or iter a tertio ad qnartum ventriculum ; Cbl, cerebeUum ; Ccl^-Ccl*, corpus callosum ; HI, septum lucidum. Com, and Coj), known respectively as the anterior, the median (or soft), and the joos^mor. The mass seen bound- ing a great part of the side of the third ventricle and united to its fellow by the soft commissure is the ojJtic thalamus. Above the aqueduct is the small median body Cn, called the pineal glajid, which contains no nervous tissue, but has an interest as being, according to Descartes, 108 THE HUMAN BODY. the seat of tlic soul. Beliiiul it come the coyjHira qnadri- (jernlna, Lq, and above the fourth ventricle the cerehcllum, Ohl, showing the primary and secondary fissures on its surface which give its section a branched appearance known as the arhor vitce. Mo is the medulla oUongdta, and P the })ons A'arolii. The canalis centrahs of the spinal cord is represented leading back from the fourth ventricle. Fig. G8 represents a vertical transverse section of the brain taken through the fore part of the corpus callosum {CcV) and altogether in front of the third ventricle. It shows the foldings of the cerebrum and its su})erficial layer of gray substance; the anterior ends of the lateral ventri- cles, VI, with a gray mass, the corpus striatum lying be- neath and on the outer side of each. If the section had been taken a little farther back the ojjtic tlialavii would have been found reaching the floor of each ventricle. The Base of the Brain and the Cranial Nerves. Twelve pairs of nerves leave the skull by apertures in its base, and are known as the cranial nerves. Most of them spring from the under side of the brain, and so they are best studied in connection with the base of that organ, which is represented in Fig. 70. H\\q first p) air, or olfactory nerves, spring from the under sides of the olfactory lobes, /, and pass out through the roof of the nose. They are the nerves of smell. The second pair, or 02)tic nerves, II, spring from the optic thalami and corjDora quadrigemina and, under the name of the op)tic tracts, run down to the base of the brain where they appear passing around the crura cerebri as represented in the figure. In the middle line the two optic tracts unite to form the opitic commissure (seen in section at //' in Fig. 69) from which an optic nerve pro- ceeds to each eyeball. Behind the optic commissure is seen the conical stalk of the pituitary lody or hypojiliysis cerebri {II in Fig. 69), and still further back a pair of hemispherical masses, about the size of split peas, known as the corpora alhicantia. All the remaining cranial nerves arise from the hind- brain. The third pair {motores oculi) arise from the front of tlie pons A'arolii, and are distributed to most of the THE CRANIAL NERVES. 169 muscles wliich move the eyeball and also to that which lifts the iipiDcr eyelid. The four-sided space bounded by the optic tracts and commissure in front and the third pair of nerves behind, and having on it the pituitary body Fig. 70.— The base of tlie brain. The cerebral hemispheres are seen over- lapping all the rest. /, olfactory lobes ; //, optic tract passing to the optic commissure from whicli the optic nerves proceed ; ///, the third nerve or jho/oc oculi ; IV, the fourtli nerve or pathcticus ; V, the fifth nerve or trigeminalia ; FZ, the sixth nerve or abducens ; F/J, the seventh or facial nerve or portio dura ; VIII, the auditory nerve or portio viollis ; IX, the ninth or glosso- phaiyngeal ; X, the tenth or pneumogastric or vagus : XI, the spinal acces- sory ; XII, the hypoglossal ; ncl, the fii'st cervical spinal nerve. and the corpora albicantia, lies beneath the third ventricle, so that a probe puslied in there would enter that cavity. The fourth pair of nerves, IV (pafJieiici), arise from the front part of the roof of the fourth ventricle. From there. 170 THE HUMA^ BODY. each curls around a cms cerebri (the cylindrical mass seen beneath it in the figure, running from the pons Varolii to enter the under surface of the cerebral hemispheres) and appears on the base of the brain. Each goes to one muscle of the eyeball. The fifth pair of nerves, V {trigeminales), resemble tlic spinal nerves in having two roots; one of these is much larger than the other and possesses a ganglion (the Gasse- rian ganglion) like the posterior root of a spinal nerve. Beyond the ganglion the two roots form a common trunk which divides into three main branches. Of these, the ophfhcdmic is the smallest and is mainly distributed to the muscles and skin over the forehead and upper eyelid; but also gives branches to the mucous membrane lining the nose, and to the integument over it. The second division {superior maxillary nerve) of the trigeminal gives branches to the skin over the temple, to the cheek between the eye- brow and the angle of the mouth, and to the upper teeth; as well as to the mucous membrane of the nose, pharynx, soft palate and roof of the mouth. The third division {inferior maxillary) is the largest branch of the trigemi- nal; it receives some fibres from the larger root and all of the smaller. It is distributed to the side of the head and the external ear, the lower lip and lower part of the face, the mucous membrane of the mouth and the anterior two thirds of the tongue, the lower teeth, the salivary glands, and the muscles which move the lower jaw in mas- tication. The sixth pair of cranial nerves ( VI, Fig. 70) or ab- dncentes arise from the posterior margin of the pons Va- rolii, and each is distributed to one muscle of the eve- ball. The seventh pair {facial nerves), VII, appear also at the posterior margin of the pons. They are distributed to most of the muscles of the face and scalp. The eiglith pair {auditory nerves) arise close to the facial. They are the nerves of hearing and are distributed entirely to the internal ear. The ninth pair {glossojjharyngeals), IX, arising close to THE CRANIAL NERVES. 171 the auditories, are distributed to the mucous memhrune of the pharynx, the posterior part of the tongue, and the middle ear. The tenth pair {pneumogastric nei'ves or vagi), X, arise from the sides of the medulla oblongata. Each gives branches to the pharynx, gullet and stomach, the larynx, windpipe and lungs, and to the heart. The vagus runs farther through the Body than any other cranial nerve. The eleventh 2^air {spinal accessory nerves), XI, do not arise mainly from the brain but by a number of roots at- tached to the lateral columns of the cervical portion of the spinal cord, between the anterior and posterior roots of the proper cervical spinal nerves. Each, however, runs into the skull cavity alongside of the spinal cord and, getting a few filaments from the medulla oblongata, passes out along with the glossopharyngeal and pneumogastric nerves. Out- side the skull it divides into two branches, one of which joins the pneumogastric trunk, while the other is distrib- uted to muscles about the shoulder. v^ The tivelfth pair of cranial nerves (hypoglossi), XII, arise from the sides of the medulla oblongata; they are distributed mainly to the muscles of the tongue and the hyoid bone. Deep Origins of the Cranial Nerves. The points re- ferred to above, at which the various cranial nerves ajipear on the surface of the brain, are known as their superficial origins. From them the nerves can be traced for a less or greater way in the substance of the brain until each is fol- lowed to one or more masses of gray matter, which con- stitute its proper starting-point and are known as its deep origin. The deep origins of all except the first and second and part of the eleventh lie in the medulla oblongata. The Ganglia and Communications of the Cranial Nerves. Besides the Gasserian ganglion above referred to, many others are found in connection with the cranial nerves. Thus for example there is one on each of the main divisions of the trigeminal, two are found on each pneumogastric and two in connection with the glosso- pharyngeal. At these ganglia and elsewhere, tlie various 172 Til?: HUMAN BODY. nerves often receive brunclies from neighboring cr;inial or spinal nerves, so that very soon after it leaves the brain hardly any one remains free from fibres derived from other trunks except the olfactory, optic, and auditory nerves. This often makes it difficult to say from Avhcre the nerves of a special part have come; for example, the nerve-fibres going to the submaxillary salivary gland from the trigemi- nal leave the brain first in the facial and only afterwards enter the fifth; and many of the fibres going apparently from the pneumogastric to tlie heart come originally from the spinal accessory. The Sympathetic System. The ganglia which form the main centres of the sympathetic nervous system lie in two rows {s, Fig. 2, and sy, Fig. 3), one on either side of the bodies of the vertebrae. Each ganglion is united by a nerve-trunk Avith the one in front of it, and so two great chains are formed reaching from the base of the skull to the coccyx. In tlie trunk region these chains lie in the ventral cavity, their relative position in which is indicated by the dots sy in the diagrammatic transverse section re- presented on p. 7 in Fig. 3. The ganglia on these chains are forty-nine in number, viz., twenty -four pairs, and a single one in front of the coccyx in which both chains terminate. Tliey are named from the regions of the vertebral column near which they lie; there being three cervical, twelve dorsal, four lumbar, and five sacral pairs. Each symj^athetic ganglion is united by communicating Iranches with the neighboring sjjinal nerves, and near the skull with various cranial nerves also; while from the gan- glia and their uniting cords arise numerous trunks, many of which, in the thoracic and abdominal cavities, form plexuses, from which in turn nerves are given off to the viscera. These plexuses frequently possess numerous small ganglia of their own; two of the most important are the cardiac plexus which lies on the dorsal side of the heart, and the solar plexus wliicli lies in the abdominal cavity and supplies nerves to the stomach, liver, kidneys, and intes- tines. Many of the sympathetic nerves finally end in the walls of the blood-vessels of various organs. To the naked HISTOLOGY OF NERVES. 1V3 eye they are commonly grayer in color than the cerebro- spinal nerves. The Sporadic Ganglia. These, for the most part very minute, nerve-centres are found scattered in nearly all parts of the Body. They are especially abundant in the neighborhood of secretorv tissues and about blood-vessels, while a very important set is found in the heart. Serves unite them with the ccrebro-spinal and sympathetic cen- tres, and probably many of them belong properly to the sympathetic system. The Histology of Nerve-Fibres. The microscope shows that in addition to connective tissue and other accessory parts, such as blood-vessels, the nervous organs contain tis- sues peculiar to themselves and known as nerve-fibres and nerve-cells. The cells are found in the centres only; while the fibres, of which there are two main varieties known as the wliite and the gray, are found in both trunks and cen- tres; the white variety predominating in the cerebro-spinal nerves and in the white substance of the centres, and the gray in the sympathetic trunks and the gray portions of the central organs. If an ordinary cerebro-spinal nerve-trunk be examined it will be found to be enveloped in a loose sheath of areolar connective tissue, Avhich forms a packing for it and unites it to neighboring parts. From this sheath, or perineurium, bands of connective tissite penetrate the nerve and divide it up into a number of smaller cords or funiculi, much as a muscle is subdivided into fasciculi; each funiculus has a sheath of its own called the neurilemma, composed of several concentric layers of a delicate membrane, with- in which the true nerve-fibres lie. These, which would be nearly all of the white kind, consist of extremely delicate threads, about 0.0125 millm. {-o-^xri) inch) in diameter, but frequently of a length which is in proportion very great. Each nerve-fibre in fact is contintious from a nerve-centre to the organ in which it ends, so that the fibres, e.g. which pass out through the sacral plexus and then run on through the sciatic nerve and its branches to the skin of the toes, are three to four feet long. If a perfectly fresh nerve-fibre 174 THE HUMAN BODY. be examined with the microscope il presents the appear- ance of a perfectly homogeneous glassy thread ; but soon it acquires a characteristic double contour (Fig. 71) from the coagulation of a portion of its substance. By proper treat- ment Avith reagents three layers may be brought into view. Outside is a tine transparent envelope (1, Fig. 72) called the jjrimitive sheath ; inside this is a fatty substance, 2, Fig. 71. — White nerve-fihres soon after removal from the Body and when they have acqmred their double contour. Fig. 72. — Diag-rani ilhi'^trating the structure of a ivhite or meduUated nerve- fihre. 1, 1, primitive sheath ; 2, 2, medidlary sheath ; 3, axis cyhnder. forming the medullarij sheath (the coagulation of which gives the fibre its double border), and in the centre is a core, the axis cylinder, 3, which is clearly the essential part of the fibre, since near its ending the primitive and medullary sheaths are frequently absent. At intervals of about one millimeter (^ inch) along the fibre are found nuclei. These are indications of the primitive cells which by their elongation, fusion and other modifications have HI8T0L0Q Y OF NER VE-CELLS. 175 built up the nerve-fibre, and around each there is a small amount of unmodified jjrotoplasm. The medullary sheath is interrupted half way between each j)air of nuclei at a point, called the node, which answers to the original bound- ary between the two cells. In the course of a nerve-trunk its fibres rarely divide; when a branch is given off some fibres merely separate from the rest, much as a skein of silk miglit be separated out at one end into smaller bundles con- iaining fewer threads. Gray Nerve-Fibres. Some of these are merely white fibres which near their j^eripheral ends have lost their me- dullary sheaths; but others have no medullary sheath throughout their whole course, consisting merely of an axis cylinder (often longitudinally striated) and primitive sheath, with nuclei. Such fibres are especially abundant in the sympathetic trunks; and they alone are found in the olfactory nerve. In the communicating branches between the sympathetic ganglia and the spinal nerves both white and gray fibres are found; the former being probably cerebro-spinal fibres passing into the sympathetic system, while the gray fibres originate in the sympathetic system and pass into the spinal cord. Another class of gray nerve- fibres may be called nerve-fihrils : they are extremely fine and result from the subdivision of axis cylinders, close to their final endings in many parts of the Body, after they have already lost both primitive and medullary sheaths. Many fine gray fibres exist in the nerve-centres. The Histology of Nerve-Cells. So far as our knoAvl- edge at present goes the only structures known with cer- tainty to be connected with the central ends of nerve-fibres are nerve-cells, and these latter may therefore be regarded as the central organs of the nerve-fibres. However, many nerve-fibres have not yet been traced into continuity with nerve-cells, and possibly end in the centres in other ways. At 1, Fig. 73, is shown a typical nerve-cell such as may be found in an anterior horn of the gray matter of the spinal cord. It consists of the cell body, or cell protoplasm, in which is a large micleus containing a niicleolns. From the body of the cell arise several branches, the great ma- 176 THE HUMAN BODY. jority of which subdivide and form fine nerve-fibres in the gray sabstanco of tlio spinal cord. One process of the cell, however, a, does not branch, but is continued into the an- terior root of a spinal nerve, acquiring a medullary and prim- itive sheath at J), and becoming thus the axis cylnider of a nerve-fibre. Other nerve-cells (as shown at 2 and 4) do not possess the peculiar axis-cylinder process; all their branches either join the branches of other cells or enter a fine net- FiG. 73. — Different forms of nerve-cells. ], a cell, one branch of which, a, becomes the axis cylinder of a nerve-fibre ; 2, two cells united by a process, c; 3, diagram of three" cells united by branches with one another, and each having an axis-cylinder process ; 4, a multipolar cell without an axis-cylinder process. work of gray nerve-fibrils. Most nerve-cells are larger than the majority of the other cells of the Body, their average diameter in the anterior horns of the gray sub- stance of the cord being 0.1 millimeter (g^^ inch). In the posterior horns they are smaller, and in the brain many minute nerve-cells are found in addition to these larger ones. In ganglia the cells as a rule are more reg- MINUTE STBUGTUBE OF SPINAL CORD. 177 ular in outline than those depicted in Fig. 73, and have fewer branches, most appearing indeed to have but two. Others have been described as possessing only one process connected with them, and some with none, but the ex- istence of these is doubtful, since in separating the cells for microscopic examination the delicate processes may readily be broken oif and so escape detection. The Structure of Nerve-Centres. These consist of white and gray nerve-fibres, of nerve-cells, and of connective tissue and blood-vessels arranged in different ways in the different centres. Ganglia are collections of nerve-cells and nerve-fibres, some of the latter being connected with the cells, while others seem merely to pass through the ganglion on their way to other parts. The whole mass is enveloped and supported by other tissues. As an illus- tration of the structure of a more complex nerve-centre we may study the spinal cord. Histology of the Spinal Cord. If a thin transverse sec- tion of the spinal cord be examined with a microscope it will be found to exhibit the following parts (Fig. 74). Enveloping the whole and adherent to the rest is the deli- cate layer of connective tissue forming the pia mater. This lines the anterior fissure, 1, and an offshoot from it fills up the posterior fissure, 2. Elsewhere fine bands of it run in and ramify through the cord, supporting the nerv- ous elements; some of the coarser of these are represented at 6, 7, and elsewhere in the figure, but from these still finer processes arise, as represented at d and e in Fig. 75, and surround the individual nerve fibres and cells. This ultimate finest connective tissue supporting the nervous tissues directly, belongs to the retiform variety (p. 106), and is called the neuroglia. In the white columns, the cord (Fig. 75) will be seen to be mainly made up of medul- lated nerve-fibres which run longitudinally and therefore appear in the transverse section as circles, with a dot in the centre, which is the axis cylinder. At h in Fig. 75 these fibres are represented, the intermediate connective tissue being omitted, while at e this latter alone is repre- sented in order to show more clearly its arrangement. At 178 THE HUMAN BODY. the levels of the nerve-ruots liorizoiitul white fibres are found (1) and 10, Fig. 74, and a, Fig. 75) running into the gray matter, and others exist at the bottom of the anterior fissure, running from one side of the cord to the other. In the gray substance the same supporting network of con- nective tissue is found, but in it the majority of the nerve- fibres are non-medullated, and at certain points nerve-cells, "Fig. 7ir^K thin triansverse section of half of the spinal cord maf^nifled about 10 diameters. 1, anterior fissure ; 2. posterior fls.sure ; 3, caualis centralis : 8, pia mater enveloping the cord ; 6, 7, bands of pia mater penetrating tlie cord and supporting its nerve elements ; 9, a posterior root ; 10, bundles of an ante rior root ; a, b, c, d, e, groups of nerve-cells in the gray matter. such as are totally absent in the white substance, are found. One collection of these nerve-cells is seen at e in Fig. 74, and others are represented at a, d, f, and elsewhere. The nerve-fibres in the gray matter are for the most part branches of these cells (see Fig. 73), and as they unite with one another freely they form, a structurally continuous HISTOLOGY OF SPIRAL COED. 179 network through the whole gray substance. The fibres of the anterior roots of the spinal nerves enter the gray mat- ter and there become continuous with the unbranchecl pro- cess of a nerve-cell; the ending of the posterior root-fibres is not quite certain, but they appear to break up and join the gray network directly, without the intervention of a cell. In any case the fundamental fact remains that every nerve-fibre joining the spinal cord is directly or indirectly in continuity Avith the gray network and so with all the Fig. 75.— a smill bit of the section representee! in Fig. 74 more magnified, a, a bundle of fibres from an anterior root passing through tlie wliite substance on its way to the gray. Towai-ds the riglit of the figure the nerve-fibi-es of the anterior "column tiave been omitted so as to render more conspicuous tlie sup- porting connective tissue, d and e. Elsewhere the nerve-fibres alone are repre- sented; c, enveloping i>ja ?waer ends are seen the cordce teiidineoe proceeding to the edges of the tlaps of tlie mitral valve. The open- ing into the auricle lies between these flaps At the beginning of the aorta are seen its three pouch-like semilunar valves. the chest, giving ofi many branches on its way. Piercing tlie diaphragm it enters the abdomen and after supplying the parts m and around that cavity with branches, it ends 210 THE HUMAN BODY. opposite the last lumbar vertebra by dividing into the right and left common iliac arteries, which carry blood to the lower limbs. We have then to consider the branches of the arch of the aorta, and those of the descending aorta, which latter is for convenience described by anatomists as consist- ing of the tlioracic aorta, extending from the end of the arch to the diaphragm, and the abdominal aorta, extending from the diaphragm to the final subdivision of the vessel. Branches of the Arch of the Aorta. From this arise first the coronary arteries {crd and crs, Figs. 79 and 80) which spring close to the h^art, just above two of the pouches of the semilunar valve, and carry blood into the substance of that organ. The remaining branches of the arch are three in number, and all arise from its convexity. The first is the imiomi7iate artery {Ab, Fig. 79), which is very short, immediately breaking up into the right subcla- vian artery, and the right common carotid. Then comes the left common carotid, Cs, and finally the left subclavian, Ssi. Each subclavian artery runs out to the arm on its own side and after giving ofE a vertebral artery (which runs up the neck to the head in the vertebral canal of the transverse processes of the cervical vertebrse), crosses the arm-pit and takes there the name of the axillary artery. This con- tinues down the arm as the brachial artery, which, giving off branches on its way, runs to the front of the arm, and just below the elbow-joint divides into the radial a.ndi ulnar arteries, the lower ends of which are seen at R and U in Fig. 77.* These supply the forearm and end in the hand by uniting to form an arch, from which branches are given ofE to the fingers. The common carotid arteries pass out of the chest into the neck, along which they ascend on the sides of the windi^ipe. Opposite the angle of the lower jaw each divides into an internal and external carotid artery, right or left as the case may be. The latter ends ma;inly in branches for the face, scalio, and salivary glands, one great subdivision of it with a tortuous course, the temporal artery, being often seen beating in thin persons on the side of the brow. The in- * P. 202. THE CAPILLARIES. 211 ternal caroti.I aitery enters the skull throngli an aperture in its base and supplies the brain, Avhich it Avill be remem- bered also gets blood through the vertebral arteries. Branches of the Thoracic Aorta. These are numerous but small. Some, the intercostal arteries, run out between the ribs and supply the chest-walls; others, the bronchial arteries, carry blood to the lungs for their nourishment, that carried to them by the pulmonary arteries being brought there for another purpose; and a few other small branches are given to other neighboring parts. Branches of the Abdominal Aorta. These are both large and numerous, supplying not only the wall of the posterior part of the trunk, but the important organs in the abdominal cavity. The larger are — the cceliac axis which supplies stomach, spleen, liver, and pancreas; the siqjerior mesenteric artery which supplies a great part of the intes- tine; the renal arteries, one for each kidney; and finally the inferior mesenteric artery which supplies the rest of the intestine. Besides these the abdominal aorta gives off very many smaller branches. Arteries of the Lower Limbs. Each common iliac di- vides into an intcDial and external iliac artery. The former mainly ends in branches to parts lying in the pelvis, but the latter passes into the thighs and there takes the name of the femoral artery. At first this lies on the ven- tral aspect of the limb, but lower down passes back round the femur, and above the knee-joint, where it is called the popliteal artery, divides into the anterior and posterior tibial arteries which supply the leg and foot. The Capillaries. As the artei'ies ai-e followed from the heart their branches become smaller and smaller, and finally cannot be traced without the aid of a microscope. Ulti- mately they pass into the capillaries, the walls of which are simpler than those of the arteries, and which form very close networks m nearly all parts of the Body; their immense number compensating for their smaller size. The average diameter of a capillary vessel is .016 mm. (:j-^ inch) so that only two or three l)lood corpuscles can pass through it abreast, and m many pnrts they are so close 213 THE HUMAN BODY. that a pin's point could not be inserted between two of them. It is while flowing in these delicate tubes that the blood does its nutritive work, the arteries being merely supply-tubes for the capillaries. The Veins. The first veins arise from the capillary net- works in various organs, and like the last arteries are very small. They soon increase in size by union and so form a I ■** Fig. 82.— a small portion of the capillary network as seen in the frog's web when magnified about 25 diameters, a, a small artery feeding the capillaries ; V, V, small veins carrying blood back from the latter. larger and larger trunks. These in many places lie near or alongside the main artery of the part, but there are many more large veins just beneath the skin than there are large arteries. This is especially the case in the limbs, the main veins of which are superficial and can in many persons be THE VEINS. 213 seen as faint ])lne lines through the skin. Fig. 83 repre- sents the arm at the front of the elbow-joint after the skin and subcutaneous areolar tissue and fat have been re- — rt has Fig. 83.— The superficial veins in front of the elbow-joint. B' , tendon of biceps muscle ; Bi, brachialis intenius muscle ; Pt, pronator teres muscle ; 1, mediau nerve ; 2, 3, 4, nerve-branches to the skin ; B, brachial artery witli its small accompanying veins ; cep, cephalic vein ; bas, basilic vein ; tii', median vein ; *, junction of a deep-lymg vem with the cephalic. moved. The brachial artery, B, colored red, is seen lying tolerably deeji and accompanied by two small veins (vence coviites) which communicate by cross-branches. The great 214 THE HUMAN BODY. median nerve, 1, a branch of the brachial plexus which supplies several muscles of the forearm and hand, the skin over a great part of the palm, and the three inner fingers, is seen alongside the artery. The larger veins of the part are seen to form a more superficial network, joined here and there, as for instance at *, by branches from deeper parts. Several small nerve-branches Avhich supply the skin (3, 3, 4) are seen among these veins. It is from the vessel, cep, called the cephalic vein, just above the point where it crosses the median nerve, that surgeons usually bleed a patient. A great part of the blood of the lower limb is brought back by the Io)ig saphenous vein, which can be seen running beneath the skin from the inner side of the ankle to the top of the thigh. All the blood which leaves the heart by the aorta, except that flowing through the coronary arte- ries, is finally collected into the superior and inferior vence cavcB {cs and ci, Figs. 79 and 80), and poured into the right auricle. The jugular veins which run down the neck, carrying back the blood which went out along the carotid arteries, unite below with the arm-vein {snhclavian) to form on each side an innominate vein {Asi and Ade, Fig. TO) and the innominates unite to form the superior cava. The coronary-artery blood after flowing through the capillaries of the heart itself, also returns to this auricle by the coronary veins. The Pulmonary Circulation. Through this the blocd gets back to the left side of the heart and so into the aorta again. The pulmonary artery, dividing into branches for each lung, ends in the capillaries of those organs. From these it is collected by the pulmonary veins which carry it back to the left auricle, whence it passes to the left ventricle to recommence its flow through the Body generally. Tho Course of the Blood. From what has been said it is clear that tho movement of the blood is a circulation. Starting from any one chamber of the heart it will in time return to it; but to do this it must pass through at least two sets of capillaries; one of these is connected with the aorta and the other with the pulmonary artery, and in its PORTAL CIRCULATION. 215 circuit the blood returns to the heart twice. Leaving the left side it returns to the right, and leaving the right it returns to the left: and there is no road for it from one side of the heart to the other except through a capillary network. Moreover it always leaves from a ventricle through an artery, and returns to an auricle through a vein. There is then really only one circulation; but it is not uncommon to speak of two, the flow from the left side of the heart to the right, through the Body generally, being called the f . , ing that it forms a single the Icit auricle lu the pumionarv closed circuit with two pumps . » t • i j t i mi •' in it, consisting of the right VCIUS 01 a bright red COlor. ihlS and left halves of the heart, i •/ • j. • x-n *i i which are represented sepa- color it maintains Until it reaches iTAi^v^i'^^'^-ell-^e, the systemic capillaries, but in these friS t!aS'tsTstem^ ^^ loscs much oxygeu to the surround- capillaries ; vc, yenaj cavaB ; i^p- tlSSUCS and gUlUS mucll CarboU po, pulmonary artery ;pc,pul- o o monary capillaries ; pv, pul- dioxidc from them. But the blood monary veins. . t i i i coloring matter which has lost its ox3'gen has a dark purple-black color, and since this un- oxidizcd or "reduced" haemoglobin is now in excess, the blood returns to the heart by the venae cavse of a dark purple-red color. This color it keeps until it reaches the lungs, when the reduced haemoglobin becomes again oxi- dized. The bright red blood, rich in oxygen and poor in carbon dioxide, is known as "arterial blood'' and the dark 8TR UCTUBE OF TEE BLOOD- VESSELS. 317 red as " venous blood:" and it must be borne in mind that the terms have this peculiar teclmical meaning, and that the pulmonary veins contain arterial blood and the pulmo- nary arteries, venous blood; the change from arterial to venous taking place in the systemic capillaries, and from venous to arterial in the pulmonary capillaries. The chambers of the heart and the great vessels containing ar- terial blood are shaded red in Figs. ?9 and 80. The Structure of the Arteries. A large artery can by careful dissection be separated into three coats; an internal, middle and outer. TL\e internal coat tears readily across the long axis of the artery and consists of an inner lining of flattened nucleated cells, and of a variable numbefof layers composed of membranes or networks of elastic tissue, outside this. The middle coat is made uji of alternating layers of elastic fibres and jilain muscular tissue; the for- mer running for the most part longitudinally and the latter across the long axis of the vessel. The outer coat is the toughest and strongest of all and is mainly made up of white fibrous connective tissue but contains a considerable amount of elastic tissue also. It gradually shades off into a loose areolar tissue which forms the slieath of the artery or the tunica adventitia, and packs it between surrounding parts. The smaller arteries have all the elastic elements less developed. The internal coat is consequently thinner, and the middle coat is made up mainly of involuntary mus- cular fibres. As a result the large arteries are highly elas- tic, the aorta being physically much like a piece of Indian- rubber tubing, while the smaller arteries are highly con- tractile, in the physiological sense of the word. Structure of the Capillaries. In the smaller arteries the outer and middle coats gradually disappear, and the elastic layers of the inner coat also go. Finally, in the capillaries the lining epithelium alone is left, with a more or less developed layer of connective-tissue corpuscles around it, representing the remnant of the tunica adven- titia. These vessels are thus extremely well adapted to al- low of filtration or diffusion taking place through their thin walls. 218 THE HUMAN BODY. Structure of the Veins. In these the same three pri- mary coats as in the arteries may be found: the inner and middle coats are less developed while the outer one remains thick, and is made up almost entirely of white fibrous tissue. Hence venous walls are much thinner than those of the corresponding arteries, and the veins collapse when empty while the stouter arteries remain ojjen. But the tenacity and toughness of their outer coats give the veins great strength. Excejot the pulmonary artery and the aorta, which pos- sess the semilunar valves at their cardiac orifices, the arteries possess no valves. Many veins on the contrary have such, formed by semilunar pouches of the inner coat, at- tached by one margin and having that turned towards the heart free. These valves, sometimes single, oftener in pairs, and sometimes three at one level, permit blood to flow only towards the heart, for a current in that direction (as in the upper diagram, Fig. 85) joresses the valve close against the side of the vessel and . meets with no oljstruction from it. Should any back-flow be attempted, however, the current closes vc^ the valve and bars its own passage as indicated in the lower figure. These valves are most numerous tJtfthelSJdf of acuofof "^ Superficial veins and those of the valves of the veins, c the muscular parts. They are absent capillary, and H^ the heart , -^ •' end of the vessel. m the veute cavne and the portal and pulmonary veins. Usually the vein is a little dilated opposite a valve and hence in parts where the valves are numerous gets a knotted look. On compressing the forearm so as to stop the flow in its sub- cutaneous veins and cause their dilatation, the points at which valves are placed can be recognized by their swollen appearance. They are most frequently found where two veins communicate. CHAPTER XV. THE WORKING OF THE HEART AND BLOOD- VESSELS. The Beat of the Heart. It is possible by methods known to physiologists to open tlie chest of a living narcotized animal, such as a rabbit, and sec its heart at work, alter- nately contracting and diminishing the cavities within it and relaxing and expanding them. It is then observed that each beat commences at the mouths of the great veins; from there runs over the rest of the auricles, and then over the ventricles; the auricles commencing to dilate the moment the ventricles commence to contract. Having finished their contraction, the ventricles also commence to dilate and so for some time neither they nor the auricles are contracting, but the whole heart expanding. The con- traction of any part of the heart is known as its systole and the relaxation as its diastole, and since the two sides of the heart work synchronously, the auricles together and the ventricles together, we may describe a whole ''cardiac period" or " heart-beat"' as made up successively of auricu- lar systole, ventricular systole, and pause. This cycle is repeated about seventy times a minute; and if the whole time occupied by it be subdivided into 100 parts, about 9 of these will be occupied by the auricular systole, about 30 by the ventricular systole, and 61 by the pause: during more than half of life, therefore, the muscles of the heart are at rest. In the pause the heart if taken be- tween the finger and thumb feels soft and flabby but dur- ing the systole it (especially in its ventricular portion) be- comes hard and rigid. Change of Form of the Heart. During its systole the 220 THE HUMAN BODY. heart becomes shorter and rounder, mainly from a change in the shape of the yentricles. A cross-section of the heart at the base of these hitter during diastole would be ellipti- cal in outline, with its long diameter from right to left: during the systole it is more circular, the long axis of the ellipse becoming shortened while the dorso-ventral diameter remains little altered. At the same time the length of the ventricles is lessened, the apex of the heart approaching the base and becoming blunter and rounder. The Cardiac Impulse, The human heart lies with its apex touching the chest-wall between the fifth and sixth ribs on the left side of the breast-bone. At every beat a sort of tap, known as the ''cardiac impulse" or "apex beat," may be felt by the finger at that point. There is, however, no actual "tapping" since the heart's apex never leaves the chest-wall. During the diastole the soft ventri- cles yield to the chest- wall where they touch it, but dur- ing the systole they become hard and tense and push it out a little between the ribs, and so cause the a]3ex beat. Since the heart becomes shorter during the ventricular systole it might be supposed that at that time the apex would move up a little in the chest. This hoAvever is not the case, the ascent of the apex towards the base of the ventricles being compensated for by a movement of the whole heart in the opposite direction. If water be pumped into an elastic tube, already tolerably full, this will be distended not only transversely but longitudinally. This is what happens in the aorta: when the left ventricle contracts and pumps blood forcibly into it, the elastic artery is elongated as well as widened, and this lengthening of that limb of its arch at- tached to the heart pushes the latter down towards the dia- phragm, and compensates for the upward movement of the apex due to the shortening of the ventricles. Hence if the exposed living heart be watched it appears as if during the systole the base of the heart moved towards the tip, rather than the reverse. Events occurring -within the Heart during a Cardiac Period, Let us commence at the end of the ventricular systole. At this moment the semilunar valves at the orifices PHENOMENA OF THE HEARTS BEAT. 321 of the aorta and the pulmonary artery are closed, so that no blood can flow back from those vessels. The whole heart, however, is soft and distensible and yields readily to blood flowing into it from the pulmonary veins and the venre cavae; this passes on through the open mitral and tricuspid valves and fills up the dilating ventricles, as well as the auricles. As the ventricles fill, back currents are set up along their walls and these carry up the flaps of the valves so that by the end of the pause they are nearly closed. At this moment the auricles contract, and since this contrac- tion commences at and narrows the mouths of the veins opening into them, and at the same time the blood in those vessels opposes some resistance to a back-flow into them, while the still flabby and dilating ventricles oppose much less resistance, the general result is that the contract- ing auricles send blood mainly into the ventricles, and hardly any back into the veins. At the same time the in- creased direct current into the ventricles produces a greater back current on the sides, which, as the auricles cease their contraction and the filled ventricles become tense and press on the blood inside them, completely close the auriculo- ventricular valves. That this increased filling of the ven- tricles, due to auricular contractions, will close the valves is seen easily in a sheep's heart. If the auricles be carefully cut away from this so as to expose the mitral and tricuspid valves, and water be then poured from a little height into the ventricles, it will be seen that as these cavities are filled the valve-flaps are floated up and close the orifices. The auricular contraction now ceases and the ventricular commences. The blood in each ventricle is imprisoned between the auriculo-ventricular valves behind and the semilunar valves in front. The former cannot yield on account of the cordse tendinese fixed to their edges: the semilunar valves, on the other hand, can open outwards from the ventricle and let the blood pass on, but they are kept tightly shut by the pressure of the blood on their other sides, just as the lock-gates of a canal are by the pressure of the water on them. In order to open the canal-gates water is let in or out of the lock until it stands at the same level 222 THE HUMAN BODY. on each side of them; but of course they might be forced open without this by applying sufficient power to overcome the higher water pressure on one side. It is in this latter way that the semilunar valves are opened. Tlie contracting ventricle tightens its grip on the blood inside it and becomes rigid to the touch. As it squeezes harder and harder, at Icist the pressure on the blood in it becomes greater than the pressure exerted on the other side of the valves by the blood in the arteries, the flajas are pushed ojoen, and the blood begins to pass out: the ventricle continues its con- traction until it has obliterated its cavity and completely emptied itself. Then it commences to relax and blood immediately to flow back into it from the highly stretched arteries. This back current, however, catches the pockets of the semilunar valves, drives them back and closes the valve so as to form an impassable barrier; and so the blood which has been forced out of either ventricle cannot flow directly back into it. Use of the Papillary Muscles. In order that the con- tracting ventricles may not force blood back into the auricles it is essential that the flaps of the mitral and tricuspid valves be maintained horizontally across the open- ings which they close, and be not pushed back into the auricles. At the commencement of the ventricular sys- tole this is provided for by the cordse tendine^e, which are of such a length as to keep the edges of the flaps in appo- sition, a i^osition which is farther secured by the fact that each set of cordse tendine« (Fig. 81*) radiating from a point in the ventricle, is not attached around the edges of one flap but on the contiguous edges of two flaps, and so tends to pull them together. But as the contracting ven- tricles shorten, the cordse tendineae, if directly fixed to their interior, would be slackened and the valve-flaps pushed up into the auricle. The little papillary muscles prevent this. Shortening as the ventricular systole proceeds, they keep the cordse taut and the valves closed. Sounds of the Heart. If the ear be placed on the chest over the region of the heart during life, two distinguish- able sounds will be heard during each cardiac cycle. They are known respectiA^ely as the first and second sounds of the * P. 209. EVENTS IX A CARDIAC CYCLE. 223 heart. The first is of lower pitch and lasts longer than the second and sharper sound: vocally their character may be tolerably imitated by the words lulh, chqj. The cause of the second sound is the closure, or as one might say the "clicking up," of the semilunar valves, since it occurs at the moment of their closure and ceases if they be hooked back in a liAdng animal. The origin of the first sound is still uncertain: it takes place during the ventricular systole and is probably due to vibrations of the tense ventricular wall at that time. It is not due, as has been supposed, to the auriculo-ventricular valves, since it may still be heard in a beatmg heart empty of blood, and m which there cotild be no closure or tension of those valves. In various forms of heart disease these sounds are modified or cloaked by additional " murmurs" which arise when the cardiac orifices are roughened or narrowed or dilated, or the valves ineffi- cient. By paying attention to the character of the new sound then heard, the exact period in the cardiac cycle at which it occurs, and the region of the chest-wall at which it is heard most distinctly, the physician can often get impor- tant information as to its catise. Diagram of the Events of a Cardiac Cycle. In the following table the phenomena of the heart's beat are rep- resented with reference to the changes of form which are seen in an exposed working heart. Events in the same vertical column occur simultaneously; on the same horizon- tal line, from left to right, successively. Auricular Sys- tole. Auricles ' Contracting and emptying. Ventricles | Dilating and filling. Impulse Auriculo - ventric- ular valves Semilunar valves. Sounds Closing. Closed. Commence- ment of Ven- tricular Sys- tole. Dilating and filling. Contracting. Apex beat. Closed. I Closed. Closed. Open. First sotmd. Ventricular Systole. Dilating and filling. Contracting and emptying. Cessation of Ventricular ! Systole. I Dilating , Dilating and filling, and filling. Dilating. , Dilating i and filling. Opening. • Open. Closing. Closed. Second sound, i 224 THE HUMAN BODY. Function of the Auricles. The ventricles have to do the work of pumping the blood througli the blood-vessels. Accordingly their walls are far thicker and more muscular Llian those of the auricles; and the left ventricle, which has TO force the blood over the Body generally, is stouter than the right, which has only to send blood around the com- paratively short pulmonary circuit. The circulation of tlic blood is in fact maintained by the ventricles, and we have to inquire what is the use of the auricles. Not un- frequently the heart's action is described as if the auricles first filled with blood and then contracted and filled the ventricles; and then the latter contracted and drove the blood into the arteries. From the account given above, however, it will be seen that the events are not accurately so represented, but that during all the pause blood flows on through the auricles into the ventricles, which latter are already nearly full when the auricles contract; this con- traction merely completing their filling and finishing the closure of the auriculo-ventricular valves. The real use of the auricles is to afford a reservoir into which the veins may empty while the comparatively long-lasting ventricular contraction is taking place: they also largely control the amount of work done by the heart. If the heart consisted of the ventricles only, with valves at the points of entry and exit of the blood, the circulation could be maintained. During diastole the ventricle would fill from the veins, and during systole empty into the ar- teries. But in order to accomplish this, during the systole the valves at the point of entry must be closed, or the ven- tricle would empty itself into the veins as well as into the arteries; and this closure would necessitate a great loss of time which might be utilized for feeding the pump. This is avoided by the auricles, which are really reser- voirs at the end of the venous system collecting blood when the ventricular pump is at work. When the ven- • tricles relax, the blood entering the auricles flows on into them: but previously, during the -^^^ of the cardiac cycle occupied by the ventricular systole, the auricles have accumulated blood, tmd when thev at last con- FUNCTION OF THE AURICLES. 225 tract they send on into the ventricles this accnmulation. Even were the flow from tlie veins stopj^ed during the auricular contraction this would be of comparatively little consequence, since that event occupies so brief a time. But, although no doubt somewhat lessened, the emptying of the veins into the heart does not seem to be, in health, stopped while the auricle is contracting. For at that mo- ment the ventricle is relaxing and receives the blood from the auricles under a less pressure than it enters the latter from the veins. The heart in fact consists of a couple of •'feed-pumps" — the auricles — and a couple of "force- pumps" — the ventricles; and so wonderfully perfect is the mechanism that the supply to the feed-pumps is never stopped. The auricles are never empty, being supplied all the time of their contraction, which is never so great as to obliterate their cavities; while the ventricles contain no blood at the end of their systole. The auricles also govern to a certain extent the amount of work done by the ventricles. These latter contract with more than sufficient force to completely drive out all the blood contained in them. If the auricles contract more })owerfully and empty themselves more completely at any given time, the ventricles will contain more blood at the commencement of their systole, and have pumped out more at its end. Now as we shall see in Chapter XVII., the contraction of the auncles is under the control of the nervous system; and through the auricles the whole work of the heart. In fact the ventricles represent the brute force concerned in maintaining the circulation, while the auricles are part of a highly developed co-ordinating mechanism, by which the rate of the circulation is governed according to the needs of the whole Body at the time. The Work Done by the Heart. This can be calculated witli approximate correctness. At each systole each ven- tricle sends out the same quantity of blood — about 180 grams (6.3 ounces); the pressure exerted by the blood in the aorta against the semilunar valves and which the ventricle has to overcome is about that which would be ex- erted on the same surface by a column of mercury 200 226 THE HUMAN BODY. millimeters (8 inches) high. The left ventricle therefore drives out, seventy times in a minute, 180 grams (6.3 ounces) of blood against this pressure. Since the specific gravity of mercury is 13,5 and that of blood may for prac- tical purposes be taken as 1, the work of each stroke of the ventricle is equivalent to raising 180 grams (6.3 ounces) of blood 200 X 12.5 = 2500 millim. (8.2 feet); or one gram 450 meters (one ounce 51.66 feet); or one kilo- gram 0.45 meters (one lb. 3.23 feet). "Work is measured by the amount of energy needed to raise a definite weight a given distance against gravity at the earth's surface, the unit, called a Icilogr ammeter, being either that necessary to raise one kilogram one meter, or, called a foot-pound, that necessary to raise one pound one foot. Expressed thus the work of the left ventricle in one minute, when the heart'.s rate is seventy strokes in that time, is 0.45 x 70 = 31.50 kilogrammeters (3.23x70 = 226.1 foot-pounds); in one hour it is 31.50 X 60 = 1890 kilogrammeters (226.1 X 60 = 13,566 footpounds); and m twenty-four hours 1890 x 24 = 45,360 kilogrammeters (325,584 foot-pounds). The pres- sure in the pulmonary artery against which the right ventricle works is about ^ of that in the aorta; hence this ventricle in twenty-four hours will do one third as much work as the left, or 15,120 kilogrammeters (108,528 foot pounds) and adding this to the amount done by the left, we get as the total work of the ventricles in a day the immense amount of 60,480 kilogrammeters (434,112 foot-jwunds). If a man weighing 75 kilograms (165 lbs.) climbed up a mountain 806 meters (2644 feet) high his skeletal muscles would probably be greatly fatigued at the end of the ascent, and yet in lifting his Body that height they would only have performed the amount of work that the ventricles of the heart do daily without fatigue. The Flow of the Blood Outside the Heart. The blood leaves the heart intermittently and not in a regular stream, a quantity being forced out at each systole of the ventri- cles: before it reaches the capillaries, however, this rhythmic movement is transformed into a steady flow as may readily be seen by examining under the microscope thin trans- BLOOD-FLOW AS SEEN WITH THE MICROSCOPE. 327 parent parts of various animals, as the web of a frog's foot, a mouse's ear, or the tail of a small fish. In consequence of the steadiness with which the capillaries supi)ly the veins the flow in these is also unaffected, directly, Ly each beat of the heart; if a vein be cut the blood wells out uniformly, while a cut artery spurts out not only with much more force, but in jets which are much more powerful at regu- lar intervals corresponding witli the systoles of the ven- tricles. The Circulation of the Blood as Seen in the Frog's Web. There is no more fascinating or instructive phenomenon than the circulation of the blood as seen with the micro- scope in the thin membrane between the toes of a frog's hind limb. Upon focusing beneath the epidermis a net- work of minute arteries, veins and capillaries, with the blood flowing through them, comes into view (Fig. 8-?*). The arteries, a, are readily recognized by the fact that the flow in ihem is fastest and from larger to smaller branches. The latter are seen ending in capillaries, which form networks, rJie channels of which are all nearly equal in size. While m the veins arising from the capillary the flow is from smaller to larger trunks, and slower than in the arteries but faster than in the capillaries. The reason of the slower flow of the capillaries is that their united area is considerably greater than that of the arteries supplying them, so that the same quantity of blood flowing through them in a given time, has a wider channel to flow m and moves more slowly. The area of the veins is smaller than that of the capillaries but gi-eater than that of the arteries, and hence the rate of movement in them IS also intermediate. Almost always when an arterv divides, the area of its branches is gi-eater than that of the mam trunk, and so the arterial current becomes slower and slower from the heart onwards. In the veins on the other hand, the area of a trunk formed by the union of two or more branches is less than that of tlie branches together, and the flow becomes quicker and quicker towards the heart. But even at the heart the united cross-sections of the veins entering the auricles is greater than that of *p.ai2- 228 THE HUMAN BODY. tjie arteries leaving the ventricles, so that, since as much blood returns to the heart in a given time as leaves it, the rate of the current in the pulmonary veins and the venae cavee is less than in the pulmonary artery and aorta. We may represent the vascular system as a double cone, widening from the ventricles to the capillaries and narrowing from the latter to the auricles. Just as water forced in at a narrow end of this would flow quickest there and slowest at the widest part, so the blood flows quickest in the aorta and slowest in the capillaries, which form together a much wider channel. The Axial Current and the Inert Layer. If a small artery in the frog's web be closely examined it will be seen that the rate of flow is not the same in all parts of it. In the centre is a very rapid current carrying along all the red corpuscles and known as the axial dream, while near the wall of the vessel the flow is much slower, as indicated by the rate at which the pale blood corpuscles are carried along in it. This is a purely physical phenomenon. If any liquid be forcibly driven through a fine tube which it wets, water for instance through a glass tube, the outermost layer of the liquid will remain motionless in contact with the tube; the next layer of molecules will move faster, the next faster still; and so on until a very rapid current is found in the centre. If solid bodies, as powdered sealing-wax, lie suspended in the water, these will all be carried on in the central faster current or aorial stream, just as the red corpuscles are in the artery. The white corpuscles, on account of their powder of executing independent amoeboid movements and their consequent irregular form, get fre- quently pushed out of the axial current, so that many of them are found in the inert layer. Internal Friction. It follows from the above-stated facts that there is no noticeable friction between the. blood and the lining of the vessel through which it flows: since the outermost blood layer in contact with the wall of the vessel is almost motionless. But there is very great fric- tion between the different concentric layers of the liquid, since each of them is moving at a different rate from those THE CAPILLARY CIRCULATION. 229 in contact with it on each side. This form of friction is known in hydro-dynamics as " internal friction" and it is of great importance in the circulation of the blood. In- ternal friction increases very fast as the calibre of the tube through which the liquid flows diminishes: so that with the same rate of flow it is disproportionately much greater in a small tube than in a larger one. Hence a given quantity of liquid forced in a minute through one large tube, would experience much less resistance from in- ternal friction than if sent in the same time through four or five smaller tubes, the united transverse sections of which were together equal to that of the single larger one. In the blood-vessels the increased total area, and consequently slower flow, in the smaller channels partly counteracts this increase of internal friction, but only to a comparatively slight extent; so that the internal friction, and conse- quently the resistance to the blood-flow, is far greater in the capillaries than in the small arteries, and in the small ar- teries than in the large ones. Practically we may regard the arteries as tubes ending in a sponge: the united areas of all the channels in the latter might be considerably larger than that of the supplying tubes, but the friction to be overcome in the flow through them would be much greater. The Conversion of the Intermittent into a Continuous Flow. Since the heart sends blood into the aorta inter- mittently, we have still to inquire how it is that the flow in the capillaries is continuous. In the larger arteries it is not, since we can feel them dilating as the "j^w^.se," by ap- plying the finger over the radial artery at the wrist, or the temporal artery on the side of the brow. The first explanation which suggests itself is that since the capacity of the blood-vessels increases from the heart to the capillaries, an acceleration of the flow during the ventricular contraction which might be very manifest in the vessels near the heart would become less and less obvi- ous in the more distant vessels. But if this were so, then when the blood was collected again fi-om the wide capillary sponge into the great veins near the heart, which together 230 THE HUM AS BODY. — cL —d' cC— B are very little bigger tlum the aorta. Ave ought to find a pulse, but we do not: the venous pulse which sometimes occurs having quite a different cause, being due to a back- flow from the auricles, or a checking of the on-flow into them, during the cardiac systole. The rhythmic flow caused by the heart is therefore not merely cloaked in the small arteries and capillaries but abolished in them. We can, however, readily contrive conditions outside the Body under which an intermittent suj^jily is transformed into a continuous flow. Sup- pose we have two vessels, A and B (Fig. 86), containing water and connected below in two ways; through the tube a on which there is a pump provided with valves so that it can only drive liquid from A to B, and through h, which may be left wide open or narrowed by the clamp c, at will. If the apjDaratus be left at rest the water will lie at the same level, d, in each vesssel. If now Ave work the pump, at each stroke a cer- tain amount of water will be conveyed from A to B, and as a result of the loAvering of the level of liquid in A and its rise in B, there will be immediately a return flow from B to A through the tube b. A, in these circumstances, Avould represent the venous system, from which the heart constantly takes blood to pump it into B, representing the arterial system; and b Avould represent the capillary vessels through which the return flow takes place: but, so far, we should have as intermittent a floAv through the capillaries, b, as through the heart-pump, a. Now imagine b to be narroAved at one point so as to oppose resistance to the back-flow, while the pump goes on working steadily. The re- sult will be an accumulation of water in B, and a fall of its level in A. But the more the difference of level in the two vessels increases, the greater is the force tending to drive water back through b to A, and more Avill floAv back, under CAUSE OF STEADY CAPILLARY BLOOD-FLOW. 331 the greatoi' difference of pressure, in a given time, until al last, when the water in B has reached a certain level, ct, and that in A has correspondingly fallen to d/', the current through b will carry hack in one minute just so much water as the pump sends the other way, and this back- flow will be nearly constant; it will not depend directly upon the strokes of the pump I)ut upon the head of water accumu- lated in B; which head of water will, it is true, be slightly increased at each stroke of the pump, but the increase will be very small compared with the whole driving force; and its influence will be inappreciable. We thus gain the idea that an incomplete impediment to the flow from the ar- teries to the veins (from Bto A in tlie diagram), such as is afforded by internal friction in the capillaries, may bring about conditions which will lead to a steady flow through the latter vessels. But in the arterial system there can be no accumulation of blood at a higher level than that in the veins, such as is supposed in the above apparatus: and we must next con- sider if the ''head of water" can be replaced by some other form of driving force. It is in fact replaced by the elas- ticity of the large arteries. Suppose an elastic bag in- stead of the vessel B connected with the pump, "a." If there be no resistance to the back-flow the current through b will be discontinuous. But if resistance be interposed. then the elastic bag will become distended, since the pum]) sends in a given time more liquid into it than it passes back through b. But the more it becomes distended the more will the bag scpieeze the liquid inside and the faster will it send that back to A, until at last its squeeze is so pow- erful that in a minute or any other unit of time it sends tniek into A as much as it receives. Thenceforth the hack-flow through b will be practically constant, being im- mediately dependent upon the elastic reaction of the bag; and only indirectly upon the action of the pump which keeps it distended. Such a state of things represents very closely the phenomena occurring in the blood-vessels. The highly elastic large arteries are kept stretched with blood by the heart; and the reaction of their elastic walls. 232 THE HUMAN BODY. steiidily squeezing on the blood in them, forces it continu- ously through the small arteries and capillaries. The steady flow in the latter depends thus on two factors: first the elasticity of the large arteries; and secondly the re- sistance to their emptying, dependent upon internal friction in the small arteries and the capillaries, which calls into play the elasticity of the large vessels. Were the capillary resistance or the arterial elasticity absent the blood -flow in the capillaries would be rhythmic. CHAPTER XVI. ARTERIAL PRESSURE. THE PULSE. V7eber's Schema. It is clear from the statements made in the last chapter that it is the pressure exerted by the elas- tic arteries upon the blood inside them which keeps up the flow through the capillaries, the heart serving to keep the bi(^ arteries tio-htly filled and so to call the elastic reaction of their walls into play. The whole circulation depends primarily of course upon the beat of the heart, but this only indirectly governs the capillary flow, and since the latter is the aim of the whole vascular apjaaratus it is of great importance to know all about arterial pressure; not only how great it is on the average but how it is altered in different vessels in various circumstances so as to make the flow through the capillaries of a given part the greater or less according to circumstances; for, as blushing and pallor of the face (which frequently occur without any change in the skin elsewhere) prove, the quantity of blood flowing through a given part is not always the same, nor is it always increased or diminished in all parts of the Body at the same time. Most of what we know about arterial pres- sure has been ascertained by experiments made upon the lower animals, from which deductions are then made con- cerning wliat happens in man, since anatomy shows that the circulatory organs are arranged upon the same plan in all the mammalia. A great deal can, however, be learnt by studying the flow of liquids through ordinary elastic tubes. Suppose we have a set of such (Fig. 87) supplied at one point Avith a pump, c, possessing valves of entry and exit which open only in the direction indicated by the arrows, and that the whole system is slightly overfilled with liquid so 234 THE HUMAN BODY. that its elastic walls are slightly stretched. These will in consequence press upon the liquid inside them and the amount of this pressure will be indicated by the gauges: so long as the pump is at rest it will be the same everywhere (and therefore equal in the gauges on B and J), since liquid in a set of horizontal tubes communicating freely, as these do at D, always distributes itself so that the pressure upon it is everywhere the same. Let the pump c now contract once, and then dilate: during the contrac- tion it will empty itself into B and during the dilatation fill itself from A. Consequently the pressure in B, indicated by the gauge x, will rise and that in A will fall. But very rapidly the liquid will redistribute itself from B to A through B, until it again exists everywhere under the same Fig. 87. — Diagram of WebPi's Schema. pressure. Every time the j^ump ^\o]'ks there will occur a similar serie.s of phenomena, and there will be a disturbance of equilibrium causing a wave to flow round the tubing; but there will be no steady maintenance of a pressure on the side B greater than that in A. Now let the ii})per tube D lie closed so that the liquid to get from B to A must flow through the narrow lower tubes D, which opi)Ose con- siderable resistance to its passage on account of their fre- quent branchings and the great internal friction in tliem; then if the pump works frcfpieiitly enough there will be produced and maintained in B a ])ressiire considerably higher than that in A, which may even become negative. If, for WEBERS SCHEMA. 235 example, the pump works 60 times a minute and at each stroke takes 180 cubic centimeters of liquid (6 ounces) from A and drives it into B, the quantity sent in at the first stroke will not (on account of the resistance to its flow oSered by the small branched tubes), have all got back into A before the next stroke takes place, sending 180 more cubic centimeters (6 oz. ) into B. Consequently at each stroke B will become more and more distended and A more ;ind more emptied, and the gauge ./; will indicate a much higher pressure than that on A. As B is more stretched, howeyer, it squeezes harder upon its contents, until at last a time comes when this squeeze is powerful enough to force through the small tubes just 180 cubic centimeters (6 oz.) in a second. Then further accumulation in B ceases. The pump sends into it 10,800 cubic centimeters (360 ounces) in a minute at one end and it squeezes out exactly that amount in the same time from its other end: and so Ions: as the pump works steadily the pressure in B will not rise, nor that in A fall, any more. But under such circumstances the flow through the small tubes will be nearly constant since it depends upon the difference in pressure prevailing between B and A, and only indirectly upon the pump which serves simjDly to keep the pressure high in B and low in ^. At each stroke of the pump it is true there will be a slight increase of pressure in B due to the fresh 180 cub. cent. (6 oz.) forced into it, but this increase will Ije but a small fraction of the total pressure and so have but an insignificant influence upon the rate of flow through the small connecting tubes. Arterial Pressure. The condition of things jii^t de- scribed represents very closely the phenomena presented in the blood-vascular system, in which the ventricles of the heart, with their auriculo-ventricular and semilunar valves, represent the pump, the smallest arteries and the capil- laries the resisttmce at B, the large arteries the elastic tube B, and the veins the tube A. The ventricles con- stantly receiving blood through the auricles from the veins, send it into the arteries, which find a difficulty in emptying themselves through the capillaries, and so blood accumu- 236 TEE 3XTMAN BODY. lates in them until the elastic reaction of the stretched ar- teries is able to squeeze in a minute through the capillaries just so much blood as the left yentricle pumps into the aorta, aud the right into tlie pulmonary artery, in the same time. Accordingly in a living animal a pressure-gauge connected with an artery shows a much higher pressure than one connected with a vein, and this persistent differ- eucc of pressure, only increased by a small fraction of the whole at each heart-beat, keeps up a steady flow from the arteries to the veins. The heart keeps the arteries stretched and the stretched arteries maintain the flow through the capillaries, and the constancy of the current in these de- pends on two factors: (1) the resistance experienced by the blood in its flow from the ventricles to the veins, and (2) the elasticity of the larger arteries which allows the blood to accumulate in them under a high pressure, in conse- quence of this resistance. The Arterial Pressure. This cannot be directly meas- ured with accuracy in man, but from measurements made on other animals it is calculated that in the human aorta its average is equal to that of a column of mercury 200 millimeters (8 inches) high. During the systole it rises about 5 millimeters (^ inch) above this and during the pause falls the same amount below it. The pressure in the venae cavae on the other hand is often negative, the blood being, to use ordinary language, often "sucked" out of them into the heart, and it rarely rises above 5 millimeters {\ inch) of mercury except under conditions (such as powerful mus- cular effort accompanied by holding the breath) which force blood on into the venae cavae and, by impeding the pulmonary circulation, interfere with the emptying of the right auricle. Hence to maintain the flow from the aorta to the vena cava we have an average difference of pressure equal to 200 — 5 = 195 millimeters. (7f inches) of mercury, rising to 205 — 5 = 200 mm. (8 inches) during the cardiac systole and falling to 195 — 5 = 190 mm. (7f inches) dur- ing the pause; but the slight alterations, only about ^ of the whole difference of aortic and vena cava pressures which maintain the blood-flow, are too slight to cause appreciable ARTERIAL PRESSURE. 237 changes in the rate of the current in the capillaries. The pressure on the Mood in the pulmonary artery is about \ of that in the aorta. Since the blood flows from the aorta to its branches and from these to the capillaries and thence to the yeins, and liquids in a set of continuous tubes flow from points of greater to those of less pressure, it is clear that the blood- pressure must constantly diminish from the aorta to the right auricle; and similarly from the pulmonary artery to the left auricle. At any point in fact the pressure is pro- portionate to the resistance in front, and since the farther the blood has gone the less of this, due to impediments at branchings and to internal friction, it has to overcome in finishing its round, the pressure on the blood always di- minishes as we follow it from the aorta to the venas cavae. In the larger arteries the fall of pressure is gradual and small, since the amount of resistance met with in the flow through them is but little. In the small arteries and capillaries the resistance passed by is (on account of the great internal friction due to their small calibre) very great, and consequently the fall of pressure between the medium- sized arteries and the veins is rapid and considerable. Modifleations of Arterial Pressure by Changes in the Rate of the Heart's Beat. A little consideration will make it clear that the pressure prevailing at any time in a given artery depends on two things — the rate at which the vessel is filled, i.e. upon the amount of work done by the heart; and the ease or difficulty with which it is emptied, that is upon the resistance in front. Returning to the system of elas- tic tubes with a pump represented in Fig. 87, let us sup- pose the pump to be driving as before 10.800 cub. cent. (360 oz.) per minute into the tubes B and that these latter are so distended that they drive out just that quantity in the same time. Under such conditions the pressure at'^any given point in B will remain constant, apart from the small variations dependent upon each stroke of the pump. Now, however, let the latter, while still sending in 180 cub. cent. (6 oz. ) at each stroke, work 80 instead of 60 times a minute and so send in that time 180 x 80 = 14,400 cub. 2??.l it.s distension i> such that \\< (jiasticiiy is able to force out in a niiiiiite thrcaigh tlit'.^mall tubes D, 14,400 cub, cent, (480oz,j, Thenceforth, so long as the pump beats with the same force and at the same rate and the peripheral resistance remains the same, the mean pressure in B will neither rise nor fall — B sending into A in a minute as much as c takes from it, and we would have a steady condition of things with a higher mean pressure jn B than before. On the other hand if the pump begins to work more slowly while the resistance remains the same, it is clear that the mean pressure in B will fall. If, for example, the pump works only forty times a minute and so sends in that time 180 X 40 = 7200 cub. cent. (240 oz.) into B, which is so stretched that it is squeezing out 10,800 cub. cent. (360 oz.) in that time, it is clear that B will gradually empty itself and its walls become less stretched and the pressure in it fall. As this takes place, however, it will force less liquid in a minute through the small tubes, until at last a pressure is reached at which the squeeze of B only sends out 7200 cub. cent. (240 oz.) in a minute; and then the fall of pressure will cease and a steady one will be main- tained, but lower than before. Applying the same reasoning to the vascular system we see that (when the iieripheral resistance remains unaltered), if the heart's force remains the same but its rate increases, arterial i)ressure will rise to a ncAV level, while a sloM'ing of the lieart's beat will bring about a fall of pressure. Modifications of Arterial Pressure Dependent on Changes in the Force of the Heart's Beat. lieturning again to Tig. 87; suppose that wliile tlie rate of the pumji remains the same, its jiower alters so that each time it sends 200 cab. cent. (6.6 oz.) insicndof 180 (6 oz.) andso in CHANGES Ilf ARTERIAL PRESSURE. 239 a minute 12,000 cub. cent. (396 oz.) instead of 10,800 (360 02.) — the quantity Avhich B is stretched enough to squeeze out in that time. Water will in consequence accunutlate in B until it becomes stretched enough to squeeze out 12,000 cub. cent. (306 oz. ) in a minute, and then a steady pressure at a new and higher level will be maintained. On the other hand if the pump, still beating sixty times a minute, ^ork* more feebly so as to send out only 160 cub. t-cut. (."). of muscles was set at work and needed an extra l)lood-snpply, this was att;iined merely by increasing the heart's activity and keeping up a faster blood-flow every- where through the Body, there would be a clear waste of force, much as if the chandeliers in a house were so ar- ranged that when a larger flame was Avanted at one burner it could only be obtained by turning more gas on at all the rest at the same time: besides the big tap at the gas-meter regulating the general supply of the house, local taps at 248 TEE HUMAN BODT. each burner are required which regulate the gas-supply to each flame independently of tlie rest, A similar arrange- ment is found in the Body. Certain nerves control the calibre of the arteries supplying different organs and, when the Litter are set at work, allow their arteries to dilate and so increase the amount of blood flowing through them while the general circulation elsewhere remains practically unaffected. The resting parts at any moment thus get Just enough blood to maintain their healthy nutrition and the working parts get more; and as certain organs come to rest and others are set in activity, the arteries of the one narrow and of the other dilate ; in this way the distri- bution of the blood in the Body is undergoing constant changes, parts which at one time contain much blood at another having but little. In addition, then, to nervous organs regulating the work of the heart and the arteries with reference to one another, we have to consider another set of vascular nerves which govern the local blood-supply of different regions of the Body. The Nerves of the Heart. The heart gets nerves from three sources. (1) From nerve-cells buried in its own sub- stance and known as its intrinsic ganglia. (2) From the tenth pair (pneumogastrics) of cranial nerves. (3) From the sympathetic nervous system. The intrinsic ganglia keep the heart beating, and the other two sets of nerves control the rate and force of the beat. The Intrinsic Heart-Nerves. The ganglia of the heart lie for the most part in the partition between the auricles and along the line of junction of the auricles and ventricles ; a few are found also in the upper parts of the latter. From some of them arise nerve-fibres which go to the muscles of the heart, while others are connected with the endings of the extrinsic nerves reaching the organ ; and probably all communicate by a network of nerve-fibres. The heart is an automatic organ : its beat, like the movement of filaments of a ciliated cell, depends on its own structure and properties and not on anything outside itself. This is proved by the fact that the heart cut out of an animal which has just been decapitated, and entirely re- CAUSE OF THE HEART'S BEAT. 249 moved from all the rest of the body, will go on beating for some time ; a time which is very short in the case of warm- blooded animals, the tissues of which die very soon when the blood-flow through them ceases, but which may extend to hours or even days in the excised heart of frog or turtle, if it be kept from drying up. Still, whether the time of its continuance be shorter or longer, the fact that the heart-beat continues after complete excision of the organ proves that it is not dependent on stimuli reaching it from other parts of the Body. In the ciliated cell we had no differentiation into muscle and nerve — its contractile and automatic parts if separated at all were not optically distin- guishable and we could only speak of the cell as still retain- ing both of those primitive protoplasmic properties. But in the heart, where we find distinct muscles and nerves, the question naturally arises in which of them does the auto- matic power reside. We have already seen (Chap. X.) that muscles elsewhere possess no automaticity : they only con- tract nnder the influence of a recognizable stimulus, and though the muscular fibres of the heart do differ somewhat from other muscular fibres in the Body, it is still aj^riori im- probable that they are automatic and we are accordingly led rather to suppose that the stimulns resides in the ganglion- cells of the heart, especially since we know that nerve-cells elsewhere are automatic. Experiment confirms this suppo- sition. If a frog's heart, removed from the body and still beating, be cut into several pieces with a sharp razor it will be found that, while bits of the auricles and the base of the ventricle go on beating, the apical portions of the ventricle lie at rest — not because the muscle there is dead and has lost its contractility, for these bits if excited by any ex- traneous muscular stimulus will still l)eat. but because that part of the heart possesses no automaticity. JSTow this is just the part of the frog's heart that has no ganglion-cells of its own, while the parts that go on beating are those which possess them ; hence we conclude that the stimulus originates in the nerve-cells of the organ and is from them carried by its nerve-fibres to the muscles. The excitant of the nerve-cells being still unknown we call them automatic 350 THE JIUMA.X BODY. in the restricted ])liy.si()logiciil sense of the word. The primary cause of the heart's beat lying thus in itself, we have next to see how this beat is controlled from outside and co- ordinated with tiie condition of the rest of tlic Body ai any given moment. Nerves Slowing the Heart's Beat. Each pneuniogas- tric trunk sends several brandies to the heart. Certain of these contain fibres which when excited slow, or even :dto- gether stop, tlie beat of tlie heart and are hence known as the cardio-inlnbitory fihrca. If one pneumogastric trunk be divided as it runs down the neck and its peripheral, or lower, end be stimulated feebly the heart's beat becomes less frequent, while a more powerful stimulation will completely stop it for a few seconds, as if its muscles were suddenly paralyzed. If the experiment be performed upon a narcotized animal. the heart of which is at the same time exposed by opening the chest, it will be seen that during the stoppage the heart lies flabby and relaxed in diastole ; the excitation of the nerve does not stop the heart's beat, as might perhaps be supjiosed, by keeping it in a state of permanent tetanic contraction, but it annuls its contractions and throws it into a state of rest ; the nerve-fibres concerned are not excitant but inhibitory, stopping instead of calling forth the activity of the part on which they act. "Whether their influence is exerted di- rectly on the muscular fibres of the heart or upon iti in- trinsic ganglia, abolishing their automatic activity and so cutting off the stimuli which normally radiate from them to the muscles, is not certainly known, but the latter view is probably the correct one. In any case the full inhibi- tory power usually lasts only a short time ; even if the pneumogastric stimulation be continued the heart will al- most always after a few seconds recover from its influence and commence to beat again. These cardio-inhibitory fibres originate in a collection of nerve-cells in the medulla oblongata known as tlie cardio- inhibitory centre. This centre is automatic and always in a state of slight excitation, feebly stimulating the fibres proceeding from it and slightly slowing the heart's beat. CARDIAV lyillDITIOX. 251 This is shown by the fact that if botli pneumogastric nerves ])Q cut in the neck the heart at once begins to beat a little faster than before; the brake, «n to speak, has been taken off it. The Influence upon Arterial Pressure of Inhibiting the Heart. If tlie heart be entirely stopped arterial jires- snre Avill of course fall very rapidly, since tlie distended arterial system Avill go on emptying itself through the capil- laries into the veins, without receiving any fresh supply at its cardiac end. So too if the heart be made to beat slower, but with the same force in each stroke, it follows from the facts pointed out in the last chapter (p. 238) that arterial pressure will fall to a new and lower level, at which the elastic arteries are only stretched enough to squeeze out in a minute as much as they receive. As a matter of fact, when the heart is made to beat slower by weak pneumo- gastric stimulation each beat is usually a little more power- ful than Ijefore. However, this extra force is not sufficient to compensate entirely for the slower rate and so the gen- eral arterial pressure falls. Use of the Cardio-Inhibitory Mechanism. Although the cardio-inhiljitory centre is automatic and always in a state of slight activity it is also greatly under the control of afferent nerve-fibres reaching it and which can arouse it to a much greater degree, and soreflexly cozitrol the heart's beat. If a frog be rendered insensible and its abdominal cavity opened, it will be found that one or two smart taps on the intestine Avill cause the heart to stop in diastole. If, how- ever, the pneumogastric nerves, or the spinal cord, or the anterior roots of the spinal nerves, or the commimicating Ijranches between the sympathetic nerves of the abdomen and the spinal nerves, be cut previously, then striking the in- testine has no influence upon the heart; nor has it if the cardio-inhibitory centre in the medulla oblongata be pre- viously destroyed. We thus get evidence that the mechani- cal stimulation of the intestinal nerves stops the heart re- ilexly through the pneumogastrics, the afferent impulses traveling from the sympathetic into the spinal nerves and passing then \\\^ the spinal cord to the cardio-inliibitory 252 THE HUMAN BODY. centre, where they are reilecled as efferent impulses down the pneumogastric trunks to tlie heart. In man and other mammals similar arrangements exist, the afferent fibres pass- ing from the alimentary canal through the solar plexus (p. 172) which lies behind the stomach. It is by exciting them and so reflexly stopping the heart, that men are sometimes killed by a severe blow on the abdomen or even occasionally by a large draught of very cold water, the sudden cold acting as a thermal stimulus, through the walls of the stomach, on the nerve-fibres outside. A hot and very thirsty jierson requir- ing a big drink should therefore not take too cold water — or if he does, swallow it only a mouthful at a time. The blood-vessels of the alimentary canal are very numer- ous and capacious and form one of the largest vascular tracts of the whole Body, and through the reflex mechanism above described we see how they may control the heart's beat. Probably if the heart is beating too frequently and keeping up too high a pressure in them, the sympathetic nerve- fibres in their coats are stimulated and then, reflexly, through the cardio-inhibitorv centre slow the heart's beat and lower the general arterial pressure ; and so we get one co-ordinating mechanism by which the heart and blood-ves- sels are made to work in unison. Some other afferent nerves are also known to be in con- nection with the cardio-inhibitory centre. For instance, some persons are made to faint by a strong smell, the olfac- tory nerves exciting the cardio-inhibitory centre and stop- ping or greatly slowing the heart. Deaths from the admin- istration of chloroform are also usually brought about in the same way, the vapor stimulating the sensory nerves of the air-passages which then excite powerfully the cardio-inhibi- tory centre and stop the heart. The Accelerator Nerves of the Heart. These originate in the spinal cord, from which tliey pass by communicating branches to the lowest cervical and upper dorsal sym- pathetic ganglia and thence to the heart. "When stimu- lated they cause the heart to beat quicker, but under what conditions they are employed in the physiological working of the Body is not known. KERVES OF THE BLOOD-VESSELS. 253 The Nerves of the Blood- Vessels. The arteries, as already pointed out, possess a muscular coat composed of fibres arranged across them, so that their contraction will narrow the vessels. This coat is most prominent in the smaller vessels, those of the size which go to supply separate organs, but disappear again in the smallest branches which are about to divide into capillaries for the individual tissue elements of an organ. These vascular muscles are under the control of certain nerves called vaso-motor (p. 186) and these latter can thus govern the amount of blood reaching any organ at a given time. The vaso-motor nerves of the arteries are, like those of the heart, intrinsic and extrinsic. The intrin- sic fibres originate from ganglion-cells in the coats of the arteries or lying alongside them, while the extrinsic origi- nate from cells in the cerebro-spinal centre, from which they commonly pass into the sympathetic system before they reach the vessels. The intrinsic ganglia, like those of the heart, are automatic and tend to keep the muscular coats of the arteries in a constant state of feeble contrac- tion so that, apart from their johysical elasticity, the arteries always hold a certain grip on the blood. The contraction, however, is as a rule persistent and steady, or tonic, instead of rhythmic like that of the heart, although slow rhythmic contractions have been seen to occur in some arteries. The difference probably depends rather on the kind of muscle concerned in each case than on the ganglion-cells, since plain muscular tissue, such as is found in the arteries, con- tracts so slowly and remains contracted so long when excited, that stimuli reaching it at intervals which would give a rhythmic beat in cardiac muscle, would keep the arterial per- manently contracted or tetanized. As in the heart, the activity of the arterial intrinsic nervous mechanism is under the control of extrinsic nerves, certain of which, the vaso-constrictors, answer to the accelerator nerves of the heart and increase the activity of the intrinsic ganglia, while others, corresponding to the cardio-inhibitory fibres, check the activity of the intrinsic vascular nerves. The Vaso-Motor Centre. The vaso-constrictor extrinsic arterial nerves are nearly always in a state of slight activity. isi 254 THE HUMAN BODY. keeping the arteries more constricted than they would be under the influence of their intrinsic nerves alone. Accord- ingly if they are cut, or paralyzed, in any region of the Body its arteries dilate and it becomes flu.slied witli blood. Those of the external ear, for example, run in the cervical sympa- thetic, from the lower part of the neck where they leave tlie spinal cord, until they reach the arterial branches for the ear and run along the smaller twigs to it. If, therefore, the cervical sympathetic be divided on one side in an anaesthetized rabbit, the ear on that side becomes red and warm from the dilatation of its arteries and the extra (^ amount of blood flowing through it. If, however, that end of the cut nerve still attached to the ear be g-jccited electri- cally or otherwise, the ear arteries contract gradually until their passage is almost closed up, and the whole organ be- comes cold and very pale. Although these vaso-constrictor fibres are thus shown to pass through the cervical sympa- thetic, other experim^ents show that they really originate in a group of nerve-cells in the medulla oblongata, and from tliere run down the spinal cord to the lower part of the neck, where they pass out in the anterior roots of some spinal nerves and reach the sympathetic system. The same is true of nearly all extrinsic vaso-constrictor nerve-fibres in the Body. Some few possibly arise from centres in the spinal cord, but the great majority come primarily from the medulla oblongata, and the collection of nerve-cells there from which they spring is known as the vaso-motor centre; a better name would be the vaso-constrictor centre. The Control of the Vaso-Motor Centre. The vaso- motor centre is automatic; that is to say it maintains a certain amount of activity of its own, indei^endently of any stimuli reaching it through afferent nerve-fibres. Never- theless, like nearly all automatic nerve-centres, it is under reflex control, so that its activity may be increased or les- sened by afferent impulses conveyed to it. Nearly every sen- sory nerve of the Body is in connection with it; any stimu- lus giving rise to pain, for example, excites it, and so constricting the arteries, increases the peripheral resist- ance to the blood-flow and raises arterial pressure. On VA80-DILAT0R NEBVE8. 255 the other hand, certain fibres conveying impulses from tlie heart inhibit the centre and dilate the arteries, lower blood-pressure, and diminish the resistance to be overcome by the heart. These fibres run in branches of the j)iieumo- gastric, and are known as the cleiyressor fibres, or in certain animals, for example the rabbit, where they are all collected into one branch, as the depressor nerve. If this nerve be divided and its cardiac end stimulated no effect is pro- duced, but if its central end (that still connected with the rest of the pneumogastric trunk and through it with the medulla oblongata) be stimulated, arterial pressure gradually falls; this result being dependent upon a dilata- tion of the small arteries, and consequent diminution of the peripheral resistance, following an inhibition of the vaso-motor centre brought about by the depressor nerve. Through the depressor nerve the heart can therefore influ- ence the calibre of the small arteries and, by lowering aortic pressure, diminish its own work if need be. Blushing. The depressor nerves control a great part of the vaso-motor centre, and so can bring about dilatation of a large number of arteries — their influence is called into play when general arterial pressure is to be lowered, but is useless for controlling local blood-supply. This is man- aged by other afferent nerves, each of which inhibits a small part only of the vaso-motor centre, governing the arteries of a limited tract of the Body; the dilatation of these increases the amount of blood flowing through the particular region to which they are distributed, but does not affect the total resistance to the blood-flow suflBciently to influence noticeably the general pressure in the arterial system. In blushing, for example, under the influence of an emotion, that part of the vaso-motor centre which sup- plies constrictor nerves to the arteries of the skin of the neck and face, is inhibited by nerve-fibres proceeding from the cerebrum to the medulla oblongata, and the face and neck consequently become full of blood and flush up. Quite similar phenomena occur under other conditions in many parts of the Body, although when not visible on the surface we do not usually call them blushes. The mucous 256 THE HUMAN BODY. membrane lining the empty stomacli is pallid and its ar- teries contracted, but as soon as food enters the organ it becomes rod and full of blood; the food stimulating afferent nerve-fibres there, which inhibit tliat part of the vaso-motor centre which governs the gastric arteries. Taking Cold. This common disease is not unfrcquent- ly caused through undue reflex excitement of the vaso- motor centre. Cold acting upon the skin stimulates, through the afferent nerves, the region of the vaso-motor centre gov- erning the skin arteries, and the latter become contracted, as shown by the jDallor of the surface. This has a two-fold influence — in the first place, more blood is thrown into in- ternal parts, and in the second, contraction of the arte- ries over so much of the Body considerably raises the gen- eral blood-pressure. Consequently the vessels of internal parts become overgorged or " congested," a condition which readily passes into inflammation. Accordingly prolonged exposure to cold or wet is apt to be followed by catarrh or inflammation of more or less of the respiratory tract caus- ing bronchitis, or of the intestines causing diarrhoea. In fact the common summer diarrhoea is far more often due to a chill of the surface, causing intestinal catarrh, than to the fruits eaten in that season which are so often blamed for it. The best preventive is to wear, when exposed to great changes of temperature, a woolen or at least a cotton garment over the trunk of the Body; linen is so good a conductor of heat that it permits any change in the exter- nal temperat»re to act almost at once upon the surface of the Body. After an unavoidable exposure to cold or wet the thing to be done is of course to maintain the cutaneous circulation; for this purpose movement should be persisted in, or a thick dry outer covering put on, until warm and dry clothing can be obtained. For healthy persons a temporary exposure to cold, as a plunge in a bath, is good, since in them the sudden contrac- tion of the cutaneous arteries soon jiasses off and is suc- ceeded by a dilatation causing a warm healthy glow on the surface. If the bather remain too long in cold water, how- ever, this reaction passes off and is succeeded by a more VASO-DILATOR NERVES. 267 persistent chilliness of the surface, which may eyen last ail day. The bath should therefore be left before this occurs, but no absolute time can be stated, as the reaction IS more marked and lasts longer in strong persons, and in those used to cold bathing, than in others. Vaso-Dilator Nerves. We have already seen, in the case of the stomach, one method by which a locally increased blood-supply may be brought about in an organ while it is at work. Usually, however, in the Body this is managed in another way; by vaso-dilator nerves which inhibit or paralyze, not the vaso-motor centre, but the intrinsic nerv- ous supply of the blood-vessels. The nerves of the skeletal muscles for example contain two sets of fibres: one motor proper and the other vaso-dilator. When the muscle con- tracts in a reflex action or under the influence of the will both sets of fibres are excited; so that when the organ is set at work its arteries are simultaneously dilated and more blood flows through it. Quite a similar thing occurs in the sali- vary glands. Their cells, which form the saliva, are aroused to activity by special nerve-fibres; but the gland nerve also contains vaso-dilator fibres which simultaneously cause a dilatation of the gland artery. Through such arrange- ments the distribution of the blood in the Body at any moment is governed: so that working parts shall have abundance and other parts less, while at the same time the general arterial pressure remains the same on the average; since the expansion of a few small local branches but little influences the total peripheral resistance in the vascular sys- tem. Moreover, commonly when one set of organs is at work with its vessels dilated, others are at rest with their arteries comparatively contracted, and so a general average blood-pressure is maintained. Few persons, for example, feel inclined to do brain work after a heavy meal: for then a great part of the blood of the whole Body is led off into the dilated vessels of the digestive organs, and the brain gets a smaller supply. On the other hand, when the brain is at work its vessels are dilated and often the whole head flushed: and so excitement or hard thought after a meal 7« tery apt to produce an attack of indigestion, by diverting 258 THE HUMAN BODY. the blood from the abdominal organs where it ought to be at that time. Young persons, whose organs have a super- abundance of energy enabling them to work under unfavor- able conditions, are less apt to suffer in such ways than their elders. One sees boys running actively about after eating, when older people feel a desire to sit quiet and ru- minate— or even go to sleep. CHAPTER XVIII. THE SECEETORY TISSUES AND ORGANS. Definition. In a strict sense of the terms every pro- cess in which substances are separated from the blood, whether they be altered or unaltered, is "secretory" and every product of such a separation is a " secretion;" in this sense secretions would be separable into three classes. " (1) Liquids or gases transuding on free surfaces of the Body, whether external or internal; (2) the liquids {lymph) moistening the various tissues of the Body directly, filling the interstices between them and not contained in definitely limited cavities ; (3) all the solid tissues of the Body since, after an early period of embryonic life, they are built up from materials derived from the blood. Secretions would thus come to include all constituents of the Body except the blood itself but, while it is well to bear in mind that the whole Body is in such a way derived from the blood, in practice the term secretion is given a narrower connota- tion, the solid tissues and the lymph being excluded; so that a secretion is a material (liquid or gaseous) derived from the blood and poured out on a free surface, whether that of the general exterior or that of an internal cavity. Such true secretions fall into two classes; one in which the product is of no further use in the Body and is merely separated for removal, as the urine; and one in which the jiroduct is intended to be used, for instance as a solvent in the diges- tion of food. The former gi-oup are sometimes distin- guished as excreiions and the latter as secretions proper, but there is no real difference between them, the organs and pro- cesses concerned being fundamentally alike in each case. A better division is into transiidata tindi secretions, atransuda- 260 THE HUMAN BODY. tion being a product which contains nothing which did not previously exist in the blood, and then in such quantity as might be derivable from it by merely physical processes; while a secretion in addition to transudation elements con- tains a specific element, due to the special physiological activity of the secretory organ; being either something which does not exist in the blood at all or something wliicli. existing in the blood in small quantity, exists in the secre- tion in such a high proportion that it must have ])een actively picked up and conveyed there by the secretoi-y tissues concerned. For instance, the gastric juice contains free hydrochloric acid which docs not exist in the blood; and the urine contains so much urea that we must sujipose its cells to have a peculiar power of removing that body from the liquids flowing near them. This subdivision is also justifiable on histological grounds ; wherever there is. a secreting surface it is covered with cells, but these where transudata are formed (as on the serous membranes) are mere flat scales, with little or no protoplasm remaining in them, while the cells which line a true secreting organ are cuboidal, spherical, or columnar, and still retain, with their high physiological activity, a good deal of their primi- tive protoplasm in a but slightly modified state. Organs of Secretion. The simplest form in which a secreting organ occurs {A, Fig. 88) is that of a flat membrane provided with a layer of cells, a, on one side (that on which the secre':ion is poured out) and with a network of capil- lary blood-vessels, c, on the other. The dividing mem- brane, h, is known as the basement memlirane and is usually made up of flat, closely fitting connective-tissue corpuscles; supporting it on its deep side is a layer of connective tissue, d, in which the blood-vessels and lymphatics are supported. Such simple forms of secreting surfaces are found on the serous membranes but are not common; in most cases an extended area is required to form the necessary amount of secretion, and if this were attained simply by spreading out plane surfaces, these from their number and extent would be hard to pack conveniently in the Body, Accordingly in most cases, the greater area is attained by folding the FORMS OF GLANDS. 261 Fig. 88.— Forms of glands. A, a simple secreting surface; a. us epithelium ; 6. basement membraue: c. capillaries: B.a. simple tubular gland : C, a secret- ing surface increased by protrusions; E. & simple racemose gland ; D and Cr, compound tubular glands: F.a. compound racemose gland. In all but i^ the capillaries are omitted for the snke of clearness. H, half of a highly devel- oped i-acemose gland: c, its main duci. The letter rf by mistake has been usea both for the basement membrane above, and tor branches of duct below 262 THE HUMAN BODY. secreting surface in various ways so that a large surface can be packed in a small bulk, just as a Chinese lantern when shut up occupies much less space than when extended, although its actual surface remains of the same extent. In a few cases the folding takes the form of protrusions into the cavity of the secreting organ as indicated at C, Fig. 88, and found on some synovial membranes; but much more commonly the surface extension is attained in another way, the basement membrane, covered by its epithelium, being pitted in or involuted as at B. Such a secreting organ is known as a gland. Forms of Glands. In some cases the surface involu- tions are uniform in diameter, or nearly so, throughout {B, Fig. 88). Such glands are known as Uihular; examples are found in the lining coat of the stomach (Fig. 97*); also in the skin (Fig.l20f), where they form the stveat-glands. In other cases the involution swells out at its deeper end and becomes more or less sacculated {B) ; such glands are racemose or aciiiotcs. The small glands which form the oily matter poured out on the hairs (Fig. 119;};) are of this t^'pe. In both kinds the lining cells near the deeper end are commonly different in character from the rest; and around that part of the gland the blood-vessels form a closer network. These deeper cells form the true secreting elements of the gland, and the passage, lined with different cells, leading from them to the surface, and serving merely to carry off the secretion, is known as the gland dud. When the duct is undivided the gland is simple; but when, as is more usual, it is branched and each branch has a true secreting part at its end, we get a compound gland, tubular {G) or racemose {F, H) as the case may be. In such cases the main duct, into which the rest open, is often of considera- ble length, so that the secretion is poured out at some dis- tance from the main mass of the gland. A fully formed gland, H, thus comes to be a complex structure, consisting primarily of a duct, c, ductules, dd, and secreting recesses, ee. The ducts and ductules are lined with epithelium which is merely protective and differs in cliurueter from tlie secreting ei^ithelium which lines the *P. 319. iP. 4ia i P. 416. PROCESSES CONCERNED IN SECRETION. 263 deepest parts. Surrounding each subdivision and bind- ing it to its neighbors is the gland stroma formed of con- nective tissue, a layer of which also commonly envelops the whole gland, as its capside. Commonly on looking at the surface of a large gland it is seen to be separated by partitions of its stroma, coarser than the rest, into lohes, each of which answers to a main division of the primar}^ duct; and the lobes are often similarly divided into smaller parts or lohules. In the connective tissue between the lobes and lobules blood-vessels jienetrate, to end in fine capillary vessels around the terminal recesses. They never pene- trate the basement membrane. Lymphatics and nerves take a similar course; but there is reason to believe the nerve-fibres penetrate the basement membrane and be- come directly united with the secreting cells. The Physical Processes in Secretion. From the struc- ture of a gland it is clear that all matter, derived from the blood and poured into its cavity, must pass not only through the walls of the capillary blood-vessels, but also, by filtra- tion or dialysis, through the basement membrane and the lining epithelium. By filtration is meant the passage of a fluid under pressure through tlie coarser mechanical pores of a membrane, as in the ordinary filtering processes of a chemical laboratory; and the higher the pressure on the liquid to be filtered the greater the amount which, other things bemg equal, will pass through in a given time. Since in the living Body the liquid pressure in the blood capillaries is nearly always higher than that outside them, filtration is apt to take place everywhere to a greater or less extent, and will be increased in amount in any region by circumstances raising bloou-pressure there, and diminished by those lowering it. To a certain extent also the nature of the liquid filtered has an influence. True solutions, as those of salt in water, pass through unchanged; but solu- tions containing substances such as boiled starch or raw egg albumen, which swell up greatly in water rather than truly dissolve, are altered by filtration; the filtrate contain- mg less of the imperfectly dissolved body than the unfil- tered liquid. The higher the pressure the greater the pro- 264 THE HUMAN BODY. portion of such substances ■which gets through; and if the pressure is shght the water or other solvent may alone pass, leaving all the rest behind on the filter. Under moderate pressure the blood may thus lose by filtration such bodies only as water and salines; while an increase of arterial pressure may lead to the passage of albumen and fibrinogen. Under healthy conditions, for example, the urine contains no albumen, but anything increasing the capillary pressure in the kidneys will cause it to appear. Dialysis or osmosis has already been considered (p. 42); by it sub- stances pass through the intermolecular pores of a mem- brane independently of the joressure on either side, and for its occurrence two liquids of ditf erent chemical constitution are required, one on each side of the membrane. At least if diffusion takes place, as is probable, between two exactly similar solutions, the amount and character of the sub- stances passing opposite ways in a given time are exactly equal, so that no change is produced by the dialysis; which practically amounts to the same thing as if none occurred. When a solution is placed on one side of a membrane allow- ing of dialysis and pure water on the other, it is found that for every molecule of the dissolved body that passes one way a definite amount of water, called the endosmotic equivalent of that body, passes in the opposite direction. Crystalline bodies as a rule (haemoglobin is an exception) have a low endosmotic equivalent or are readily dialyzable; while colloids such as gum and proteids, have a very high one, so that to get, by dialysis, a small amount of albumen through a membrane, a practically infinite amount of water must pass the other way. Accordingly, if we find such bodies in a secretion we cannot suppose that they have been derived from the blood by osmosis. The Chemical Processes of Secretion. As above point- ed out certain secretions, called transudata, seem to be pro- ducts of filtration and dialysis alone, containing only such substances as those which are found in the blood plasma, more or less altered in relative quantity by the ease or diffi- culty with which they severally passed through the layers met with on their way to the sui-face. But in many cases THE SPECIFIC ELEMENTS OF SECRETIONS. 265 the composition of a secretion cannot be acconntcd for in this way; it contains some specific element, either a substance which does not exist in the blood at all and must therefore have been added by the secreting membrane, or some body which, although existing in the blood, does so in such minute proportion compared with that in which it is found in the secretion, that some special activity of the secreting cells is indicated; some affinity in them for these bodies by which they actively pick them up. Each living cell, we have seen, is the seat of constant chemical activity, taking u]) materials from the medium about it, transforming and utilizing them, and sooner or later restoring their elements, differently combined, to the medium again. By such means it builds up and maintains its living substance, and obtains energy to carry on its daily work. While this is true of all cells in the Body, we find certain groups in which chemical metabolism is the promi- nent fact; cells vrhich are specialized for this purpose just as muscular fibre is for contraction or a nerve-fibre for con- duction, and certain of these prominently metaholic tissues, exist in the true glands and produce or collect the specific elements of their secretions. Their chemical processes are no doubt primarily directed to their own nutritive mainte- nance; they live primarily for themselves, but their nutritive processes are such that the bodies formed in them and sent . into the secretion are such as to be useful to the rest of the cells of the community; or the bodies which they specially collect, and in a certain sense feed on, are those the re- moval of which from the blood is essential for the general good. Their individual nutritive peculiarities are utilized for the welfare of the whole Body. The Mode of Activity of Secretory Cells. If we con- sider the modes of activity of living cells in general, it be- comes clear that secretory cells may produce the specific element of a secretion in either of two ways. They may, as a by-result of their living play of forces, produce chemi- cal changes in the surrounding medium; or they may build up certain substances in themselves and then set them free as specific elements. Yeast, for example, in a saccharine 266 THE HUMAN BODY. solution causes the rearrangement into carbon dioxide, alcohol, glycerine and succinic acid, of many atoms of car- bon, hydrogen and oxygen Avhich previously existed as sugar; and which during the metamorphosis were probably not j)assed through the living cell. How the latter acts we do not know with certainty, but most likely by picking certain atoms out of the sugar molecule, and leaving the rest to fall down iruto simpler compounds. On the other hand, we find cells forming and storing up in themselves large quan- tities of substances, which they afterwards liberate; starch, for instance, being formed and laid by in many fruit- cells, and afterwards rendered soluble and passed out to nourish the young jDlant. Ghmd-cells might a priori give rise to the specific ele- ments of secretions in either of these ways and we have to seek in which manner they work. Do they simply act as ferments (however that is) ujDon the surrounding medium; or do they form the special bodies which charac- terize their secretion, first within their own substance, and then liberate them, either disintegrating themselves or not at the same time? At present there is a large and an increasing mass of evidence in favor of the second view. There is, no doubt, some reason to believe that every living cell can act more or less as a ferment upon certain solu- tions should they come into contact with it. Not always, of course, as an alcoholic ferment, though even as regards that one fermentative power it seems very generally pos- sessed by vegetable cells, and there is some evidence that alcohol is normally produced in small amount (and presum- ably by the fermentation of sugar) under the influence of certain of the living tissues of the Human Body. As re- gards distinctively secretory cells, however, the evidence is all the other way, and in many cases we can see the specific element collecting in the gland-cells before it is set free in the secretion. For example, in the oil-glands of the skin (Chap. XXVII.) we find tlie secreting cells, at first granular, nucleated and protoplasmic, gradually undergoing changes by which their protoplasm disappears and is rejDlaced by oil-droplets, until finally the whole cell falls to bits and its THE ACTION OF GLAND-CELLS. 267 detritus forms the secretion; the cells being replaced by new- ones constantly formed within the gland. In such cases the secretion is the ultimate product of the cell life; the result of degenerative changes of old age occurring in it. In other cases, however, the liberation of the specific element is not attended with the destruction of the secret- ing cell; as an example we may take the pancreas, wdiich is a large gland lying in the abdomen and forming a secre- tion used in digestion. Among others, this secretion pos- sesses the power, under certain conditions, of dissolving proteids and converting them into dialyzable peptones (p. 11). This it owes to a specific element known as tryp- sin, the formation of which within the gland-cells can be ' traced with the microscope. The pancreas, like the majority of the glands connected with the alimentary canal, has an intermittent activity; determined by the presence or absence of food in various parts of the digestive tract. If the organ be taken from a recently killed dog which has fasted thirty hours and, after proper preparation, be stained with carmine and examined microscopically, we get specimens of what we may call the " resting gland '' — a gland which has not been secreting for some time. In these it will be seen that the cells lining the secreting recesses present two very distinct zones; an outer next the basement membrane which does not combine wath the coloring matter and is granular, and an inner which is '^^/c^y not granular but picks up the carmine. The granules w^e shall find to be indications of the presence of a trypsin- vieldino- substance, formed in the cells. ^yl If another dog be kept fasting until he has a good appe- "^ti, tite and be then allowed to eat as much meat as he will, he will commonly take so much that the stomach will only be emptied at the end of about twenty hours. This period may, so far as the pancreas is concerned, be divided into two. From the time the food enters the stomach and on for about ten hours, the gland secretes abundantly; after that the secretion dwindles, and by the end of the second ten hours has nearly ceased. "We have, then, a time during which the pancreas is working hard, followed by a period =».> 268 TEE HUMAN BODY. in which its activity is very little, but during which it is abundantly supijlied with food materials. The pancreas taken from an animal at the end of the first period and prepared for microscopic examination will be found dif- ferent from that taken from a dog killed at the end of the second digestion period, and also from the resting gland. Towards the end of the period of active work, the gland-cells are diminished in size and the proportions of the granular and non-granular zones are quite altered. The latter now occupies most of the cell, while the granular non-staining inner zone is greatly diminished. During the secretion there is, therefore, a growth of the non-granular and a de- struction of the granular zone; and the latter process rather exceeding the former, the whole secreting cell is diminished in size. During the second digestive period, when secre- tion is languid, exactly a reverse process takes place. The cells increase in size so as to become larger than those of the resting gland; and this growth is almost entirely due to the granular zone which now occupies most of the cell. These facts suggest that during secretion the granular part of the cells is used up: but that, simultaneously, the deeper non-granular zone, being formed from materials yielded by the blood, gradually gives rise to the granular. During active secretion the breaking doAvn of the lat- ter to yield the sjiecific elements occurs faster than its re- generation; in a later period, however, when the secretion is ceasing, the whole cell grows and, especially, the granular zone is formed faster than it is disintegrated; hence the great increase of that part of the cell. If this be so, then we ought to find some relationship between the diges- tive activity of an infusion or extract of the gland and the size of the granular zones of the cells; and it has been shown that such exists; the quantity of trypsin which can be obtained from a pancreas being proportionate to the size of that portion of its cells. The trypsin, however, does not exist in the cells ready formed, but only a body which yields it under certain cir- cumstances, and called zymogen. If a perfectly fresh pancreas be divided into halves and INFLTJENCE OF NERVES ON SECRETION. 269 one portion immediately minced and extracted with glyce- rine, while the other is laid aside for twenty-four hours in a warm place and then similarly treated, it will be found that the first glycerine extract has no power of digesting proteids, while the second is very active. In other words the fresh gland does not contain trypsin, but only sonu^- thing which yields it under some conditions; among others, on being kept. The inactive glycerine extract of the fresh gland is however rich in zymogen: for if a little acetic acid be added to it, trypsin is formed and the extract becomes powerfully digestive. f We may then sum up the life of pancreas cell in this way. It grows by materials derived from the blood and first laid down in the non-granular zone. This latter, in the ordinary course of the cell-life, gives rise to the granu- lar zone; and in this is a store of zymogen produced by the nutritive metabolisms of the cell. When the gland secretes, the zymogen is converted into try^isin and set free in the secretion; but in the resting gland this transforma- tion does not occur. During secretory activity therefore the chemical processes taking place in the cell, are different from those at other periods; and we have next to consider how this change in the mode of life of the cells is brought about. Influence of the Nervous System upon Secretion. When the gland is active it is fuller of blood than when at rest: its arteries are dilated and its capillaries gorged so that it gets a brighter pink color; this extra blood-supply might be the primary cause of the altered metabolism. Again, the activity of the pancreas is under the influence of the nervous system, as evinced not only by the reflex secretion called forth when food enters the stomach, but also by the fact that electrical stimulation of the medulla oblongata will causo the gland to secrete. The nervous system may, however, only act through the nerves governing the calibre of the gland arteries, and so but indirectly on the secreting cells; while on the other hand, it is possible that nerve-fibres act directly upon tlie gland-cells and, con- trolling their nutritive processes, govern the production of 270 THE HUMAN BODY. the trypsin. To decide between the relative importance of these possible agencies we must pass to the consideration of other glands; since the question can only be decided by experiment upon the lower animals, and the position of the pancreas and the difficulty of getting at its nerves with- out such severe operations as upset the physiological condi- tion of the animal, furnish obstacles to its study which have not yet been overcome. In certain other glands, however, we find conclusive evi- dence of a direct action of nerve-fibres upon the secreting elements. If the sciatic nerve of a cat be stimulated elec- trically the balls of its feet will sweat. Under ordinary circumstances they become at the same time red and full of blood; but that this congestion is a factor of subsidiary importance as regards secretion is proved by the facts that stimulation of the nerve is still able to excite the gland- cells and cause sweating in a limb which has been ampu- tated ten or fifteen minutes (and in which therefore no cir- culatory changes can occur) and also by the cold sweats, with a pallid skin, of phthisis and the death agony. It is, however, with reference to the submaxillary and parotid salivary glands that our information is most precise. When the mouth is empty and the jaws at rest the sali- vary secretion is comparatively small: but a sapid substance placed on the tongue will cause a copious flow. The phe- nomenon is closely comparable to the production of a reflex muscular contraction. A stimulus acting upon an irritable tissue excites through it certain afferent nerve-fibres; these excite a nerve-centre, which in turn stimulates efferent fibres; going to a muscle in the one case, to a gland in the other. It will be useful to consider again for a moment what occurs in the case of the muscle, taking account only of the efferent fibres and the j)arts they act upon. When a muscle in the Body is made to contract reflexly, through its nerve, two events occur in it. One is the shortening of the muscular fibres; the other is the dilata- tion of the muscular arteries; every muscular nerve con- tains two sets of fibres, one motor and one vaso-dilator, and normally botli act together. In tliis case, however, INFLUENCE OF NERVES ON SECRETION 271 it is clear that the actiYities of both, though correlated, are essentially independent. The contraction is not due to the greater blood-flow for, not only can an excised muscle en- tirely deprived of blood, be made to contract by stimulating its nerves, but in an animal to which a small dose of curari — ^the arrow poison of certain South American Indians — has been given, stimulation of the nerve will cause the vascu- lar dilatation but no muscular contraction: the curari par- alyzing the motor fibres, but, unless in large doses, leaving the vaso-dilators intact. The muscular fibres themselves are quite unacted upon by the poison, as evinced by their ready contraction when directly stimulated by an electric shock. Now let us return to the salivary glands and see how far the facts are comparable. The main nerve of the submax- illary gland is known as the cliorda iijmiiani. If it be di- vided in a narcotized dog, and a tube placed in the gland- duct, no saliva will be found to flow. But on stimulating the peripheral end of the nerve (that end still connected with the gland) an abundant secretion takes place. At the same time there is a great dilatation of the arteries of the organ, much more blood than before flowing through it in a given time: the cliorda obviously then contains vaso- dilator fibres. Now in this case it might very well be that the process was different from that in a muscle. It is con- ceivable that the secretion may be but a filtration due to increased pressure in the gland capillaries, consequent on dilatation of the arteries supplying them. If a greater filtration into the lymph spaces of the gland took place, this liquid might then merely ooze on through the secreting cells into the commencing ducts and, as it passed through, dis- solve out and carry on from the cells the specific organic elements of the secretion. Of these, in the submaxillary of the dog at least, mucin is the most important and abundant. That, however, the process is quite different, and that there are in the gland true secretory fibres in ad- dition to the vaso-dilator, just as in the muscle there are true motor fibres, is proved by other experiments. If the flow of liquid from the excited gland were merely 272 THE HUMAN BODY. the outcome of a filtration dependent on increased blood pressure in it, then it is clear that the pressure of the secretion in the duct could never rise above the pressure in the blood-vessels of the gland. Now it is found, not only that the gland can be made to secrete in a recently decapi- tated animal, in which of course there is no blood-pressure, ])ut that, when the circulation is going on, the pressure of the secretion in the duct can rise far beyond that in the gland arteries. Obviously, then, the secretion is no ques- tion of mere filtration, since a liquid cannot filter against a higher pressure. Finally, the proof that the vascular dila- tation is quite a subsidiary phenomenon has been com- pleted by showing that we can produce all the increased blood-flow through the gland without getting any secretion — that just as in a muscle nerve we can, by curari, paralyze the motor fibres and leave the vaso-dilators intact, so we can by atropin, the active principle of deadly night-shade, get similar phenomena in the gland. In an atropized animal stimulation of the chorda jn-oduces vascular dila- tation but not a droj? of secretion. Bringing blood to the cells abundantly, will not make them drink; we must seek something more in the chorda than the vaso-dilator fibres — some proper secretory fibres; that the poison acts upon them and not upon the gland-cells, is shown, as in the muscle, by the fact that the cells still are capable of activity when stimulated otherwise than through the chorda tympani. For exami)le, by stimulation of the sym- pathetic fibres going to the gland. So far then we seem to have good evidence of a direct action of nerve-fibres upon the gland-cells. But even that is not the whole matter. It is extremely probable, if not certain, that there are two sets of secretory fibres in the gland-nerves: a set which so acts ujoon the cells as to make them pass on more abundantly the transudation elements of the secretion (the water and mineral salts), and another, quite different, which governs the chemical transformations of the cells so as to make them produce mucin from matters previously stored in them, in a comparable way to the ])ro- duction of trypsin from zymogen in the active pancreas. INFLUENCE OF NEBVES ON SECRETION. 273 These latter fibres may be called ''trophic," since they directly control the cell metabolism: while the former may be called -'transudatory" fibres. Some of the evidence which leads to this conclusion is a little complex, but it is worth while to consider it briefly. In the first place, on stimulation of the chorda of an unexhausted gland (that is a gland not over-fatigued by previous work) the following points can be noted: — With increasing strength of the stimulus the quantity of the secretion, that is of the water poured out in a unit of time, increases; at the same time the mineral salts also increase, but more rapidly, so that their percentage in a rapidly formed secretion is greater than in a more slowly formed, up to a certain limit. The percentage of organic constituents of the secretion also increases \\^ to a limit; but soon ceases to rise, or even falls again, while the water and salts still increase. This of course is readily intelligible; since the water and salts can be derived continually from the blood, while the specific elements, coming from the gland-cells, may be soon exhausted; and so far the experi- ment gives no evidence of the existence of distinct nerve- fibres for the salts and water, and for the specific elements: all vary together with the strength of the stimulus applied to the nerve. But under slightly different circumstances their quantities do not run parallel. The proportion of specific elements in the secretion is largely dependent on whether the gland has been previously excited or not. Prior stimulation, not carried on of course to exhaustion, largely increases the percentage of organic matters in the secretion produced by a subsequent stimulation; but has no effect whatever on the quantity of water or salts. These are governed entirely by the strength of the second stimu- lation. Here, then, we find that under similar circumstances the transudatory and specific elements of the secretion do not vary together; and are therefore probably dependent upon different exciting causes. And the facts might lead us to suspect that there are in the chorda, besides the vaso- dilator, two other sets of fibres: one governing the salts and water, and the other the specific elements of the secre- 274 THE HUMAN BODY. tiou. The evidence is, perhai)s, not quite conclusive, but experiments upon the parotid gland of the dog put the matter beyond a doubt. The submaxillary gland receives fibres from the sympa- thetic system, as well as the cliorda tympani from the cerebro-s])inal. Excitation of the sympathetic filn'es causes the gland to secrete, but the saliva poured out is differ- ent from that following chorda stimulation, which is in the dog abundant and comparatively i)oor in organic constituents, and accompanied by vascular dilatation: while the "sympathetic saliva," as it is called, is less abundant, very rich in mucin, and accompanied with constriction of the gland arteries. According to the above view we might suppose that the chorda contains many transuda- tory and few trophic fibres, and the sympathetic many trophic and few transudatory. It might, however, well be objected that the greater richness in organic bodies of the symiiathetic saliva was really due to the small quantity of blood reaching the gland, when that nerve was stimulated. This might alter the nutritive phenomena of the cells and cause them to form mucin in unusual abundance, in which case the tro|)hic influence of the nerve would be only in- direct. Experiments on the parotid preclude this explan- ation. That gland like the submaxillary gets nerve-fibres from two sources: a cerebral and a sympathetic. The latter enter the gland along its artery, while the former, origin- ating from the glosso-phar^mgeal, run in a roundabout course to the gland. Stimulation of the cerebral fibres causes an abundant secreticm, rich in water and salts, but with hardly any organic constituents. At the same time it produces dilatation of the gland arteries. Stimulation of the sympathetic causes contraction of the parotid gland arteries and no secretion at all. Nevertheless it causes great changes in the gland-cells. If it be first stimulated for a Avhile and then the cerebral gland-nerve, the resulting secretion may be ten times as rich in organic bodies as that obtained without previous stimulation of the sympathetic; and a similar phenomenon is observed if the two nerves be stimulated simultaneously. So that the sympathetic nerve. INFLUENCE OF NEEVES ON SECRETION. 275 though, unable of itself to cause a secretion, brings about great chemical changes in the gland-cells. It is a distinct trophic nerve. This conclusion is confirmed by histology. Sections of the gland after prolonged stimulation of the sym- pathetic show its cells to be quite altered in appearance, and in their tendency to combine with carmine, when com- pared either with those of the resting gland or of the gland which has been made to secrete by stimulating its glosso- pharyngeal branch alone. We have still to meet the objection that the sympathetic fibres may be only indirectly trophic, governing the meta- bolism of the cells through the blood-vessels. If this be so, cutting off or diminishing the blood-supply of the gland, in any way. ought to have the same result as stimula- tion of its sympathetic fibres. Experiment shows that such is not the case and reduces us to a direct trophic influ- ence of the nerve. When the arteries are closed and the cerebral gland-nerve stimulated, it is found that the per- centage of organic constituents in the secretion is as low as usual; it remains almost exactly the same whether the arteries are open or closed or have been previously open or closed. We must conclude that the peculiar influence of the sympathetic does not depend upon its vaso-con stricter fibres. These observations make it clear that the phenomena of secretion are dependent on very complex conditions, at least in the salivary glands and presumably in all others. Primarily dependent upon filtration and dialysis from the blood-vessels and the physiological character of the gland- cells, both of these factors are controlled by the nervous system, the secretory tissues being no more automatic than the muscular; and the facts also give us important evidence of power of the nervous system to influence cell nutrition directly. Summary. By secretion is meant the separation of such substances from the blood as are poured out on free surfaces of the Body, whether external or internal. In its simplest form it is merely a physical process dependent on filtra- tion and dialysis; for example, the elimination of carbon dioxide from the surfaces of the lungs, and the watery 276 THE HUMAN BODY. li(iuid poured out on the surfaces of the serous membranes. Such secretions are known as transiidata and their amount is only indirectly controlled by the nervous system, through the influence of the latter upon tlie circulation of the blood. The cells lining such surfaces are not secretory tissues in any true sense of the Avord, being merely flat, inactive, thin scales protecting the surfaces. In other cases the lining cells are thicker, and actively concerned in the process; they are then usually spread over the recesses of a much folded membrane, so that the whole is rolled up into a compact organ called a gland, the secretion of which may contain only transudation elements (as for example that of the lachrymal glands which form the tears) or may contain a specific element, formed in the gland by its cells, in addition to transudation elements. In either case the activity of the organ is directly influenced by the nervous system, usually in a reflex manner {e.g. the watering of the eyes when the eyeball is touched and the saliva poured into the mouth when food is tasted) but may also be otherwise ex- cited, as for example the flow of tears under the influence of those changes of the central nervous system which are associated with sad emotions, or the watering of the mouth at the thought of dainty food. The nerves going to such glands, besides controlling their blood-vessels, act upon the gland-cells; one set governing the amount of transudation of water and salines which shall take place through them, and another (in the case of glands producing secretions with one or more specific elements) controlling the produc- tion of these, by starting new chemical processes in the cells by which a substance built up in them during rest is con- verted into the specific element, which is soluble in and carried off by the transudation elements. What the speci- fic element of gland shall be, or whether its secretion con- tain any, is dependent on the nature of its special cells; how much transudation and how much specific element shall be secreted at any time is controlled by the nervous system; just as the contractility of a muscle depends on the endowments of muscular tissue, and whether it shall rest or contract — and if the latter how powerfully — upon its nerve. CHAPTER XIX. THE INCOME AND EXPENDITUKE OF THE BODY. The Material Losses of the Body. All day long while life lasts each of us is losing something from his Body. The air breathed into the lungs becomes in them laden with carbon dioxide and water vajoor, which are carried off with it when it is expired. The skin is as constantly giv- ing off moisture, the total quantity in twenty-four hours being a good deal, even when the amount passed out at any one time is so small as to be evaporated at once and so does not collect as drops of visible perspiration. The kidneys again are constantly at work separating water and certain crystalline nitrogeneous bodies from the blood, along with some mineral salts. The product of kidney activity, how- ever, not being forthwith carried to the surface but to a reservoir, in which it accumulates and Avhich is only emptied at intervals, the activity of those organs appears at first sight intermittent. If to these losses we add cer- tain other waste substances added to the undigested residue of the food passed out from the alimentary canal, and the loss of hairs and of dried cells from the surface of the skin, it is clear that the total amount of matter removed from the Body daily is considerable. The actual quantity varies with the individual, with the work done, and with the nature of the food eaten; but the following table gives approximately that of the more important daily material losses of an averairo man. 278 TEE HUMAN BODY. o W o W H P O M & |z; o <^ ^-^ M Q > 1 5 ci <^ rt lO o CO 1—1 ^* -"k 00 Ci (N o ^^ CO ^ s ->1 o CO B o o o o o ; 1> 1> ■^ ^ iO 1-1 *> 1 o 0« (M w o o w lO o ' -*■ i6 od CO' "*" 1 «o CO CO TjH 00 !5 0 Ci CO t-H OJ : 1 00 O O O 00 o d -*■ CO 1- « CO CD 1 ■r-l O CO O- 1— ( 1-1 05 lO -* CO fr^ O Tj^ i- CO r-( -^ CO m iO lO IC c o in a O 't '^ CI o i~ c3 T— 1 T— ot c- ^ CO ■» Oi ^ o cc '^ a o ;5 O o oj o sr C o o a B OD ^ h: & "o a. a 'C -§ o 1 i ' ^ - Tl 03 H !3 " c • — ' c ^ a ^ 1— o 1 THE DAILY LOSSES OF TEE BODY. 379 The living Body thus loses daily in round numbers 4 kilograms of matter (8 lbs.) and, since it is unable to create new matter, this loss must be compensated for from the exterior or the tissues would soon dwindle away alto- gether; or at least until they were so impaired that life came to an end. After death the losses would be of a differ- ent kind, and their quantity much more dependent upon surrounding conditions; but except under very unusual circumstances the wasting away would still continue in the dead Body. Finally, the composition of the daily wastes of the living Body is tolerably constant; it does not simply lose a quantity of matter weighing so -much, but a certain amount of definite kinds of matter, carbon, nitrogen, oxygen, and so on; and these same substances must be restored to it from outside, in order that life may be continued. To give one asking for bread a stone might, no doubt, if it were swallowed, compensate in weight for the matter he lost in twenty-four hours; but bread would be needed to keep him alive. In other words, the Body not only requires a supply of matter from outside, but a supply of certain definite kinds of matter. The Losses of the Body in Energy. The daily expendi- ture of matter by the living Body is not the only one: as continuously it loses in some form or another energy, or the power of doing work; often as mechanical work ex- pended in moving external objects, but even when at rest energy is constantly being lost to the Body in the form of lieat, by radiation and conduction to surrounding objects, by the evaporation of water from the lungs and skin, and by removal in warm excretions. Unless the Body can make energy it must therefore receive a certain supply of it also from the exterior, or it would very soon cease to carry on any of its vital work ; it would be unable to move and would cool down to the temperature of surround- ing objects. The discoveries of this century having shown that energy is as indestructible and uncreatable (see Pliy- sics) as matter, we are led to look for the sources of the supply of it to the Body; and findijig that the living Body daily receives it and dies when the supply is cut off, we no •:i80 THE HUMAN BODY. longer suppose, with the older physiologists, that it works by means of a mysterious vital force existing in or created by it; but that getting energy from the outside it utilizes it for its purposes — for the performance of its nutritive and other living work — and then returns it to the exterior in what the physicists know as a degraded state; that is in a less utilizable condition. While energy like matter is in- destructible it is, unlike matter, transmutable; iron is always iron and gold always gold; neither can by any means which we possess be converted into any other form of matter; and so the Body, needing carbon, hydrogen, oxygen, and nitro- gen to build it and. to cover its daily losses, must be sup- plied with those very substances. As regards energy this is not the case. While the total amount of it in the uni- verse is constant, its form is constantly subject to change — and that one in which it enters the Body need not be that in which it exists while in it, nor that in which it leaves it. Daily losing heat and mechanical work the Body does not need, could not in fact much utilize energy, supplied to it ■ in these forms; but it does need energy of some form and in amount equivalent to that which it loses. The Conservation of Energy. The forms of energy known to us are not nearly so numerous as the kinds of matter. Still we all know several of them; such as light, heat, sound, electricity, and mechanical work; and most people nowadays know that some of these forms are inter- convertible, so that directly or indirectly we can turn one into another. In such changes it is found that a definite amount of one kind ahvays disajijiears to give rise to a certain quantity of the other; or, in other words, that so much of the first form is equivalent to so much of the second. In a steam-engine, heat is produced in the fur- nace; when the engine is at work all of this energy does not leave it as heat; some goes as mechanical work, and the more work the engine does the greater is the difference be- tween the heat generated in the furnace and that leaving the machine. If, however, we used the work for rubbing two rough surfaces together we could get the heat back again, and if (which of course is impossible in practice) THE CONSERVATION OF ENERGY. 281 we could avoid all friction in the moving parts of the machine, the quantity thus restored would be exactly equal to the excess of the heat generated in the furnace over that leaving the engine. Having turned some of the heat into mechanical work we could thus turn the work back into heat again, and find it yield exactly the amount which seemed lost. Or we might use the engine to drive an elec- tro-magnetic machine and so turn part of the heat liber- ated in its furnace first into mechanical work and this into electricity; and if we chose to use the latter with the proper apparatus, we could turn more or less of it into light, and so have a great part of the energy which first became conspicuous as heat in the engine furnace, now manifested in the form of light at some distant point. In fact, starting with a given quantity of one kind of energy, we may by proper contrivances turn all or some of it into one or more other forms; and if we collected all the final forms and retransformed them into the first, we should have exactly the amount of it which had disappeared when the other kinds appeared. This law, that energy can change its form but that its amount is invariable, that it cannot be created or destroyed but simply transmuted, is known as the Icvw of the Conservation of Energy (see Phy- sics), and, like the indestructibility of matter, lies at the basis of all scientific conceptions of the universe, whether concerned with animate or inanimate objects. Since all forms of energy are interconvertible it is con- venient in comparing amounts of different kinds to express thein in terms of some one kind, by saying how much of that standard form the given amount of the kind spoken of would give rise to it were all converted into it. Since the most easily measured form of energy is mechanical work this is commonly taken as the standard form, and the quantities of others are expressed by saying how great a distance against the force of gravity at the earth's sur- face a given weight could be raised by the energy in ques- tion, if it were all spent in lifting the weight. The units of mechanical work being the kilogrammeter, or the foot- pound, the mechanical equivalent of any given kind of cnei-gv 282 THE HUMAN BODY. is the number of kilogram meters or foot-pounds of work its unit quantity would perform, if converted into mechanical work and used to raise a weight. For exam^jle the unit quantity of heat is that necessary to raise one kilogram of water one degree centigrade in temperature; or sometimes, in books written in English, the quantity necessary to warm one pound of water one degree Fuhronhcit. When there- fore we say that the mechanical equivalent of heat is 423 kilogrammeters we mean that the quantity of heat which would raise one kilogram of water in temperature . from 4° 0. to 5° C. would, if all turned into mechanical work, be able to raise one kilogram 423 meters against the attrac- tion of the earth; and conversely that this amount of me- chanical work if turned into heat would warm a kilogram of water one degree centigrade. The mechanical equiva- lent of heat, taking the Fahrenheit thermometric scale and using feet and pounds as measures, is 772 foot-pounds. Potential and Kinetic Energy. At times energy seems to be lost. Ordinarily we only observe it when it is doing work and producing some change in matter: but sometimes it is at rest, stored away and producing no changes that we recognize and thus seems to have been destroyed. Energy at work is known as hinetic energy; energy at rest, not producing changes in matter, is called j'^o^eniiV// energy. Suppose a stone pulled uji by a string and left suspended in the air. We know a certain amount of energy was used to lift it; but while it hangs we haA'^e neither heat nor light nor mechanical work to represent it. Still the energv" is not lost; we know we have only to cut the string and the weight will fall, and striking something give rise to heat. Or we may wind up a spring and keep it so by ii catch. In winding it up a certain amount of energy in the form of mechanical work was used to alter the form of the spring. Until the catch is removed this energy re- mains stored away as potential energy: but we know it is not lost. Once the spring is let loose again it may drive a clock or a watch, and in so doing will jjcrform again just so much work as was spent in coiling it; and when the watch has run down this energy will all have been turned POTENTIAL EimRGT OF CHEMICAL AFFINITY. 283 into other forms — mainly heat deyeloped in the friction of the parts of the watch against one another: but partly also in producing movements of the air, a portion of which Vv-e can readily observe in the sound of its ticking. The law of the conservation of energy does not say, then, that either the total potential or the total kinetic energy in the universe is constant in amount: but that the sum of the two is inva- riable, while constantly undergoing changes from kinetic to potential and vice versa: and from one form uf kinetic to another. The Energy of Chemical AflQ.mty. Between every two chemical atoms which are capable of entering into combi- nation there exists a certain amount of potential energy; when they unite this energy is liberated, usually in the form of heat, and once they have combined a certain amount of kinetic energy must be spent to pull them apart again; this being exactly the amount which was liberated when they united. The more stable the compound formed the more kinetic energy appears during its formation, and the more must be spent to break it up again. One may imagine the separated atoms as two balls pushed together by sjjrings, the strength of the spring being jDroportionate to the de- gree of their chemical affinity. Once they are let loose and permitted to strike together the potential energy pre- viously represented by "the compressed springs disappears, and in its place we have the kinetic energy, represented by the heat developed when the balls strike together. To pull them apart again, against the springs, to their original positions, just so much mechanical work must be spent as is the equivalent of that amount of heat which appeared when they struck; and thus kinetic energy will again be- come latent in breaking up the compound represented by the two in contact. The energy liberated in chemical com- bination is the most important source of that used in our machines: and also of that spent by the living Body. The Relation between the Matters Removed from the Body daily and the Energy Spent by it. A working locomotive is, we know, constantly losing matter to the exterior in the form of ashes and gaseous products of com- 284 THE HUMAK BODY. busliion, tlic latter being mainly carbon dioxide ana* water vapor. The engine also expends energy, not only in the form of heat radiated to the air, but as mechanical work in drawing the cars against the resistance offered ])y fric- tion or sometimes, up an incline, by gravity. Now the en- gine-driver knows that there is a close relationship between the losses of matter and the expenditure of energy, so that he has to stoke his furnace more frequently and allow a greater draft of air through it in going up a gradient than when running on the level. The more work the en- gine does the more coals and air it needs to make up for its jri eater waste. If we seek the cause of this relation- ship between work and waste, the first answer naturally is that the engine is a machine the special object of which is to convert heat into mechanical work, and so the more work it has to do the more heat is required for conversion, and consequently the more coals must be burnt. This, however, opens the question of the source of the heat — of all that vast amount of kinetic energy which is liberated in the furnace; and to answer this we must consider in what forms matter and energy enter the furnace, since the energy liberated there must be carried in somehow from outside. For present purposes coals may be considered as consisting of carbon and hydrogen, both of which sub- stances tend to forcibly combine with oxygen at high tem- peratures, forming in the one case carbon dioxide and in the other water. The oxygen necessary to form these com- ])Ounds being supplied by the air entering the furnace, all the potential energy of chemical affinity which existed be- tAveen the uncombined elements becomes kinetic, and is liberated as heat when the combination takes place. The energy utilized by the engine is therefore supplied to it in the form of potential energy, associated with the uncom- bined forms of matter whicn reach the furnace. Once the carbon and hydrogen have combined with oxygen they are no longer of any use a-s liberators of energy; and the com- pounds formed if retained in the furnace would only clog it and impede farther combustion; they are therefore got rid of as wastes througli the smoke-stack. Tlie engine. SOURCES OF EXERGY. 285 in short, receives uncombined elements associated with })otential energy; and loses combined elements (which have lost the energy previonsh* associated with them) and kinetic energy: it so to speak separates the energy from the mat- ter with which it was connected and, utilizing it, gets rid of the exhausted matter. The amount of kinetic energy liberated during such chemical combinations is very great; a kilogram of carbon uniting with oxygen to form car- bon dioxide sets free 8080 units of heat, or calories. Dur- ing the combination of oxygen and hydrogen to form water even more energy is liberated, one kilogi-am of hydro- gen when completely burnt liberating more than thirty-four thousand of the same units. The mechanical equivalent of this can be calculated if it is remembered that one heat unit = 423 kilogrammeters. Turning now to the living Body we find that its income and expenditure agree very closely with those of the steam- engine. It receives from the exterior substances capable of entering into chemical union; these combine in it and liberate energy; and it loses kinetic energy and the products of combination. From the outside it takes oxygen through the lungs, and oxidizable substances (in the form of foods) through the alimentary canal; these combine under the conditions prevailing in the living cells just as the carbon and oxygen, which will not unite at ordinary temperatures, combine under the conditions existing in the furnace of the engine; the energy liberated is employed in work of the Body, while the useless products of combination are got rid of. To explain, then, the fact that our Bodies go on working we have no need to invoke some special mysterious power resident in them and capable of creating energy, a i'ital force having no relation with other natural forces, such as the older physiologists used to imagine. The Body needs and gets a supply of energy from the exterior jnst as the steam-engine does, food and air being to one what coals and air are to the other; eacli is a machine in which energy is liberated by chemical combinations and then used for special work; the character of which depends upon the peculiarities of mechanism which utilizes it in each case, 286 777^ HUMAN BODY. and not upon any peculiarity in the energy utilized or in its source, "^riic ]5ody is, however, a far more economical machine than any steam-engine; of all the energy liberated in the latter only a small fraction, about one eighth, is use- fully employed, while our Bodies can utilize for the perform- ance of muscular work alone one fifth of the whole energy supplied to them; leaving out of account altogether the nutritive and other work carried on in them, and the heat lost from them. The Conditions of Oxidation in the Living Body. Al- though the general principles uijon which the Body and the steam-engine get their working power are the same, still in minor points very obvious differences are found between them. In the first place the coals of an engine are oxidized only at a very high temperature, one which would be instantly fatal to our Bodies which, although warm when compared with the bulk of inanimate objects, are very slow fires when compared with a furnace. Chem- istry and physics, however, teach us that this diffei'ence i.s quite unimportant so far as concerns the amount of energy liberated. If magnesium wire be ignited in the air it will become white-hot, flame, and leave at the end of a few seconds only a certain amount of incombustible rust or magnesia, which consists of the metal combined with oxygen. The heat and light evolved in the process repre- sent of course the energy which, in a potential form, was associated with the magnesium and oxygen before their combination. We can, however, oxidize the metal in a differ- eut way, attended with no evolution of light and no very per- ceptible rise of temperature. If, for instance, we leave it in the air it will become gradually turned into magnesia with- out having ever been hot to the touch or luminous to the eye. The process will, however, take days or weeks; anti while in this slow oxidation just as much energy is liberated as in the former case it now all takes the form of heat; and instead of being liberated in a short time is sj)read over a much longer one. as the gradual chemical combination takes place. The slowly oxidizing magnesium is, therefore, at no moment noticeably hot since it loses its heat to surrounding SOURCE OF BODILY ENERGY. 287 objects as fast as it is generated. The oxidations occurring in our Bodies are of this slow kind. An ounce of arrow- root oxidized in a fire, and in the Human Body, would liberate exactly as much energy in one case as the other, but the oxidation would take place in a few minutes and at a high temperature in the former, and slowly, at a lower temperature, in the latter. In the second place, the engine differs from the living Body in the fact that the oxidations in it all take place in a small area, the furnace, and so the temperature there becomes very high; while in our Bodies the oxidations take place all over, in each of the living cells; there is no one furnace or hearth where all the energy is liberated for the Avhole and transferred thence in one form or another to distant parts: and this is another reason why no one part of tlie Body attains a very high temperature. The Fuel of the Body, This is clearly different from tbat of an ordinary engine: no one could live by eating coals. This difference again is subsidiary; a gas-engine requires different fuel from an ordinary locomotive; and the Body requires a somewhat different one from either. It needs as foods, substances which can, in the first place, be absorbed from the alimentary canal and carried to the various tissues; and, in the second, can there be oxidized at a low temperature or, perhaps more probably, can be converted by the living cells into compounds which can be so oxidized. With some trivial exceptions, all substances which fulfill these conditions are complex chemical com- pounds, and to understand their utilization in the Body we must extend a little the statements above made as to the liberation of energy in chemical combinations. The general law may be stated thus — Energy is liberated tuhenever cJicmi- ral union takes place : and whenever more stable compounds are formed from less stable ones, in which the constituent atoms were less firmhj held toijplhcr. Of the liberation by simj^le combination we have already seen an instance in the oxidation of carbon in a furnace: but the union need not be an oxidation. Everyone knows how hot quicklime becomes when it is slaked; the water combining strongly with the lime, and energy being liberated in the form of 288 THE HUMAN BODY. heat, during the process. Of the liberation of energy by the breaking down of a complex compound, in which the atoms are only feebly united, into simpler and stabler ones, we get an example in alcoholic fermentation. During that process grape sugar is broken down into more stable com- ])ounds, mainly carbon dioxide and alcohol, while oxygen is at the same time taken up. To pull apart the carbon, hydrogen and oxygen of the sugar molecule requires a cer- tain expenditure of kinetic energy: but in the simultaneous formation of the new and stabler compounds a greater amount of energy is set free, and the difference appears as heat, so that the brewer has to cool his vats with ice. It is by processes like this latter, rather than by direct com- binations, that most of the kinetic energy of the Body is obtained; the complex proteids and fats and starches lyid sugar taken as food being broken down (usually with con- comitant oxidation) into simpler and more stable com- pounds. Oxidation by Successive Steps. In the furnace of an engine the oxidation takes place completely at once. The carbon and hydrogen leaving it, if it is well managed, are each in the state of their most stable oxygen compound. But this need not be so: we might first oxidize the carbon so as to form carbon monoxide, CO, and get a certain amount of heat; and then oxidize the carbon monoxide farther so as to form carbon dioxide, CO2, and get more heat. If w^e add together the amounts of heat liberated in each stage, the sum will be exactly the quantity which would have been obtained if the carbon had been completely burnt to the state of carbon dioxide at first. Every one wdio has studied chemistry will think of many similar cases. As the process is important physiologically we may take an- other example; say the oxidation of alcohol. This may be burnt completely and directly, giving rise to carbon dioxide and Avater — CHeO + 0« = 2C0i + 3H2O 1 Alcohol. C Oxygen. 2 Carbon dioxide. 3 water. But instead of this we can oxidize the alcohol by stages. UTILIZATION OF ENERGY IN THE BODY. 289 getting at each stage only a comparatively small amount of heat evolved. By combining it first with one atom of oxy- gen, we get aldehyde and water — C.HeO + 0 = C-'H40 + H=0 1 Alcohol. 1 Oxygen. 1 Aldehyde. 1 Water. Then we add an atom of oxygen to the aldehyde and get acetic acid (vinegar) — CiH40 + 0 = C-2H402 1 Aldehyde. 1 Oxygeu. 1 Acetic acid. And finally we may oxidize the acetic acid so as to get car- bon dioxide and water — C.HiO. + 04 = 2CO2 + 2H2O We get, in either case, from one molecule of alcohol, two of carbon dioxide and three of water; and six atoms of oxygen are taken up. In each stage of the gradual oxidation a certain amount of heat is evolved; and the sum of these is exactly the amount which would have been evolved by burning the alcohol completely at once. The food taken into the Body is for the most part oxi- dized in this gradual manner; the products of imperfect combustion in one set of cells being carried off and more completely oxidized in another set, until the final pro- ducts, no longer capable of further oxidation in the Body, are carried to the lungs, or kidneys, or skin, and got rid of. A great object of physiology is to trace all intermediate compounds between the food which enters and the waste products which leave; to find out Just how far chemical degradation is carried in each organ; and what substances are thus formed in various parts: but at present this j^art of the science is very imperfect. The Utilization of Energy in the Living Body. In the steam-engine energy is liberated as heat; some of the heat is used to evaporate water and expand the resulting steam; and then the steam to drive a piston. But in the living Body it is very probable (indeed almost certain) that a great part of the energy liberated by chemical transfor- 290 THE HUMAN BODY. mations docs not first take the form of heat; tlioiigli some of it does. This, again, does not affect the general prin- ciple: the source of energy is essentially the same in both cases; it is merely the form which it takes that is dif- ferent. In a galvanic cell energy is liberated during the union of zinc and sulphuric acid, and we may so arrange matters as to get this energy as heat; but on the other hand we may lead it off, as a so-called galvanic current, and use it to drive a magneto-electric machine before it has taken the form of heat at all. In fact, that heat may be used to do mechanical work we must reduce some of it to a lower temperature: an engine needs a condenser of some kind as well as a furnace; and, other things being equal, the cooler the condenser the greater the proportion of the whole heat liberated in the furnace which can be used to do work. Now in a muscle there is no condenser; its temperature is uniform throughout. So when it contracts and lifts a weight, the energy employed must be liberated"* in some other form than heat — some form which the muscu- lar fibre can use without a condenser. Summary. The living Body is continually losing mat- ter and expending energy. So long as we regard it as working by virtue of some vital force, the power of gener- ating which it has inherited, the waste is difficult to account for, since it is far more than we can imagine as due merely to wear and tear of the working j^arts. When, however, we consider the nature of the income of the Body, and of its expenditure, from a chemico-physical point of view, we get the clue to the puzzle. The Body does not waste because it works but works because it wastes. The working power is obtained by chemical changes occurring in it, associated with the liberation of energy which the living cells utilize; and the products of these chemical changes, being no longer available as sources of energy, are passed out. The chemical changes concerned are mainly the breaking down of complex and unstable chemical com- pounds into simpler and more stable ones, with concomi- tant oxidation. Accordingly the material losses of the SOURCES OF EXERGY IJV THE BODY. 291 Body are highly or completely oxidized, tolerably simple chemical compounds; and its material income is mainly uncombined oxygen, and oxidizable substances, the former obtained through the lungs, the latter through the alimen- tary canal. In energy, its income is the potential energy of uncombined or feebly combined elements, which are capable of combining or forming more stable combinations; and its final expenditure, is kinetic energy almost entirely in the form of mechanical work and heat. Given oxygen, all oxidizable bodies will not serve to keep the Body alive and working, but only those which (1) are capable of ab- sorption from the alimentary canal and (3) those which are oxidizable at the temperature of the Body under the influ- ence of protoplasm. Just as carbon and oxj'gen will not unite in the furnace of an engine unless the "fire be lighted" by the application of a match but, when once started, the heat evolved at one point will serve to carry on the conditions of combination through the rest of the mass, so the oxidations of the Body only occur under special con- ditions; and these are transmitted from parent to offspring. Every new Human Being starts as a portion of protoplasm separated from a parent and affording the conditions for those chemical combinations which supply to living matter its working power : this serves, like the energy of the burning ])art of a fire, to start similar processes in other portions of matter. At present we know nothing in physi- ology answering to the match which lights a furnace; those manifestations of energy which we call life are handed down from generation to generation, as the sacred fire in the temple of Vesta from one watcher to another. Science may at some time teach us how to bring the chemical con- stituents of protoplasm into that combination in which they possess the faculty of starting oxidations under those conditions which characterize life; then we will have learnt how to strike the vital match. For the present we must be content to study the i)ro]ierties of that form of matter which possesses living faculties; since there is no satisfactory proof that it has ever been produced, within 292 THE HUMAN BODY. our experience, apart from the influence of matter already living. How the vital spark first originated, how mole- cules of carbon, hydrogen, nitrogen and oxygen first united with water and salts to form protoplasm, we have no scien- tific data to ground a positive opinion upon, and such as we may have must rest upon other grounds. CHAPTER XX. FOODS. Poods as Tissue Formers. Hitherto we have considered foods merely as sources of energy, but they are also re- quired to build up the substance of the Body. From birth to manhood we increase in bnlk and weight, and that, not merely by accnmnlating water and sncli substances, liut by forming more bone, more muscle, more brain, and so on, from materials wdiich are not necessarily bone or muscle or nerve tissue. Alongside of the processes by which com- plex substances are broken down and oxidized and energy liberated, constructive processes take place by Avhicli new complex bodies are formed from simpler substances taken as food. A great part of the energy liberated in the Body is in fact ntilized first for this purpose, since to construct com- plex unstable molecules, like those of protoplasm, from the simpler compounds taken into the Body, needs an expendi- ture of kinetic energy. Even after full growth, when the Body ceases to gain weight, the same synthetic processes go on; the living tissues are steadily broken down and con- stantly reconstructed, as we see illustrated by the condition of a man who has been starved for some time, and who loses not only his power of doing work and of maintaining his bodily temperature but also a great part of his living tissues. If again fed properly he soon makes new fat and new muscle and regains his original mass. Another illustration of the continuance of constructive powers during the whole of life is afforded by the growth of the muscles when exercised properly. Since the tissues, on ultimate analysis, yield mainly car- bon, hydrogen, nitrogen and oxygen, it might be supposed a priori tlvdt a supplv of these elements in the uncombined 294 THE HVMAH BODY. state would serve as material iui- t he constructive forces of the Body to Avork with. Ex})eriL'uce, however, teaches u^ that this is not the case, hut thai ihe aninuil hody requires, I'or the most part, highly complex compounds for the con- struction of new tissue elements. All the active tissues yield on analysis large quantities of proteids which, as pointed out in Chapter I., enter always into the structure of protoplasm. Now, so far as we know at jsresent, the animal body is unable to build ujj proteids from simpler com- pounds of nitrogen, although when given one variety of them it can convert that one into others, and combine them with other things to form protoplasm. Hence proteids are an essential article of diet, in order to replace that portion of the living cells which is daily broken down and elimi- nated m the form of urea and other waste substances. Even albuminoids (p. 11), although so nearly allied to pro- teids, will not serve to replace them entirely in a diet; a man fed abundantly on gelatin, fats, and starches, would starve as certainly, though not so quickly, as if he got no nitrogenous food at all; his tissue waste would not be made good, and he Avould at last be no more able to utilize the energy-yielding materials supplied to him, than a worn-out steam-engine could emjiloy the heat of a fire in its furnace. So, too, the animal is unable to take the carbon for the construction of its tissues, from such simple compounds as carbon dioxide. Its constructive power is limited to the utilization of the carbon contained in more comjDlex and less stable compounds, such as proteids, fats, or sugars. The main bulk of all useful foods must therefore be made up of complex substances, and of these a part must be proteids, since the Body can utilize nitrogen for tissue formation only when supplied with it in that form. The bodies thus taken in are sooner or later broken down into simpler and eliminated; some at once in order to yield energy, others only after having first been built up into part of a living cell. The partial exceptions afforded by such losses to the Body as milk for suckling the young, or the albuminous and fatty bodies stored for the same purpose in the egg of a bird, are only apparent: the chemi- FOOD OF PLANTS. 295 cal degradation is only postponed, taking place in the body of the ofEsi^ring instead of that of the parent. In all cases animals are tlius, essentially, proteid consumers or wasters, and breakers down of complex bodies; the carbon, hydro- gen and nitrogen which they take as foods in the form of complex unstable bodies, ultimately leaving them in the simpler compounds, carbon dioxide, water, and urea: which are incapable of either yielding energy or building- tissue for any other animal and so of serving it as food. The question immediately suggests itself. How, since animals are constantly breaking up these complex bodies and can- not again build them, is the supply kept up ? For exam- ple, the supply of proteids, which cannot be made artificially by any process which we know, and yet are necessary foods for all animals, and daily destroyed by them. The Pood of Plants. As regards our own Bodies the question at the end of the last paragraph might perhaps be answered by saying that we get our proteids from the flesh of the other animals which we eat. But, then, we have to account for the possession of them by those animals; since they cannot make them from urea and carbon dioxide and water any more than we can. The animals eaten get them, in fact, from plants which are the great proteid formers of the world, so that the most carnivorous animal really de- pends for its most essential foods upon the vegetable king- dom; the fox that devours a hare in the long run lives on the proteids of the herbs that the hare had previously eaten. All animals are thus, in a certain sense, parasites; they only do half o£ their own nutritive work, just the final stages, leaving all the rest to the vegetable kingdom and using the products of its labor; and plants are able to meet this demand because they can live on the simple compounds of carbon, hydrogen, and nitrogen eliminated by animals, building up out of them new complex substances which animals can use as food. A green plant, supplied with am- monia salts, carbon dioxide, water, and some minerals, will grow and build up large quantities of proteids, fats, starches, and similar things; it will pull the stable compounds eli- minated by animals to pieces, and build tliem up into com- 296 THE HUMAN BODY. plex unstable oodies, capable of yielding energy when again broken down. However, to do such work, to break up stable combinations and make from them less stable, needs a supply of kinetic energy, which disai)pcars in the process being stored away as potential energy in the new compound; and we may ask whence it is that the plant gets the suj)})ly of energy which it thus utilizes for chemical construction, since its simple and highly oxidized foods can yield it none. It has been proved that for this purpose the green plant uses the energy of sunlight: those of its cells which contain the substance called chlorophyl (leaf green) have the power of utilizing energy in the form of light for the perform- ance of chemical work, just as a steam-engine can utilize heat for the performance of mechanical work. Exposed to light, and receiving carbon dioxide from the air, and water and ammonia (which is produced by the decomposition of urea) from the soil, the plant builds them ujo again, with the elimination of oxygen, into complex bodies like those which animals broke down, with fixation of oxygen. Some of the bodies thus formed it uses for its own growth and the formation of new protoplasm, just as an animal does; but in sunlight it forms more than it uses, and the excess stored up m its tissues is used by animals. In the long run, then, all the energy spent by our Bodies comes through millions of miles of space from the sun; but to seek the source of its supply there would take us far out of the domain of Physiology (see Astronomy). Non-Oxidizable Foods. Besides our oxidizable foods, a large number of necessary food materials are not oxidiza- ble, or at least are not oxidized in the Body. Typical in- stances are afforded by water and common salt. The use of these is in great part physical; the water, for instance, dissolves materials in the alimentary canal, and carries the solutions through the walls of the digestive tube into the blood and lymph vessels, so that they can be carried from part to part; and it permits interchanges to go on by diffusion. The salines also influence the solubility and chemical interchanges of other things present with them. Serum albumen, tlie chief proteid of the blood, for example, NON-OXIDIZABLE FOODS. 297 is insoluble in pure water, but dissolves readily if a small quantity of neutral salts is present. Besides such uses the non-oxidizable foods have probably others, in vt^hat we may call machinery formation. In the salts which give their hardness to the bones and teeth, we have an example of such an emjiloyment of them: and to a less extent the same may be true of other tissues. The Body, in fact, is not a mere store of potential energy but something more — it is a machine for the disposal of it in certain ways; and, wherever practicable, it is clearly advantageous to have the purely energy-expending parts made of non-oxidizable mat- ters, and so protected from change and the necessity of frequent renewal. The Body is a self-building and self- repairing machine, and the material for this building and repair must be supplied in the food, as well as the fuels, or oxidizable foods, which yield the energy the machine ex- pends; and while experience shows us, that even for ma- chinery construction, oxidizable matters are largely needed, nevertheless it is a gain to replace them by non-oxidizable substances when possible; just as if practicable it would be advantageous to construct an engine out of materials which would not rust, although other conditions determine the use of iron for the greater part of it. Definition of Poods. Foods may be defined as sub- stances ichich are taken into the alimentary canal, and which, vjhen absorbed from it, serve either to supply material for the growtli of the Body, or for the replacement of matter which has been removed from it, either after oxidation or without having been oxidized. Foods to replace matters which have been oxidized must be themselves oxidizable; they are force generators, but may be and generally are also tissue formers; and are nearly always complex organic sub- stances derived from other animals or from plants. Foods to replace matters not oxidized in the Body are force regu- lators, and are for the most j^art tolerably simple inorganic compounds. Among the force regulators we must, how- ever, include certain organic foods which, although oxidized in the Body and serving as liberators of energy, yet produce effects totally disproportionate to the energy they set free. 398 THE UUMAJS- BODY. iind for which effects they are taken. In other words, theii- iuiluence as stimuli in exciting certain tissues to liberate energy, or as inhibitory agents checking the activity of parts, is more marked than their direct action as force gen- erators. As examples, we may take condiments: mustard and pepper are not of much use as sources of energy, al- though they no doubt yield some; we take them for their stimulating effect oii the mouth and other parts of the alimentary canal, by which they promote an increased flow of the digestive secretions or an increased appetite for food. Thein, again, the active princiiDle of tea and coffee, is taken for its stimulating effect on the nervous system, rather than for the amount of energy which is yielded by its own oxidation. Conditions which a Food must Fulfill. ' (1) A food must contain the elements Avhich it is to replace in the Body: but that alone is not sufficient. The elements leav- ing the Body being usually derived from the breaking down of com^Dlex substances in it, the food must contain them either in the form of such complex substances, or in forms which the Body can build up into them. Free nitrogen and hydrogen are no use as foods, since they are neither cxi- dizable under the conditions prevailing in the Body (and consequently cannot yield it energy), nor are they capable of construction by it into its tissues. (2) Food after it has been swallowed is still in a strict sense outside the Body; the alimentary canal is merely a tube running through it, and so long as food lies there it is not forming any part of the Body proper. Hence foods must be capa- ble of absorption from the alimentary canal; either directly, or after they have been changed by the processes of diges- tion. Carbon, for example, is no use as a food, not merely because the Body could not build it n^ into its own tissues, but because it cannot be absorbed from the alimentary canal. (3) Neither the substance itself nor any of the products of its transformation in the Body must be inju- rious to the structure or activity of any organ. If so it is di poison, not a food. Alimentary Principles. What m common language we ALIMENTARY PRINCIPLES. 299 commonly call foods are, in nearly all cases, mixtures of several foodstuffs, with substances which are not foods at all. Bread, for example, contains water, salts, gluten (a proteid), some fats, much starch, and a little sugar; all true foodstuffs: but mixed with these is a quantity of cellulose (the chief chemical constituent of the walls which surround vegetable cells), and this is not a food since it is incapable of absorption from the alimentary canal. Chemical exami- nation of all the common articles of diet shows that the actual number of important foodstuffs is but small: they are repeated in various proportions in the different things we eat, mixed with small quantities of different flavoring substances, and so give us a pleasing variety in our meals; but the essential substances are much the same in the fare of the workman and in the "delicacies of the season." These primary foodstuffs, which are found repeated in so many different foods, are known as " alimentary principles f and the physiological value of any article of diet depends on them far more than on the traces of flavoring matters which cause certain things to be especially sought after and so raise their market value. The alimentary principles may be conveniently classified into proteids, albuminoids, hydrocarbons, carbohydrates, and inorganic bodies, Proteid Alimentary Principles. Of the nitrogenous foodstuffs the most important are proteids: they form an essential part of all diets, and are obtained both from animals and plants. The most common and abundant are myosin and syntonin which exist in the lean of all meats; egg albumen; casein, found in milk and cheese; gluten and vegetable casein from various plants. Albuminoid Alimentary Principles. These also con- tain nitrogen, but cannot replace the proteids entirely as foods; though a man can get on with less proteids when he has some albuminoids in addition. The most important is gelatin, which is yielded by the white fibrous tissue of animals when cooked. On the whole the albuminoids are not foods of high value, and the calf's-foot jelly and such compounds, often given to invalids, have not nearly the nutritive value they are commonly supposed to possess. 300 THE HUMAN BODY. Hydrocarbons {Fals (iiul Oils). The most inii)or1:i,ni. are stearin, palmatin, margarin and olein, wliicli exist m various proportions in animal fats and vegetable oils; the more fluid containing most olein. Butter consists chiefly of a fat known as butyrin. All are neutral compounds of gly- cerine and fatty acids and, speaking generally, any such sub- stance which is fusible at the tem])erature of the Body will serve as a food. The stearin of beef and mutton fats is not by itself fusible at the body temperature, but is mixed m those foods with so miich olein as to be melted in the alimentary canal. Beeswax, on the other hand, is a fatty body which will not melt in the intestines and so passes on unabsorbed; although from its composition it Avould be useful as a food could it be digested. It is convenient to distinguish fats proper (the adipose tissue of animals consisting of fatty compounds inclosed in albuminous cell- walls) from oils, or fatty bodies which are not so surrounded. Carbohydrates. These arc mainly of vegetable origin. The most important are starch, found in nearly all vege- table foods; dextrin; gums; //ra^^e -m^/ar (into which starch is converted during digestion); and cane sugar. Sugar of 7nilh and glycogen are alimentary principles of this group, derived from animals. All of them, like the fats, consist of carbon, hydrogen and oxygen; but the percentage of oxvgen in them is much higher, there being one atom of oxvgen for every two of hydrogen in their molecule. Inorganic Foods. Water; common salts; and the chlo- rides, phosphates, and sulphates of potassium, magnesium and calcium. More or less of these bodies, or the materials for their formation, exists in all ordinary articles of diet, so that we do not swallow them in a separate form. Phos- ])hates, for example, exist in nearly all animal and yegetabk; foods; while other foods, as casein, contain phosphorus in combinations Avhich in the Body yield it up to be oxidized to form i^hosphoric acid. The same is true of sulphates, which are partially swallowed as such in various articles of diet, and are partly formed in the Body by the oxidation of the sulphur of various proteids. Calcium salts are abun- dant in bread, and arc also found in many drinking waters. FLESH FOODS. 301 Water and table salt form exceptions to tlie rule that in- organic bodies are eaten imperceptibly along with other things, since the Body loses more of each daily than is usu- ally supplied in that way. It has, however, been main- tained that salt, as such, is an unnecessary luxury; and there seems some evidence that certain savage tribes live without more than they get in the meat and vegetables they eat. Such tribes are, however, said to suffer especially from intestinal parasites; and there is no doubt that to civilized man the absence of salt is a great deprivation. Mixed Foods. These, as already pointed out, include nearly all common articles of diet; they contain more than one alimentary principle. Among them we find great differences; some being rich in proteids, others in starch, others in fats, and so on. The formation of a scientific die- tary depends on a knowledge of these characteristics. The foods eaten by man are, however, so varied that we cannot do more than consider the most important. Flesh. This, whether derived from bird, beast, or fish, consists essentially of the same things — muscular fibres, tendons, fats, blood-vessels, and nerves. It contains several proteids and especially myosin; gelatin-yielding matters in the white fibrous tissue; stearin, palmatin, margariu, and olein among the fats; and a small amount of carbohydrates in the form of glycogen and grajje sugar; also inosite, a kind of sugar found only in muscles. Flesh also contains much water and a considerable number of salines, the most imi^ortant and abundant being jjotassium phosphate. Os- mazome is a crystalline nitrogenous body which gives much of its taste to flesh; and small quantities of various similar substances exist in different kinds of meat. There is also more or less yellow elastic tissue in flesh; it is indigestible and useless as food. When meat is cooked its white fibrous tissue is turned into gelatin, and the whole mass becomes thus softer and more easily disintegrated by the teeth. When boiled much of the proteid matters of the meat pass out into the broth, and there in part coagulate and form the scum : this loss may be prevented in great part, if it is not intended to use 303 THE HUMAN BODY. ; lie broth, by putting the raw meat at once into boiling •.vater, which coagulates the surface albumin before it dis- solves out, and this keeps in the rest. In any case the myosin, being insoluble in water, remains behind in the boiled meat. In baking or roasting, all the solid parts of the flesh are preserved and certain agreeably flavored bodies are produced, as to the nature of which little is known. Eggs. These contain a large amount of egg albumen and, in the yolk, another proteid, known as vitellin. Also fats, and a substance known as lecithin (p. 14), which is important as containing a considerable quantity of phos- l^horus. Milk. This contains the proteid known as casein; several fats forming the butter; a carbohydrate, the milh sugar; and much water and salts, especially phosphates. Butter is mainly made of butyrin, a compound of butyric acid and glycerine. In the milk it is disseminated in the form of minute globules which, for the most part, float up to the top when the milk is let stand and then form the cream. In this each fat droplet is surrounded by a pellicle of albumi- nous matter; by churning, these pellicles are broken up and the fat droplets run together to form the butter. Casein is insoluble in water, and in the milk it is dissolved by the alkaline salts present. When milk is kept its sugar fer- ments and gives rise to lactic acid, which neutralizes the alkali and precipitates the casein as curds. In cheese- making the casein is similarly precipitated by the addition of an acid and (the whey being pressed out) it constitutes the main bulk of the cheese. Vegetable Foods. Of these wheat affords the best. In 1000 parts it contains 135 of proteids, 568 of starch, 46 of dextrin (a carbohydrate), 49 of grape sugar, 19 of fats, and 32 of cellulose, the remainder being water and salts. The proteid of wheat is mainly gluten, which when moist- ened with water forms a tenacious mass, and this it is to which wh eaten bread owes its superiority. When the dough is made yeast is added to it, and produces a fermen- tation by which, among other things, carbon dioxide gas is produced. This gas, imprisoned in the tenacious dough. VEGETABLE FOODS. 303 and expauded during baking, forms cavities in it and causes it to ''rise" and make "light bread," which is not only more pleasant to eat but more digestible than heavy. Other cereals may contain a larger percentage of starch but none have so much gluten as wheat; when bread is made from them the caibon dioxide gas escapes so readily from the less tenacious dough that it does not expand the mass properly. Corn contains in 1000 parts, 79 of proteids, 637 of starch, and from 50 to 87 of fats; much more than any other kind of grain. Rice is poor in proteids (50 parts in 1000) but very rich in starch (823 parts in 1000). Peas and beans are rich in proteids (from 220 to 260 parts in 1000), and contain about half their weight of starch. Potatoes are a poor food. They contain a great deal of water and cellulose, and only about 13 parts of proteids and 154 of starch in 1000. Other fresh vegetables, as carrots, turnips and cabbages, are valuable mainly for the salts they contain; their weight is mainly due to water, and they contain but little starch, proteids, or fats. Fruits, like most fresh vegetables, are mainly valu- able for their saline constituents, the other foodstuffs in them being only j)resent in small proportion. Some form of fresh vegetables is, however, a necessary article of diet; as shown by the scurvy which used to prevail among sailors before fresh vegetables or lime-juice wore supplied to them. The Cooking of Vegetables. Tliisis of more importance even than the cooking of flesh, since in most the main ali- mentary principle is starch, and raw starch is difficult of digestion. In plants starch is nearly always stored up in the form of solid granules, which consist of alternating layers of starch ceUnlose and starch granulose. The diges- tive fluids turn the starch into grape sugar which is soluble and can be absorbed from the alimentary canal, while starch itself cannot. Now these fluids act very slowly and imper- fectly on raw starch, and then only on the granulose; but when boiled, the starch gi-anules swell up, and are more readily converted into grape sugar; and the starch cellulose is so altered that it too undergoes that change. When starch is roasted it is turned into a substance known as soluble starch which is readily dissolved in the alimentary canal. 304 THE HUMAN BODY. There is therefore a scientific foundation for the common belief that tlie crust of a loaf is more digestible than the crumb, and toast than ordinary bread. Alcohol. Tliere are perhaj^s no common articles of diet concerning which more contradictory statements have been made than alcoliolic drinks. This depends upon their l)eculiar position: according to quantity or circumstances alcohol may be a poison or a food; and as a food it may be i-egarded either as a force regulator or a force generator. There is no doubt that alcohol in certain doses may be pro])- perly called a food. If not more tlian two ounces (which would be contained in about four ounces of whiskey or two quarts of lager beer) are taken in the twenty-four hours, it is completely oxidized in the Body and excreted as water and carbon dioxide. In this oxidation energy is of course liberated and can be utilized. Commonly, however, alco- hol is not taken for this purpose but, as a force regulator, for its influence on the nervous system or digestive organs, and it is in this capacity that it becomes dangerous. For not only may it be taken in quantities so great that it is not all oxidized in the Body but is passed through it as alco- hol, or even that it acts as a narcotic poison instead of a stimulant, but when taken in what is called moderation there can be no doubt that the constant " whipping up" of the flagging organs, if continued, must be dangerous to their integrity. Hence the daily use of alcohol merely ni such quantities as to produce slight exhilaration or to facili- tate work is by no means safe; though in disease when the system wants rousing to make some special effort, the phy- sician cannot dispense with it or some other similarly act- ing substance. In fact, as a force generator alcohol maybe advantageously replaced by other foods in nearly all cases; and there is no evidence that it heljis in the construction of the working tissues, thougli its excessive use often leads to an abnormal accumulation of fat. Its proper use is as a '•' Avhip," and, one has no more right to use it to the healthy Body than the lash to overdrive a Avilling horse. The jdiysician is the proper jjerson to determine whether it is wanted under any given circumstances. ALCOHOL. 305 If alcohol is to be used as a daily article of diet it sliould he borne in mind that when concentrated it coagulates the proteids of tlie cells of the stomach witJi Avhich it comes in contact, in the same sort of way, though of course to a much less degree, as it shrivels and dries up an animal pre- served in it. Dilute alcoholic drinks, such as claret and beer, are therefore far less baneful than whiskey or brandy, and these are worst of all in the almost undiluted form of most " mixed drinks." For the same reason alcohol \a far more injurious on an empty stomach than after a meal. When the stomach is full it is diluted, is more slowly absorbed, and, moreover, is largely used up in coagulating the proteids of the food instead of those of the lining membrane of the stomach. The old '"three bottle" men who drank their port-wine after a heavy dinner, got off far more safely than the modern tippler who is taking ''nips" all day long, although the latter may imbibe a smaller quantity of alco- hol in the twenty-four hours. By far the best way, how- ever, is to avoid alcohol altogether m health. If the facts lead us to conclude, against the extremists, that it is to a certain extent a food, it is nevertheless a dangerous one; even in what we may call "physiological " quantities, or such amounts as can be totally oxidized in the Body. The Advantage of a Mixed Diet. The necessary quan- tity of daily food depends upon that of the material daily lost from the Body, and this varies both in kind and amount with the energy expended and the organs most used. In children a certain excess beyond this is required to furnish materials for growth. Although it is impossible to lay down with perfect accuracy how much daily food any individual requires, still the average quantity may be de- rived from the table of daily losses given on page 278, which shows that a healthy man needs daily in assimilable forms about 274 grams (4220 grains) of carbon and 19 grams (292 grains) of nitrogen. The daily loss of hydrogen which is very great (352 grams or 5428 grains), is nearly all that contained in water which has been drunk and, so to speak, merely filtered through the Body, after having assisted in the solution and transference through it of other 306 THE HUMAN BODY substances. About 300 grams (4820 grains) of water (containing 33.3 grams (513 grains) of hydrogen are, how- ever, formed in tlie Body by oxidation, and the liydrogenfor this purpose must be supplied m the form of some oxidiza- ble foodstuff, wliether proteid, fat, or carbohydrate. The oxygen wanted is mainly received from the air through the lungs, but some is taken in the food. Since protcid foods contain carbon, nitrogen and hydro- gen, life may be kept up on them alone, with the necessary salts, Avater and oxygen; but such a form of feeding would be anything but economical. Ordinary protcids contain in 100 parts (p. 10) about 52 of carbon and 15 of nitrogen, so a man fed on them alone would get about 3| parts of carbon for every 1 of nitrogen. His daily losses are not in this ratio, but about that of 274 grams (4220 grains) of carbon to 20 grams (308 grains) of nitrogen, or as 13.7 to 1; and so to get enough carbon from proteids far more tlian the necessary amount of nitrogen must be taken. Of dry proteids 527 grams (8116 grains) would yield the necessary carbon, but would contain 79 gi'ams (1217 grains) of nitrogen; or four times more than is necessary to cover the daily losses of that element from the Body. Fed -on a purely proteid diet a man would, therefore, have to digest a vast quantity to get enough carbon, and in eating and absorbing it, as well as in getting rid of the extra nitrogen which is useless to him, a great deal of unnecessary labor would be thrown upon the various organs of his Body. Similarly, if a man were to live on bread alone he would burden his organs with much useless work. For bread contains but little nitrogen in proportion to its carbon, and so, to get enough of the former, far more carbon than was utilized would have to be eaten, digested, and eliminated daily. Accordingly, we find that mankind in general employ a mixed diet when they can get it, using richly proteid sul)- stances to supply the nitrogen needed, but deriving the carbon mainly from non -nitrogenous foods of the fatty or carbohydrate groups, and so avoiding excess of either. For instance, lean beef contains about \ of its weight of dry ADVANTAGE OF A MIXED DIET. 307 proteid, which contaiDS 15 per cent of nitrogen. Conse quently the 133 grams (2048 grains) of proteid which would be foiind in 532 grams (1 lb 3 oz.) of lean meat would supply all the nitrogen needed to compensate for a day's losses. But the proteid contains 52 per cent of carbon, so the amount of it in the above weight of fatless meat would be 69 grams (1062 grains) of carbon, leaving 205 grams (3157 grains) to be got either from fats or car- bohydrates. The necessary amount would be contained m about 256 grams (3942 grains) of ordinary fats or 460 grams (7084 grains^ of starch; hence either of these, with the above quantity of lean meat, would form a far better diet, both for the purse and the system, than the meat alone. As already pointed out, nearly all common foods contain several foodstuffs. Ordinary butcher's meat, for example, contains nearly half its weight of fat; and bread, besides proteids, contains starch, fats, and sugar. In none of them, however, are the foodstuffs mixed m the i)hysiologically best proportions, and the practice of employing several of them at each meal or different ones at different meals during the day, is thus not only agreeable to the palate but in a high degree advantageous to the Body. The strict vegetarians who do not employ even such substances as eggs, cheese, and milk, but confine themselves to a purely vegetable diet (such as is always poor in proteids), daily take far more carbon than they require, and are to be congratu- lated on their excellent digestions which are able to stand the strain. Those who use eggs, cheese, etc., can of course get on very well, since such substances are extremely rich m proteids, and supply the nitrogen needed without the necessity of swallowing the vast bulk of food which mast be eaten in order to get it from plants directly. / CHAPTER XXI. THE ALIMENTARY CANAL AND ITS APPEN- DAGES. General Arrangement. The alimentary canal is essen- tially an involuted jiortion of the skin, specially set apart for absorption, and forming a tube which runs through the Body (Fig. 2); it communicates with the exterior at three points (the nose, the mouth, and the anal aperture), at which this modified skin, or mucous memhrane, is con- tinuous with the general outer integument. Supporting the lining a])SorptiYe membrane are other layers which strengthen the tube, and are also in part muscular and, by their contractions, serve to pass materials along it from one end to the other. In the walls of the canal are numerous blood and lymphatic vessels which carry off the matters absorbed from its cavity; and there also exist in connection with it numerous glands, whose function it is to pour into it various secretions which exert a solvent influ- ence on such foodstuffs as would otherwise escape absor])- tion. Some of these glands are minute and imbedded in the walls of the alimentary tube itself, but others (such as the salivary glands) are larger and lie away from the main channel, into which their products are carried by ducts of various lengths. The alimentary tube is not uniform but presents several dilatations on its course; nor is it straight, since, being much longer than the Body, it is packed away by being coiled up in the abdominal cavity. Subdivisions of the Alimentary Canal. The mouth opening leads into a chamber containing the teeth and tongue, the mouth chamber or buccal cavity. This is sue- MOUTH CAVITY. 309 ceeded by the j)Iiari/nx or throat cavity, which narrows at the top of the ueck into the gullet or oesophagus; this runs down through the thorax and, passing through the dia- phragm, dilates in the upper part of the abdominal cavitr into the stomach. Beyond the stomach the channel again narrows to form a long and greatlv coiled tube, the small intestine, which terminates by opening into the large intes- tine, much shorter although wider than the small, and ter- minating by an opening on the exterior. The Mouth Cavity (Fig. 89) is bounded in front and on the sides by the lips and cheeks, below by the tongue, k, and above by the palate; Avhich latter consists of an an- terior part, 7, supported by bone and called the hard pal- ate, and a jDosterior, /, con- taining no bone, and called the soft 2^alate. The two can readily be distinguished by applying the tip of the tongue to the roof of the mouth and drawing it backwards. The hard palate forms the parti- tion between the mouth and nose. The soft palate arches down over the back of the mouth, hanging like a cur- tain between it and the pharynx, as can be seen by holding the mouth open in front of a looking-glass. From the middle of its free border a conical process, i\iQ uvula, hangs doAvn. The Teeth. Immediately within the cheeks and lips are Fig. 89.— The mouth, nose and pha- rynx, with the commencement of the gullet and larynx, as exposed by a section, a little to the left of the me- dian plane of the head. a. vertebral column : b, gullet : c, windpipe ; c1. larynx : e, epiglottis ; /. soft palate ; (J. opening of Eustachian tube ; k. tongue : ?, hard palate ; m. the sphe- noid bone on the base of the skull; n. the fore part of the cranial cavity : o. p. g, the tubinate bones of the out- er side of the left nostril chamber. 310 THE HUMAN BODY. two semicircles, formed by the borders of the upper and lower jaw-bones, which arc covered by tlie gums, except at intervals along their edges where they contain sockets in which the teeth are implanted. During life two sets of teeth are developed; the first or milk set appears soon after birth and is shed during childhood, when the second or permanent set appears. The teeth differ in minor points from one another, but in all three parts are distinguishable; one, seen in the mouth and called the croiun of the tooth; a second, im- bedded in the jaw-bone and called the root or fang; and between the two, embraced by the edge of the gum, is a narrowed portion, the neck or cervix. From differences in their forms and uses the teeth are divided into incisors, canines, bicnsjjids, and molars, arranged in a definite order in each jaw. Beginning at the middle line we meet in each half of each jaw with, successively, two incisors, one canine, and two molars in the milk-set; making twenty altogether in the two jaws. The teeth of the permanent set are thirty-two in number, eight in each half of each jaw, viz. — beginning at the middle line — two incisors, one canine, two bicuspids, and three molars. The bicuspids, or premolars, of the permanent set replace the milk molars, while the permanent molars are new teeth added on as the jaw grows, and not substituting any of the milk teeth. The hindmost permanent molars are often called the tvis- doin teeth. Characters of Individual Teeth. The incisors (Fig. 90) are adapted for cutting the food. Their crowns are chisel - shaped and have sharp horizontal cutting edges, which become worn away by use so that they are beveled off behind in the upper row, and in the opposite direction in the lower. Each has a single long fang. The canines (Fig. 91) are somewhat larger than the incisors. Their crowns are thick and somewhat conical, having a central point or cus}) on the cutting edge. In dogs, cats, and other carnivora the canines are very large and adapted for seizing and holding prey. The bicuspids or premolars (Fig. 93) are rather shorter than the canines and their TEE TEETH. 311 crowns are somewhat cnboidal. Each has two cusps, an outer towards the cheek, and an inner on the side turned towards the interior of the mouth. The fang is compressed laterally, and has usually a groove partially subdividing it Fig. • Fig. 92. Fig. 93. Fig. 90.— An incisor tooth. Fig. 91.— a canine or eye tooth. Fig. 92.— a bicuspid tooth seen from its outer side; the inner cusp is, accord- ingly, not visible. Fig. 93.— a molar tooth. into two. At its tip the separation is often complete. The molar teeth or grinders (Fig. 93) have large crowns with broad surfaces, on which are four or five projecting tubercles, which roughen them and make them better adapt- ed to crush the food. Each has usually several fangs. The milk teeth only differ in subsidiary points from those of the same names hi the permanent set. The Structure of a Tooth. If a tooth be broken open a cavity extending through both crown and fang will be found in it. This is filled during Hfe with a soft vascular pulp, and hence is known as the "pulp cavity" {c, Fig. 94). The hard parts of the tooth disposed around the pulp cavity consist of three different tissues. Of these one im- mediately surrounds the cavity and makes uj) most of the bulk of the tooth; it is dentine (8, Fig. 94); covering the dentine on the crown is the enamel (1, Fig. 94) and, on the fang, the cement (3, Fig. 94). The pulp cavity opens below by a narrow aperture at the tip of the fang, or at the tip of each if the tooth has more than one. The pulp consists mainly of connective tissue, but its surface next the dentine is covered by a layer of 312 THE HUMAN nODT. columnar cells. Through tlic opening on the fang blood- vessels and nerves enter the i)ulp. The dentine vields on clieniical analysis the same mate- FiG. 0-4. —Section through a premolar tooth of the cat still imbedded iu its socket. 1. enamel: 2, dentine; 3, cement; 4, the gum, 5, the bone of the lower jaw ; c, the pulp cavity. rials as bone but is somewhat harder, earthy matters con- stituting 72 per cent of it a? against 6G per cent in bone. Under the microscope it is recognized by the fine dentinal THE TONGUE. 313 tiidnles which, radiating from the pulp cayity, perforate it throughout, finally ending in minute branches which open into irregular cavities, the interglobular spaces, which lie Just beneath the enamel or cement. At their widest ends, close to the pulp cavity, the dentinal tubules are only about 0.005 millimeter (j-gVo of an inch) in diameter. The cement is much like bone in structure and composition, possessing lacuna? and canaliculi but rarely any Haversian canals. It is thickest at the tip of the fang and thins away towards the cervix. Enamel is the hardest tissue m the Body, yielding on analysis only from two per cent to three per cent of organic matter, the rest being mainly calcium. phosi)hate and carbonate. Its histological ele- ments are minute hexagonal prisms, closely packed, and set on vertically to the surface of the subjacent dentine. It is thickest over the free end of the crown, until worn away by use. Covering the enamel in unworn teeth is a thin struc- tureless horny layer, the enamel cuticle. The Tongue (Fig- 95) is a muscular organ covered with a mucous membrane, extremely mobile, and endowed not only with a delicate tactile sensibility but with the ter- minal organs of the special sense of taste; it is attached by its root to the hyoid bone. Its upper surface is covered with small eminences or j»?«/?i//«9, much like those more highly developed on the tongue of a cat, where they are readily felt. On the human tongue there are three kinds of papillffi, the circum vallate, the fu7igiform, and the fili- form. The circum vallate papillae, 1 and 2, the largest and least numerous, are from seven to twelve in number and lie near the root of the tongue arranged in the form of a V with its open angle turned forwards. Each is an ele- vation of the mucous membrane, covered by epithelium, and surrounded by a trench. On the sides of the papillae, imbedded in the epithelium, are small oval bodies (Fig. 155) richly supplied with nerves and supposed to be concerned in tlie sense of taste, and hence called the taste buds (Chap. XXXIV.). The fungiform paj)ill(B,d, are rounded elevations attached by somewhat narrower stalks, and found all over the middle and fore part of the upper surface of 314 THE HUMAN BODY. the tongue. They are easily recognized on the living tongue by their bright red color. The filiforni papiUce, most numerous and smallest, are scattered all over the dor- '\3 Fig. 05.— The upper surface of the tongue. 1, 2. oircumvallate papillas; 3, fun- giform papillae: 4, filiforni papillas; 6, mucous glands ; 7, tonsils; 8, part of epi- £■10111?. sum of the tongue except near its base. Each is a conical eminence covered by a thick horny layer of epithelium. It is these papillfe which are so highly developed on the tongues of Carmvoi'a, and serve them to scrape bones clean TEE SALIVARY GfL4ND8. 315 of even such tongli structures as ligaments. Tamed tigers have been known to draw blood by licking the'^hand of their master. li'ote. In health the surface of the tongue is moist, covered by little '^fur.'" and in childhood of a red color. In adult life the natural color of the tongue is less red, ex- cept around the edges and tip; a bright red glistening tongue being, then, usually a symptom of disease. When the digestive organs are deranged the tongue is commonly covered with a thick yellowish coat, composed of a little mucus, a few cells of epithelium shed from the surface, and numerous microscopic organisms known as bacteria; and there is frequently a "bad taste" in the mouth. The whole alimentary mucous membrane is in close physio- logical relationship; and anything disordering the sto- mach is likely to produce a "furred tongue." The Salivary Glands. The saliva, which is poured into the mouth and which, mixed with the secretion of minute glands mibeddcd in its lining membrane, moistens it, is secreted by three pairs of glands, the parotid, the submaxil- lary and the snUinguaJ. The parotid glands lie in front of the ear behind the ramus of the lower jaw; each sends its secretion into the mouth by a tube known as Stenon^fi duct, which crosses the cheek and opens opposite the second upper molar tooth. In the disease known as mumps the parotid glands are inflamed and enlarged. The submaxillary glands lie between the halves of the lower jaw-bone, near Its angles, and their ducts open beneath the tongue near the middle line. The sublingual glands lie beneath the floor of the mouth, covered by its mucous membrane, between the back part of the tongue and the lower jaw-bone. Each has many ducts (8 to 20), some of which join the submaxil- lary duct, while the rest open separately in the floor of the mouth. The Fauces is the name given to the aperture which can be seen at the back of the mouth (Fig. 89), leading from it into the pharynx below the soft palate. It is bounded above by the soft palate and uvula, below by the root of the tongue, and on the sides by muscular elevations, covered by 316 77//; HUMAN BODY. mucous membrane, which reach from the soft palate to tlie tongue. Tliese elevations are the pillar's of the fauces. Each bifurcates below, and in the liollow between its divi- sions lies a tonsil (7, Fig 95), a soft rounded body about the size of an almond, and containing numerous minute glands which form mucus. Note. 1'he tonsils not unfrequently become enlarged during a cold or sore throat, and then pressing on the Eustachian tube (Chap. XXXIII. ), which leads from the pharynx to the middle ear, keep it closed and produce tem- porary deafness. Sometimes the enlargement is permanent and causes much annoyance. Tlie tonsils can, however, be readily removed without danger, and this is the treatment usually adopted in such cases. The Pharynx or Throat Cavity (Fig. 89). This por- tion of the alimentary canal may be described as a conical bag with its broad end turned upwards towards the base of the skull, and its narrow end downwards and passing into the gullet. Its front is imi)erfect, presenting apertures which lead into the nose, the mouth, and (through the larynx and windpipe) into the lungs. Except when food is being swallowed the soft palate hangs down between the mouth and pharynx; during deglutition it is raised into a horizontal joosition and separates an upper or resjnratory portion of the pharynx from the rest. Through this upper part, therefore, air alone passes, entering it from the posterior ends of the two nostril chambers; wliile through th3 lower portion both food and air j^ass, one on its way to the gullet, 1), Fig. 89, the other through the larynx, d, to the windpipe, c ; when a morsel of food "goes the wrong way" it takes the latter course. Opening into the upper portion of the pharynx on each side is an Eustachian tube, g-. so that the apertures leading out of it are seven in num- ber; the two posterior nares, the two Eustachian tubes, the fauces, the opening of the larynx, and that of the gul- let. At the root of the tongue, over the opening of the larynx, is a plate of cartilage, the ejnglottis, e, which can be seen if the mouth is widely opened and the back of the tongue pressed down by some such thing as the handle of THE STOMACH 317 a s[)Oon. During swallowing the epiglottis is i^ressed clown like a lid over the air-tube and helps to keep food or saliva from entering it. In structure the pharynx consists essen- tially of a bag of connective tissue lined by mucous mem- brane, and having muscles in its walls which, by their con- tiactions, drive the food on. The CEsophagus or Giillet is a tube commencing at the lower termination of the pharynx and which, passing on through the neck and chest, ends in the stomach below the diaphragm. In the neck it lies close behind the windpipe. It consists of three coats — a mucous membrane within ; next, a submucous coat of areolar connective tissue; and, outside, a muscular coat made up of two layers, an inner with trans- verse and an outer with longitudinally arranged fibres. In and beneath its mucous membrane are numerous small glands whose ducts open into the tube. The Stomach (Fig. 96) is a somewhat conical bag placed transversely m the upper part of the abdominal cavity. Its larger end is turned to y the left and lies close beneath the diaphragm; opening into its upper border, through the car- diac orifice at a, is the gullet, d. The narrower right end is continuous at c Avitli the small intes- tine; the communication between the two is the pyloric orifice The py- loric end of the stomach lies lower in the abdomen than the cardiac, and is separated from the diapliragm by the liver (see Fig. 1). The concave border between the two orifices is known as the small curvature, and the convex as the great curvature, of the stomach. From the latter hangs down a fold of peritoneum {ne, Fig. 1) known as the great omentum. It IS spread over the rest of the abdominal contents like an .^/ Fig. 9R.— The stomach, cf, lower end of the gullet; a. position of the cardiac aper tiii'n : 6, the fundus : c, the pylorus: e. the commencement of the small intestine along a, 6, c, the great curvature; between the pylorus and d, the lesser curvature. 318 THE HUMAN BODY. apron. After middle life much fat frequently accumulates in the omentum, so that it is largely responsible for the "fair round belly with good capon hn'd." The protrusion h to the left side of the cardiac orifice, Fig. 96, is i\\c fun- dus or (jrctd cul de sac. The size of the stomach varies greatly with the amount of food in it; just after a mode- rate meal it is about ten inches long, by five wide at its Ijroadest part. Structure of the Stomach. This organ has four coats, known successively from without in as the serous, the mus- cular, the submucous, and the mucous. The serous coat is formed by a reflexion of the peritoneum, a double fold of which slings the stomach; after separating to envelop it the two layers again unite and, hanging down beyond it, form the great omentum. The muscular coat (Fig. 54*) consists of un.striped muscilar tissue arranged in three layers: an outer, longitudinal, most developed about the curvatures; a circu- lar, evenly spread over the whole organ, except around the pyloric orifice where it forms a thick ring; and an inner, oblique and very incomplete, radiating from the cardiac orifice. The suhmicous coat is made up of lax areolar tissue and binds loosely the mucous coat to the muscular. The mucous coat is a moist pink membrane which is inelastic, and large enough to line the stomach evenly when it is fully distended. Accordingly, when the organ is empty and shrunk, this coat is thrown into folds. During digestion the arteries supplying the stomach become dilated and, its capillaries being gorged, its mucous membrane is then much redder than when the organ is empty. The blood-vessels of the stomach run to it between the folds of peritotieuni which sling it. After giving off a few branches to the outer layers, most of the arteries break up into small branches in the submucous coat, from wiiich twigs proceed to supply the close capillary network of the mucous membrane. The pneiimogastric nerves (p. 171) end in the stomach, and it also gets branches from the sympathetic system. Histology of the Gastric Mucous Membrane. Exami- nation of the inner surface of the stomach with a hand *P. 124. STRUCTURE OF STOMACH. 319 lens shows it to be covered with minute shallow pits. Into these open the mouths of minute tubes, the gastric glands, which are closely packed side by side in the mucous membrane; there being between them only a small amount of connective tissue, a close netAvork of lymph-channels, and capillary blood-vessels. The whole surface of the mucous membrane is lined by a single layer of columnar epithehum cells (Fig. 97). These dip down and line the tubular glands, being in some (especially those about the pyloric end of the stomach) but little modified in appear- ance (c, Fig. 97). In others the epithelial cells become shorter and cuboidal, and have beneath them {a and b. Fig. 97) a second incomplete layer of much larger oval cells, d. The glands of this second kind are the most numerous, and have been called peptic glands from the idea that they alone formed pepsin, the essential digestive ingredient of the gastric juice; this is how- ever by no means certain. The peptic glands frequently branch at their deeper ends. The Pylorus. If the stomach be opened it is seen that the mucous membrane projects in a fold around the pyloric orifice and narrows it. This is due to a thick ring of the circular muscular layer there developed, and forming a spTiincter muscle around the orifice, which in life, by its contraction, keeps the passage to the small intestines closed except when portions of food are to be passed on from the stomach to succeeding divisions of the alimentary canal. Xote. The cardiac end of the stomach lying immediately beneath the diaphragm, which has the heart on its upper side, over-distension of the stomach, due to indigestion or flatulence, may impede the action of the thoracic organs, and Fig. 97.— a thin section through the gastric mucous membrane, perpendi- cular to its surface, magnified about 25 diameters, a. a simple peptic gland : 6, a c 'mpoimd peptic gland ; c, a mu- cous gland; d, oval, chief, or so-called peptic cells. 32G THE HUMAN BODY. cauRO feelings of oppression in (lie chest, or palpitation of the heart. The Small Intestine, eommeneiiig at the pylorus, ends. after maiiv Avindiiigs, in the large. It is about six meters (twenty feet) long, and about five centimeters (two inches) ^vide at its gastric end, narrowing to about two thirds of t hat width at its lower portion. Externally there are no lines of subdivision on the small intestine, l)ut anatomists ;;rbitrarily describe it as consisting of three parts; the first I welve inches being the duodenum, the succeeding two fifths of the remainder the jejunum, and the rest the ileum. Like the stomach, the small intestine i)osscsses four coats; a serous, a muscular, a submucous, and a mucous. The serous coat is formed by a duplicature of the peritoneum, but presents nothing answering to the great omentum; this double fold, slinging the intestine as the small omentum slings the stomach, is called the mesentery. The muscular coat is composed of plain muscular tissue arranged in two strata, an outer longitudinal, and an inner transverse or circular. The submtccous coat is like that of the stomach; consisting of loose areolar tissue, binding together the mu- cous and muscular coats, and forming a bed in which the blood and lymphatic vessels (which reach the intestine in the fold of the mesentery) break up into minute branches before entering the mucous membrane. The Mucous Coat of the Small Intestine. This is pink, soft, and extremely vascular. It does not present tempo- rary or effaceable folds like those of the stomach, but is. throughout a great portion of its length, raised up into j)er- manent transverse folds in the form of crescentic ridges, each of which runs transversely for a greater or less way round the tube (Fig. 98). These folds are the valvuhr C07iniventes. They are first found about two inches from the pylorus, and are most thickly set and largest in the upper half of the jejunum, in the lower half of which they become gradually less conspicuous; and they finally disappear altogether about the middle of the ileum. The folds serve greatly to increase the surface of the mucous mem- brane both for absorption and secretion, and they also de- SMALL INTESTINE. 321 lay the food somewliat in its passage, since it must collect in the hollows between them, and so be longer exposed to the action of the digestive litpiids. Examined closely with the eye or, better, with a hand lens, the mucous membrane of the small intestine is seen to be not smooth but shaggy, be- ing covered everywhere (both over the valvulae conniventetf Mild between them) with closely packed minute processes, standing up somewhat like the "• pile" on velvet, and known as the villi. Each villus is from 0.5 to 0.7 millimeters (^i_ to -3»^ inch) long ; some are conical and rounded, but the majority are compressed at the base in one diameter (Fig. 99). In structure a villus is somewhat complex. Covering it is a single layer of columnar epithelial cells, be- neath which the villus may be regarded as made up of a Fig. 98. — A portion of the small intestine opened to show the valvulae conni- ventes. framework of connective tissue supporting the more essen- tial constituents. Near the surface is an incomplete layer of plain muscular tissue, continuous below with a muscular layer found on the deep side of the mucous membrane. In the centre is an offshoot of the lymphatic system; some- times in the form of a single vessel with a closed dilated end, and sometimes as a network formed by two main ves- sels with cross-branches. During digestion these lym- phatics are filled with a milky white liquid absorbed from the intestines and they are accordingly called the lacteals. They communicate with larger branches in the submucous coat which end in trunks that pass out in the mesentery to join t'.ie main lymi)hatic system. Finally, in each villus, 322 THE HUMAN BODY. outside the lacteals find beneath the muscular layer, is a close network of blood-vessels. Opening on the surface of the small intestine, between the bases of the villi, are small glands, the crypts of Lieber- kilhn. Each is a simple unbranched tube lined by a layer of columnar cells similar to that wliich covers the villi and the surface of the mucous membrane between them. In Fig. 99.— Villi of the small intestine; magnified about 80 diameters. In the left-hand figure the lacteals, a, b, c, are filled with white injection; (/, blood-ves- sels. In the right-hand ligure the lacteals alone are represented, filled with a dark injection. The epith. hum covering the vilh, and their muscular fibres, are omitted. structure they greatly resemble the mucous glands of the stomach {c, Fig. 97). In the duodenum are found other minute glands, the glands of Brunner. They lie in the submucous coat and send their ducts through the mucous membrane to open on its inner side. The Large Intestine, forming the final portion of the alimentary canal, is about 1.5 meters (5 feet) long, and varies in diameter from about 6 to 4 centimeters {2^ to 1|- inches). Anatomists describe it as consisting of the ccecum Avith the vermiform appendix, the colon, and the rectum. The small intestine does not open into the commencement of the large but into its side, some distance from its closed THE LIVER. 323 upper end, and the caecum is tliut part of the large intes- tine which extends beyond the communication. From it projects the vermiform appendix, a narrow tube not thicker than a cedar pencil, and about 10 centimeters (4 inches) long. The colon commences on the right side of the abdominal cavity where the small intestine communicates with the large, runs up for some way on that side {ascend- ing colon), then crosses the middle line {transverse colon) below the stomach, and turns down {descending colon) on the left side and there makes an S-shaped bend known as the sigmoid flexure; from this the rectum, the termi- nal straight portion of the intestine, proceeds to the anal opening, by which the alimentary canal communicates with the exterior. In structure the large intestine presents the same coats as the small. The external stratum of the muscular coat is not, however, developed uniformly around it, except on the rectum, but occurs in three bands separated by intervals in which it is wanting. These bands being shorter than the lest of the tube cause it to be puckered, or sacculated, between them. The mucous coat possesses no villi or valvule conniventes, but is usually thrown into effaceable folds, like those of the stomach but smaller. It contains numerous closely set glands much like the crypts of Lieberkiihn of the small intestine. The neo-Colic Valve. Where the small intestine joins the large there is a valve, formed by two flaps of the mucous membrane sloping down into the colon, and so disposed as to allow matters to pass readily from the ileum into the large intestine but not the other way. The Liver. Besides the secretions formed by the glands imbedded in its walls, the small intestine receives those of two other large glands, the liver and pancreas, which lie in the abdominal cavity. The ducts of both open by a common aperture into the duodenum about 10 centimeters (4 inches) from the pylorus. The liver is the largest gland in the Body, weighing from 1400 to ITOO gi-ams (50 to 64 ounces). It is situated in the upper part of the abdominal cavity {le, le. Fig. 1), rather more on the riglit than on the left side and immediately below 3U THE HmfAiV BODY. the diaphragm, into the concavity <>f wliich its upper sur- face fits; it reaches across the middle line above the pyloric end of the stomach. It is of dark reddish-brown coloi, and of a soft friable texture. A deep fissure incompletely divides the organ into r?'_(//// and left lohes, of which the right is much the larger; on its under surface (Fig. 100) shallower grooves mark olf several minor lobes. Its upper surface is smooth and convex. The vessels carr3ing blood to the liver are i\\e 2)ortal vein, Vp, and the Jiepatic artery; both enter it at a fissure {the portal fissure) on its under side, and there also a duct passes out from each half of the organ. Fig. 100.- Til e imrler surface of the liver. cJ. right, and s, left lobe ; Vli, hepatic vein; Vp, portal vein; Fc, vena cava inferior; Dch, common bile-duct; Dc cystic duct; Dh, hepatic duct; Vf, gall-bladder. The ducts unite to form the hepatic duct, Dh, which meets at an acute angle, the cystic duct, Dc, proceeding from the gall-bladder, Vf, a pear-shaped sac in which the bile, or gall, formed by the liver, accumulates when food is not being di- gested in the intestine. The common hile-duct, Dch, formed by the union of the hepatic and cystic ducts, opens into the duodenum. The blood which enters the liver by the ])ortal vein and hepatic artery passes out by the hepatic veins, Mt, which leave the posterior border of the organ HISTOLOGY OF LIVEB. 335 close to the vertebral column, and there open into the infe- rior vena cava, Vc, jnst before it passes up through the diaphragm. The Structure of the Liver. On closely examining the surface of the liver, it will be seen to be marked out into small angular areas from one to two millimeters {^ to y^j inch) in diameter. These are the outer sides of the superficial layer of a vast number of minute polygonal masses, or lohules, of which the liver is built up; similar areas are seen on the surface of any section made through the organ. Each lobule (Fig. 101) consists of a number of «^H -^o-^tr "Q-^ihy-^ "^^MHi Fig. 101. — A lobule of the liver, magrnifled, showing the hepatic cells radiately arrange J around thecenti-al Latralobular vein, and the lobulai- capillaries inter- laced with them. liepatic cells supjiorted by a close network of capillaries; and is separated from neighboring lobules by connective tissue, larger blood-vessels, and branches of the hepatic duct. The hepatic cells are the proper tissue elements of the liver, all the rest being subsidiary arrangements for their nutrition and protection. Each is polygonal, nucle- ated and very granular, and has a diameter of about .025 millimeter (yo'ito" of ^^^ inch). In each lobule they are ar- ranged in rows or strings, which intercommunicate and form a network, in the meshes of which the blood capilla- 326 THE HUMAN BODY. rios run. Covering the surface of the liver is a layer of the peritoneum, beneath which is a dense connective-tissue layer, forming the capsule of Glisson. At the portal fissure offsets from this capsule run in, and line canals, the portal canals, which are tunneled through the organs. These, becoming smaller and smaller as they branch, finally be- come indistinguishable close to the ultimate lobules. From their walls and from the external capsule, connective- tissue partitions radiate in all directions through the organ and support its other parts. In each portal canal lie three ves- sels— a branch of the portal vein, a branch of the hepatic Fig. 102.— a small portion of the liver, injected, and magmifled about twenty diameters. The blood-vessels are represented wlute ; the large vessel is a sub- lobnlar vein, receiving the intralobular veins, which m turn are derived from the capillaries of the lobules. artery, and a branch of the hepatic duct; the division of the portal vein being much the largest of the three. These vessels break up as the portal canals do, and all end in minute branches around the lobules. The blood carried in by the portal vein (which has already circulated through the capillaries of the stomach, spleen, intestines and pan- creas) is thus conveyed to a fine vascular interlohtilar plexus around the liver lobules, from which it flows on through the capillaries {lohular plexus^ of the lobules them- selves. These (Fig. 101) unite in the centre of the lobule HEPATIO CIRCULATION. 327 \fO form a small intralohular vein, which carries the blood out and pours it into one of the branches of origin of the Fig. 103.— The stomach, pancreas, liver, and duodenum, -with part of the rest of the small intestine and the mesentery; the stomach and hver have been turned up so as to expose the pancreas. V, stomach; D. ly . D" , duodenum; L, spleen ; i^, pancreas ; R. right kidney ; T, jejunum ; Vf, gall-bladder ; /i, hepatic duct ; c, cystic duct ; ch. common bile-duct ; 1, aorta ; 2, an artery (left coronary) of the stomach ; 3, hepatic artery; 4, splenic artery ; 5, superior mesenteric artery ; 6, superior mesenteric veui ; 7, splenic vein ; Vp, portal vein. hepatic vein, called the sublohular vein. Each of the latter has many lobules emptying blood into it, and if dissected out 328 THE nUMAN BODY. with them (Fig. 102) would look sonietliing like a branch of a tree with apples attached to it by short stalks, repre- sented by the intralobular veins. The blood is finally car- ried, as above pointed out, by the he^iatic veins into the inferior vena cava. The hepatic artery, a branch of the cceliac axis (p. 211), ends mainly in Glisson's capsule and the walls of the blood-vessels and bile-ducts, but some of its blood reaches the lobular plexuses; it all finally leaves the liver by the hepatic veins. The bile-ducts can be readily traced to the periphery of the lobules, and there proba])ly communicate with a minute network of commencing bile-ducts ramifying in the lobule between the hepatic cells composing it. The Pancreas or Sweetbread. This is an elongated soft organ of a pinkish yellow color, lying along the great curvature of the stomach. Its right end is larger, and is embraced by the duodenum (Fig. 103), which there makes a curve to the left. A duct traverses the gland and joins the common bile-duct close to its intestinal opening. The pancreas forms a watery-looking secretion wiiich is of great importance m digestion. CHAPTER XXII. THE LYMPHATIC SYSTEM AND THE DUCT- LESS GLANDS. The Lymphatics or Absorbents form close networks in nearly all parts of the Body. Most organs, as has been pointed out (p. 62), possess a sort of internal skeleton made up of connective tissue, which consists mainly of bundles of fibres, united together and coyered in by a "cement" sub- stance. In this substance are found numerous cavities, usually branched, and communicating with one another by their branches. They frequently contain connective-tissue corpuscles, which, however, do not completely fill them; and they thus, with their branches, form a set of intercom- municating channels known as the "serous canalicidij" these are filled with lymph and constitute the origin of lymphatic vessels in many organs. Elsewhere the com- mencing lymphatics seem to be merely interstices {lacunm) between the constituent tissues of an organ; this is esjie- cially the case in glands. Such spaces differ from the se- rous canaliculi in being lined by a definite epithelium. Structure of Lymph-Vessels. The serous canaliculi and lym})h-spaces open into better defined channels, lined with a single layer of wavy-edged flattened epithelial cells. These form networks in most parts of the Body and are knowTi as the lymph capillaries. They are usually wider til an blood capillaries. From the capillary networks larger vessels arise which in structure resemble veins, and have similar, but more numerous, valves. The Thoracic Duct. All the lymphatics end finally in two main trunks which open into the venous system .on each side of the neck, at the point of junction of the jugular and 330 THE HUMAN BODY. subclavian. The trank on the right side is much smaller than the other and is known as the "right lymphatic duct." It collects lymph from the right side of the thorax, from the right side of the head and neck, and the right arm. The lymph from all the rest of the Body is collected into the thoracic duct. It commences at the uj)2oer part of tlie abdominal cavity in a dilated reservoir (the receptaculum chyli), into which the lacteals from the intestines, and the lymphatics of the rest of the lower part of the Body, open. From thence the thoracic duct, receiving tributaries on its course, runs up the thorax alongside of the aorta and, pass- ing on into the neck, ends on the left side at the point already indicated; receiving on its way the main stems from the left arm and the left side of the head and neck. The thoracic duct, thus, brings back much more lymph than the right lymphatic duct. The Serous Cavities. These are great dependencies of the lymphatic system and may be regarded as large lacunae. Each of them (peritoneal, pleural, arachnoidal and peri- cardiac) is lined by a definite epithelioid layer of close-fit- ting, hexagonal cells. At certain points, however, o^jenings or stomata occur, surrounded by a ring of smaller cells, and leading into tubes which open into subjacent lymphatic vessels. The liquid moistening these cavities is, then, really lymph. The Lymphatic Glands. These are roundish masses in- terposed, at various points, on the course of the lymph- ves- sels. They are especially numerous in the mesentery, groin, and neck. In the latter position they often inflame and give rise to abscesses, especially in scrofulous persons; and still more often enlarge, harden, and become more or less tender, so as to attract attention to them. In common par- lance it is then frequently said that the person's " kertiels have come down," or that he has " waxing kernels." Each lymphatic gland is enveloped in a connective-tissue capsule, and is pervaded by a connective-tissue framework. In the meshes of this lie numerous lymph corpuscles, which appear to multiply there by division. "Afferent" lymphatic ves- sels open into the periphery of the so-called gland, and ef- MOVEMEWl OF TEE LYMPH. 331 ferent vessels arise in its centre. Hence, the lymiDh in its flow traverses tlie cellular gland substance, and in its course picks up extra corpuscles which it carries on to the blood. In the giand there is a close network of blood capil- laries. It is clear that these organs are not glands at all, in the proper sense of the word. They are sometimes called lymphatic ganglia; but that suggests a false connection with nerve-centres. The Movement of the Lymph. This is no doubt some- what irregular in the commenciug vessels, but, on the whole, sets on to the larger trunks and through them to the veins. In many animals (as the frog) at points where the lymphatics communicate with the veins, there are found regularly contractile "lymph-hearts" which beat with a rhythm inde- pendent of that of the blood-heart, and pump the lymph into a vein. In the Human Body, however, there are no such hearts, and the flow of the lymph is dependent on less definite arrangements. It seems to be maintained mainly by three things. (1) The pressure on the blood plasma in the capillaries is greater than that in the great veins of the neck; hence any plasma filtered through the capillary walls will be under a pressure Avliich will tend to make it flow to the venous termination of the thoracic or the right lym- phatic duct. (2) On account of the numerous valves in the lymphatic vessels (which all only allow the lymph to flow past them to larger vessels) any movement compress- ing a lymph-vessel will cause an onward flow of its contents. The influence thus exerted is very important. If a tube be put in a large lymph-vessel, say at the top of the leg of an animal, it will be seen that the lymph only flows out very sloMdy when the animal is quiet; but as soon as it moves its leg the flow is greatly accelerated. (3) During each inspiration the pressure on the thoracic duct is less than that in the lymphatics in parts of the Body outside the thorax (see Chap. XXIV. ). Accordingly, at that time, lymph is pressed, or, in common jihrase, is " sucked," into the thoracic duct. During the succeeding expiration the pres- sure on the thoracic duct becomes greater again, and some of its contents are pressed out: 1>ut on account of the valves 332 THE nUMAX BODY. tliey can only ac forwards, tliat is, towards the ending of the duct in the veins of the neck. During digestion, moreover, the contraction of the villi will press on the lymph or chyle; and in certain parts of the Body gravity, of course, aids the flow, though it will impede it in others. The Spleen. There are in the Body several organs of such consideraljle size, and of so great constancy in a large number of vertebrate animals, that they would a j^^'iori appear to be of considerable functional importance. What their use may be is still, however, unknown or uncertain. They are commonly spoken of collectively, along with the lymphatic ganglia, as the ductless glands; but they are not glands in the proper sense of the word. The spleen is the largest of them. It is a red organ situated at the left end of the stomach and about 170 grams (6 oz.) in weight. Its size is however very variable; it enlarges during diges- tion and shrinks again after it until the next meal. In malarial diseases it also becomes enlarged, frequently to a very great extent, and then constitutes the so-called '* ague- cake," In structure, the spleen consists of a connective- ti?sne capsule, rich in elastic fibres, and giving off processes whicli ramify through the organ and form a framework for its pidj). The latter contains numerous blood corpuscles; and maiiy bodies which seem to be red corpuscles in pro- cess of decay or destruction. Hence the spleen has been supposed to be a sort of gi-aveyard for their bodies — a place where they are broken up and their materials utilized when they have run their life cycle. Others, however, consider that in the spleen new red blood corpuscles are pro- duced from colorless; and others, again, that the main function of the organ is the formation of substances which are carried off to the stomach and pancreas, to be there finally elaborated into digestive ferments. The arteries of the spleen open directly into the pulp cavities, from which the veins arise. On their walls are rounded whitish nodules about the size of a millet-seed, and known as the THYMUS AND- THYROID. 333 Malpighian corpuscles. They resemble tiny lymphatic glands in structure. The Thymus is an organ which only exists in childhood. At birth it is found lying around the windpipe, in the upper part of the chest cavity and the lower part of the neck. It increases in size until the end of the second year, and then begins to dwindle away. It is grayish pink in color, of a soft texture, and, in microscopic structui'e, resembles some- what a lymphatic gland. Hence its function has been sup- posed to be the formation of ne^v lymph corpuscles. The "sweetbread" of butchers is sometimes the pancreas and sometimes the thymus of young animals — neck and belly sweetbread. The Thyroid Body and the Suprarenal Capsules. The former of these structures lies in the neck on the sides of and below "Adam's apj^le." It is dark red-brown in color, and sometimes becomes very much enlarged, as in the disease known as goitre. This enlargement ajjpears to be often due to drinking water containing magnesian limestone in solution. In England, for instance, it is known as "Der- byshire neck" from being especially frequent in parts of that county, where the hills are mainly composed of magnesian limestone rocks; and the same geological formation is found in those districts of Switzerland where cretinism (one of the symptoms of which is an enlarged thyroid body) prevails. The suprarenal bodies lie one over the top of each kid- ney. Their use, like that of the thyroid, is quite pro- blematical. In what is known as Addison's disease (in which the skin becomes of a bronze color) it is said that these bodies are altered; but it is very improbable that the change in them is the actual cause, rather than another symptom, of the disease. CHAPTER XL DIGESTION. The Object of Digestion. Of the various foodstufEe swallowed, some are already in solution and ready to dialyze at once into the lymphatics and blood-vessels of the alimen- tary canal; others, such as a lumjj of sugar, though not dissolved when put into the mouth, are readily soluble in the liquids found in the alimentary canal, and need no fur- ther digestion. In the case of many most imijortant food- stuffs, however, special chemical changes have to be wrought, either with the object of converting insoluble bodies into soluble, or non-dialyzable into dialyzable, or both. The different secretions poured into the alimentary tube act in various ways upon different foodstuffs, and at last get them nito a state in which they can pass into the circulating medium and be carried to all parts of the Body. The Saliva. The first solvent that the food meets with is the saliva, which, as found in the mouth, is a mixture of pure saliva, formed in parotid, submaxillary, and sublingual glands, with the mucus secreted by small glands of the oral mucous membrane. This mixed saliva is a colorless, cloudy, feebly alkaline liquid, "ropy" from the mucin present in it, and usually containing air-bubbles. Pure saliva, as ob- tained by putting a fine tube in the duct of one of the sali- vary glands, is less tenacious aud contains no imprisoned air. The uses of the saliva are for the most part physical and mechanical. It keeps the mouth moist and allows us to speak with comfort; most young orators know the distress occasioned by the suppression of the salivary secretion through nervousness, and the imperfect efl&cacy under such USES OF SALIVA. 335 circumstances of the traditional glass of water placed be- side public speakers. The saliva, also, enables us to swallow dry food; such a thing as a cracker when chewed would give rise merely to a heap of dust, impossible to swallow, were not the mouth cavity kept moist. This fact used to be taken advantage of in the East Indian rice ordeal for the detection of criminals. The guilty person, believing firmly that he cannot swallow the parched rice given him and sure of detection, is apt to have his salivary glands paralyzed by fear, and so does actually become unable to swallow the rice; while in those with clear consciences the nervous system, acting normally, excites the usual reflex secretion, and the dry food causes no difficulty of degluti- tion. The saliva, also, dissolves such bodies as salt and sugar, when taken into the mouth in a solid form, and enables us to taste them; undissolved substances are not tasted, a fact which any one can verify for himself by Aviping his tongue dry and placing a fragment of sugar upon it. No sweetness will be felt until a little moisture has exuded and dissolved part of the sugar. In addition to such actions the saliva, however, exerts a chemical one on an important foodstuff. Starch (although it swells up greatly in hot water) is insoluble, and could not be absorbed from the alimentary canal. The saliva contains a specific element, ptyalin, which has the power of turning starch into the readily soluble and dialyzable grape sugar. In effecting this change the ptyalin is not altered; at least a very small amount of it can cause the conversion of a vast amount of starch, and it does not seem to have its activity impaired in the process, being still ready at the end of it to act upon more. The starch is made to combine with the elements of a molecule of water, and the ptyalin remains behind as it was — starch. Water. Grape sugar. Substances acting in this way, producing chemical changes without being themselves noticeably altered, are found in many of the digestive secretions, and are called 336 THE TTr.VA y BOD Y. ferments or enzymes. Tlicy differ from the true ferments, such as yeast, in tlie fact tliat they are not living organisms, and do not multiply during the occurrence of the change whicli tlicy sot up; tlieir activity belongs to the obscure chemical category of contact actions. In order that the i^tyalin may act upon starch certain conditions are essential. Water must be present, and the liquid must be neutral or feebly alkaline; acids retard, or if stronger, entirely stop the process. The change takes place most quickly at about the temperature of the Human Body, and is greatly checked by cold. Boiling the saliva destroys its ptyalin and renders it quite incapable of converting starch. Boiled starch is changed more rapidly and com- pletely than raw. Saliva has another important but indirect influence in promoting digestion. Weak alkalies stimulate the mucous membrane of the stomach and cause it to pour forth more gastric juice. Hence the efficac}' of a little carbonate of soda, taken before meals, in some forms of dyspepsia. The saliva by its alkalinity exerts such an action; and this is one reason why food should be well chewed before being swallowed; for then its taste, and the movements of the Jaws, cause the secretion of more saliva. Deglutition. A mouthful of solid food is broken uj? by the teeth, and rolled about the mouth by the tongue, until it is thoroughly mixed with saliva and made into a soft pasty mass. The muscles of the cheeks keep this from getting between them and the gums; persons with facial paralysis have, from time to time, to press out with the finger food which has collected outside the gums, where it can neither be chewed nor swallowed. The mass is finally sent on from the mouth to the stomach by the process of derjlntition, which is described as occurring in three stages. The first stage includes the passage from the mouth into the pharynx. The food being collected into a heap on the tongue, the tip of that organ is placed against the front of the hard palate and then the rest of its dorsum is raised from before back, 30 as to press the food mass between it and the palate, and drive it back through the fauces. This j)ortion of the act of SWALLOWT^^G. 337 swallowing is volimtary, or at least is under the control of the will, ulthougli it commonly takes place unconsciously. The second stage of deglutition is that in which the food passes through the pharynx; it is the most rapid part of its progress, since the pharynx has to be emptied quickly so as to clear the opening of the air-passages for breathing pur- poses. The food mass, passing back over the root of the tongue, pushes down the epiglottis; at the same time the larvnx (or voice-box at the top of the windpipe) is raised, so as to meet this and thus the passage to the lungs is closed; muscles around the aperture probably also contract and narrow the opening. The raising of the larynx can be readily felt by placing the finger on its large cartilage form- ing " Adam's apple" in the neck, and then swallowing something. The soft palate is at the same time raised and stretched horizontally across the pharynx, thus cutting off communication with its upper, or respiratory portion, lead- ing to the nostrils and Eustachian tubes. Finally, the isthmus of the fauces is closed, as soon as the food has passed through, by the contraction of the muscles on its sides and the elevation of the root of the tongue. All pas- sages out of the pharynx except the gullet are thus blocked, and when the pharyngeal muscles contract the food can only he squeezed in+o the a3sophagus. The muscular move- ments concerned in this part of deglutition are all reflexly excited; food coming in contact with the mucous mem- brane of the pharynx stimulates afferent nerve-fibres in it; these excite the centre of deglutition which is placed in the medulla ohlongata, and from it efferent nerve-fibres proceed to the muscles concerned and (under the co-ordinating in- fluence of the centre) cause them to contract in proper se- quence. The pharyngeal muscles, although of the striped variety, are but little under the control of the will; it is ex- tremely difiicult to go through the movements of swallow- ing without something (if only a little saliva) to swallow, and excite the movements reflexly. Many i^ersons after having got the mouth completely empty cannot perform the movements of the second stage of deglutition at all. On account of the reflex nature of the contractions of the 338 THE HUMAN BODY. pliarynx, any food wliich lias oucc entered it must be swal- lowed: the isthmus of the fauces forms a sort of Kubicon; food that has passed it must continue its course to the stomach although the swallower learnt immediately that he was taking poison. The third stage of deglutition is that in which the food is passing along the gullet, and is compara- tively slow. Even liquid substances do not fall or flow down this tube, but have their passage controlled by its muscular coats, which grip the successive portions swallowed and pass them on. Hence the possibility of performing the ajiparently wonderful feat of drinking a glass of water while standing upon the head, often exhibited by jugglers; people forget- ting that one sees the same thing done every day by horses, and other animals, which drink with the pharyngeal end of the gullet lower than the stomach. The movements of the oesophagus are of the kind known as vermicular or peri- staltic. Its circular fibres (p. 317) contract behind the morsel and narrow the passage there; and the constriction then travels along to the stomach, pushing the food in front of it. Simultaneously the longitudinal fibres, at the jioint where the food-mass is at any moment and immediately in front of that, contracting, shorten and widen the passage. The Gastric Juice. The food having entered the sto- mach is exposed to the action of the gastric juice, which is a thin, colorless, or pale yellow liquid, of a strongly acid re- action. It contains as sj^ecific elements free liydrocliloric acid (about .€)2 per cent), and an enz3"me called pepsin which, in acid liquids, has the power of converting the or- dinary non-dialyzablei^roteids which we eat, into the closely allied but dialyzable bodies called peptones. It also dis- solves solid j)roteids, changing them too into pej^tones. Dilute acids will by themselves produce the same changes in the course of several days, but in the iDresence of pejDsin and at the temperature of the Body the conversion is far more rapid. In neutral or alkaline media the pepsin is inactive; and cold check sits activity. Boiling destro3\s it. In addition to pepsin, gastric juice contains another enzyme which has the power of coagulating the casein of milk, as illustrated by the use of ^'rennet," prejjared from the mu- DIGESTIoy IN THE STOMACH. 339 cons membrane of the calf's digestive stomach, in cheese- making. The acid of the natural gastric juice might itself, it is true, coagulate the casein, but neutralized gastric juice still possesses this power; and, since pure solutions of pepsin do not, it must be due to some third body, which has, however, not yet been isolated. The curdled condition of the milk regurgitated so often by infants is, therefore, not any sign of a disordered state of the stomach, as nurses commonly suppose. It is natural and proper for milk to undergo this change, before the pepsin and acid of the gastric juice convert its casein into peptone. Gastric Digestion. The jDrocess of swallowing is con- tinuous, but in the stomach the onward progress of the food is stayed for some time. The pyloric sphincter, re- maining contracted, closes the aperture leading into the intestine, and the irregularly disposed muscular layers of the stomach keep its semi-liquid contents in constant movement, maintaining a sort of churning by which all portions are brought into contact with the mucous mem- brane and thoroughly mixed with the secretion of its glands. The gelatin-yielding connective tissue of meats is dissolved away, and the proteid-containing fibres, left loose, are dis- solved and turned into peptones. The albuminous walls of the fat-cells are dissolved and their oily contents set free; but the gastric juice does not act upon the latter. Certain mineral salts (as phosphate of lime, of which there is always some in bread) which are insoluble in water but soluble in dilate acids, are also dissolved in the stomach. On the other hand the gastric juice has itself no action upon starch, and since ptyalin does not act at all, or only imperfectly, in an acid medium, the activity of the saliva in converting starch is stayed in the stomach. By the solu- tion of the white fibrous connective tissue, that disintegra- tion of animal foods commenced by the teeth, is carried much farther in the stomach, and the food-mass, mixed with much gastric secretion, becomes reduced to the con- sistency of a thick soup, usually of a grayish color. In this state it is called chyme. This contains, after an ordi- nary meal, a considerable quantity of peptones which are 340 THE HUMAN BODY. in great part gradually dialyzod into the blood and lympha- tic vessels of the gastric mucous membrane, and carried off, along with other dissolved dialyzable bodies, such as salts and sugar. After the food has remained in the stomach some time (one and a half to two hours) the chyme begins to be passed on into the intestine in successive portions. The pyloric sphincter relaxes at intervals, and tlie rest of the stomach, contracting at the same moment, injects a ((uantity of chyme into the duodenum ; this is repeated frequently, the larger undigested fragments being at first unable to pass the orifice. At the end of about three or four hours after a meal the stomach is again quite emptied, the pyloric sphincter finally relaxing to a greater extent and allowing any larger indigestible masses, which the gas- tric juice cannot break down, to be squeezed into the intes- tine. The Chyle. When tlie chyme passes into the duodenum it finds preparation made for it. The jiancreas is in reflex connection with the stomach, and its nerves cause it to commence secreting so soon as food enters the latter; hence a quantity of its secretion is already accumulated in the intestine when food enters. The gall-bladder is distended with bile, secreted since the last meal; this passing down the hepatic duct has been turned back up the cystic duct {Dc, Fig. 100*) on account of the closure of the common bile-duct. The acid chyme, stimulating nerve-endings in the duodenal mucous meml)rane, causes reflex contrac- tion of the muscular coat of the gall-bladder, and a relaxa- tion of the orifice of the common bile-duct; and so a gush of bile is poured out on the chyme. From this time on, both liver and pancreas continue secreting actively for some hours, and pour their products into the intestine. The glands of Brunner and the crypts of Lieberkiihn are also set at work, but concerning their physiology we know very little. All of these secretions are alkaline, and they suffice very soon to more than neutralize the acidity of the gastric juice, and so to convert the acid chyme into alka- line chyle, which, after an ordinary meal, will contain a great variety of things: mucus derived from the alimen- *P. 324 PANCREATIC DIGESTION. 341 tary canal; ptyalin from the saliva; pepsin from the sto- mach; water, partly swallowed and partly derived from the salivary and other secretions; the .peculiar constitiients of the bile and pancreatic Juice and of the intestinal secretions; some undigested proteids; unchanged starch; oils from the fats eaten; peptones formed in tlie stomacli but not yet absorbed; possibly salines and sugar which have also escaped absorption in the stomach; and indigestible substances taken with the food. The Pancreatic Secretion is clear, watery, alkaline, and much like saliva in appearance. The Germans call the pancreas the '• abdominal salivary gland." In digestive properties, however, the pancreatic secretion is far more important than the saliva, acting not only on starch but, also, on proteids and fats. On starch it acts like the saliva, but more energetically. It produces changes in jiroteids similar to those effected in the stomach, but by the agency of a different ferment, trypsin; which differs from pepsin in acting only in an alkaline instead of an acid medium. On fats it has a double action. To a certain extent it breaks them up, with hydration, into free fatty acids and glycerine; for examjDle — (C"H.^O)> I 0. + 3H.0 = 3(C-H-.0 | o ) +C'II; | 0. 1 Stearin + 3 Water = 3 Stearic acid + 1 Glycerine. The fatty acid then combines with some of the alkali pres- ent to make a soap, which being soluble in water is cai)able of absorption. Glycerine, also, is soluble in water and diah^- zable. The greater part of the fats are not, however, so broken up, but are simply mechanically separated into little droplets, which remain suspended in the chyle and give it a whitish color, just as the cream-drops are suspended in milk, or the olive-oil in mayonnaise sauce. This is effected by the help of a quantity of albumin which exists dissolved in the pancreatic secretion. In the stomach, the animal fats eaten have lost their cell-walls, and have become melted by the temperature to which they are exposed. Hence their oily part floats free in the chyme when it enters the 342 THE HUMAN liOBY. duodenum. If oil be shaken up with water, the two can- not be got to mix; immediately the shaking ceases the oil floats up to the top; but if some raw Q'^g be added, a creamy mixLurc is readily formed, in which the oil remains for a long time evenly suspended in the watery menstruum. The reason of this is that eacli oil-droplet becomes sur- rounded by a delicate pellicle of albumen, and is thus pre- vented from fusing with its neighbors to make large drops, which would soon float to the toji. Such a mixture is called an emulsion, and the albumin of the pancreatic secretion emulsifies the oils in the chyle, which becomes white (for the same reason as milk is that color) because the innumerable tiny oil-drops floating in it reflect all the light which falls on its surface. The pancreatic secretion thus converts starch into grape sugar, dissolves proteids (if necessary) and converts them into peptones, emulsifies fats, and, to a certain extent, breaks them up into glycerine and fatty acids, which latter are saponified by the alkalies present. The Bile. Human bile when quite fresh is a golden brown liquid; it becomes green when kept. As formed in the liver it contains hardly any mucin, but if it makes any stay in the gall-bladder it acquires a great deal from, the lining membrane of that sac, and becomes very ''ropy." It is alkaline in reaction and, besides coloring matters, min- eral salts, and water, contains the sodium salts of two nitro- genized acids, taurocJiolic and glycliocholic, the former j^re- dominating in human bile. Pettenkofer's Bile Test. If a small fragment of cane sugar be added to some bile, and then a large quantity of strong sulphuric acid, a brilliant purple color is develo^ied, by certain products of the decomposition of its acids; the physician can in this way, in disease, detect their presence in the urine or other secretions of the Body. Gmelvii's Bile Test. The bile-coloring matters, treated with yellow nitric acid, go through a series of oxidations, accompanied with changes of color from yellow-brown to green, then to blue, violet, purple, red, and dirty yellow, in succession. Bile has no digestive action upon starch or proteids. It DIGESTIVE ACTION OF BILE. 343 does not break up fats, but to a limited extent emulsifies them, though far less perfectly tlian the iDancreatic secre- tion. It is even doubtful if this action is exerted in the intestines at all. In many animals, as in man, the bile and pancreatic ducts open together iuto the duodenum, so that, on killing them during digestion and finding emulsified fats in the chyle, it is impossible to say whether or no the bile had a share in the process. In the rabbit, however, the pancreatic duct opens into the intestine about a foot farther from the stomach than the bile-duct, and it is fonnd that if a rabbit be killed after being fed with oil, no milky chyle is found down to the point where the pancreatic duct opens. In this animal, therefore, the bile alone does not emulsify fats, and, since the bile is j^retty much the same in it and other mammals, it probably does not emulsify fats in them either. From the inertness of bile with respect to most foodstuffs it has been doubted if it is of any digestive use at all, and whether it should not be regarded merely as an excretion, poured into the alimentary canal to be got rid of. But there are many reasons against such a view. In the first place, the entry of the bile into the upper end of the small intestine where it has to traverse a course of more than twenty feet before getting out of the Body, instead of its being sent into the rectum close to the final opening of the alimentary canal, makes it probable that it has some function to fulfill in the intestine. Moreover, a great part of the bile poured into the intestines is again absorbed from them, only a small part being finally excreted; and this also seems to sIioav that part of it at least, is secreted for some other purpose than mere elimination from the Body. One use is, no doubt, to assist, by its alkalinity, in overcoming the acidity of the chyme, and so to allow the trypsin of the pancreatic secretion to act upon proteids. Constipation is, also, apt to occur in cases where the bile-duct is temporarily stopped, so that the bile probal^ly helps to excite the con- tractions of the muscular coats of the intestines; and it is said that under similar circumstances putrefactive decom- positions are extremely apt to occur in the intestinal con- tents. Apart from such secondary actions, however, the 344 THE nUMAN BODY. bile probably has some iiifliieiice in proiiioting the absorp- tion of fats. If one end of a capillary glass tube, moistened with water, be dipped in oil, the latter will not ascend in it, or but a short way; but if the tube be moistened with bile, instead of water, the oil will ascend higher in it. So, too, oil i)asses through a plug of porous clay kept moist with bile, under a much lower pressure than tlirough one wet with water. Hence bile, by soaking the epithelial cells lining the intestine, may facilitate the passage into the villi of oily sub- stances. At any rate, experiment shows that if the bile be prevented from entering the intestine of a dog, the animal eats an enormous amount of food compared with that amount which it needed previously; and that of this food a great proportion of the fatty parts passes out of the ali- mentary canal unabsorbed. There is no doubt, therefore, that the bile somehow aids in the absorption of fats, but exactly how is uncertain. Its possible action in exciting the muscles of the villi to contract will be referred to jires- ently. Bile precipitates from solution, not only pepsin, but any peptones contained in the chyme which enters the in- testine from the stomach. The Intestinal Secretions or Succus Entericus. This consists of the secretions of the glands of Brunner and the crypts of Lieberkiihn. It is difficult to obtain pure; in- deed the product of Brunner's glands has never been ob- tained unmixed. That of the crypts of Lieberkiihn is watery and alkaline, and poured out more abundantly dur- ing digestion than at other times. It has no special action on starches, most proteids, or on fats; but is said to dissolve blood fibrin and convert it into peptone, and to change cane into grape sugar, a transformation the object of which is not very clear, since cane sugar is itself readily soluble and diffusible. Intestinal Digestion. Having considered separately the actions of the secretions with which the food meets in the small intestine we may now consider their combined effect. The neutralization of the chyme, followed by its conver- sion into alkaline cliyle. will prevent any further action of the pepsin on pi'oteids, but v,i!l allow tlic pty;;lin of the IXTESTiyAL digestion: 345 saliva (the actiyity of which was stopped by the acidity of the gastric Juice) to recommence its action upon starch. Moreover, in the .stomach there is produced, alongside of the true pe^Jtones, a body called parapeptone, which agrees very closely with syntonin (p. 126) in its projierties, and this passes into the duodenum in the chyme. As soon as the bile meets the chyme it precipitates the parapeptone, and this carries down with it any peptones which, having es- caped absorj)tion in the stomach, may be present; it also precipitates the pepsin. In consequence, one commonly finds, during digestion, a sticky granular precipitate over the villi, and in the folds between the valvulse conniventes of the duodenum. This is soon redissolved by the pancre- atic secretion, which also changes into peptones the pro- teids (usually a considerable proportion of those eaten at a meal) which have passed through the stomach unchanged, or in the form of joaraiieptones. The conversion of starch into grape sugar will go on rapidly under the influence of the pancreatic secretion. Fats will be split up and saponi- fied to a certain extent, but a far larger proportion will be emulsified and give the chyle a whitish appearance. Cane sugar, which may have escaped absorjition in the stomach, will be converted into grape sugar and absorbed, along with any salines which may, also, have hitherto escaped. Elastic tissue from animal substances eaten, cellulose from plants, and mucin from the secretions of the alimentary tract, will all remain unchanged. Absorption from the Small Intestine. The chyme leav- ing the stomach is a semi-liquid mass which, being mixed in the duodenum with considerable quantities of pancreatic secretion and bile, is still further diluted. Thenceforth it gets the intestinal secretion added to it but, the absorjition more than counterbalancing the addition of liquid, the food- mass becomes more and more solid as it approaches the ileo-colic valve. At the same time it becomes poorer in nutritive constituents, these being gradually removed from it in its progress; most dialyze through the epithelium into the subjacent blood and lymphatic vessels, and are carried off. Those passing into the blood capillaries are taken by 346 THE HUMAN BODY. the portal vein to the liver; while tliose entering the hicteuls are carried into the left jugular vein by the thoracic duct. As to which foodstuffs go one road and which the other, there is still much doubt; sugars jirobably go by the portal system, while the fats, mainly, if not entirely, go through the lacteals. How the fats are absorbed is not clear, since oils will not dialyze through membranes, such as that lining the intestine, moistened with Avatery liquids. Most of them, however, certainly get into the lacteals as oils and not as solulilo soaps; for one finds these vessels, in a digest- ing animal, filled with a beautifully white milky chyle; Avhile at other jDeriods their contents are watery and colorless like the lymph elsewhere in the Body. The little fat-drops of the emulsion formed in the intestine, go through the epi- thelial cells and not between them, for during digestion one finds these cells loaded wnth oil-droplets. Now the free ends of these cells arc striated and probably devoid of any definite cell-wall, and it is jiossible that the intestinal movements squeeze oil-drops into them, which the cell then passes on to its deeper end and, thence, out into the sub- jacent lymph-spaces, which communicate with the central lacteal of a villus. Possibly, too, these cells are amoeboid and can thrust out processes from their free ends and actively pick up the oil-drops. In the villus there are all the anatomical arrangements for a mechanism which shall actively suck u]? substances into it. Each is more or less elastic, and, moreover, its capillary network when filled with blood wall distend it. If therefore the muscular stratum (p. 321) contracts and compresses it, emptying its lacteals into the vessels lying deeper in the intestinal wall, the villus will actively expand again so soon as its muscles relax. In so doing it could not fill its lacteals from the deeper ves- sels, on account of the valves in the latter, and, accordingly, would tend to draw into itself materials from the intestines; much like a sponge re-expanding in Avater, after having been squeezed dry. The liquid tlius sucked up may draw oil- drops with it, into the free ends of the cells and between them; and by repetitions of the process it is possible that considerable quantities of liquid, with suspended oil-drops. DI9ESTI02ir IN THE LARGE INTESTINE. 347 might be carried into the vilhis iudependently of any pro- cess of dialysis. Tlie bile moistening the surface of the Yillns may facilitate the passage of oil, as it does through a paper filter or a plate of plaster-of-Paris, and it is also said to stimulate the contractions of the villi; if so, its efficacy in promoting the absorption of fats will be explained, in si)ite of its chemical inertness with respect to those bodies. Digestion in the Large Intestine. The contractions of the small intestine drive on its continually diminishing contents, until they reach the ileo-colic valve, through which they are ultimately pressed. As a rule, when the mass enters the large intestine its nutritive portions have been almost entirely absorbed, and it consists merely of some water, with the indigestible portion of the food and of the secretions of the alimentary canal. It contains cellulose, elastic tissue, mucin, and somewhat altered bile pigments; commonly some fat if a large quantity has been eaten; and some starch, if raw vegetables have formed part of the diet. In its progress through the large intestine it loses more water, and the digestion of starch and the absorj)tion of fats is continued. Finally the residue, with some excretory matters added to it in the large intestine, collects in the sigmoid flexure of the colon and in the rectum, and is finally sent out of the Body from the latter. The Digestion of an Ordinary Meal. We may best sum up the facts stated in this chaj^ter by considering the diges- tion of a common meal; say a breakfast consisting of bread and butter, beefsteak, potatoes and milk. Many of these substances contain several alimentary principles, and, since these are digested in different ways and in different parts of the alimentary tract, the first thing to be done is to con- sider what are the proximate constituents of each. We then separate the materials of the breakfast as in the fol- lowing table — 348 THE HUMAN BODY. eg © d •(H O •iH o •iH u & g ■^ o ^■/. .2 1 tn f - o ® M it? !§ H w C^-r- H o ;>'. .2 .a c *» as =-a OS O fi'o « V. <1 o U o K o o 1i 3 V CD a ~ c s c .-S ■S ■« o — -g 3 £. e 2; 3 it -a P-( _IL cj =" a. 3 O o s •a ^ . Cob 1) da 1-11:5 fc 1 O : ii li oj a BJ 'J _o c 1" 3 tf y. •-,:d || 01 "i tr O c -< t^ b « -» -tJ M M a , tl aT ^ D « __ ^ _^ ^ •s ; ^C w a )ttle closed air-tight by a cork through which two tubes pass, one of which, b, leads into an elastic bag, d, and the other, c, provided with a stop- cock, opens freely below into the bottle. If the stop-cock, c, is open the air will enter the bottle and press there on the outside of the bag, as well as on its in- side through b. The bag will therefore 'ollapse, as the lungs do when the chest cavity is opened. But if some air be sucked out of c the pressure of that remain- ing in the bottle will diminish, while that inside the bag will be the same, and r.he hni will thus be blown up, because the atmospheric pressure on its interior will not be balanced by that on its exterioiv At last, when all the air is sucked out of the bottle and the ,^top-cock on c closed, the bag, if sufficiently distensible, will be expanded until it completely 61ls the bottle and presses against its inside, and the state Ftg. 107.— Diagram illustrating the pres- sure relationships of the lungs in the tho rax. 3^S THE HUMAN BODY. of things will then answer to that naturally found in the chest. If the bottle were now increased in size without letting air into it, the bag would exjiand still more, so as to fill it, and in so doing would receive air from outside through b\ and if the bottle then returned to its original size, its walls would press on the bag and cause it to shrink and expel some of its air through b. Exactly the same must of course happen, under similar circumstances, in the chest, the windpipe answering to the tube b through which air enters or leaves the elastic sac. The Respiratory Movements. The air taken into the lungs soon becomes laden in them with carbon dioxide, and at the same time loses much of its oxygen; these inter- changes taking place mainly in the deep recesses of the alveoli, far from the exterior, and only communicating with it through a long tract of narrow tubes. The alveolar air, thus become unfit to any longer convert venous blood into arterial, could only very slowly be renewed by gaseous dif- fusion with the outer air through the long air-passages — not nearly fast enough for the requirements of the Body, as any one readily experiences through the sensation of suffo- cation which follows holding the breath for a very short time. Consequently, added on to the breathing lungs is a respiratory mechanism, by which the air within them is periodically mixed with fresh air taken from the outside, and also the air in the alveoli is stirred up so as to bring fresh layers of it in contact with the Avails of the air-cells. This mixing is brought about by the breathing movements, consisting of regularly alternating inspiratiofis, during Avhich the chest cavity is enlarged and fresh air enters the lungs, and expirations, in which the cavity is diminished and air expelled from the lungs. When the chest is enlarged the air the lungs contain immediately distends them so as to fill the larger s})ace; in so doing it become rarefied and less dense than the external air; and since gases flow from points of greater to those of less pressure, some outside air at once flows in by the air-passages and enters the lungs. In expiration the reverse takes place. The chest cavity, liminishing, presses on the lungs and makes the air inside MOVEMENTS OF THE THORAX. 359 them denser than the external air, and so some passes out until an equilibrium of pressure is restored. The chest, in fact, acts very much like a bellows. When the bellows are opened air enters in con- sequence of the rarefaction of that in the interior, which is expanding to fill the larger space; and when the bellows are closed ... n 1 m Fig. 108.— Diagram to illustrate the en- again it is expelled. iO try of air to the lungs when the thoracic make the bellows quite ^^^^y ^^^^^s^^- like the lungs we must, however, as in Fig 108, have only one opening in them, that of the nozzle, for both the entry and exit of the air; and this opening should lead, not directly into the bellows cavity, but into an elastic bag lying in it, and tied to the inner end of the nozzle-pipe. This sac would represent the lungs and the space between its outside, and the inside of the bellows, the pleural cavi- ties. We have next to see how the expansion and contraction of the chest cavity are brought about. The Structure of the Thorax. The thoracic cavity has a conical form determined by tlie .shape of its skeleton (Fig. 109), its narrower end being turned upwards. Dorsally, ventrally, and on the sides, it is supported by the rigid framework afforded by the dorsal vertebrae, the breast-bone, and the ribs. Between and over these lie muscles, and the whole is covered in, air-tight, by the skin externally, and the parietal layers of the pie urge inside. Above, its aperture is closed by muscles and by various organs passing between the thorax and the neck; and below it is bounded by the diaiJtliragm, which forms a movable bottom to the, other- v/ise, tolerably rigid box. In inspiration this box is in- creased in all its diameters — dorso-ventrally, laterally, and from above down. The Vertical Enlargement of the Thorax. This is !>rought about by the contraction of the diaphragm which (Figs. 1 and 110) is a thin sheet-like muscle, with a fibrous membrane, serving as a tendon, in its centre. In rest, the 3G0 THE HUMAN BODY. diaphragm is dome-shaped, its concavity being turned towards the ahdoinen. From the tendon on the crown of the dome stri])ed muscuhir fibres radiate, downwards and outwards, to all sides; and are fixed by their inferior ends to the lower ribs, the breast-bone, and the vertebral column. In expiration the lower lateral portions of the diaphraj:;m lie close against the chest-walls, no lung inten'cning between them. In inspiration the muscular fibres, shortening, flat- FiG. 109.— The skeleton of the thorax, clavicle; d, third rib; i, glenoid fossa. a, g, vertebral coliunn; h, first rib; c. ten the dome and so enlarge the thoracic cavity at the ex- pense of the abdominal; and at the same time its lateral portions are pulled away from the chest-walls, leaving a space into which the lower ends of the lungs expand. The contraction of the diaphragm thus increases greatly the size of the thorax chamber by adding to its lowest and widest part. The Dorso-Ventral Enlargement of the Thorax. The ribs on the whole slope downwards (t, Fig. 25) from the vertebral column to the breast-bone, the slope being most 2HE EyLARGEMENT OF THE THORAX. 361 marked in the lower ones. During inspiration the breast- bone and the sternal ends of the ribs attached to it are ' imm\ ^, 4 Fig. 110.— The diaphragm seen from below. raised, and so the distance between the sternum and the vertebral column is increased. That this must be so will readily be seen by examining the diagram Fig. Ill, where ah represents the vertebral column, c and d two ribs, and st the ster- num. The continuous lines repre- sent the natural position of the ribs at rest in expiration, and the dotted lines the position in insi^iration. It is clear that when their lower ends are raised, so as to make the bars lie in a more horizontal plane, the ster- num is pushed away from the spine. and so the chest cavity is increased dorso- ventrally. The inspiratory elevation of the ribs is mainly due to the action of the scalene and exter- nal intercostal muscles. The scalene muscles, three on each side, arise from the cervical vertebra and are inserted into Fig. 111.— Diagram illus- trating the dorso-ventral in- crease in the diameter of the thorax when the ribs are raised. 3g;> TlIK II UMAX BODY. tlie nppcr ribs. Tlie oxternul iutercostuls (Fig. 11'3, A)h(i between the ribs and extend from the vertebral eolunm to the costal cartilages; their fibres slope downwards and forwards. Dui'ing an inspiration the scalenes contract and fix the u])per ribs firmly; then the external intercostals shorten and each raises the rib below it. The mnscle, in fact, tends to pull together the pair of ribs between which it lies, but as the upper one of these is held tight by the Fig. 112. — Portions of four ribs of a dog with the muscles between them, a, a. ventral ends of the ribs, joining at c the rib cartilages, b, which are fixed to cartilaginous portions, d. of the sternum. A, external intercostal muscle, ceas- ing between tlie rib cartilages, where the internal intercostal, B. is seen. Between the middle two ribs the external intercostal muscle has been dissected away, so as to displaj' the internal which was covered by it. scalenes and other muscles above, the result is that the lower rib is pulled up, and not the upper down. In this way the lower ribs are raised much more than the upjoer, for the whole external intercostal muscles on one side may be re- garded as one great muscle with many bellies, each belly separated from the next by a tendon, represented by the rib. When the whole muscular sheet is fixed above and MECHANISM OF EXPIRATION. 363 contracts, it is clear that its lower end will be raised more than any intermediate point, since there is a greater length of contractile tissue above it. The elevation of the ribs tends to diminish the vertical diameter of the chest; this is more than compensated for by the simultaneous descent of the diaphragm. The Lateral Enlargement of the Chest is mainly due to the diaphragm, which, when it contracts, adds to the lowest and widest part of the conical chest cavity. Some small widening is, however, brought about by a rotation of some of the middle ribs which, as they are raised, roll round a little at their vertebral a-rticulations and twist their car- tilages. Each rib is curved and, if the bones be examined in their natural position iu a skeleton, it will be seen that the most curved part lies below the level of a straight line drawn from the vertebral to the sternal attachment of the bone. By the rotation of the rib, during inspiration, this curved j^art is raised and turned out, and the chest widened. The mechanism can be understood by clasping the hands opposite the lower end of the sternum and a few inches in front of it, with the elbows bent and pointing downwards. Each arm will then answer, in an exaggerated way, to a curved rib, and the clasped hands to the breast-bone. If the hands be simply raised a few inches by movement at the shoulder-Joints only, they will be separated farther from the front of the Body, and rib elevation and the consequent dorso-ventral enlargement of the cavity surrounded will be represented. But if, simultaneously, the arms be rotated at the shoulder-joints so as to raise the elbows and turn them out a little, it will be seen that tlie space surrounded by the two arms is considerably increased from side to side, as the chest cavity is in inspiration by the similar elevation of the most curved part or " angle" of the middle ribs. Expiration. To produce an inspiration requires con- siderable muscular effort. The ribs and sternum have to be raised; the elastic rib cartilages bent and somewhat twisted; the abdominal viscera pushed down; and the ab- dominal wall pushed out to make room for them. In ex- piration, on the contrary, but little, if any, muscular effort 364 THE nUMAN BODY. is needed. As soon as the muscles which have raised the ribs and sternum rehix, these tend to return to their natural unconstrained position, and the rib cartilages, also, to un- twist themselves and bring the ribs back to their position of rest; the elastic abdominal wall presses the contained viscera against the under side of the diaphragm, and pushes that up again as soon as its muscular fibres cease contract- ing. By these means the chest cavity is restored to its original capacity and the air sent out of the lungs, rather by the elasticity of the parts which were stretched in inspir- ation, than by any special expiratory muscles. Forced Respiration. When a very deep breath is drawn or expelled, or when there is some impediment to the entry or exit of the air, a great many muscles take part in pro- ducing the respiratory movements, and exjiiration then be- comes, in part, an actively muscular act. The main expira- tory muscles are the internal intercostals which lie beneath the external between each pair of ribs (Fig. 112, B), and have an opposite direction, their fibres running upwards and forwards. In forced exijiration the lower ribs are fixed or pulled down by muscles running in the abdominal wall from the pelvis to them and to the breast-bone. The in- ternal intercostals then contracting, pull down the upper ribs and the sternum, and so diminish the thoracic cavity dorso-ventrally. At the same time, the contracted abdomi- nal muscles press the walls of that cavity against the viscera within it, and pushing these up forcibly against the dia- phragm make it very convex towards the chest, and so diminish the latter in its vertical diameter. In very violent expiration many other muscles may co-operate, tending to fix points on which those muscles which can directly dimin- ish the thoracic cavity, pull. In violent inspiration, also, many extra muscles are called into play. The neck is held rigid to give the scalenes a firm attachment; the shoulder- joint is held fixed and muscles going from it to the chest- wall, and commonly serving to move the arm, are then used to elevate the ribs; the head is held firm on the verte- bral column by the muscles going between the two, and then other muscles, which pass from the collar-bone and CAPACITY OF TEE LUNGS. 365 sternum to the skull, are used to pull up the former. The muscles which are thus called into play in labored but not in quiet breathing are called extraordinary muscles of res- piration. The Respiratory Sounds. The entry and exit of air are accompanied by resi)iratory sounds or innrmurs, which can be heard on a])plying the ear to the chest wall. The character of these sounds is different and characteristic over the trachea, the larger bronchial tubes, and portions of lung from which large bronchial tubes are absent. They are yariously modified in pulmonary affections and hence the value of auscultation of the lungs in assisting the phy- sician to form a diagnosis. The Capacity of the Lungs. Since the chest cavity never even approximately collaiDses, the lungs are never completely emptied of air: the space they have to occupy is larger in inspiration than during expiration but is always considerable, so that after a forced exjDiration they still con- tain a large amount of air which can only be expelled from them by opening the pleural cavities; then they entirely collapse, just as the bag in Fig. 107 would if the bottle in- closing it were broken. The capacity of the chest, and therefore of the lungs, varies much in different individuals, but in a man of medium height there remains in the lungs after the most violent possible expiration, about 16-iO cub. cent. (100 cub. inches) of air, called the residual air. After an ordinary expiration there will be in addition to this about as much more supplemental air; the residual and supplemental together forming the stationary air, which remains in the chest during quiet breathing. In an ordi- nary inspiration 500 cub. cent, (30 cub. inches) of tidal air are taken in, and about the same amount is expelled in nat- ural expiration. By a forced inspiration about 1600 cub. cent. (98 cub. inches) of complemental air can be added to the tidal air. After a forced inspiration therefore the chest will contain 1640 + 1640 + 500 + 1600 = 5380 cubic centi- meters (328 cubic inches) of air. The amount which can be taken in by the most violent possible inspiration after the strongest possible expiration, that is, the supplemental, 366 rilE HUMAN BODY. tidal and complemental air together, is known as the vital capacity. For a healthy man 1. 7 meters (5 feet 8 inches) high it IS about 3700 cub. cent. (225 cub. inches) and increases GO cub. cent, for each additional centime- ter of stature; or about 9 cubic inches for each inch of height. The Quantity of Air Breathed Daily. Knowing the quantity of air taken in at each breath and expelled again (after more or less thorough mixture with the stationary air) we have only to know, in addition, the rate at which the breathing movements occur, to be able to calculate how much air passes through the lungs in twenty-four hours. The average number of respirations in a minute is found by counting on persons sitting quietly, and not knowing that their breathing rate is under observation, to be fifteen in a minute. In each of these half a liter (30 cubic inches) of air is concerned; therefore 0.5 x 15 X 60 X 24 = 10,800 liters (374 cubic feet) is the quantity of air breathed under ordinary circumstances by each person in a day. Hygienic Remarks. Since the diaphragm when it con- tracts pu-slies down the abdominal viscera beneath it, these have to make room for themselves by pushing out the soft front of the abdomen which, accordingly, protrudes when the diaphragm descends. Hence breathing by the dia- phragm, being indicated on the exterior by movements of the abdomen, is often called "abdominal respiration," as distinguished from breathing by the ribs, called "costal'" or "chest breathing." In both sexes the diaphragmatic breathing is the most important, but, as a rule, men and children use the ribs less than adult women. Since both abdomen and chest alternately ex})and and contract in healthy breathing anything which impedes their free move- ment is to be avoided; and the tight lacing which used to be thought elegant a few years back, and is still indulged in by some who think a distorted form beautiful, seriously impedes one of the most important functions of the Body, leading, if nothing worse, to shortness of breath and an in- capacity for muscular exertion. In extreme cases of tight lacing some organs are often directly injured, weals of ASPIRATION OF THE THORAX. 3G7 fibrous tissue being, for exam^ile, uot uufretjueutl}' found developed on tlie liver, from the pressure of the lower ribs forced against it by a tight corset. The Aspiration of the Thorax, As already pointed out, in consccpience of the rigid framework which supjiorts its walls, the external air cannot jjress directly \x\)0\\ the con- tents of the thoracic cavity. It still, however, presses on them indirectly, through the lungs. Acting on the interior of these with a pressure ecpial to that exerted on the same area by a column of mercury 760 mm. (30 inches) high, it distends them, and pushes them against the inside of the chest-walls, the heart, the great thoracic blood-vessels, the thoracic duct, and the other contents of the cavity. The pressure thus exerted is not equal to that of the external air, since some of the total air-pressure on the inside of the lungs is used up in overcoming their elasticity, and it is only the residue which pushes them against the things outside them. In expiration this residue is about equal to that ex- erted by a column of mercury 754 mm. (| inch less than be- fore) high. On most parts of the Body the atmospheric pressure can, however, act with full force. Pressing on a limb it pushes the skin against the soft parts beneath, and these against the blood and lymph vessels among them; and the yielding abdominal walls do not, like the rigid thoracic walls, carry the atmospheric pressure themselves but transmit it to the contents of the cavity. It thus comes to pass that the blood and lymph in most parts of the Body are under a higher atmospheric pressure than in the chest, and consequently these liquids tend to flow into the thorax, until the extra distension of the vessels in which they there accumulate compensates for the less external pressure to which those vessels are exposed. An e(|uili- brium would thus very soon be brought about were it not for the respiratory movements, in consequence of which the mtra-thoracic pressure is alternately increased and dimin- ished, and the thorax comes to act as a sort of suction-pump on the contents of the vessels of the Body outside it; thus the respiratory movements come to influence the cir- culation of the blood and the flow of the lymph. 368 THE HUMAN BODY. Influence of the Respiratory Movements upon the Circulation. Suppose the chest in a condition oi' natural expiration and the external pressure on the blood in the blood-vessels within it, and in the heart, to have come, in the manner pointed out in the last paragi-aph, into equili- brium with the atmospheric pressure on the blood-vessels of the neck and abdomen. If an insjoiration now occurs, the chest cavity being enlarged the pressure on all of its con- tents will be diminished. In consequence, air enters the lungs from the windpipe, and blood enters the venae cavae and the right auricle of the heart. Not only the lungs, then, but the right side of the heart, and the intra-thoracic portions of the systemic veins leading to it, are expanded during an inspiration; but the lungs being much the most distensible take far the greatest part in filling up the in- creased space. The left side of the heart is not much in- fluenced as it is filled from the pulmonary veins; and the Avhole vessels of the lesser circulation lying within the chest, and being all affected in the same way at the same time, the blood-fiow in them is not influenced by the aspi- ration of the thorax. Distension of the lungs seems, how- ever, to diminish the capacity of their vessels, and so to a certain extent the flow is influenced; as the lungs expand blood is forced out of their vessels into the left auricle, and when they again contract their vessels fill up from the right ventricle. The pressure on the thoracic aorta being dimin- ished in inspiration, blood tends to flow back into it from the abdominal portion of the vessel, but cannot enter the heart on account of the semilunar valves; and the back-flow does not in any case equal the onflow due to the beat of the heart; so the what happens in the aorta is but a slight slowing of the current. The general result of all this is that the circulation is considerably assisted. When the next expiration occnrs, and the pressure in the thorax again rises, air and blood both tend to be expelled from the cavity. The aorta thus regains what it lost during insjiiration; the pressure on it is increased and it empties itself faster into its abdominal portion. The semilunar valves having pre- EFFECT OF RESPIRATION ON BLOOD-FLOW. 369 vented any regurgitation into the heart, there is neither gam noi loss so far as it is concerned. With the systemic intrathoracic veins, however, this is not the case; the extra blood entering them has already in great par*" gone on be- vond the tricuspid valve, and cannot flow back during ex- piration; and the pressure in the auricle being constantly kept low by its emptying into the ventricle, the increased pressure on the venee cava? tends as much to send the blood on into the heart, as back into the extra-thoracic veins. Moreover, whatever blood tends to take the latter course cannot do it effectually since, although the vense cava? themselves contain no valves, the more distant veins which open into them do. Consequently, whatever extra blood has, to use the common phrase, been ''sucked" into the intra-tlioracic veiife cav^e in inspiration and has not been sent already on into the right ventricle before expiration occurs, is, on account of the venous valves, imprisoned in the cavge under an increased pressure during expiration; and this tends to make it flow faster into the auricle during the diastole of the lattei". How much the alternating res^ piratory movements assist the venous flow is shown by the dilation of the veins of the head and neck which occur.s when a person is holding his breath; and the blackness for the face, from distension of the veins and stagnation of the capillary flow, which occurs during a prolonged fit of cough- ing, which is a series of expiratory efforts without any in- spirations. In still another way the aspiration of the thorax assists the heart. The heart and lungs are both extensible, though in different degrees, and each is stretched in the chest somewhat beyond its natural size; the one by the atmos- pheric pressure directly, the other by that pressure in- directly, exerted through the blood exposed to it in the extra-thoracic veins. Supposing, therefore, the heart sud- denly to shrink it would leave more space in the chest to be filled by the lungs, and these, accordingly, at each cardiac systole expand a little to fill the extra room, just as thevdo when the space around them is otherwise enlarged during 370 THE HUMAN BODY, an inspiration. Tlie elasticity of the lungs, however, causes them to resist this distension and oppose the cardiac systole. The matter may be made clear by an arrangement like that in Fig. 113. A is an air-tight vessel with a tube, e, provided with a stop-cock, leading from it; J is a highly distensible elastic bag in free communication through d with the exterior; and c, representing the heart, is a less extensible sac, from which a tube leads and dips under water in the vessel B. If air be pumped out through e both bags will dilate, h filling with air, and c with water driven up by atmospheric pres- sure. Ultimately, if sufficiently ex- tensible, they would fill the whole space, the thinner walled, h, occupy- ing most of it. If then the stop-cock be closed, things will remain in equi- librium, each bag striving to collapse and so exerting a pull on the other, for if h shrinks c must expand and vice versa. If c suddenly shrink, as the heart does in its systole, b will dilate; but as soon as the systole of c ceases, h will shrink again and pull c out to its previous size. In the same way, after the cardiac sys- tole, when the heart-walls relax, the lungs pull them out again and dilate the organ. The contracting heart thus expends some of its work in overcoming the elasticity of the lungs, which ppposes their expansion to fill the space left by the smaller heart; but during the diastole of the heart this work is utilized to pull out its walls again, and draw blood into it. Since the normal heart has muscular power, and to spare, for its systole, this arrangement, by which some of the work then spent is stored away to assist the diastole, which cannot be directly performed by cardiac muscles, is of service to it on the whole. It is a physio- logical though not a mechanical advantage; no work power is gained, but what there is, is better distributed. Fig. 113.— Diagram illus- trating the influence of as- piration of the thorax on the circulation of the blood. INFLUENCE OF RESPIBATION OX I.T^IPHFLO^V. 371 Influence of the Respiration on the Lymph-Flow. During inspiration, when intra-thoracic pressure is lowered, lympli is pressed mto the thoracic duct from the abdomi- nal lymphatics. In expiration, when thoracic pressure rises again, the extra IjTuph cannot flow back on account of the valves in the lymphatic vessels, and \t ^s consequently driven on to ths cervical ending of the thoracic duct. The breathing movements thus pump the lymph on. CHAPTER XXV. THE CHEMISTllY OF EESPIEATION. Nature of the Problems. Tlie study of the respira- tory process from a chemical standpoint has for its object to discover, first, wliat are, in kind and extent, tlie inter- changes between the air in the kings and the blood in pul- monary cajDillaries; and, in the second place, the nature and amount of the corresponding gaseous changes between the various living tissues and the blood in the systemic capillaries. These processes are the reverse of one another and in the long run balance, the blood losing as much car- bon dioxide gas in the pulmonary circulation as it gains in the systemic, and gaining as much oxygen in the former as it loses in the latter. To thoroughly comprehend the matter it is, moreover, necessary to know the jjliysical and chemical conditions of these gases in the lungs, in the blood, and in the tissues genei'ally; for only so can we understand how it is that in different localities of the Body such exactly contrary processes occur. So far as the prob- lems connected with the external respiration are concerned our knowledge is tolerably complete; but as regards the internal respiration, taking place all through tlie Body, much has yet to be learnt; for example, we know that a muscle at work gives more carbon dioxide to the blood than one at rest and takes more oxygen from it, but exactly how much of the one it gives and of the other it takes is only known approximately; as are also the conditions under which this greater interchange during the activity of the muscular tissue is effected; and concerning nearly all the other tissues we know even less than about muscle. In fact, as regards the Body as a whole, it is comparatively easy to CHANGES PB on UCED IM AIB ONCE BREATHED. 373 find how great its gaseous interchauges with, the air are during work and rest, waking and sleeping, while fasting or digesting, and so on; but when it comes to be decided what organs are concerned in each case in producing the greater or less exchange, and how much of the whole is due to each of them, the question is one far more difficult to settle and still very far from comj)letely answered. The Changes produced in Air by being once Breathed. These are fourfold — changes in its temperature, in its moisture, in its chemical composition, and its volume. The air taken into the lungs is nearly always cooler than that expired, which has a temperature of about 36° C. (97° F. ). The temperature of a room is usuially about 21° C. (70° F. ). The warmer the inspired air the less, of course, the heat which is lost to the Body in the breathing process; its average amount is calculated as about equal to*3.5 calories in twenty-four hours; a calorie (see Physics) being as much heat as will raise the temperature of one kilogram (2.2 lbs) of water one degree centigrade (1.8° F.). The inspired air always contains more or less w^ater vapor, but is rarely saturated; that is, rarely contains so much but it can take up more without showing it as mist; the warmer air is, the more water vapor it requires to saturate it. The expired air is nearly saturated for the temperature at which it leaves the Body, as is readily shoAvn by the water deposited when it is slightly cooled, as when a mirror is breathed upon; or by the clouds seen issuing from the nostrils on a frosty day, these being due to the fact that the air, as soon as it is cooled, cannot hold all the water vapor which it took up when warmed in the Body. Air, therefore, when breathed once, gains water vapor and carries it off from the .lungs; the actual amount being subject to variation witli the tem- perature and saturation of the inspired air: the cooler and drier this is, the more water will it gain when breathed. On an average the amount thus carried off in twenty-four hours is about 255 grams (9 ounces). To evaporate this water in the lungs an amount of heat is required, which disappears for this purpose in the Body, to apj^ear again outside it when the water vapor condenses (see Physics). 374 THE HUMAN BODY. The amount of heat taken oil in ihis way during the day is about 7.2 calories. The total daily loss of heat from the Body througli tlie lungs is therefore 10.7 calories, 3.5 in warming the inspired air and 7.2 in evaporating water. The most important changes brought about in the breathed air are those in its chemical composition. Pure air when completely dried consist in 100 parts of — By Volume. By Weight. Oxygen 20.8 23 Nitrogen 79.2 77 Ordinary atmospheric air contains in addition 4 volumes of carbon dioxide in 10,000, or 0,04 in 100, a quantity which, for practical purposes, may be neglected. When breathed once, such air gains rather more than 4 volumes in 100 of carbon dioxide, and loses rather more than 5 of oxygen. More accurately, 100 volumes of expired air when dried con- sist of — Oxygen 15.4 Nitrogen 79.2 Carbon dioxide 4.3 The expired air also contains volatile organic substances in quantities too minute for chemical analysi-s, but readily detected by the nose upon coming into a close room in which a number of persons have been collected. Since 10,800 liters (346 cubic feet) of air are breathed in twenty-four hours and lose 5.4 per cent of oxygen, the total quantity of this gas taken up in the lungs daily is 10,800 X 5.4-f- 100 =583.2 liters (20.4 cubic feet). One liter of oxvgen measured at 0°C (32° F.) and under a pressure equal to one atmosphere, weighs 1.43 grams (see Chemistry), so the total weight of oxygen taken up l)y the lungs daily is 583.2 X 1.43 = 833.9 grams. Or, using inches and grains as standards, 44.5 cubic inches of oxygen at the above tem- perature and pressure weigh almost exactly 16 grains, so the 20,4 culjic feet absorbed in the lungs daily weigh 20,4 X 1728 --44,5X16 = 12,818 grains. The amount of carbon dioxide excreted from the luno^s VEJS^TILA riON. 3 To being 4.3 ^jer cent of the volume of the air breathed daily, is— 10,800 X 4.3 -j- 100 = 4G4.4 liters (1G.,:25 cubic feet) measured at the normal temperature and pressure. This volume -weighs 910 grams, or 14,li35 grains. If the expired air be measured as it leaves the Body its bulk will be found greater than that of the inspired air, since it not only has water vapor added to it, but is expanded in consequence of its higher temperature. If, however, it be dried and reduced to the same temperature as the inspired air its volume will be found diminished, since it has lost 5.4 volumes per cent of oxygen and gained only 4.3 of carbon dioxide. In round numbers, 100 volumes of dry inspired air at zero, give 99 volumes of dry expired air measured at the same temperature and pres- sure. Ventilation. Since at every breath some oxygen is taken from the an" and some carbon dioxide given to it, were the atmosphere around a living man not renewed he would, at last, be una'ole to get from the air the oxygen he required; he would die of oxygen starvation or be suffocated, as such a mode of death is called, as surely, though not quite so fast, as if he were put under the receiver of an air-pump and all the air around him removed. Hence the necessity of ven- tilation to supply fresh air in place of that breathed, and clearly the amount of fresh air requisite must be deter- mined by the number of persons collected in a room; the supply which would be ample for one person would be in- sufficient for two. Moreover fires, gas, and lamps, all use up the oxygen of the air and give carbon dioxide to it, and hence calculation must be made for them in arranging for the ventilation of a building in which they are to be em- ployed. In order that air be unwholesome to breathe, it is by no means necessary that it have lost so much of its oxygen as to make it difficult for the Body to get what it wants of that gas. The evil results of insufficient air-supply are rarely, if ever, due to that cause even in the worst ventilated rooms for, as we shall see in the next paragraph, the blood can take what oxygen it wants from air containing compara- 376 THE limfAX BODY. lively little of that gas. The lieadache and drowsiness wliich come on from silling in a badly venlihitcd room, and the want of energy and general ill-health wliich result from permanently living in such, are dependent on a slow ijoison^ ing of the Body by the reabsorption of the things elimi- nated from the lungs in previous respirations. What these are is not accurately known; they doubtless belong to those volatile bodies mentioned above, as carried off in minute quantities in each breath; since observation shows that the air becomes injurious long before the amount of carbon dioxide in it is sufficient to do any harm. Breathing air containing one or two per cent of that gas produced by ordinary chemical methods does no particular injury, but breathing air containing one per cent of it produced by respiration is decidedly injurious, because of the other things sent out of the lungs at the same time. Carbon dioxide itself, at least in any such percentage as is com- monly found in a room, is not poisonous, as used to be believed, but, since it is tolerably easily estimated in air, while the actually injurious substances evolved in breath- ing are not, the purity or foulness of the air in a room is usually determined by finding the percentage of carbon dioxide in it; but it must be borne in mind that to mean much this must have been produced by breathing; other- wise the amount of it present is no guide to the cpumtity of really important injurious substances jjresent. Of course when a great deal of carbon dioxide is present the air is irrespirable: as for examj)le sometimes at the bottom of wells or brewing-vats. In one minute as we have seen (p. 3GG) .5 X 15 = 7.5 liters (0.25-4 cubic feet) of air are breathed and vitiated with carbon dioxide to the extent of rather more than four per cent; this, mixed with three times its volume of extermxl air, would give thirty liters (a little over one cubic foot) vitiated to the extent of one per cent, and such air is no longer respirable for any length of time with safety. The result of breathing it for an evening is headache and gen- eral malaise; of breathing it for weeks or months a lowered tone of the whole Body — less power of work, physical or VENTILA nON. 377 mental, and less power of resisting disease; the ill effects may not show themselves at once, and may accordinciy be overlooked, or considered scientific fancies, by the careless; but they are there ready to manifest themselves neverthe- less. In order to have air to breathe in a fairly pure state every man should have for his own allowance at least •^3,000 liters of space to begin with (about 800 cubic feet) and the arrangements for ventilation should, at the very least, renew this at the rate of 30 liters (one cubic foot) per minute. The nose is, however, the best guide, and it is found that at least live times this supply of fresh air is necessary to keep free from any odor the room inhabited bv one adult. In the more recently constructed hospitals, as a result of experience, twice the above minimum cubic space IS allowed for each bed in a ward, and the replacement of the old air at a far more rapid rate, is also provided for. Ventilation does not necessarily mean draughts of cold air, as is too often supposed. In warming by indirect radia- tion it may readily be secured by fixing, in addition to the registers from which the new warmed air reaches the room, corresponding openings at the opposite side, by which the old air may pass off to make room for the fresh. An open fire in a room will always keep up a current of air through it, and is one of the healthiest, though not the most economi- cal, methods of warming an apartment. Stoves in a room, unless constantly supplied with fresh air from without, dry its air to an unwholesome extent. If no a^jpliance for providing this supply exists in a room, it can usually be got, without a draught, by fixing a board about four inches wide under the lower sash and shutting the window down on it. Fresh air then comes in by the open- ing between the two sashes and in a current directed upwards, which gradually diffuses itself over the room with- out being felt as a draught at any one point. In the method of heating by direct radiation, the apparatus em- ployed provides of itself no means of drawing fresh air into a room, as the draught up the chimney of an open fireplace or of a stove does; and therefore special inlet and outlet openings are very necessary. Since few doors and windows, 378 THE lir}rA^ BODY. rurtunaicly, fit quite tight, fresh air gets even into closed rooms, in tolerable abundance for one or two inhabitants, if there be outlets for the air already in them. Changes undergone by the Blood in the Lungs. These are the exact reverse of tliose exliiljited by the breathed air — what the air gains the blood Uiscs, and vice versa. Con- sequently, the blood loses heat, and water, nnd carbon dioxide in the pulmonary capillaries; and gains oxygen. These gains and losses are accompanied by a change of color from the dark purple wliich the blood exhibits in the pul- monary artery, to the bright scarlet it jDOSsesses in the pul- monary veins. The dependence of this color change upon the access of fresh air to the lungs while the blood is flowing througli them, can be readily demonstrated. If a rabbit be rendered unconscious by chloroform, and its chest be opened, after a pair of bellows has been connected with its windpipe, it is seen that, so long as the bellows are worked to keep up artificial respiration, the blood in the right side of the heart (as seen through the thin auricle) and that in the pulmo- nary artery, is dark colored, while that in the pulmonary veins and the left auricle is bright red. Let, however, the artificial respiration be stopped for a few seconds and, consequently, the renewal of the air in the lungs (since an animal cannot breathe for itself when its chest is opened), and very soon the blood returns to the left auricle as dark as it left the right. In a very short time symptoms of suffocation show themselves and the animal dies, unless the bellows be again set at work. The Blood Gases. If fresh blood be rapidly exposed to as complete a vacuum as can be obtained it gives off certain gases, known as the gases of the blood. These are the same in kind, but differ in proportion, in venous and arteriiil blood; there being more carbon dioxide and less oxygen obtainable from the venous blood going to the lungs by the pulmonary artery, than from the arterial blood coming back to the heart by the pulmonary veins. The gases given off by venous and arterial blood, measured under the normal pressure and at the normal temperature (see Physics), THE BLOOD GASES. 379 amouTit to about 72 Yolumes for every 100 volumes of bloody and iu the two cases are as follows — Venous Blood. Arterial Blood. Oxygen 10 20 Carbon dioxide 60 50 Nitrogen 2 2 It is important to bear in mind that Avhilo arterial blood contains some carbon dioxide that can be removed by the air-pumj), venous blood also contains some ox3'gen, remova- ble in the same way; so that the difference between the two is only one of degree. When an animal is killed by f'lffocation, however, the last trace of oxygen which can be yielded up in a vacuum, disappears from the blood before the heart ceases to beat. All the blood of such an animal is what might be called suffocation blood; and has a far darker color than ordinary venous blood. The Cause of the Bright Color of Arterial Blood. The color of the blood depends on its red corpuscles, since pure blood plasma or blood serum is colorless, or at most a very faint straw yellow. Hence the color change which the blood experiences in circulating through the lungs must be due to some chanige in its red corpuscles. Now, minute solid bodies suspended in a liquid reflect more light when they are more dense, other things being equal; and the first thing that suggests itself as the cause of the change in color of the blood is that, its red corpuscles have shrunk in the pulmonary circulation, and so reflect more light and give the blood a brighter look. This idea gains some support from the fact that, as seen under the microscope, the red blood corpuscles of some animals, as the frog, do expand somewhat when exposed to carbon dioxide gas and shrink up a little in oxygen. But that this is not the chief cause of the color change is readily proved. By diluting blood with water the coloring matter of the red corpuscles can be made to pass out of them and go into solution in the plasma (p. 46) and it is found that such a solution, in which there can be no question as to the reflecting powers of colored solid bodies suspended in it, is brighter red when supplied with oxygen than when deprived of that gas. 380 THE HUMAN BODY. This suggests that tlio c )loring iiiiittcr or hmmoglohi n of the red corpuscles combines Avith oxygen to form a scarlet compound, and when deprived of that gas has a darker and more purple color; and further experiments confirm this. Hannoglohin combined with oxygen is known as oxylimmo- (jjiihin and it is on its predominance that the color of arterial blood dei)ends. H;emoglol)in uncombined with oxy- gen is reduced hoenioglobin; it predominates in venous blood, and is alone found in the blood of a suffocated mammal. The Laws Governing the Absorption of Gases by a Liquid. In order to understand the condition of the gases in the blood liquid it is necessary to recall the general laws in accordance with which liquids absorb gases. They are as follows: — 1. A given volume of a liquid at a definite temperatnTe if it absorbs any of a gas to which it is ex2:)0sed, and yet does not combine chemically with it, takes up a definite volume of the gas. If the gas be compressed the liquid will still, at the same temperature, take ujt the same volume asljefore, but now it takes up a greater weight; and a weight exactly as much greater as the pressure is greater, since one volume of a gas under any pressure contains exactly twice as much of the gas by weight as the same volume under half the pressure, and so on. A liter or a quart of water, for example, exposed to the air will dissolve a certain amount of oxygen. If the air (and therefore the oxygen in it) he compressed to one fourth its bulk tlien the water will dissolve exactly the same volume of oxygen as before, but this volume of the compressed gas will contain exactly four times as much oxygen as did the same volume of the gas under the origi- nal pressure; and if now the pressure be again diminished the oxygen Avill be given off exactly in proportion as its pressure on the surface of the water decreases. Finally, when a complete vacuum is formed above the surface of the water it will be found that the latter has given off all its dissolved oxygen. This law, that the quantity of a gas dis- solved by a lifjuid varies directly as the pressure of that gas on the surface of the liquid is known as Dalton's law (see Physics). THE ABSORPTION OF GASES BY LIQUIDS. "-SI , 2. The amount of a gas dissolyed by a liquid depends, not on the total pressure exerted by all the gases pressing on its surface, but on the fraction of the total pressure which IS exerted by the particular gas in question. For example, the total atmospheric pressure is equal to that of a column of mercury 760 mm. (30 inches) high. But 100> volumes of air contain approximately 80 volumes of nitrogen and 20 of oxygen: therefore \ of the total pressure is due to oxygen and |- to nitrogen: and the amount of oxygen absorbed by water is Just the same as if all the nitrogen were removed from the air and its total pressure there- fore reduced to -t of 760 mm. (30 inches) of mercury; that is to 152 mm. (6 inches) of mercury pressure. It is only the fraction of the total pressure exerted by the oxygen itself which affects the quantity absorbed by water at any given temperature. So, too, of all the atmospheric pressure 4 is due to nitrogen, and all the oxj'gen might be removed from the air without affecting the quantity of nitrogen which would be absorbed from it by a given volume of water. The atmospheric pressure would then be f of 760 mm. of mercury, or 608 mm, (24 inches), but it would all be due to nitrogen ga? — and be exactly equal to the fraction of the total pressure due to that gas before the oxygen was removed from the air. When several gases are mixed to- gether the fpaction -of the total pressure exerted by each one is known as the partial pressure of that gas; and it is this partial pressure which determines the amount of each individual gas dissolved by a liquid. If a liquid exposed to the air for some time had taken up all the oxygen and nitrogen it could at the partial pressures of those gases in the air, and were then put in an atmosphere in which the oxygen had all been replaced by nitrogen, it would now give off all its oxygen since, although the total gaseous pressure on it was the same, no part of it was any longer due to oxygen; and at the same time it would take up \ more nitrogen, since the whole gaseous pressure on its sur- face was now due to that gas while before only \ of the total was exerted by it. If, on the contrary, the liquid were exposed to pure hydrogen under a pressure of one atmos- 382 THE HUMAN BODY. pliere it would give ofE nil its previously dissolved oxygen and nitrogen, since none of ihj pressure on its surface woulcT now bo due to tlioso gases; and would take up as much liydrogen as corresponded to a pressure of that gas equal to 7G0 mm. of mercury (30 inches). 3. A liquid may be such as to combine chemically with a gas. Tlien the amount of the gas absorbed is indepen- dent of the partial jjressure of the gas on the surface of tlie liquid. The quantity absorbed will depend upon how much the liquid can combine with. Or, a liquid may partly be composed of things which simply dissolve a gas and partly of things which chemically combine with it. Then the amount of the gas taken uj? under a given partial pressure will depend on two things; a certain portion, that merely dissolved, will vary with the pressure of the gas in question; but another portion, that chemically combined, will remain the same under difEerent pressures. The amount of this second portion depends only on the amount of the sub- stance in the liquid which can chemically combine with it, and is totally independent of the partial pressure of the gas. 4. Bodies are known which chemically combine with certain gases when the partial pressure of these is consider- able; but the compounds thus formed are broken up, and the gas liberated, when its partial pressure on the surface of the liquid falls below a certain limit. 5. A membrane, moistened by a liquid in which a gas is soluble, does not essentially alter the laws of absorption, by a liquid on one side of it, of a gas present on its other side, * whether the absorption be due to mere solution or to chemical combinations or to both. The Absorption of Oxygen by the Blood. Applying the physical and chemical facts stated in the joreceding paragraph to the blood, we find that the blood contains (1 ) plasma, which simply dissolves oxygen, and (2) licemoglohin. which combines with it under some partial pressures of that gas, but gives it up under lower. Blood plasma or, what comes to the same thing, fresh serum, exposed to the air, takes up no more oxygen than so ABSOBPTIOY OF OXYGEy BY THE BLOOD. 383 much water, that is about 3 volumes of the gas for every 100 of the liquid. This quantity obeys Dalton's law. If instead of blood serum fresh whipped blood be em- ployed, the quantity of oxygen taken up is much greater; this extra quantity must therefore be taken up by the red corpuscles (in possessing which whipped blood alone differs from blood serum) and it does not obey Dalton's law. If the partial pressure of oxygen on the surface of the whipped blood be doubled, only as much more oxygen will be taken up as corresponds to that dissolved in the plasma; and if the partial pressure of oxygen on its surface be re- duced to one half only a very small amount of oxygen (^ of that dissolved by the serum) will be given off. All the much larger quantity taken up by the red corpuscles will be unaffected and must therefore be chemically combined with something in them. Since 90 per cent of their dry weight is made up by h^Bmoglobin, and this body when pre- pared pure is found capable of combining with oxygen, there is no doubt that it is the haemoglobin in the circulating blood which carries around nearly all the oxygen found in it. The red corpuscles are so many little packages in which oxygen is stowed away. The compound formed between oxygen and hsemoglobin is, however, a very feeble one; the two easily separate, and always do so when the oxygen pressure in the liquid or gas to which the oxyhoemoglobin is exposed falls below 25 mil- limeters of mercury. Hence, in an air-pump, the blood only gives off some of its small portion of merely dissolved oxy- gen, until the pressure falls to about ^ of an atmosphere, that is to -^Q- = 125 mm. (5 inches) of mercury, of which total pressure one fifth (25 millimeters or 1 inch) is due to the oxygen present. As soon as this limit is reached the haemoglobin gives up its oxygen. Consequences of the Peculiar "Way in which the Oxygen of the Blood is Held. The first, and most im- portant, is that the blood can take up far more oxygen in the lungs than would otherwise be possible. Since blood serum exposed to pure oxygen takes up only 3 volumes for 100, blood exposed to the air would take up \ only of that 384 THE HUMAN BODY. amount at ordinary temperatures, and still less at the tcm- l)erature of the Body, were it not for its haemoglobin. In the lungs even less would be taken up, since tlie air in the air-cells of those organs is poorer in oxygen than the external air; and consequently the partial pressure of that gas in it is lower. The tidal air taken in at each breath serves merely to renew directly the air in the big bronchi; the deeper one examined the pulmonary air the less oxygen and more carbon dioxide would be found, till, in the layers farthest from the exterior and only re- newed by difEusion with the air of the large bronchi, it is estimated that the oxygen only exists in such quantity that its partial pressure is equal to 130 millimeters of mercury, instead of 153 as in ordinary air. In the second place, on account of the way in which haemoglobin combines with oxygen, the quantity of that gas taken up by the blood is independent of such variations of its partial pressure in the atmosphere as we are subjected to in daily life. At the top of a high mountain, for example, the atmospheric pressure is gi'eatly diminished, but still we can breathe freely and get all the oxygen we want. So long as the partial pressure of tliat gas remains above 25 millimeters (1 inch) of mercury, the amount of it taken up by the blood will depend on how much haemoglobin there is in that liquid and not on how much oxygen there is in the air. So, too, breathing pure oxygen under a pressure of one atmosphere, or air com- l^ressed to ^ or \ its bulk, does not increase the quantity of that gas taken up by the blood, apart from the very small extra quantity which would be dissolved by the i")lasma. All the widespread statements found as to the exhilaration and excitement caused by breathing pure oxygen are, as a matter of fact, erroneous, being founded, on early experi- ments made with impure gas, and corrected by many com- petent observers since. The General Oxygen Interchanges in the Blood. We may now try to depict what hap})ens to the blood oxygen in a complete circulation. Sup}iose we have a quantity of arterial blood in the aorta. This, fresh from the lungs, will have its haemoglobin almost fully combined with oxygen THE BLOOD GASES. 385 and in lii-c state of oxylisaiogiobiu. lu the blood jjlusma some more oxygen will be dissolved and so much as answers to a pressure of that gas ec^ual to 130 mm. {o.'Z inches) of mercury, which is the partial pressure of oxygen in the pulmonary air-cells. This teyisiou of the gas in the jjlasma will be more than sufficient to keep the hgemogiobin from giving off its oxygen. Suppose the blood now enters the capillaries of a muscle. In the liquid moistening this organ the oxygen tension is almost nil, since the tissue elements are steadily taking the gas up from the lymph around them. Consequently, through the capillary walls, the plasma will give oft' oxygen until the tension of tliat gas m it falls below 35 millimeters of mercury. Immediately some of the oxy- liJBmoglobin is decomposed, and the oxygen liberated is dis- solved in the plasma, and from there again passed on to the lymph outside; and so the tension in the plasma is once more lowered and more oxyhasmoglobin decomposed. This goes on so long as the blood is in the capillaries of tlie muscle, or at any rate so long as the muscular fibres kee}) on taking oxygen from the lymph bathing them; if they cease to do so of course the tension of that gas in the lymph will soon come to equal that in the plasma: the latter will therefore cease to yield oxygen to the former; and so main- tain its tension (by the oxygen received from the last de- composed oxyhemoglobin) at a point Avhich will preven' the liberation of anymore oxygen from such red cori^uscles as have not yet given all theirs up. The blood will now go on as ordinary venous blood into the veins of the muscle and so back to the lungs. It will consist of (1) plasma with oxygen dissolved in it at a tension of about 25 milli- meters (1 inch) of mercury. (2) A number of red cor- puscles containing reduced hajmoglobin. (3) A number of rod corpuscles containing oxyha^mogiobin. Or perhaps all of the red corpuscles will contain some reduced and some oxidized haemoglobin. The relative proportion of reduced aud unreduced haemoglobin will depend on how active the muscle was; if it worked while the blood flowed through it it Avill have used up more oxygen, and the blood leaving it will consequently be more venous, than if it rested. This 386 THE HUMAN BODY. venous blood, returning to the heart, is sent on to the pul- monary capillaries. Here, Lhe partial pressure of oxygen in the air-cells being loU mm. (5.;i inches) and that in the blood plasma much less, oxygen will be taken \\\) by the latter, and the tension of that gas in the jjlasma tend to be raised above tiie limit at which haemoglobin combines with it. Hence, as fast as the plasma gets oxygen those red cor- puscles which contain any reduced haemoglobin rob it, and so its oxygen tension is kept down below that in the air- cells until all the hasmogiobin is satisfied. Then the oxygen tension of the plasma rises to that of the gas in the air-cells; no more oxygen is absorbed, and the blood returns to the left auricle of the heart in the same condition, so far as oxygen is concerned, as when we commenced to fol- low it. The Carbon Dioxide of the Blood. The same general laws ajjply to this as to the blood oxygen. The gas is partly merely dissolved and partly in a loose chemical com- bination much like that of oxygen with haemoglobin, but the body Avith which it combines in this way exists in the plasma and not in the red corpuscles; what it may be is not certainly known. Besides this, some more carbon dioxide is stably combined and is only given off on the addition of a stronger acid. The partial pressure of carbon dioxide in the pulmonary air-cells is about 40mm. (1.6 inches) of mercury. Therefore the tension of that gas in the pulmonary capillaries must be more than this. On the other hand its tension in arterial blood must be less than that in the lymph around the tissues; otherwise it could not enter the blood in the systemic circulation, which it does, as proved by the fact that 100 vols, of venous blood give off 60 of this gas, and 100 vols, of arterial only 50. The nitrogen contained in the blood is, so far as we know, quite unimportant. Internal Respiration. As to the amount of oxygen used by each tissue and the quantity of carbon dioxide pro- duced by it we know but little; the following points seem, however, tolerably certain: — 1. The amount of carbon dioxide produced in an organ mTERNAL RESPIRATION. 387 in a given time bears no constant ratio to the amount of oxygen taken np by it simultaneously. This is certainly true of muscle, for experiment shows that muscular work, while it continues, leads to an elimination of carbon dioxide containing more oxygen than the total oxygen taken up from the lungs in the same time. The balance is of course made up in subsequent periods of rest, when more free oxygen is taken up than is eliminated in combination during the same time. Moreover, a frog's muscle excised from the body and put in an atmosphere containing no oxygen and made there to contract, will evolve with each contraction considerable quantities of carbon dioxide — although from the conditions of the experiment it can receive from outside no uncombined oxygen, and other experiments show that it contains none. Hence the living muscular fibre must contain a substance which is decom- posed during activity and yields carbon dioxide as one pro- duct of decomposition; and this quite independent of any simultaneous direct oxidation. 2. What is true of muscle is probably true of most of the tissues. During rest they take up oxygen and fix it in the form of complex compounds, bodies which, like gun- powder, are readily decomposed into simpler, and in such decompositions liberate energy which is used by the work- ing tissue. One product of the decomposition is the highly oxidized carbon dioxide, c^nd this is eliminated; other products are less oxidized, and possibly are not elimi- nated but built up again, with fresh oxygen taken from the blood and fresh carbon from the food, into the decomposa- ble substance. 3. During the day a man gives off from his lungs more oxygen in carbon dioxide, tlian he takes up by the same organs from the air. During the night the reverse is the case. This, however, has nothing to do with the alternating jjeriods of light and darkness, as it has in the case of a green plant, which in the light evolves more oxygen than it consumes and in the dark the contrary. It depends, rather, on the fact that during the day more muscular effort is exerted than at night, and the meals are then taken 388 TITM HUMAN BODY. and digested. The activity of tlic muscles and the digestive glands is dependent on processes which give rise to a large production of carbon dioxide and, during the night, when botli are at rest, more oxygen is taken up than is contained in the carbon dioxide eliminated. If a man works and takes his meals at night, and slecj^s in the d:iy, the usual ratios of his gaseous exchanges with the exterior are entirely reversed. 4. The amount of work that a man's organs do, is not dependent on the amount of oxygen supplied to them, but the amount of oxygen used by him depends on how much he uses his organs. The quantity of oxygen supj)lied must of course always be, at least, that required to prevent suffoca- tion; but an excess above this limit will not make the tissues work. Just as a man must have a certain amount of food to keej^ him alive, so he must have a certain amount of oxygen; but as extra food will not make his tissues or Mm (who is physiologically the sum of all his tissues) work, apart from some stimulus to exertion, so it is with oxygen. Highly arterialized blood, or an abnormal amount of blood, flowing through an organ will not arouse it to activity; the working organ, muscle (p. 257) or gland (p. 269), for ex- ample, usually gets more blood to supply its extra needs — just as a healthy man who works will have a better appe- tite than an idle one; but as taking more food by an idle man will not of itself make him more energetic, so neither will sending more arterial blood through an organ excite it to activity. 5. The preceding statement is confirmed by experiments which show that an aninud uses no more oxygen in an hour when made to breathe that gas in a pure state, than when allowed to breathe ordinary air. In other words, the amount of oxygen an animal uses (provided it gets the minimum necessary for health) is dependent only on how much it uses its tissues. These (the rest in most cases sul>- Ject to a certain amount of control from the nervous) de- termine their own activity, and this, in turn, how much oxygen shall be used in the systemic circulation and re- INTERNAL RESPIRATION. 389 stored in the pulmonary. In other words, the physiological work of an animal, which in turn is largely dependent upon how external forces act upon it, determines how much oxygen it uses daily; and not the supjily of oxygen how much its tissue activity shall be, unless the supply sinks below the starvation limit. CHAPTER XXVI. THE NERVOUS FACTORS OF THE RESPIRA- TORY MECHANISM. ASPHYXIA. The Respiratory Centre. The respiratory movements are to a certain extent under the control of the will; we can breathe faster or slower, shallower or more deejAj, as we wish, and can also " hold the breath" for some time — but the voluntary control thus exerted is limited in extent; no one can commit suicide by holding his breath. In ordinary quiet breathing the movements are quite involun- tary; they go on perfectly without the least attention on our part, and, not only in sleep, but during the unconscious- ness of fainting or of an apoplectic fit. The natural breath- ing movements are therefore either reflex or automatic. The muscles concerned in producing the changes in the chest which lead to the entry or exit of air are of the ordinary striped kind; and these, as we have seen, only con- tract in the Body under the influence of the nerves going to them; the nerves of the diaphragm are the two phrenic nerves (p. 161), one for each side of it; the external inter- costal muscles are supplied by certain branches of the dor- sal spinal nerves, called the intercostal nerves. If the phrenic nerves be cut the diaphragm ceases its contractions, and a similar paralysis of the external intercostals follows section of the intercostal nerves. Since the inspiratory muscles only act when stimulated by nervous impulses reaching them, we have next to seek where these impulses originate; and experiment shows that it is in tlie medulla oblongata. All the brain of a cat or a rabbit in front of the medulla can be removed, and it will still go on breathing; and children are sometimes born with THE RESPIRATORY CENTRE. 391 the medulla oblongata only, the rest of the brain being un- dereloped, and yet they breathe perfectly well. If, on the other hand, the spinal cord be divided immediately below the medulla of an animal all breathing movements of the chest cease at once. We conclude, therefore, that the nervous impulses calling forth contractions of the respira- tory muscles arise in the medulla oblongata, and travel down the spinal cord and thence out along the phrenic and intercostal nerves. This is conlirmed by the fact that if the spinal cord be cut across below the origin of the fourth ]-)air of cervical spinal nerves (from which the phrenics mainly arise) but above the first dorsal spinal nerves, the respiratory movements of the diaphragm continue but those of the intercostal muscles cease; this phenomenon has sometimes been observed in men stabbed in the back, so as to divide the spinal cord in the region indicated. Finally, that the nervous impulses exciting the inspiratory muscles originate in the medulla, is proved by the fact that if a small portion of that organ, the so-called vital point, be destroyed, all the respiratory movements cease at once and forever, although all the rest of the brain and spinal cord may be left uninjured. This part of the medulla is known as the respiratory centre. In the above statements, for the sake of simplicity, atten- tion has been chiefly confined to the diaphragm and the intercostal muscles; but what is said of them is true of the respiratory innervation of all other breathing muscles, whether expiratory or insjoiratory, normal or extraordinary; in all cases the impulse giving rise to a resjDiratory move- ment starts from the centre placed in the medulla oblon- gata. Is the Respiratory Centre Reflex ? Since this centre goes on working independently of the will we have next to inquire is it a reflex centre or not; are the efferent dis- charges it sends along the respiratory nerves due to afferent impulses reaching it by centripetal nerve-fibres; or does it originate efferent nervous impulses independently of excita- tion through afferent nerves? We know, in the first place, that the respiratory centre is 392 ■ THE HUMAN BODY. largely tinder reflex control; ji dash of coldwatei on the skin, the irritation of the nasal mucous membrane by snuff, or of the larynx by a foreign body, Avill eacli cause a modification in the respiratory movements — along indrawn breath, a sneeze, or a cough. But, although tluis subject to influences reaching it by afferent nerves, tlie respiratory centre seems essentially independent of such. In many animals, as rabbits, (and in some men,) marked breathing movements take place in the nostrils, which dilate during inspiration; and when the spinal cord of a rabbit is cut close to the medulla, thus cutting off all afferent nervous im- pulses to the respiratory centre except such as may reach it through cranial nerves, the resjiiratory movements of the nostrils still continue until death. The movements of the ribs and diaphragm of course cease, and so the animal dies very soon unless artificial respiration be maintained. Moreover, if after cutting the spinal cord as above described, all afferent cranial nerves be divided, so as to cut off the respiratory centre from all possible afferent nervous im- pulses, the regular breathing movements of the nostrils continue. It is, therefoi'e, obvious that the activity of the respiratory centre, however much it may be capable of modification through sensory nerves, is essentially inde- pendent of them; in other words the normal respiratory movements are not reflex. What it is that Excites the Respiratory Centre. The thing that, above all others, influences the respiratory centre is the greater or less venosity of the blood flowing through it. If this blood be very rich in oxygen and comparatively poor in carbon dioxide the inspiratory centre acts but feebly, and the respirations are shallow. If, on the other hand, this blood be highly venous the respiratory movements are more rapid than normal, and forced, the extraordinary muscles of respiration being called into play; this state of violent labored respiration, due to deficient aeration of the blood, is called dyspncea. IsTormal quiet breathing is eupncea. If the blood be highly aerated, as by keeping up forced artificial respiration for a time, all respiratory movements cease; the highly oxygenated blood does not excite the respiratory STIMULATIOX OF THE RESPIRATOBT CENTRE. 393 centre, tiiid the auimal therefore remains without breathing at all for some time; tliis condition is apncea, though phy- sicians by the word apno^a commonly mean merely extreme dyspnoea. If an animal be made apnoeic and the artificial respiration stopped, its blood, during the cessation of the respiratory movements, gradually losing oxygen and receiv- ing carbon dioxide, passes into the state of ordinary blood and again stimulates the respiratory centre, and the breath- ing movements then recommence. How it is that highly venous blood causes great excita- tion of the respiratory centre, and highly arterial cessation of its activity, is not certainly known; but we may make the following provisional hypothesis. The chemical changes occurring in the. respiratory centre give rise to a substance or substances which stimulate its nerve-cells. When the blood is richly supplied with oxygen this substance is oxi- dized and removed as fast as it is formed, and so the centre is not excited. When the blood, on the other hand, is un- usually poor in ox3'gen, this stimulating body accumulates and the respiratory discharges become more powerful. Under normal circumstances the blood oxygen is not kept quite up to the point of entirely removing this exciting substance, and the centre is stimulated so far as to pro- duce the natural breathing movements but not the more forced ones of dyspna3a. That the stimulating cause, what- ever it is, acts upon the respiratory centre and not upon the various organs of the Body and through their sensory nerves, in turn, upon the medulla, is proved not only by the facts above cited showing that the resjjiratory centre con- tinues to act when all afferent nerves are cut off from it, but also by experiments which show that the circulation of venous blood through the body of an animal, while at the same time its respiratory centre is supplied with arterial blood, does not produce dyspnoea; while sending venous blood to the medulla and arterial to all the rest of the Body does cause dyspncea. Why are the Respiratory Discharges Rhythmic ? Every complete resj)iratory act consists of an inspiration, an ex2)ira- tion and a pause; and then follows the inspiration of the 394 THE HUMAN BOD I*. next act. In natural quiet breathing there is no essential difference between the expiration and the pause. The in- spiration is the only active part (p. 303); the expiration and the pause are dependent on muscular inactivity and, there- fore, on the cessation of the discharge of nervous impulses from the resjjiratory centre. But then, we may ask, if in accordance with the hypothesis made in the last paragrai)h, the respiratory centre is constantly being excited, why is it not always discharging? why does it only send out nervous impulses at intervals? This question, which is essentially the same as that why the heart beats rhythmically, belongs to the higher regions of Ph}' siology and can only at present be hypothetically answered. Let us consider, for a moment, ordinary mechanical circumstances under which a steady supply is turned into an intermittent discharge. Suppose a tube closed water-tight below by a hinged plate, which is kept shut by a spring. If a steady stream of water is poured into the tube from above, the water will rise until its weight is able to overcome the pressure of the spring, and the plate will then be forced down and some water flow out. Tlie spring will then press the plate up again, and the water accumulate until its weight again forces open the bottom of the tube, and there is another outrush; and so on. By opposing a certain resistance to the exit we could thus turn a steady inflow into a rhythmic outflow. Or, take the case of a tube with one end immersed in water and a steady stream of air sent into its other end. The air will emerge from the immersed end, not in a steady current, but in a series of bubbles. Its pressure in the tube must rise until it is able to overcome the cohesive force of the water, and then a bubble bursts forth; after this the air has again to get up the requisite pressure in the tube before another bubble is ejected; and so the continuous suj^ply is trans^ formed into an intermittent delivery. Physiologists sup- pose something of the same kind to occur in the respira- tory centre. Its nerve-cells are always, under usual circumstances, being excited; but, to discharge a nervous impulse along the efferent respiratory nerves, they have to overcome a certain resistance. The nervous impulses have CAUSE OF THE RESPIRATOIiT RHYTHM. 395 to accumulate, or ''gain a head," before they travel out from the centre, and, after their discharge, time is required to attain once more the necessary level of irruption before a fresh innervation is sent to the muscles. This method of accounting for the respiratory rhythm is known as the "resistance theory " If not altogether satisfactory it is at least far preferable to the older mode of considering the question solved by assuming a rhythmic character or prop- erty of the respiratory centre. It gives a definite hypothe- sis, which accords with what is known of general natural laws outside of the Body, and the truth or falsity of which can be tested by experiment: and so serves very well to show how scientific differs from pre-scientific, or mediseval, physiology. The latter was content with observing things in the Body and considered it explained a phenomenon when it gave it a ijame. Now we call a phenomenon ex- plained, when we have found to what general catego^'y of natural laws it can be reduced as a special example; and this reducing a special case to a particular manifesta- tion of some one or more general properties of matter already known is, of course, all that we ever mean wlien we say we explain anything. We explain the fall of an apple and the rise of the tides by referring them to the class of general results of the Law of Gravitation; but the why of the law of gravitation we do not know at all; it is merely a fact which we have found out. So with regard to Physi- ology; we are working scientifically when we try to reduce the activities of the living Body to special instances of mechanical, physical, or chemical laws otherwise known to us, and unscientifically when we lose sight of that aim. Cerfeain vital phenomena, as those of blood-pressure, we can thus explain, as much as we can explain anything; others, as the rhythm of the respiratory movements, we can provision- ally explain, although not yet certain that our explana- tion is the right one; and still others, as the phenomena of consciousness, jwo_cannot explain at all, and possibly _never will, by referring them to general properties of matter, since they may be properties only of that particular kind-of 306 THE HUMAX BODY. matter called protoplasm, and i)erlia])s only of some varie- ties of it. The Relation of the Pneumogastric Nerves to tlie Res- piratory Centre, ^^'e have iiexi lo CDiisidcr if any })lieno- mena presented by the living Body give support to the resistance theory of the res})iratoi"y rhythm. A very ini- porti^nt collateral proj) to it is given by the relation of the pneumogastric nerves to the rate and force of the respira- tory movements. These nerves give branches to the larynx, the windpipe, and the lungs, in addition to numerous other parts, and might therefore be suspected to have something to do with breathing. That they are not concerned in in- fluencing the resjiiratory muscles directly is shown by the fact that all of these muscles (except certain small ones in the larynx) contract as usual in breathing after both pneumogastric nerves have been divided. Still, the section of both nerves has a considerable influence on the respira- tory movements; i\iQj\)eGOXi\e sloiver and deeper. We may understand this by supjjosing that the resistance to the dis- charges of the respiratory centre is liable to variation. It may be increased, and then the discharges will be fewer and larger; or diminished, and then they will be more frequent but each one less powerful. If the spring, in the illustra- tion used in the preceding paragrajih, be made stronger, while the inflow of water to the tube remains the same, the outflows will be less frequent but each one greater; and vice verna. The effect of section of the pneumogastric trunk ^ may, therefore, be explained if we suppose that, normally, it carries up, from its lung branches, nervous impulses which diminish the resistance to the discharges of the respiratory centre; when the nerves are cut these helping impul:;es are lost to the centre, and its impulses must gather more head before they break out, but will be greater when they do. This view is confirmed by the fact that stimulation of the central ends of the divided pneumogastrics, if weak, brings back the respirations to their normal rate and force; if strcaiger makes them more rapid and shallower; and when stronger still, abolishes the respiratory rhythm altogether, with the inspiratory muscles in a steady state of feeble con- THE EXPTRArORY CENTRE. 397 traction. That is to say, tlie resistance to the discharges of the centre being entirely taken away (which is equivalent to tlie total removal of the spring m our example), the cen- tre sends out uninterrupted and non-rhythmic stimuli to tlio ins])iratory muscles. The pneumogastric nerve gives two branches to the larynx: known respectively as the superior and inferior {recurrent) laryngeal nerves; the action of these on the respiratory centre is opposite to that of the fibres from the lungs coming up in the main pneumogastric trunk. If the superior laryngeal branch be divided and its central end stimulated, the respirations become less frequent but each one more powerful; hence this nerve is supposed to increase the resistance to the discharges of the respiratory centre. The same, but to a less degree, is true of the inferior laryn- geal branch. The Expiratory Centre. Hitherto we have considered Ijreathing as due to the rhythmically alternating activity and rest of an inspiratory centre — and such is the case in normal quiet breathing, in which the expirations are pas- sive. But in dyspnoea expiration is a muscular act, and so there must be a section of the respiratory centre control- ling the expiratory muscles. This part of the respiratory centre, howe^'er, is less irritable than the inspiratory part, and hence when the blood is in a normal state of aeration never gets stimulated up to the discharging ]3oint. In dysp- ntea the stimulus becomes sufficient to cause it also to discharge, but only after the more irritable inspiratory centre; hence the expiration follows the inspiration. This alternation of activity is, moreover, promoted by the fact that the pneumogastric nerve-fibres coming ^^.^ from the lungs are of two kinds. The predominant sort are those already referred to, which diminish the resistance to dis- charge of the inspiratory centre, and perhaps also increase the resistance to the expiratory discharge. This set is ex- cited when the lungs diminish in bulk, as in expiration; and when the whole nerve is stimulated electrically they usually get the better of the other set, which carry up to the medulla impulses which increase the resistance to in- 398 THE Hi' MAX BODY. spiratory discliarges and diminish that to expiratory, and are stimulated when the lungs expand. Hence, every ex- pansion of the lungs (inspiration) tends to promote an expiration, and every collapse of the lungs (expiration) tends to produce an inspiration; and so, through the pneumo- gastric nerves, the respiratory mechanism is largely self- regulating. Asphyxia. Asphyxia is death from suffocation, or want of oxygen by the tissues. It may be brought about in various ways; as by strangulation, which prevents the entry of air into the lungs; or by exposure in an atmosphere con- taining no oxygen; or by putting an animal in a vacuum; or by making it breathe air containing a gas which has a stronger affinity for haemoglobin than oxygen has, and Avhich, therefore, turns the oxygen out of the red corpuscles and takes its place. The gases which do the latter are very interesting since they serve to prove conclusively that the Body can only live by the oxygen carried round by the haemoglobin of the red corpuscles; that amount dissolved in the blood plasma being insufficient for its needs. Of such gases carbon monoxide is the most important and best studied; in the favorite French mode of committing suicide by stopping up all the ventilation holes of a room and burning charcoal in it, it is poisoning by carbon monoxide which causes death. The Relations of Carbon Monoxide to Hsemoglobin. If aerated whipped blood, or a solution of oxyhemoglobin, be exposed to a gaseous mixture containing carbon mon- oxide, the liquid will absorb the latter gas and give off oxygen. The amount of carbon monoxide taken up will (apart from a small amount dissolved in the plasma) be inde- pendent of the partial pressure of ihat gas in the gaseous mixture to which the blood is exposed; the quantity absorbed depends on the quantity of haemoglobin in the liquid, and is replaced by an equal volume of oxygen liberated. This equivalence of volume, of itself, proves that the phe- nomenon is due to the chemical rej)lacement of oxygen in some compound, by the carbon monoxide; for if the carbon monoxide were merely dissolved in the liquid in CABBOX-MOyOXIDE HAEMOGLOBIN. 399 proportion to its partial pressure on the surface, it would turn out no oxygen; the quantity of dissolved gases held by a liquid being dependent only on the partial pressure of each individual gas on its surface, and unaffected by that of all others. During the taking up of carbon monoxide the blood changes color in a way that can be recognized by a practiced eye; it becomes cherry red instead of scarlet. This shows that some new chemical compound has been formed in it; examination with the spectroscope confi]-ms this, and shows the color change to be due to the formation of carbon-monoxide ha?moglol)in which has a different color from oxyhsemogiobin. A dilute solution of reduced hsemogiobin absorbs all the rays of light in one region about the green of the solar spectrum (see Physics), and so produces there a dark band; a thin layer of the blood of an asphyxiated animal does the same. Dilute solution of oxyhemoglobin absorbs the rays in two narrow regions of the solar spectrum at the confines of the yellow and green, and arterial blood does the same. Dilute solution of carbon-monoxide haemoglobin, or blood which has been exposed to this gas, also absorbs the light in two narrow bands of the solar spectrum; but these are nearer the blue end of the spectrum than the absorption bands of oxyha3moglobin. Pure blood serum saturated with oxy- gen gas or with carbon monoxide does not specially absorb any part of the spectrum; therefore the absorjitions when haemoglobin is present, must be due to chemical compounds of those gases with that body. Since carbon-monoxide-hremoglobin has a bright red color, we find in tlie Bodies of persons poisoned by that gas, the blood all through the Body cherry red; the tissues being unable to take carbon monoxide from haemoglobin in the sys- temic circulation. Hence the curious fact that, while death is really due to asphyxia, the blood is almost the color of arterial blood, instead of very dark purple, as in ordinary cases of death by suffocation. Experiments with animals show that in poisoning by carbon monoxide persistent ex- posure of the blood to oxygen, by means of artificial respi- ration, will cause the poisonous gas to be slowly replaced 400 THE HmrAN BODY. again by oxygen; hence if the heart has not yet quite stopped beating, artificial respiration, kept up patiently, should be emi)loyed for the restoration of i^ersous poisoned l)y carlion monoxide. The Phenomena of Asphyxia. As soon as the oxygen in the blood falls below the normal amount the breathing becomes hurried and deeper, and tlie extraordinary muscles of respiration are called into activity. The dyspnoea Ijc- comes more and more marked, and this is especially the case with the expirations which, almost or quite passively performed in natural breathing, become violently muscular. At last nearly all the muscles in the Body are set at work; the rhythmic character of the respiratory acts is lost, and general convulsions occur, but, on the whole, the contrac- tions of the expiratory muscles are more violent than those of the inspiratory. Thus undue want of oxygen at first merely brings about an increased activity of the respiratory centre, and especially of its expiratory division which is not excited in normal breathing. Then it stimulates other jior- tions (the convuUive centre) of the medulla oblongata also, and gives rise to violent and irregular muscular spasms. That the convulsions are due to excitation of nerve-centres in the medulla (;ind not, as might be supposed, to poisoning of the muscles by the extremely venous blood) is shown by the facts (1) that they do not occur in the trunk of an animal when the spinal cord has been divided in the neck so as to cut off the muscles from the medulla; and (2) that they still occur if (the si)inal cord remaining undivided) all the parts of the brain in front of the medulla have been re- moved. The violent excitation of the nerve-centres soon exhausts them, and all the more readily since their oxygen supply (which they like all other tissues need in order to continue their activity) is cut off. The convulsions therefore gradu- ally cease, and the animal becomes calm again, save for an occasional act of breathing when the oxygen want becomes so gi-eat as to cause efficient stimulation even of the dying respiratory centre: these final movements are inspira- tions and, becoming less and less frequent, at last cease. ASPHYXIA. 401 and the animal appears dead. If, however, its chest be opened the heart will be found gorged with extremely dark venous blood and making its last few slow feeble pulsations. So long as it beats artificial respiration can restore the ani- mal, but once the heart has finally stopped restoration is impossible. There are thus three distinguishable stages in death from asphyxia. (1) The stage of dyspna?a. (2) The stage of convulsions. (3) The stage of exhaustion; the convulsions having ceased but there being from time to time an inspiration. The end of the third stage occurs in a mammal about five minutes after the oxygen supply has been totally cut off. If the asphyxia be due to deficiency, and not absolute want, of oxygen of course all the stages take, longer. Circulatory Changes in Asphyxia. During death by suffocation characteristic changes occur in the working of the heart and blood-vessels. The heart at first beats quicker, btit very soon, before the end of the dyspnceic stage, more slowly, though, at first, more powerftilly. This slowing is due to the fact that the itnusual want of oxygen leads to stimulation of the cardio-inhibitory centre in tlie medulla (p. 250) and this, through the pneumogastric nerves, slows the heart's beat. Soon, however, the want of oxygen affects the heart itself and it begins to beat more feebly, and also more slowly, from exhaustion, until its final stoppage. During the second and third stages the heart and the venge cavfB become greatly overfilled with blood, because the violent mttscular contractions facilitate the flow of blood to the heart, while its beats become too feeble to send it oitt again. The overfilling is most marked on the right side of the heart which receives the venous blood from the Body generally. During the first and second stages of asphyxia arterial pressure rises in a marked degree. This is due to excitation of the vaso-motor centre (p. 254) by the venous blood, and the consequent constriction of the nmscular coats of the arteries and increase of the peripheral resistance. In the third stage the blood-pressure falls very rapidly, becaitse the feebly acting heart then fails to keep the arteries 402 THE HUMA^ BODY. tense, even although their diminished cahbre greatly slows the rate at which they empty themselves into the capilla- ries. Another medullary centre unduly excited during asphyxia IS that from which j)roceed the nerve-fibres governing (hose muscular fibres of the eye which enlarge the pupil. During suffocation, therefore, the pupils become widely dilated. At the same time all reflex irritability is lost, and touching the eyeball causes no wink; the reflex centres all over the Body being rendered, through want of oxygen, in- capable of activity. The same is true of the higher nerve- centres; unconsciousness comes on during the convulsive stage, whicli, liorrible as it looks, is unattended with suffer- ing. Modified Bespiratory Movements. SigMng is a deep long-drawn inspiration followed by a shorter but corresj^ond- ingly large expiration. Yuuming is similar, but the air is mainly taken in by the mouth instead of the nose, and the lower jaw is drawn down in a characteristic manner. Hic- cough depends upon a sudden contraction of the dia])hragm, while the aperture of the larynx closes; the entering air, drawn through the narrowing opening, causes the peculiar sound. Coug]ii7ig consists of a full inspiration followed by a violent and rapid expiration, during the first part of which the laryngeal opening is kept closed; being afterwards sud- denly opened, the air issues forth with a rush, tending to carry out with it anything lodged in the windpipe or larynx. Sneezing is much like coughing, except that, while in a cough the isthmus of the fauces is held open and the air mainly passes out through the mouth, in sneezing the fauces are closed and the blast is driven through the nostrils. It is commonly excited by irritation of the nasal mucous membrane, bat in many persons a sudden bright light falling into the eye will produce a sneeze. Laughing consists of a series of short expirations following a single inspiration; . the larynx is open all the time, and the vocal cords (Chap. XXXVI.) are set in vibration. Crying is, phy- siologically, much like laughing and, as we all know, one often jjasses into the other. The accomjjanyiug contrac- MODIFIED RmPIRATORT MOVEMENTS. 403 tions of the face muscles giving expression to the counten- ance are, however, different in the two. All these modified resjiiratory acts are essentially reflex, but, with the exception of hiccough, they are to a certain extent, like natural breathing, under the control of the will. Most of them, too, can be imitated more or less perfectly by voluntary muscular movements; though a good stage sneeze or cough is rare. CHAPTER XXVII. THE KIDNEYS AND SKIN. General Arrangement of the Urinary Organs. These consist of (1) the Tculneys, the ghmds which secrete the urine; (2) the ureters or ducts of the kidneys, which carry their secretion to (3) the urinary bladder, a reservoir in Avhich it accumulates and from which it is expelled from time to time through (4) an exit tube, the urethra. The general arrangement of these parts, as seen from behind, is shown in the figure opposite. The kidneys, i?, lie in the dorsal part of the lumbar region of the abdominal cavity, one on each side of the middle line. Each is a solid mass, with a convex outer and a concave inner border, and its upper end a little larger than the lower. From the abdominal aorta. A, a renal artery, Ar, enters the inner border of each kidney, to break up within it into finer branches, ultimately ending in capillaries. The blood is collected from these into the renal veins, Vr, one of which leaves each kidney and opens into the inferior vena cava, Vc. From the concave border of each kidney proceeds also the tcreter, U, a slender tube from 28 to 34 cm. (11 to 13.5 inches) long, opening below into the bladder, Vu, on its dorsal aspect, and near its lower end. From the bladder proceeds the urethra, at Ua. The channel of each ureter passes very obliquely through the T^all of the bladder to open into it; accordingly if the pressure inside the latter organ rises above that of the liquid in the ureter, the walls of the oblique passage are pressed together and THE RENAL OBGANS. 405 Fig. 114.— The renal organs, viewed from behind. R, right kidney; A, aorta; Ar, right renal artery; Vc, inferior vena cava; TV, right renal vein; U, i-ight ureter; Fm, bladder; C/a, commencement of ui'ethra. 406 THE HUMAN BODY. it is closed. Usually the bladder, which has a thick coat of unstriped muscular tissue lined by a mucous membrane, is relaxed, and the urine flows readily into it from the ureters. The commencement of the urethra being kept closed by clastic tissue around it (which can voluntarily be reinforced by muscles which compress the tube) the urine accumulates in the bladder. When this latter contracts and presses on its contents, the ureters are closed in the way above indicated, the elastic fibres closing the urethral exit from the bladder are overcome, and the liquid forced out. Naked Eye Structure of the Kidneys. These organs have externally a red-brown color, which can be seen through the transparent capsule of peritoneum which en- velops them. When a section is carried through a kidney from its outer to its inner border (Fig. 115) it is seen that a deep fissure, the hilus, leads into the latter. In theJiilus the ureter widens out to form the ^;eZy/.s, which breaks up again into a number of smaller divisions, the cups or calices. The cut surface of the kidney proper is seen to consist of two distinct parts; an outer or cortical portio7i, and an inner or medullary. The medullary portion is less red and more glistening to the eye, is finely striated in a radial direction, and does not consist of one continuous mass but of a number of conical portions, the pyramids of MalpigJii, 2', each of which is separated from its neighbors Ijy an in- ward prolongation,*, of the cortical substance. This, how- ever, does not reach to the inner end of the pyramid, which projects, as the papilla, into a calyx of the ureter. At its outer end each pyramid separates into smaller i^or- tions, the pyramids of Ferrein, 2", separated by thin layers of cortex and gradually spreading everywhere into the lat- ter. The cortical substance is redder and more granular looking and less shiny than the medullary, and forms every- where the outer layer of the organ next its capsule, besides dipping in between the j^yramids in the way described. The renal artery divides in the hilus into branches (5) which run into the kidney between the pyramids, giving off a few twigs to the latter and ending finally in a much HISTOLOGY OF THE KIDyETS. 401 richer vascular network in the cortex. The branches of the renal vein have a similar course. The Minute Structure of the Kidney. The kidneys are compound tubular glands, composed essentially of Fig. 115— Section througrh the li-htkidnev from its-outer to its inner border 1. cortex; 2, medulla; e. pyramirt of Blalpighi: 2'. pyramid of Ferrein: ^- small branches of the renal artery entering between the pyramids; A, a branch or the renal artery ; C, the pelvis of the kidney ; U, ureter. branched microscopic iiriniferous tubules, lined by epithe- lium. Each tubule commences at a small opening on a papilla and from thence has a very complex course to its other extremity. Usually about twenty open, side by side, on one papilla. There they have a diameter of about 408 THE HUMAN BODY. 0.125 mm. (^-^ inch). Running into the i^jTamid from this point each tuhule divides several times. At first the hranclies are smaller than the main tube; but as soon as they have come down to about 0.04 mm. (-g^ inch) this diminution in size ceases, and the division continuing while the tubules retain the same diameter, the pyramid thus gets, in part, its conical form. Ultimately each branch runs somewhere out of the pyramid, either from its base or side, into the cortex and there dilates and is twisted. It then narrows and doubles back again into the pyramid and runs as a straight tube towards the papilla, but before reach- ing it makes a loop, and turns back again as a straight tube to the base of the pp'amid;, where it once more enters the cortex, dilates and becomes contorted, and then ends in a sioherical capsule, containing a tuft of small blood-vessels. Or, followed the other way, each tubule commences in the cortex with a globular dilatation, the MaljngMan capsule. From this it continues as a convoluted tubule in the cortex; this passes into a pyramid, becomes straight, and runs on as the descending limb of a loop of Henle. Turn- ing at the loop, it continues as its ascending limb, and this passes out again into the cortex and becomes the convoluted junctional tubule, which passes as a straight collecting tubule into the pyramid and there joins others to form an excretory tnbule which opens on the papilla. Throughout its course the tubule is lined by a single layer of epithelium cells differing in character in its different sections. All the tubes are bound together by connective tissue and blood- vessels to form the gland. The Blood-Plow through the Kidney. The final twigs of the renal artery in the cortex, giving off a few branches which end in a capillary network around the convoluted tubules, are continued as the afferent vessels of Malpi- ghian capsules, the walls of which are doubled in before them (Fig. 116); there each breaks up into a little knot of capillary vessels called the glomerulus, from which ulti- mately an efferent vessel proceeds, and outside the capsule this breaks up into a close capillary network among the con- voluted tubes. From the capillaries the blood is collected THE RENAL SECRETION. 409 into the renal vein. Most of the blood flowing through the kidney thus goes through two sets of capillaries; one in the capsules, and a second formed by the breaking uj) of the efferent vein of the latter. The capillary network in the pyramids is much less close than that in the cortex, which gives reason to suspect that most of the secretory work of the kidneys is done in the cap- sules and convoluted tubules. The pyramidal blood flows only through one set of capil- laries, there being no glome- ruli in the kidney medulla. The Renal Secretion. The amount of this carried off from the Body in 24 hours is subject to considerable varia- tion, being especially dimin- ished by anything which pro- motes perspiration, and increased by conditions, as cold to the surface, which diminish the skin excretion, its average daily quantity varies from 1200 to 1750 cub. cent. (40 to 60 fluid ounces). The urine is a clear amber-colored liquid, of a slightly acid reaction; its specific gravity is about 1040, being higher when the total quantity excreted is small than when it is greater, since the amount of solids dissolved in it remains nearly the same in health; the changes in its bulk being dependent mainly on changes in the amount of water separated from the blood by the kidneys. Normal urine consists, in 1000 parts, of about 9G0 water and 40 solids. The' latter are mainly crystalline nitro- genous bodies {iirea and tiric acid), but small quantities of pigments and of non- nitrogenous organic bodies are also present, and a considerable quantity of mineral salts. The following table gives approximately, in the first column, the Fig. 116.— The termination of a uriniferous tubule, with its glomeru- lus, a. the glomerulus or Malphighian corpuscle; 6. the convoluted ending of the tubule :d, its lining epithelium; /, the afferent blood-vessel of the glomerulus; (/, the efferent vessel ; c, h, the blood-vessels forming the tuft in the glomerulus. 410 THE HUMAN BODY. average composition of the urine excreted in twenty-four hours expressed in grams; in the second column the same expressed in grains. The third column gives the composi- tion of 1000 parts of urine. The solids consisls of — Urea (CN2H4O) Uric acid (C5H4N4O:0. Hippuric acid ^. . Kreatinin T . . Pigments and fats. . . . Sulphiu'ic acid Pliosplioric acid Chlorine Ammonia Potassium Sodium Calcium Magnesium Urine in 24 hours. 1500 grams. 23,250 grains. In'lOOO parts. Water Solids 142S.00 73.00 23,134.00 1116.00 952.00 48 00 33.00 0.50 0.40 1.00 10.00 2.00 3.00 7.00 0.75 2.50 11.00 0.25 0.20 511.50 7.75 6.20 15.50 155.00 31.00 46.50 108.50 13.00 38.75 170.50 3.80 3.00 71.60 1110.00 22.00 0.83 0.27 10.33 6.66 1.33 2.00 4.70 0.50 1.70 7.33 0.16 0.13 47.44 The urine, however, even in health is subject to consid- erable variation in composition; not only as regards the amount of water in it, but also in its solid constituents; the latter are especially modified by the quantity and nature of the food taken. 100 volumes of urine contain in solution about 14 vol- umes of gas, consisting of about 13 of carbon dioxide, 1 of nitrogen, and mere traces of oxygen. Mechanism of the Benal Secretion. The kidneys, as secreting organs, consist of two distinct parts; (1) the glome- ruli through which a filtration of water, probably with salts in solution, takes 2:)lace; and (2) an actively secretory appa- ratus, formed by all parts of the urinif erous tubules between their terminal capsules and the collecting tubes. Accord- inglv, we find in the urine bodies, as water and common salt, which already exist in the blood and can be removed from it SOURCES OF UREA. 411 merely by dialysis or filtration ; and others (the specific ele- ments of the secretion), especially urea, which are selected or made by a special activity of the renal gland-cells. The total quantity of the twenty-four hours' urine thus depends largely on the pressure in the renal arteries, since the higlier this is the greater will the amount of filtered liquid be. Under ordinary pressures such substances as albumen will not filter, but they do under higher; accordingly in healthy conditions none of the albumen of the blood jjlasma passes into the urine, but if the pressure in the capillaries of the glomeruli is considerably raised it does; its presence in the urine being the most prominent symjotom of that danger- ous class of maladies grouped together under the name of Briglifs disease. Filtration in the glomeruli is largely pro- moted by the fact that the calibre of the efferent vessel of each is rather less than that of the afferent; and thus the pressure m the thin-walled vessels of the vascular tuft is raised. The Role of the Renal Epithelium. Water and salines being passed out mainly through the glomeruli, we have now to consider what part the secreting cells of the kidney play; and especially as regards urea, the most important constituent of the urine. Urea represents the final state in which the proteids taken into the Body from the alimen- tary canal (or at- least their nitrogen) leave, after having 3delded up, by chemical changes, a certain amount of energy. In this process the proteids are oxidized and broken down into carbon dioxide and water and urea; and the kidneys get rid of the latter. Since the life and activity of every tissue is accompanied by a breaking down of proteids (though not necessarily at once into urea, as many intermediate stages may, and doubt- less do, occur in different tissues), there is no doubt that the main chemical degradation of albuminous compounds takes place outside the kidneys. Whether the final steps by which urea is formed occur in those organs or elsewhere is not yet certainly known. According to one view the urea is carried to the kidney m the blood of the renal artery, and there merely picked up and passed on by the excreting cells; while, according to another, not urea, but the penul- 412 THE HUMAN BODY. timate products of protcid degradiition from which urea is made, are carried to the kidneys, and tlie final formation of urea occurs in tliese organs. The results of hlood analy- sis are conflicting, but on the whole it seems proved that more urea exists in renal-artery blood than in renal-vein blood, which indicates that urea is not made in the kidneys. In death, too, from suppression of the renal reaction, urea is found to accumulate in the blood which would not be the case unless it were normally formed elsewhere and car- ried off by the kidneys. The whole urea question, which is one of great importance, will l)o more fully considered in Chapter XXVIII., in connection with the chemistry of nutrition in general. The Skin» which covers the whole exterior of the Body, consists everywhere of two distinct layers; an outer, the cuticle or ejjulet'rnis, and a deeper, the dermis, cutis vera, or corium. A blister is due to the accumulation of liquid between these 'two layers. The hairs and nails are excessively develoj)ed parts of the epidermis. The Epidermis, Fig. 117, consists of cells, arranged in many layers, and united by a small amount of cementing substance. The deepest layer, d, is composed of elongated or columnar cells, set on with their long axes perpendicular to the corium beneath. To it succeed several strata of roundish cells, h, which in the outer layers become more and more flattened in a plane parallel to the surface. The outermost epidermic stratum is comi)osed of many layers of extremely flattened cells from which the nuclei (conspicu- ous in the deeper layers) have disappeared. These super- ficial cells are dead and are constantly being shed from the surface of the Body, while their place is taken by new cells, formed in the deeper layers, and pushed up to the surface and flattened in their progress. The change in the form of the cells as they travel outwards is accompanied l)y chemical changes, and they finally constitute a semitrans- parent dry lior^iy stratmn, a, distinct from the deeper, more ojiaque and softer Ifalpighian or mucoiis layer, h and d, of the epidermis. The cells of this latter are soluble in acetic acid; those of the horny stratum, not. THE EPIDERMIS. 413 The rolls of material which are peeled off the skin in the " shampooing" of the Turkish bath, or by rubbing with a rough towel after an ordinary warm bath, are the dead outer scales of the horny stratum of the epidermis. Fig. 117.— a section through the epidermis, somewhat diagrammatic, highly magnified. Below is seen a papilla of the dermis, with its artery, /, and veins. Of/; a, the horny layer of the epidermis; b, the rete mucosum or Malpighian layer; d, the layer of columnar ei)idermic cells in immediate contact with the dermis; A, the duct of a sweat-glaud. In dark races the color of the skin depends mainly on minute pigment granules lying in the deeper cells of the Malpighian layer. 414 THE HUMAN BODY. H^o blood-vessels or lymphatics enter the epidermis, which is entirely nourished by matters derived from the subjacent corium. Fine nerve-fibres run into it and end there among the cells, in various ways. The Corium, Cutis Vera, or True Skin, Fig. 118, con- sists fundamentally of a close feltwork of elastic and white fibrous tissue, which, becoming wider nie.'shed below, passes .'<« L.^ ...d Fig. 118. — A sectiuu through the skin and subcutaneous areolar tissue, a, norny stratum, and 6, Malpighian layer of the epidermis; c, dermis, passing below into, d, loose areolar tissue, with fat,/, in its meslies: above, dermic papillae are seen, projecting into the epidermis which is moulded on them, i, opening of a sweat-gland ; /i, duct of ditto ; g, the gland itself. gradually into the subcutaneous areolar tissue (p. 102) which attaches the skin loosely to parts beneath. In tanning it is the cutis vera which is turned into leather, its white fibrous tissue forming an insoluble and tough com- pound with the tannin of the oak-bark employed. Wherever there are hairs, bundles of plain muscular tissue are found in the corium; it contains also a close capillary network HAIRS. 415 and numerous lymphatics and neryes. In shaving, so long as the razor keejss in the epidermis there is no bleed- ing; but a deeper cut shows at once the vascularity of the true skin. The outer surface of the corium is almost everywhere raised mto minute elevations, called the papillm, on which the epidermis is moulded, so that its deep side presents pits corresponding to the projections of the dermis. In Fig. 117 is a papilla of tho, corium containing a knot of blood-vessels, supplied by the small artery,/, and having the blood carried off from them by the two little veins, g g. Other papillse contain no capillary loops but special organs connected with nerve-fibres, and supposed to be concerned in the sense of touch. On the palmar surface of the hand the dermic papillae are especially well developed (as they are in most parts where the sense of touch is acute) and are frequently comjjoioid or branched at the tip. On the front of the hand, they are arranged in rows; the epidermis fills up the hollows between the papillop, of the same row, but dips down between adjacent rows, and thus are produced the epidermic ridges seen on the palms. In many places the corium is furrowed, as opposite the finger-joints and on the palm. Elsewhere such furrows are commonly less marked, but they exist over the whole skin. The epidermis closely follows all the hollows, and thus they are made visible from tlie surface. The wrinkles of old persons are due to the absorption of subcutaneous fat and of other soft parts beneath the skin, which, not shrinking itself at the same rate, becomes thrown into folds. Hairs. Each hair is a long filament of epidermis devel- oped on the top of a special dermic papi,lla, seated at the bottom of a depression reaching down from the skin into the tissue beneath and called the Mir follicle. The portion of a hair buried m the skin is called its root, this is succeeded by a stem which, in an uncut hair, tapers off to apoint. The stem is covered by a single layer of overlapping scales form- ing the Jiair cuticle; the projecting edges of these scales are directed towards the top of the hair. Beneath the hair cuticle comes the cortex, made up of greatly elongated cells 416 THE HUMAN BODY. united to form fibres; and in the centre of the shaft there is found, in many hairs, a meduUa, ni.ide np of more or less rounded cells. The color of hair is mainly dependent upon pigment granules lying lietween the fibres of the cortex. All hairs contain some air cavities, especially in the medulla. They are very abundant in white hairs and cause the white- ness by reflecting all the incident light, just as a liquid beaten into fine foam looks white because of the light reflected from the walls of all the little air cavities in it. In dark hairs the air cavities are few. The hair follicle (Fig. 119) is a narrow pit of the dermis, projecting down into the subcutaneous areolar tissue, and lined by an involution of the epidermis. At the bottom of the follicle is a j^apilla and tlie 3, ^.^ •• ^.^„' . epidermis, turning up over this, becomes continuous with the hair. On the papilla epidermic cells multi})ly rapidly so long as the hair is growing, and the whole hair is there made up of roundish cells. Firr. 119.— Parts of two hairs imbedded As thcSC get pushed Up by in their follicles, o, the skin, which is seen « -i „,„^ ^„ „n v^ „„j-i to dip down and line tiie follicle; 6, the trcsh oucs tomied beneath subontaneous tissue; c, the muscles of the ^-i,„„-, thp nntprmnsf livor hair follicle, which by their contraction inem, tlie OLlieimoSL laytr can erect the hair. become flattened and form the hair cuticle; several succeeding layers elongate and form the cortex; while, in hairs with a medulla, the middle cells retain pretty much their original form and size. Pulled apart by tlie elongating cortical cells, these central ones then form the medulla with its air cavities. The innermost layer of the epidermis, lining the follicle, has its cells projecting, with overlapping edges turned down- wards. Accordingly these interlock with the upward directed edges of the cells of the hair cuticle; consequently when a hair is pulled out the epidermic lining of the follicle IS usually brought with it. So long as the dermic papilla is left intact a new hair will be formed, but not otherwise. Slender bundles of unstriped muscle {c, Fig. 119) run from NAILS. 417 the dermis to the side of the hair follicles. The latter are obliquely implanted in the skin so that the hairs lie down on the surface of the Body, and the muscles are so fixed that, when they shorten, tlioy erect the hair and cause it to bristle, as may be seen in an angry cat, or sometimes in a greatly terrified man. Opening into each hair follicle are usually a couple of sebaceous glands (p. 418). Hairs arc found on all regions of the skin except the palms of the hands and the soles of the feet; the back of the last phalanx of the fingers and toes, the upper eyelids, and one or two other regions. Nails. Each nail is a part of the epidermis, with its horny stratum greatly developed. The back part of the nail fits behind into a furrow of the dermis and is called its root. The yisible part consists of a iody, fixed to the dermis beneath (which forms the led of the nail), and of a free edge. Near the root is a little area whiter than the rest of the nail and called the lunula. The whiteness is due in part to the nail being really more opaque there and partly to the fact that its bed, which seen through the nail causes its pink color, is in this region less yascular. The portion of the corium on which the nail is formed is called its matrix. Behind, this forms a furrow lodging the root, and it is by new cells added on there that the nail grows in length. The part of the matrix lying beneath the body of the nail, and called its bed, is highly vascular and raised up into papillfe which, except in the region of the lunula, are arranged in longitudinal rows, slightly diverging as they run towards the tip of the finger or toe. It is by new cells formed on its bed and added to its under surface that the nail grows in thickness, as it is i)ushed forward by the new growth in length at its root. The free end of a nail is therefore its thickest part. If a nail is " cast" in consequence of an injury, ortornoif, a new one is produced, provided the matrix is left. The Glands of the Skin are of two kinds, the sudori- jjarous or sweat glands, and the sebaceous or oil glands. The former belong to the tubular, the latter to the race- mose type. The sweat-glands, Fig. 120, lie in the subcu- 418 THE HUMAN BODY. taneons tissue, where they form little globular masses com- posed of a coiled tube. From the coil a duct (sometimes double) leads to the surface, being usually spirally coiled as it passes througli the epidermis. The secret- ing part of the gland consists of a connective-tissue tube, continuous along the duct with the dermis; Avithin this is a basement membrane^ and the final secretory lining consists of several layers of gland-cells. A close capillary network intertwines with the coils of the gland. Sweat- glands are found on all regions of the skin, but more closely set in some places, as the palms of the hands and the brow, than elsewhere. There arc altogether about two and a half millions of them opening on the surface of the Body. The sebaceous glands nearly always open into hair follicles, and are found wherever there are hairs. Each con- sists of a duct opening near the mouth of a hair follicle and branching at its other end : the final branches lead into globu- lar secreting saccules, which, like the ducts, are lined with epithelium. In the saccules the substance of the cells becomes charged with oil-drops, the protoplasm disappearing; and finally the whole cell falls to pieces, its detritus constitut- ing the secretion. New cells are, meanwhile, formed to take the place of those destroyed. Usually two glands are con- nected with each hair follicle, but there may be three or only one. A jiair of sebaceous glands are represented on the sides of each of the hair follicles in Fig. 119. The Skin Secretions. The skin besides forming a pro- tective covering and serving as a sense-organ (Chap. XXXIV.) also plays an important part in regulating the temperature of the Body, and, as an excretory organ, in carrying off certain waste products from it. Fig. 120.— a sweat gland, d, horny layer of cuticle; c, Malpighian layer: h. dermis. The coils of the gland proper, imbedded in the suljcu- taneous f at,areseenbelow the dermis. PERSPIRATIOX. . 419 The sweat poured out by tlie sudoriparous glands is a transparent colorless liquid, with a peculiar odor, yaryiug in different races, and in the same individual in different regions of the Body. Its quantity in twenty-four hours is subject to great variations, but usually lies between 700 and 'lOQO grams (10,850 and 31,000 grains). The amount is in- fluenced mainly by the surrounding temperature, being greater when this is high; but it is also increased by other things tending to raise the temperature of the Body, as muscular exercise. The sweat may or may not evaporate as fast as it is secreted ; in the former case it is known as insensiUe, in the latter as sensible jperspiration. By far the most passes off in the insensible form, drops of sweat onh' accumulating when the secretion is very profuse, or the surrounding atmosphere so humid that it does not readily take up more moisture. The perspiration is acid, and in 1000 parts contains 990 of water to 10 of solids. Among the latter are found urea (1.5 in 1000), fatty acids, sodium chloride, and other salts. In diseased conditions of the kidneys the urea may be greatly increased, the skin supple- menting to a certain extent deficiencies of those organs. The Nervous and Circulatory Factors in the Sweat Secretion. It used to be believed that an increased flow ot blood through the skin would suffice of itself to cause in- creased perspiration; but against this view are the facts that, in terror for example, there may be profuse sweating with a cold pallid skin; and that in many febrile states the skin may be hot and its vessels full of blood, and yet there may be no sweating. Eecent experiments show that the secretory activity of the sweat-glands is under the direct control of nerve-fibres, and is only indirectly dependent on the blood-supply in their neighborhood. Stimulating the sciatic nerve of the freshly amputated leg of a cat will cause the balls of its feet to sweat, although there is no blood flowing through the limb. On the other hand, if the sciatic nerve be cut, so as to paralyze it, in a living animal, the skin arteries di- late and the foot gets more blood and becomes warmer; but it does not sweat. The sweat-fibres originate in certain 430 THE HUMAN BODY. sweat-centres in the spinal cord, which may either be di- rectly excited by blood of a higher temperature than usual flowing through thcni or, reflexly, by Avarmth acting on the exterior of the Body and stimulating the sensory nerves there. Both of these agencies commonly also excite the vaso-dilator nerves of the sweating part, and so the increased blood-supply goes along with the secretion; but the two phenomena are fundamentally independent. The Sebaceous Secretion. This is oily, semifluid, and of a sjjecial odor. It contains about 50 per cent of fats (olein and palmatin). It lubricates the hairs and usually renders them glossy, even in persons who use none of the various compounds sold as "hair-oil." No doubt, too, it gets S})read more or less over the skin and makes the cuticle less permeable by water. Water poured on a healthy skin does not wet it readily but runs off it, as ''off a duck's back" though to a less marked degree. Hygiene of the Skin. The sebaceous secretion, and the solid residue left by evaporating sweat, constantly form a solid film over the skin, which must tend to choke up the mouths of the sweat-glands (the so-called "pores" of the skin) and impede their activity. Hence the value to health of keeping the skin clean: a daily bath should be taken by every one. Women cannot well wash their hair daily as it takes so long to dry, but a man should immerse his head when he takes his bath. As a general rule, soap should only be used occasionally; it is quite unnecessary for cleanliness, except on exposed parts of the Body, if frequent bathing is a habit and the skin be well rubbed afterwards until dry. Soap nearly always contains an excess of alkali which in itself injures some skins, and, besides, is apt to com- bine chemically with the sebaceous secretion and carry it too freely away. Persons whose skin will not stand soap can find a good substitute, for washing the hands and face, in a little cornmeal. ISTo doubt many folk go about in very good health with very little washing ; contact with the clothes and other external objects keeps its excretions from accumulating on the skin to any very great extent. But apart from the duty of personal cleanliness imposed on BATHII^G. 421 man as a social animal in daily intercourse with others, the mere fact that the healthy Body can manage to get along under unfavorable conditions is no reason for expos- ing it to them. A clogged skin throws more work on the lungs and kidneys than their fair share, and the evil con- sequences may be experienced any day when something else throws another extra strain on them. Animals, a considerable portion of whose skin has been varnished, die within a few hours. This used to be thought due to poisoning by retained ingredients of the sweat. But the real cause of death seems to be an excessive radia- tion of heat from the surface of the body, which the vital oxidative processes cannot keep up with, so the bodily tem- perature falls until it reaches a fatal point, about 20° C. (68° F.) for rabbits. If the animal be packed in raw cotton or kef)t in an atmosphere at a temperature of 30° C. (86° F.) it will not die from the varnishing. Bathing. The general subject of bathing may be con- sidered here. One object of it is that above mentioned, to cleanse the skin; but it is also useful to strengthen and invigorate the whole frame. For strong healthy persons a cold bath is the best, except in extremely severe weather, when the temperature of the water should be raised to 15° C. (about 60° F.), at which it still feels quite cold to the surface. The first effect of a cold bath is to contract all the skin -vessels and make the surface pallid. This is soon followed by a reaction, in which the skin becomes red and congested, and a glow of warmth is felt in it. The proper time to come out is while this reaction lasts, and after emersion it should be promoted by a good rub. If the stay in the cold water be too prolonged the state of reaction passes off, the skin again becomes pallid, and the person probably feels cold, uncomfortable, and depressed all d[iy. Then bathing is injurious instead of beneficial; it lowers instead of stimulating the activities of the Body. Hov/ long a stay in the cold water may be made with benefit, depends greatly on the individual; a vigorous man can bear it and set uj) a healthy reaction after much longer immer- sion than a feeble one; moreover, being used to cold bathing 433 THE HUMAN BODY. renders a longer stay safe, and, of course, the temperature of the water has a great influence: water called "cold'* may vary within very wide limits of temperature, as indi- cated by the thermometer; and the colder it is the shorter is the time which it is wise to remain in it. Persons who in the comparatively warm water of Narragansett dur- ing the summer months stay with benefit and pleasure in the sea, have to content themselves with a single plunge on parts of the coast where the water is colder. The nature of the water has some influence; the salts contained in sea-water stimulate the skin-nerves and pi-omote the afterglow. Many persons who cannot stand a simple cold fresh-water bath take one with benefit when some salines are previously dissolved in the water. The best for this jiurpose are probably those sold in the shops under the name of "sea-salts." It is perfectly safe to bathe when warm, provided the skin is not perspiring profusely, the notion commonly pre- valent to the contrary notwithstanding. On the other hand, no one should enter a cold bath when feeling chilly, or in a depressed vital condition. It is not wise to take a bath immediately after a meal, since the afterglow tends to draw away too much blood from the digestive organs, which are then actively at work. The best time for a long bath is about three hours after breakfast; but for an ordinary daily dip, lasting but a short time, there is no better period than on rising and while still warm from bed. The shower-bath abstracts less heat from the skin than an ordinary cold bath and, at the same time, gives it a greater stimulus: hence it has certain advantages. Persons in feeble health may diminish the shock to the system by raising the temi3erature of the water they bathe in up to any point at which it still feels cool to the skin. Bathing in water which feels warm is not advisable: it tends generally to diminish the vital activity of the Body. Hence Avarm baths should only be taken occasionally and for special purposes. CHAPTER XXVIII. NUTEITION. The Problems of Animal Nutrition. We have in pre- ceding chapters traced certain materials, consisting of foods more or less changed by digestion, into the Body from the alimentary canal, and oxygen into it from the lungs. We have also detected the elements thus taken into the Body in their passage out of it again by lungs, kidneys and skin; and found that for the most part their chemical state was different from that in which they entered; the difference being expressible in general terms by saying that more oxidized forms of matter leave the Body than enter it. We have now to consider what hapj)ens to each food during the journey through the Body: is it changed at all? is it oxidized? if so where? what products does its oxidation give rise to? Is the oxidation direct and complete at once or does it occur in successive steps? Has the food been used first to make part of a living tissue and is this then oxidized; or has it been oxidized without forming part of a living tissue? if so, where? in the blood stream or outside of it? Finally, if the chemical changes undergone in the Body are such as to liberate energy, how has this energy been utilized? to maintain the temperature of the Body or to give rise to muscular work, or for other purposes? This is a long string of questions, the answers to many of which Physiology has still to seek. The Seat of the Oxidations of the Body. According to older views oxidation mainly took place in the blood while flowing through the lungs. Those organs were con- sidered a sort of furnace in which heat was liberated by blood oxidation, and then distributed bv the circulation. 4:24 THE HUMAN BODY. But if this were so the lungs ouglit U> be the hottest part.- of the Body, and the blood leaving them by the pulmonale veins much hotter than that brought to them by the pulmo- nary artery after it had been cooled by warming all the tis- sues; and neither of these things is true. A small amount of heat is liberated when haemoglobin combines with oxygen in the pulmonary capillaries, but the affinities thus satisfied are so feeble that the energy liberated is trivial in amount when compared with that set free when this oxygen subse- quently forms stabler compounds elsewhei-e. It is now, more- over, tolerably certain that hardly any of this latter class of oxidations occurs in the living circulating blood at all; its cells do, no doubt, use up some oxygen and set free some car- bon dioxide; but not enough to be detected by ordinary methods of analysis. The percentage of oxygen liberated in a vacuum by two specimens of the blood of an animal, taken one from an artery near the heart, and the other from a distant one, are practically the same; showing that during the time occupied in flowing two or three feet through an artery the blood uses up no appreciable quantity of its own oxygen; while in the short time occu2:)ied in its brief capillary transit it loses so much oxygen as to become venous. The differ- ence is explained by the fact that the blood gives off oxygen gas through the thin capillary walls to the surrounding tissues; and in them the oxidation takes place. As we have already seen, a freshly excised muscle deprived of blood can still be made to contract; and for some consider- able time if it be the muscle of a cold-blooded animal. During its contraction it evolves large amounts of carbon dioxide, although the resting fresh muscle contains hardly any of that gas. Here Ave have direct evidence of oxidation taking place in a living tissue and in connection with its fuuctional activity; and what is true of a muscle is prob- ably true of all tissues; the oxidations which supply them with energ)' take place withm the living cells themselves. The statement frequently made that the oxygen m the cir- culating blood exists as ozone, rests on no sufficient basis; decomposing haemoglobin does seem to form ozone when exposed to the air, but fresh blood yields no sign of it. SYS TRESES IX THE BODY. 425 Experiments made by adding yarious combustible sub- stances, as sugar, to fresh blood, also fail to prove the oc- currence of any oxidation of such bodies in tbat liquid. Tissu.e-Building and Energy- Yielding Foods. The Human Body, like that of other animals, is, on the whole, chemically destructive; it takes in highly complex sub- stances as food, and eliminates their elements in much simpler compounds, which can again be built up to their original condition by plants. Nevertheless the Body has certain constructive powers; it, at least, builds up protoplasm from proteids and other substances received from the exterior; and there is reason to believe it does a good deal more of the same kind of work, though never an amount equaling its chemical destructions. Given one single pro- teid in its food, say egg albumen, the Body can do very well; making serum albumen and fibrin factors out of it for the blood, myosin for the muscles, and so on: in such cases the original proteid must have been taken more or less to pieces, remodeled, and built up again by the living tissues; and it is extremely doubtful if anything different occurs in other cases, when the proteid eaten happens to be one found in the Body. In fact, during digestion the pro- teids are broken down somewhat, and turned into peptones; in this state they enter the blood and must again be built up into proteids, either there or in the solid tissues. The constructive powers of the Body used to be rather too much ignored. Foods were divided into assimilahle and combustible, the former serviug directly to renew the organs or tissues as they were used up, or to supply materials for growth; these were mainly proteids and fats; no special chemical synthesis was thus supposed to take place, the living cells being nourished by the reception from outside of molecules similar to those they had lost. Fat-cells grew by picking up fatty molecules, like their contents, received from the food; and protoplasmic tissues by the reception of ready-made proteid molecules, needing no further manufacture in the cell. The combustible foods, on the other hand, were the carbohydrates and some fats: these, according to the Inqiothesis, were incapable of being made 4:■^6 THE HUMAN BODY. into parts of a living tissue, and were simply burnt at once in order to maintain the bodily warmth. It having been proved, however, that more fat might accumulate in the body of an animal than was taken in its food, this excess was accounted for by supposing it was due to excess of com- bustible foods, converted into fats and stored away as oil- droplets in various cells; but not actually built up into true living adipose tissue. Liebig, somewhat similarly, classed all foods into plastic, concerned in making new tissues, and respiratory, directly oxidized before they ever constituted a tissue. The plastic foods were the proteids, but these also indirectly gave rise to the energy expended in muscu- lar work, and to some heat: the proteid muscular fibre being broken first into a highly nitrogenous part (urea, or some body well on the road to become urea) and a noii- nitrogenized richly hydrocarbonous part; and this latter was then oxidized and gave rise to heat. Several facts may be urged against this view — (1) Men in tropical climates live mainly on non-jDroteid foods, yet their chief needs are not heat production, but tissue formation and muscular work : according to Liebig's view their diet should be mainly nitrogenous. (2) Carnivorous animals live on a diet very rich in proteids, nevertheless develop plenty of animal heat, and that without doing the excessive muscu- lar work which, on Liebig's theory, must first be gone through in order to break vq) the proteids, with the jiroduc- tion of a non-azotized part which could then be oxidized for heat-jDroduction. (3) Great muscular work can be done on a diet poor in proteids; beasts of burden are for the most part herbivorous. (4) Further, we know exactly how much energy can be liberated by the oxidation of proteids to that stage which occurs in the Body; and it is perfectly possible to estimate pretty accurately the amount of urea and uric acid excreted in a given time; from their sum the amount of proteid oxidized and the amount of energy liberated in that oxidation can be calculated; if this be done it is found that, nearly always, the muscular work done during the same period represents far more energy expended than could be yielded by the proteids broken down. SOURCE OF MUSCULAR WORK. 427 The Source of the Energy Expended in Muscular Work. This question, which was postponed in tlie chap- ters dealing with the muscular tissues, on account of its importance demands here a discussion. It may be put thus : — Does a muscle-fibre work by the oxidation of its proteids, i.e. by breaking them down into compounds which are then removed from it and conveyed out of the Body? or does it work by the energy liberated by the oxidation of carbon and hydrogen compounds only? The problem mav be attacked in two ways; first, by examining the excretions of a man, or other animal, during work and rest; second, by examining directly the chemical changes produced in a muscle when it contracts. Both methods point to the same conclusion, viz. that proteid oxidation is not the source of the mechanical energy expended by the Body. One gram (15.5 grains) of pure albumen when completely burnt liberates, as heat, an amount of energy equal to 2117 kilogrammeters (15,270 foot-pounds). But in the Body proteids are not fully oxidized; part of their carbon is, to form carbon dioxide, and part of the hydrogen, to form water; but some carbon and hydrogen pass out, combined with nitrogen and oxygen, in the incompletely oxidized state of urea. Therefore all of the energy theoretically ob- tainable is not derived from proteids in the Body: from the above full amount for each gram of proteid we must take the quantity carried off in the urea, which will be the amount liberated when that urea is completely oxidized. Each gram (15.5 grains) of proteid oxidized in the Body gives ^ of a gram (5.14 grains) of urea; and since one gram of urea liberates, on oxidation, energy amounting to 934 kilo- grammeters (6740 foot-pounds) each gram of proteid oxidized, so far as is possible in the Body, yields during the proccess 2117- ^^ = 1805.7 kilogrammeters (13..037 foot- pounds) of energy. Knowing that urea carries off practi- cally all the nitrogen of proteids broken up in the Body, and contains 46.6 per cent of nitrogen, while proteids contain 16 per cent, it is easy to find that each gram of urea repre- sents the decomposition of about 2.80 grams of proteid and, therefore, the liberation of 5060.00 kilogrammeters (36,533.0 438 THE nUMAN BODY. foot-ponnds) of cnerg}'. If, therefore, we know how much urea a man excretes during a given time, and iiow much mechanical work lie does during the same time, we can readily discoyer if the latter could possibly have been done by the energy set free by proteid decomposition. Let us take a sj^ecial case. Fick and Wisleccnus, two German observers, climbed the Faulhorn mountain, which is 1956 meters (about 6415 feet) high. Fick weighed 06 kilograms and, therefore, in lifting his Body alone, did during the ascent 129,096 kilogrammeters (932,073 foot-pounds) of work. "Wislecenus, who weighed 76 kilograms, did similarly 148,656 kilogrameters (1,073,296 foot-pounds) of work. But during the ascent, and for five hours afterwards, Fick secreted urine containing urea answering only to 37.17 grams of proteid, and Wislecenus urea answering to 37 grams. Since each gram of jjroteid broken up in the Body liberates 1805.7 kilogrammeters (13,037 foot-pounds) of energy, the amount that Fick could possibly have obtained from such a source is 1805.7 X 37.17 = 07,117 kilogram- meters (484,584 foot-pounds), and Wislecenus 1805.7 X 37 = 66,810 kilogrammeters. If to the muscular work done in actually raising their bodies, we add that done simul- taneously by the heart and the respiratory muscles, and in such movements of the limbs as were not actually concerned in litiug their weight, we should have, at least, to double the above total muscular Avork done; and the amount of energy liberated meanwhile by proteid oxidation, becomes utterly inadequate for its execution. It is thus clear that \ muscular work is not wholly done at the expense of the 1 oxidation of muscle proteid, and it is very probable that I none is so done under ordinary circumstances, for the urea excretion during rest is about as great as that during work, if the diet remain the same; if the work is very | violent, as in long-distance walking matches, the urea quan- 1 , tity is sometimes temporarily raised but this increase, / which no doubt represents an abnormal wear and tear of muscle-fibre, is probably independent of the liberation of energy in the form in which a muscle can use it, more likely taking the form of heat; and is, moreover, compen- UltEA AXD MUSCULAR WOBK. 429 sated for afterwards by a diminished urea excretion. Thus, hourly, before the ascent Fick and Wislecenus each ex- creted on the average about 4 grams (63 grains) of urea; during the ascent between 7 and 8 grams (108-124 grains); but during the subsequent 16 hours, when any urea formed in the work would certainly have reached the urine, only an average of about 3 grams (46. 5 grains) per hour. It may still be objected, however, that a good deal of the muscle work may be done by the energy of oxidized muscle proteid; that the amount of this oxidation occurring in a muscle during rest or ordinary work is pretty constant and simply takes different forms in the two cases, much as a steam-engine with its furnance in full blast will burn as much coal when working as when resting, but in the former case lose all the energy generated in the form of heat, and in the latter partly as mechanical work. Thus the want of increase in urea during muscular activity would be explained, while still a good deal of utilizable energy might come from proteid degradation. But if this were so, then the work- ing Body should eliminate no more carbon dioxide than the resting; the amount of chemical changes in its muscles being by hypothesis the same, the carbon dioxide eliminated should not be increased. Experiment, however, shows that it is, and that to a very large extent, even when the work done is quite moderate and falls within the limits which could be performed by the normal proteid degrada- tion of the Body. Quite easy work doubles the carbon dioxide excreted in twenty-four hours, and in a short period of very hard work it may rise to five times the amount eliminated during rest. Since the urea is not increased, or but very slightly increased, at the same time, this carbon dioxide cannot be due to increased joroteid metamorphosis; and it therefore indicates that a muscle works by the oxidation of carbonaceous non-nitrogenous compounds. Since all the carbon compounds oxidized in the Body contain hydro- gen this element is also no doubt oxidized during muscu- lar work; but its estimation is difficult and has not been attempted, because the Body contains so much water ready formed that a large -quantity is always ready for increased 430 TEE HUMAN BODY. evaporation from the lungs and skin, whenever the respira- tions are quickened, as they are by exercise. It, thus, is very difficult to say how much of the extra water eliminated from the Body during work is due merely to this cause and how much to increased hydrogen oxidation. The conclusion we are led to is, then, that a muscle works hy the oxidation mainlj^ if not entirely, of carbon and hydrogen; much as a steam-engine does: the j)roteid con- stituents of the muscle answer roughly to the metallic parts of the engine, to the machinery using the energy liberated by the oxidations, but in itself only suffering wear and tear bearing no direct proportion to the work done; as an engine may rust, so muscle proteid may and does oxidize, but not to supply the organ with energy for use. This conclusion, arrived at by a study of the excretions of the whole Body, is confirmed by the results obtained by the chemical study of a single muscle. A fresh frog's muscle (which agrees in all essential points with a man's) contains practically no car- bon dioxide, yet, made to work in a vacuum, gives off that gas, and more the more it works. Some carbon dioxide is therefore formed in the working muscle. If the muscle, after contracting as long as it will, be thrown into death rigor it gives off more carbon dioxide; and if taken per- fectly fresh and sent into rigor mortis without contracting it gives off carbon dioxide also, in amount exactly equal to the sum of that which it would have given off in two stages, if first worked and then sent into rigor. The muscle must, therefore, contain a certain store of a carbon-dioxide-yield- ing body, and the decomjiosition of this is associated with the occurrence both of muscular activity and death stiffen- ing. Similar things are true of the acid simultaneously developed ; the muscle when it works produces some sarco- lactic acid, and when thrown into rigor mortis still more. N"o increase of urea or kreatin or any similar product of nitrogenous decomposition is found in a worked muscle when compared with a rested one, but the total carbohy- drates are rather less in the former. These facts make it clear that muscular work is not done at the expense of proteid oxidation; and we have already seen {]). 387) that CHEMISTRY OF WORKING MUSCLE. 431 the oxygen a muscle uses in contracting is not taken up by it at the time it is used, since a muscle containing no oxy- gen will still contract and evolve carbon dioxide in a vacuum. It is probable that the chemical phenomena occurring in con- traction and rigor are essentially the same; the death stiffen- ing results when they occur to an extreme degi'ee. Pro- visionally one may explain the facts as follows — A muscle in the Body takes up from the blood, oxygen, proteids, and non-nitrogenous (carbohydrate or fatty) substances. These it builds up into a highly complex and very unstable com- pound, comparable, for example, to nitro-glycerine. When the muscle is stimulated this falls down into simpler sub- stances in which stronger affinities are satisfied; among these are carbon dioxide and sarcolactic acid and a proteid (myosin). The energy liberated is thus independent of any simultaneous taking up of oxygen; the amount possible depends only on how much of the decomposable body exists in the muscle. Under natural conditions the carbon dioxide is carried off in the blood and perhaps the sarco- lactic acid also, the latter to be elsewhere oxidized further to form water and more carbon dioxide. The myosin remains in the muscle-fibre and is combined with more oxygen, and compounds of carbon and hydrogen taken from the blood, and built up into the unstable energy-yielding body again; none of it, under ordinary circumstances, leaves the muscle. If, however, the blood-supply be defi- cient, the myosin clots (p. 125) before this restitution takes place and it cannot then be rebuilt; and in excessive work the same thing partially occurs, the decomposition occur- ring faster than the recomposition; the clotted myosin is then broken up into simpler bodies and yields a certain mcrease of the urea excreted. In rigor mortis all the myosin passes into the clotted state and causes the rigidity. A working muscle takes up more oxygen from the blood than a resting one, as is shown by a comparison of the gases of its venous blood in the two cases; this oxygen assumption is not necessarily proportionate to the carbon-dioxide elimina- tion at the same time; for the latter depends on the break- ing down of a body already accumulated in the muscle dur- 432 THE HUMAN BODY. ing rest, and this breaking down may occur faster than the reconstruction. We are thus enabled, also, to understand how, during exercise, the carbon dioxide evolved from the lungs may contain more oxygen than that taken up at the same time; for it is largely oxygen previously stored dur- ing rest, that then appears in the carbon dioxide of the ex- pired air. Are any Foods Respiratory in Liebig's Sense of the Term? AVo find, then, tliut Liebig's classification of food.'i cannot be accepted in an absolute sense. There is no doubt that the substance broken down in muscular contraction is proper living muscular tissue; and if this (its proteid con- stituent being retained) be reconstructed from foods con- taining no nitrogen (whether carbohydrates or fats) then the term plastic or tissue-forming cannot be restricted to the proteids of the diet. We must rather conclude that any alimentary princi^ile containing carbon may be used to replace the oxidized carbon, and any containing hydrogen to replace the oxidized hydrogen, of a tissue; and so even non-proteid foods may be plastic. A certain proportion of the foods digested may i:)erhai)s be oxidized to yield energy, before they ever form part of a tissue; and so correspond jiretty much to Liebig's respiratory foods; but no hard and fast line can be drawn, making all proteid foods jolastic and all oxidizaljle non-proteid foods respiratory. Luxus Consumption. Not only, as above pointed out, may non-nitrogenous foods be ])lastic, but it is certain that if any foods are oxidized at once before being organized into ji tissue, proteids are under certain circumstances; namely, when -they are contained in excess in a diet. If an tinimal be starved it is found that its non-nitrogenous tissues go first; an insufficiently fed animal loses its fat first, and if it ultimately dies of starvation, is found to have lost 97 per cent of its adipose tissue and only about 30 per cent of its proteid-rich muscular tissue, and almost none of its brain and spinal cord; all of course reckoned by their dry weight. It is thus clear that the proteids of the tissues resist oxida- tion much better than fat does. But, on the other hand, if a well-fed animal be given a very rich proteid diet all the SOURCES OF UREA. 433 nitrogen of its food reappears in its urine, and that when it is laying up fat; so that then we get a state of things in which proteids are broken up more easily than fats. This indicates that proteid in the Body may exist under two conditions; one, when it forms part of a living tissue and is protected to a great extent from oxidation, and another, in which it is oxidized with readiness and is presumably in ;! different condition from the first, and not yet built up into part of a living cell. The use of proteids for direct oxidation is known as luxus consumption ; how far ic occurs under ordinary circumstances will be considered presently. The main point now to be borne in mind is that while all organic non-nitrogenous foods cannot be called respiratory, neither can proteids under all circumstances be called ^j/as/ic, in Liebig's sense. The Antecedents of Urea. In the long run the pro- genitors of the urea excreted from the Body are the proteids taken in the food; but it remains still to be considered what intermediate steps these take before excretion in the urine; and whether urea itself is finally formed in the kid- neys or merely separated by them from the blood. In seeking antecedents of urea one naturally turns first to the muscles, which form by far the largest mass of pro- teid tissues in the Body. Analysis shows that they always contain kreatin, a body intermediate chemically between proteids and urea. The quantity of this in muscles is prac- tically unaffected by work, and is from 0.2 to 0.4 per cent. Since it is readily soluble and dialyzable, and therefore fit- ted to pass rapidly out of the muscles into the blood stream, it is a fair conclusion that a good deal of it is formed in the muscles daily and carried off from them. Kreatin, too, exists in the brain, and probably there, and elsewhere in the nervous system, is produced by chemical degrada- tion of protoplasm; the spleen also contains a good deal of kreatin, and so do many glands. This substance would therefore seem to be constantly produced in considerable quantities by the protoplasmic tissues generally; and since it belongs to a group of nitrogenous compounds which the Body is unable to utilize for reconstruction into proteids. 434 THE HUMAN BODY. it must be carried off somehow. The urine, however, con- tains very little kreatin, or its immediate derivative, krea- tinin, and what it does contain depends mainly on the feeding, since it varies with the diet and vanishes during starvation; so it is probable that this substance is con- verted into urea and excreted in that form. This conver- sion must occur elsewhere than in the muscles, which con- tain no urea; also, very little, if any, exists in the brain. Where the kreatin is finally changed into urea is doubt- ful. It may be in the kidneys by the renal epithelium, or it may be elsewhere, and the urea produced be merely picked up from the blood and passed out by the kidney- cells; or both may occur; histologically the distinctly secretory epitheliums of the convoluted parts of the tubules and of Henle's loops, differ so much as to suggest an en- tirely different function for them. On the whole, the evidence seems to show that urea is merely separated and not produced in the kidneys; a priori this is more probable, since in the degradation of kreatin to yield urea energy is liberated and this might very well be utilized in some organ; while if the process took place in the kidney tubules the force set free would be wasted. The blood always contains urea, and renal-artery blood aj)parently more than renal- vein blood, which shows that urea is removed from the blood in the kidneys. Moreover, if a mammal's kidneys be extirpated urea accumulates in its blood, which could not be the case if urea were nor- mally only produced in the kidneys; and if urea be injected into a vein it is rapidly picked up and carried off in the urine, showing that the kidney-cells have a selective power with respect to it. While the urea resulting from further changes in the kreatin formed in the tissues is a measure of the wear and tear of their protoplasm, part of the urea excreted has probably a different source; being due to the oxidation of proteids, as energy liberators or respiratory foods, before they have ever formed a tissue. When plenty of proteid food is taken the urea excretion is largely increased and ihat very rapidly, within a couple of hours for example, LUXUS CONSUMPTION. 435 and before we can well suppose the proteids eaten to have been built up into tissues, and these in turn broken down; in fact there need be, and usually is, under such circumstances no sign of any special activity of any group of tissues, such as one would expect to see if the urea always came from the breaking down of formed histological elements. This urea is thus indicative of a utilization of proteids for other than plastic purposes; and the same fact is indicated by the storage of carbon and elimination of all the nitrogen of the food (p. 444) when a diet very rich in proteid alimentary principles is taken. This hixus consumption may be com- pared to the paying out of gold by a merchant instead of greenbacks when he has an abundance of both. Only the gold can be used for certain purposes, as settling foreign debts, but any quantity above that needed for such a pur- pose is harder to store than the paper money and not so convenient to handle; so it is paid out in preference to the paper money, which is really somewhat less valuable, as available at par only for the settlement of domestic debts. In artificial pancreatic digestions, when long carried on, two bodies, called lencin and tyrosin, are produced from proteids. It is found that when leucin is given to an animal in its food it reappears in the urine as urea; so the Body can turn leucin into that substance. Hence a possible source of some of the luxus-consumjition urea is leucin produced during intestinal digestion; and this is very likely turned into urea in the liver. At any rate the liver, to which the portal vein might carry all leucin thus formed, contains urea, which no other gland does; and when the liver is greatly altered, as in phosphorus poison- ing and the disease known as acute yellow atrophy, urea almost entirely disappears from the urine. This latter fact seems to point to a final production of urea in the liver, what- ever its immediate antecedents may be; whether muscle kreatin, or intestinal leucin, or excess of peptones in the diet. The latter might perhaps be broken up there into a nitrogenous part (urea) and a non-nitrogenous part; and we shall find that a non-nitrogenous substance (glycogen) is stored in the liver. 436 THE HUMAN BODY. Proteid Starvation and Overfeeding. When an ani- mal is fed on food deficient in proteids, or containing none of them at all, its urea exci'etion falls very rapidly during the first day or two, but then much more slowly until death: there is thus indicated a double source of urea, a part resulting from tissue wear and tear, and always present; and a part resulting from the breaking down of proteids not built up into tissue, and ceasing when the amount of this 2)roteid in the Body (in tlie blood for example) falls below a certain limit as a result of the starvation. As tlie nitrogen-starved Body wastes, its bulk of proteid tissues is slowdy reduced and the urea I'esulting from their degrad- ation diminishes also. How well jirotcid built up into a tissue resists removal is shown by the facts already men- tioned (p. 432) as to the relative losses of the proteid-rich and proteid-poor tissues in starvation. On tlie other hand, if an animal be taken while starving and losing weight and have a small amount of flesh giA^en it, it will continue to lose weight, and more urea than before will appear in the urine; increased proteid diet in- creases the proteid metamorphosis, and the animal still loses, though less rapidly than it did. A little more proteid still increases proteid metamorphosis in the body, and the urea elimination, and so on for some time; but each increment of proteid in the food increases the nitrogcnou^' metamorphosis somewhat less than the last did, until, finally, a point is reached at which the nitrogeii egesta and ingesta balance: in a dog this occurs when it gets daily gijj- its weight of meat, and no other solid food. More food if then given is at first stored up and the annual increases in Aveight; but very soon the greater wear and tear of the larger mass of tissues shows itself as increased urea ex- cretion; again the egesta and ingesta balance, and the ani- mal comes to a new weight equilibrium at the higher level. More meat now causes a repetition of the phenomenon: at first increase of tissue, and nitrogen storage; and then a cessation of the gain in weight, and an excretion m twenty- four hours of all the nitrogen taken. And so on, until the animal refuses to eat any more. STORAGE TISSUES. 43? These facts seem, very clearly, to show that proteids can- not be built up quickly into tissues. Meat given to the starving animal has its proteids, at first, used up mainly in luxus cofisumption — while a little is stored as tissue, though at first not enough to counterbalance the daily tissue waste. When a good deal more proteid is given than answers to the nitrogen excretion during starvation, the animal builds up as much into living tissues as it breaks down in the vital vrocessesot these, therest going ml H.rus consnmpf ion ; it thus neither gains nor loses. More proteid does not all appear in the urine at once; some is used to build up new tissue, but only slowly; then, after some days, the increased metabolism of the increased flesh balances the excess of nitrogen in the diet, and equilibrium is again attained. But, all through, it seems clear that the tissue formation is slow and gradual; and so it becomes additionally probable that the increased urea excretion soon after a meal is not due to rapidly in- creased tissue formation and degradation, but to a more direct proteid oxidation. The Storage Tissues. Every healthy cell of the Body contains at any moment some little excess of material laid by in itself, above what is required for its immediate neces- sities. The capacity of contracting, and the concomitant evolution of carbon dioxide, exhibited by an excised muscle in a vacuum, seem to show that even oxygen, of which warm-blooded animals have but a small reserve, may be stored up in the living tissues in such forms that they can utilize it, even when the air-pump fails to extract any from them. But in addition to the supplies for immediate spend- ing, contained in all the cells, we find special food reserves in the Body, on which any of the tissues can call at need. These, especially the oxygen and proteid reserves, are found largely in the blood. Special oxygen storage is, however, rendered unnecessary by the fact that the Body can, except under very unusual circumstances, get more from the air at any time, so the quantity of this substance laid by is only small; hence death from asphyxia follows very rapidly when the air-passages are stopped, while, on account of the reserves laid up, death from other forms of starva- 438 THE HUMAN BODY. tion is a much slower occurrence. Proteids, also, we have learnt from the study of muscle, are probably but little con- cerned in energy-production in the tissues. Speaking broadly, the work of the Body is carried on by the oxidation of carbon and hydrogen, and we find in the Body, in corre- spondence with this fact, two great storehouses of fatty and carl)ohydrate foods, which serve to supply the materials for the performance of work and the maintenance of the bodily temperature in the intervals between meals, and during longer periods of starvation. One such store, that of car- bohydrate material, is found in the liver-cells; the other, or fatty reserve, is found in the adipose tissue. That such substances are true reserves, not for any special local purpose but for the use of the Body generally, is shown by the way they disappear in starvation; the liver reserve in a few days, and the fat somewhat later and more slowly, but very largely before any of the other tissues have been seriously affected. By using these accumulated matters the Body can work and keep warm during several days of more or less deficient feeding; and the fatter an animal is at the beginning of a starvation period the longer will it live; which would not be the case could not its fat be utilized by the working tissues. Hybernating animals prove the same thing; bears, before their winter sleep, are very fat, and at the end of it commonly very thin; while their muscular and nervous systems are not noticeably diminished in mass. During the whole winter, then, the energy needed to keep the heart and respiratory muscles at work, and to maintain the tempera- ture of the body, must have been obtained from the oxida- tion of the fat reserve with which the animal started. Glycogen. It may perhaps have struck the reader as curious that so large an organ as the liver should be set apart for the formation of so comparatively unimportant a digestive secretion as the bile; and were this the sole use of the liver the size of the organ would certainly be anomalous. The main function of the liver is, however, quite a different one, the formation and storage of a carbohydrate called glycogen, from the abundant food materials carried through it by the portal vein after a meal; in the times between GLYCOOEN. 439 meals this substance is then doled out gradually, and sent round tlie Body in the blood. If a liver be cut up two or three hours after removal from the body of a healthy well- fed animal, and thoroughly extracted with water, it will yield up much grape sugar. If, on the other hand, a per- fectly fresh liver be heated rapidly to the temperature of boiling water, and then pounded up and extracted, it will yield a milky solution, containing little grape sugar but much glycogen; a substance which chemically has the same empirical formula as starch (CeHioOs), and in other ways is closely allied to that body. The salivary and pancreatic secretions rapidly convert it into sugar, as they do starch, the elements of a molecule of water being taken up at the same time — CeHioOs + H.2O = CeHiiOe Glycogen. Water. Glucose. The same transformation is rapidly effected by ferments present in the blood and liver, and hence the first thing to be done in preparing glycogen is to heat the organ at once to a temperature high enougli to destroy these ferments. Pure glycogen is a white amorphous inodorous powder, readily soluble in water, forming an opalescent milky solu- tion; insoluble in alcohol, and giving with iodine a red coloration which disappears on heating and reappears on cooling again. About four per cent of glycogen can be obtained from the liver of a well-nourished animal (dog or rabbit). This for the human liver, which weighs about 1500 grams (53 oz.), would give about 60 grams (3.1 oz.) of glycogen at any one moment. The quantity actually formed daily is, however, much in excess of that, since glycogen is constantly being removed from the liver and carried elsewhere, while a fresh supply is formed in the organ. Its quantity is subject, also, to considerable fluctuations; being greatest about two hours after a good meal, and falling from that time until the next digestion period commences, when it begins to rise until it reattains its maximum. "When a warm-blooded animal is starved the glycogen entirely disappears from its liver in the course of four or five days. Glycogen is, thus, clearly 440 THE HUMAN BODY. being constantly used up, and its maintenance in normal quantity depends on food. The Source and Destination of Liver Glycogen. All foods arc not equally efficacious in keeping up the stock of glycogen in the liver; fats by themselves ai-e useless; pro- teids by themselves give a little; but by far the most is formed on a diet rich in starch and sugar; so it would seem that glycogen is mainly formed from cai-bohydrate materials absorbed from the alimentary canal and carried to the hepatic cells by the portal vein. These materials are mainly glucose, since the starch eaten is changed into that substance before absorption. This view of' the matter is supported by several facts. (1) Grape sugar if it exist in the blood in above a certain small percentage passes out by the kidneys and appears in the urine, constituting the characteristic symptom of the disease called diahetes. In health, however, even after a meal very rich in carbohy- drates, no sugar appears in the urine; so that the large quantity of it absorbed from the alimentary canal, within a brief time under such circumstances, must be stopped somewhere before it reaches the general blood current. (2) Glucose injected into one of the general veins of an animal, if in any quantity, soon appears in the urine; but the same amount injected into the portal vein, or one of its radicles, causes no diabetes, but an accumulation of glycogen in the liver. We may therefore conclude that the grape sugar absorbed from the alimentary canal is taken by the portal vein to the liver; there stayed and converted into glycogen; which is then more slowly passed on into the hepatic veins during the intervals between meals. Thus in spite of the intervals which elapse between meals the carbo- hydrate content of the blood is kept pretty constant: dur- ing digestion it is not suffered to rise very high, nor dur- ing ordinary periods of fasting to fall very much ])elow the average. In what form glycogen leaves the liver is not certain; it might be dissolved out and carried off as such, or previously turned again into glucose and sent on in that form: since the blood and the liver both seem to contain GLYCOGEN. 441 ferments capable of changing glycogen into^^cose the latter view is the more probable. Analj'ses'of portal and hepatic bloods, made with the yiew of determming whether more sugar was carried out of the liver during fasting than into it, are conflicting. The main fact, how- ever, remains that somehow this carbohydrate reserve in the liver is steadily carried off to be used elsewhere: and animal glycogen thus answers pretty much to vegetable starch, which, made in the green leaves, is dissolved and carried away by the sap currents to distant and not green parts (as the grains of corn or tubers of a potato, which cannot make starch for themselves) and in them is again laid down in the form of solid starch grains, which are subsequently dissolved and used for the growth of tlie ger- minating seed or potato. Reasons have already been given (p. 423) for believing that the carbohydrate leaving the liver is not oxidized in the blood, but first after it has jiassed out of that into a living tissue. Among these the muscles at least seem to get some, since a fresh muscle always contains glycogen, and even in normal amount when an animal is starved for some time; the muscle-fibres then, so to sj^eak, calling on the balance with their banker (the liver) so long as there is any. When a muscle contracts this glycogen disappears and some glucose appears, but not an amount equivalent to the glycogen used up; so that the working muscle would appear, probably for its repair after each con- traction (see p. 431), to utilize this substance. How it is that the glycogen, which is so rapidly con- verted into grape sugar by the liver ferment after death, escapes such rapid conversion during life has not been satisfactorily answered. Two possible reasons readily sug- gest themselves; the liver ferment may be only produced by dying hepatic cells; or the glycogen in the living cell may not exist free, but combined with other portions of the cell substance so as to be protected; while, after death, post-mortem changes may rapidly liberate it in a condition to be acted upon by the ferment. Diabetes. The study of this disease throws some light upon the history of glycogen. Two distinct varieties of it 442 THE HUMAN BODY. arc known; one in which sngar appears in the urine only when the patient takes carbohydrate foods; the other in which it is still excreted when he takes no such foods, and must therefore form sugar in his Body from substances not at all chemically allied to it. The most probable source of the sugar in the latter case is proteids; since some glycogen is found in the livers of animals fed on proteids only, while fats alone give none of it. In some complex way the proteid mole- cule would appear to split up in the liver into a highly nitro- genized part (urea or an antecedent of urea) and a non- azotized part, glycogen. On this view the more severe form of diabetes would be due to an increased activity of a normal proteid-decomposing function of the hepatic cells; and sometimes the urea and sugar in the urine of diabetics rise and fall together, thus seeming to indicate a com- munity of origin. Diabetes dependent on carbohydrate food might be produced in several ways. The liver-cells might cease to stop the sugar and, letting it all pass on into tlie general circulation, suffer it to rise to such a percentage in the blood after a meal, that it attained the proportion in which the kidneys pass it out; or the tissues might cease to use their natural amount of sugar, and this, sent on steadily out of the liver, at last rise in the blood to the point of ex- cretion. Or the liver might transform (into glucose) and pass on its glycogen faster than the other tissues used it. and so diabetes might arise; but this would only be tem- porary, lasting until the liver stock was used up by the rapid conversion. Artificially we can, in fact, produce diabetes in several of these ways; curari poisoning, for ex- ample, joaralyzing the motor nerves, makes the skeletal muscles lie completely at rest, and so diminishes the glyco- gen consumption of the Body and produces diabetes. Car- bon monoxide poisoning produces diabetes also, presumably l)y checking bodily oxidation. Finally, pricking a certain spot in the medulla oblongata causes a temporary diabetes. This may be due to the fact that the operation injures that part of the vaso-motor centre which controls the mus- cular coat of the hepatic artery; this artery, then dilating, carries so much blood through the liver that an excess of FATS. 443 glycogen is turned into glucose in a given time, and carried off by the hepatic veins. If the sj^lanchnic nerves be cut the whole arteries of the abdominal viscera dilate and no diabetes follows, because so many vessels being dilated a great par- of the blood of the Body accumulates in them, and there is no noticeably increased flow through the liver. Others, however, maintain that the " j^iqfire" diabetes (as that due to pricking the medulla is called) is due to irritation of trophic nerve-fibres originating there, and governing the rate at which the liver-cells produce glycogen or convert it into glucose. This latter view, though perhaps the less commonly accepted, is probably the more correct. The hepatic cells do not merely hold back glucose carried through the liver so that it is there to be washed out by a greater blood-flow, but they feed on glucose and proteids and make glycogen; and this is later converted into glucose and carried off. Glycogen is thus comparable to the zymo- gen of the pancreas and other glands (Cluq). XVIII. ); and the transformation of such bodies into the specific element of a secretion we have already seen to be directly under the control of the nervous system, and almost entirely or quite independent of the blood-flow. The History of Fats. "While glycogen forms a reserve store of material that is subject to rapid alterations, deter- mined by meal-times, the fats are much more stable; their periods of fluctuation are regulated by days, weeks, or months of good or bad nutrition, and during starvation they are not so readily, or at least so rapidly, called upon as the hepatic glycogen. If we carry on the simile by which we compared the reserve in each cell to pocket-money (p. 31), tlie glycogen would answer somewhat to a balance on the right side with a man's banker ; while the fat would represent assets or securities not so rapidly realizable; as capital in business, or the cargoes afloat in the argosies of Antonio, the "Merchant of Venice." Fat, in fact, is slowly laid down in fat-cells and surrounded in these by a cell-wall, and, being itself insoluble in blood plasma or lymph, it must undergo chemical changes, which no doubt 444 THE HUMAN BODY. require some time, before it cuu be taken into the blood and carried off to other parts. When adipose tissue is developing it is seen that undif- ferentiated cells in the connective tissues (especially areolar) show minute oil-drops in their protoplasms. These increase in size and, ultimately, fuse together and form one larger oil-droplet, while most of the original protoplasm dis- ap])cars. The oily matter would thus seem due to a chemical metamorphosis of the cell protoplasm, during which it gives rise to a non-azotized fatty residue which remains behind, and a highly nitrogenous part which is carried off. In many parts of the Body protoi)lasmic masses are subject to a similar but less complete metamorphosis; fatty degenera- tion of the heart, for example, is a more or less extensive replacement of the proper substance of its muscular fibres by fat-droplets; and the cream of milk and the oily matter of the sebaceous secretion are due to a similar fatty degeneration in gland-cells. Moreover, careful feeding ex- periments undoubtedly show that fat can come from pro- teids; when an animal is very richly supplied with these all the nitrogen taken in them reappears in its execretions, but all the carbon does not; it is in part stored in the Body: and, since such feeding produces but little glycogen, this carbon can only be stored as fat. While there is, then, no doubt that some fat may have a proteid origin, it is not certain that all has such. During digestion a great deal of fat is ordinarily absorbed, in a chemically unchanged state, from the alimentary canal; it is merely emulsified and carried off in minute drops by the chyle to be poured into the blood: and this fat might be directly deposited, as such, in adipose tissue. There are, however, good reasons for supposing that all the fat in the Body is manufactured. The fat of a man, of a dog, and of a cat varies in the proportions of palmatin, stearin, margarin, iuid olein in it; and varies in just the same way if all be fed on the same kind of food, which could not be the case if the fat eaten were simply deposited unchanged. Moreover, if an animal be fed on a diet containing one kind of fat only. ORIGIX OF FATS. 445 say olein, but a very slightly increased percentage of that particular fatty substance is found in its adij^ose tissue, which goes to show that if fats come from fats eaten, these latter are first pulled to bits by the living cells and built up again into the forms normal to the animal; so that, even with fatty food, the fats stored up seem to be in most part manufactured in the Body. In still another way it is proved that fats can be constructed in the Body. In animals fed for slaughter, the total fat stored up in them during the process is greatly in excess of that taken with their food during the same time. For ex- ample, a fattening pig may store up nearly five hundred parts of fat for every hundred in its food, and this fat must be made from proteids or carbohydrates. Whether it can come from the latter is still perhaps an open question; for, while all fattening foods are rich in starch or similar bodies, there are considerable chemical difficulties in sup- / posing an origin of fats from such; and it is on the whole I more probable that they simply act by sparing from use fats simultaneously formed or stored in the body, and which would have otherwise been called upon. They make glycogen, and this shelters the fats. Liebig, indeed, in a very celebrated discussion, maintained that fats were formed from carbohydrates. He showed that a cow gave out more butter in its milk than it received fats in its food; and Huber, the blind naturalist, showed that bees still made wax (a fatty body) for a time when fed on pure sugar; and indefinitely when fed on honey. Consequently, for a long time, an origin of fats from carbohydrates was supposed to be proved; but their possible origin from proteids (a possi- bility now shown to be a certainty) was neglected, and the validity of the above proofs of their carbohydrate origin is thus upset. The cow may have made its butter from proteids; the bees, fed on sugar, their wax for a time from proteids in their bodies already; and, indefinitely, when fed on lioney, from the proteids in that substance. Moreover, animals (ducks) fed on abundant rice, which contains much carbo- hydrate but very little proteid or fat, remain lean; while if some fat be added they lay up fat. 446 THE HUMAN BODY. Persons who fatten cattle for the butcher find that the foods useful for the purpose all contain proteids, carbohy- drates, and fats, and that rapid fattening is only obtained with foods containing a good deal of fat; as oilcake, milk, or Indian corn. Taking all the facts into account we shall probably not be wrong in concluding that nearly all the bodily fat is manufactured either from fats or proteid.--; from fats easier than from anything else, but when much proteid is eaten some is made from it also. Carbohydrates alone do not fatten; the animal body cannot make its pal- matin, etc., out of them. Nevertheless they are, indirectly, important fattening foods when given with others, since, being oxidized instead of it, they protect the fat formed. Dietetics. That ^' one man's meat may be another man's poison" is a familiar saying, and one that, no doubt, expresses a certain amount of truth; but the difference probably depends on the varying digestive powers of indi- viduals rather than on peculiarities in their laws of cell nutrition: all need pretty much the same amount of pro- teids, fats, and carbohydrates for each kilogram of body weight; but all cannot digest the same varieties of them equally well : while many foods have peculiar, almost poisonous, effects on some persons. A good many people are made ill by mutton, which the majority digest better than beef. The proper diet, too, will necessarily vary, at least as to amount,with the work done; whether it should vary in kind with the nature of the work is not so certain. Provided a man gets enough proteids to balance those lost in the wear and tear of his tissues, it probably matters little whether he gets for oxidation and the liberation of energy either fats or car- bohydrates, or even excess of proteids themselves; any one of the three will allow him to work either his brain or his muscles, and to maintain his temperature. Proteids, however, are wasteful foods for mere energy-yielding23uri:)oses: in the first place, they are more costly than the others; secondly, they are incompletely oxidized in the Body; and, thirdly, it is probably more laborious to the system to get rid of urea than of the carbon dioxide and water, which alone are yielded by DIETETICS. 447 the oxidation of fats arid carbohydrates. Between fats and carboliydrates simihir considerations lead to a use of the latter when practicable: starch is more easily utilized in the Body than fats, as shown by the manner in which it protects the latter from oxidation; and a given weight of starch fully oxidized in the Body will liberate one and a half times as much energy as the same amount of butter, while it costs consider- * • \ ably less than half the money. Probably, too, starch is . •., more easily digested than fats; at least by many people: - "^ children especially are apt to be fond of starchy or sac- charine foods and to loathe fats; and the appetite in such cases is a good guide. As a race, too, the American people differ markedly from the English in their love of sweet foods of all kinds; whether this is correlated with their characteristic activity, calling for some food that can be rapidly used, is an interesting question, to which, however, it would be rash to give at present an affirmative answer. It is clear, therefore, that no general rules for every one's diet can be laid down; but still on broad principles the best diet would be that which contained just the amount of proteid necessary for tissue repair, and so much carbo- hydrates as could be well digested, the balance needed, if any, being made up by fats. Such a food would be the cheapest; that is the supplying of it would call for less of the time and energy of the nation using it, and leave more work to spare for other pursuits than food production — for all the arts which make life agreeable and worth living, and which elevate civilized man above the merely material life of the savage whose time is devoted to catching and eat- ing. We have high authority for saying that man does not live by bread alone; m other words his highest develop- ment is impossible when he is totally absorbed in *' keeping body and soul together," and the more labor that can be spared from getting enough food the better chance has he, if he use his leisure rightly, of becoming a more worthy man. While there is, thus, a theoretically best diet, it is nevertheless impossible to say what that is for each indi- vidual; but what the general experience is may be approxi- mately gathered by taking an average of the dietaries of a 448 THE HUMAN BODY. number of iniblic institutions in which the health of many people is maintained as economically as ])ossible. Such an examination made by Moleschott, gives us as its result a diet containing daily — Proteids /SO grams or 465 grains. Fats 84 " or 1,300 Amyloids 404 " or 6,263 " Salts 30 " or 465 Water.... 2800 " or 43,400 " People in easy circumstances take as a rule more proteids and fats and less amyloids; and this selection, when a choice is possible, probably indicates that such a diet is the better one: the proteids in the above table seem especially deficient. CHAPTER XXIX. THE PRODUCTION AND RECfULATIOX OF THE HEAT OF THE BODY. Cold- and Warm-Blooded Animals. All animals, so long as they are alive, are tiie seat of ehemical changes by "which heat is liberated; hence all tend to be somewhat warmer than their ordinary surroundings, though the difference may not be noticeable unless the heat production is considerable. A frog or a fish is a little hotter than the air or water in which it lives, but not much; the little heat that it pro- duces is lost, by radiation or conduction, almost at once. Hence such animals have no proper temperature of their own; on a warm day they are warm, on a cold day cold, and are accordingly known as cliangeable-temperatured {poihilo-thermous) or,in ordinary language, "'' cold-blooded" animals. Man and other mammals, as well as birds, on the contrary, are the seat of very active chemical changes by which much heat is produced, and so maintain a tolerably uniform temperature of their own, much as a fire does whether it be burning in a warm or a cold room; the heat production at any given time balancing the loss a nor- mal body temperature is maintained, and usually one con- siderably higher than that of the medium in which they live; such animals are therefore known as animals of con- stant temperature {Jiomo-thermous), or more commonly " warm-blooded" animals. The latter name, however, does not properly express the facts; a lizard basking in the sun on a Avarm summer's day may be nearly as hot as a man usually is; but on the cold day the lizard becomes cold, while the average temperature of the healthy Human Body 450 THE HUMAN BODY. is, within a degree, the same in winter or summer; within the iirctic circle or on the equator. Moderate warmth accelerates protoplasmic activity; com- pare a frog dormant in the winter with the same animal active in the warm months: what is true of the whole frog is true of each of its living cells. Its muscles contract more rapidly when warmed, and the white corpuscles of its blood when heated up to the temperature of the Human Body are seen (with the microscope) to exhibit much more active amoeboid movements than they do at the tempera- ture of frog's blood. In summer a frog or other cold- blooded animal uses much more oxygen and evolves much more carbon dioxide than in winter, as shown not only by direct measurements of its gaseous exchanges, but by the fact that in winter a frog can live a long time after its lungs have been removed (being able to breathe sufficieiitly through its moist skin), while in warm weather it dies of asphyxia very soon after the same loss. The warmer weather puts its tissues in a more active state; and so the amount of work the animal does, and therefore the amount of oxygen it needs, depend to a great extent upon the tem- perature of the medium in which it is living. With the warm-blooded animal the reverse is the case. It always keeps up its temperature to that at which its tissues live best, and accordingly in cold weather uses more oxygen and sets free more carbon dioxide because it needs a more active internal combustion to compensate for its greater loss of heat to the exterior. In fact the living tissues of a man may be compared to hothouse plants, living in an artifici- ally maintained temperature; but they differ from the ])lants in the fact that they themselves are the seats of the combustions by which the temperature is kept up. Since, within wide limits, the Human Body retains the same tem- ]ierature no matter whether it be in cold or warm surround- ings, it is clear that it must jDOSsess an accurate arrangement for heat regulation; either by controlling the production of heat in it, or the loss of heat from it, or both. The Temperature of the Body. The parts of the Body are all eitlior in contact with one another directly or, if SOURCES OF BODILY HEAT. 451 not, at least indirectly through the blood, which, flowing from part to part, carries heat from warmer to colder regions. Thus, although at one time one group of muscles ma}' especially work, liberating heat, and at other times another, or the muscles may be at rest and the glands the seat of active oxidation, the temperature of the whole Body is kept pretty much the same. The skin, howeyer, which is in direct contact with external bodies, usually colder than itself, is cooler than the internal organs; its temperature in health is from 36° to 37° C. (96.8-98.5° F.), being warmer in more protected parts, as the hollow of the armpit. In internal organs, as the liver and brain, the temjierature is higher; about 42° 0. (107° F.) in health. In the lungs, though there is a certain quantity of heat liberated when oxygen combines with haemoglobin, this is more than counterbalanced by loss of the heat carried out by the ex- pired air, and that used up in evaporating the water carried out in the breath; so blood returned to the heart by the pulmonary veins is slightly colder than that carried from the right side of the heart to the lungs. The Sources of Animal Heat. These are two-fold; direct and indirect. Heat is directly produced wherever oxidation is taking place; so that all the living tissues at rest produce heat as the result of the chemical changes sup- plying them with energy for the maintenance of their vitality; and whenever an organ is active and its chemical metamorphoses are increased it becomes hotter: a secreting gland or a contracting muscle is warmer than a resting one. Indirectly, heat is developed by the transformation of other forms of energy; mainly mechanical work, but, to a less extent, also of electricity. All movements of parts of the Body which do not move it in space or move external objects, are transformed into heat within it; and the energy they represent is lost in that form. Every cardiac contraction sets the blood in movement, and this motion is for the most part turned into heat within the Body by friction within the blood-vessels. The same transformation of energy occurs with respect to the movements of the alimentary canal, ex- cept in so far as they expel matters from the Body; and 452 THE HUMxiN BODY. every muscle in contracting has part of the mechanical energy expended by it turned into heat by friction against neighboring parts. Similarly the movements of cilia and of amoeboid cells are for the most part converted in the Body into heat. The muscles and nerves are also the seats of manifestations of electricity, which, though small in amount, for the most part do not leave the Body in that form but are first converted into heat. A certain amount of heat is also carried into the Body with hot foods and drinks. The Energy Lost by the Body in Twenty -four Hours. Practically speaking, the Body only loses energy in two forms; as heat and mechanical work: by applying conduc- tors to different parts of its surface small amounts of elec- tricity can be carried off, but the amount is quite trivial in comparison with the total daily energy expenditure. Dur- ing complete rest, that is when no more work is done than that necessary for the maintenance of life, nearly all the loss takes the form of heat. The absolute amount of this will vary with the surrounding temperature and other con- ditions, but on an average a man loses, during a day of rest, 2700 calories; that is enough to raise 2700 kilograms (5-940 lbs.) of water from 0° to 1° C. (from 32° to 33.8° F.); otherwise expressed, this amount of heat would boil 27 kilos (59.4 lbs.) of ice-cold water. This does not quite represent all the energy lost by the Body in that time: since a small proportion is lost as mechanical work in moving the clothes and air by the respiratory movements, and even by the beat of the heart, which at each systole pushes out the chest-wall a little and moves the things in contact with it. The working Body liberates and loses much more energy; part as mechanical work done on external objects, part as increased heat radiated or conducted from the surface, or carried off by the expired air in the quickened respirations. Every one knows that he becomes warmer Avhen he takes exercise, and measurements made on men show that the heat produced and lost in a day of moderate work is about one third greater than that in a day of rest. The follow- ing table gives more accurate numbers — BODILY ENERGY LOST PER DAY. 453 Day of Rest. Day of Work. Rest 16 hrs. Sleep 8 hrs. Rest 8 hrs. Work 8 hrs. Sleep 8 hrs! Heat units (calor- 1 oj^q^ goQ jggg g 2169.6 320. les) produced.. ) o-on ^ / 10,885l^ah.-lb. \ q^^ « /^ 14,528 Fall. -lb. \ ^^■^ \ heat units. } '^''^■^ \ heat units. ) The mechanical work done on the working day, repre- sented in addition an expenditure of energy of 213,344 kilogrammeters, which is equal to 502 calories. Of the ex- cess heat in the working day, part is directly produced by the increased chemical changes in the quicker working heart and respiratory muscles, and the other muscles set at work; while part is indirectly due to heat arising from increased friction in the blood-vessels as the blood is driven faster around them, and to friction of the various muscles used. The average cardiac work in twenty-four hours is about 60,000 kilogrammeters ; that of the respiratory muscles about 14,000 ; and since nearly all of both is turned finally into heat within the Body, we have 74,000 kilogrammeters of energy answering to about 174 calories (6786 Fah.-lb. units) indirectly produced in the resting Body daily from these sources. Of 100 parts of heat lost from the resting Body, about 73 are carried off in radiation or conduction from the skin. 14.5 are carried off m evaporation from the skin. 7.2 " *' " " '^ '^ lungs. 3.0 " " '* expired air. 1.8 " " " in the excretions. In a day of average work, of every 100 parts of energy lost i 11 any form from the Body — 1-2 go as heat in the excreta. 3-4 in heating the expired air. 20-30 in evaporating water from the lungs and skin. 60-75 in heat radiated or conducted from the surfaces and in external mechanical work. The Superiority of the Body as a Working Machine. During eight hours of work, we find (table at top of page) the Body loses 2169.6 calories of energy as heat; and can do 454 THE HUMAN BODY. simnltaneonsly work equivalent to 502 calories. So of all the energy lost from it in that time tibont \ may take the form of mechanical work; this is a very large proportion of the total energy expended, being a much liiglier per- centage than that given by ordinary machines. The best steam-engines can utilize as mechanical work only libout ■^ of the total energy liberated in them and lost from them m a given time; the remainder is transmitted directly as heat to the exterior, and is lost to the engine for all useful purpo.-es. The Maintenance of an Average Temperature. This is necessary for the continuance of the life of a warm-blooded animal; should the temperature rise above certain limits chemical changes, incompatible with life, occur in the tis- sues ; for example at about 49° C. (120° F.) the muscles be- gin to become rigid. On the other hand death ensues if the Body be cooled down to about 19° C. (66° F.). Hence the need for means of getting rid of excess heat, and of protec- tion from excessive cooling. Either end may be gained in two ways; by altering the rate at which heat, is lost or that at which it is produced. As regards heat-loss, by far the most important regulating organ is the skin: under ordi' nary circumstances nearly 90 per cent of the total heat given off from the Body in 24 hours goes by the skin (73 by radi- ation and conduction, 14.5 by evaporation; see above table). This loss may be controlled — 1. Jiy clothing; we naturally wear more in cold and less in warm weather ; the effect of clothes being, of course, not to Avarm the Body but to diminish the rate at which the heat formed in it is lost. 2. Increased temperature of the surrounding medium in- creases the activity of the heart and lungs. A hastened circulation by itself does not as already pointed out (p. 388) increase tlie general tissue activity of the Body, or the oxida- tions occurring in it, and, so. a})art from the harder working heart itself, does not influence the amount of heat liberated in the Body during a given time: but the more rapid blood-flow through the skin carries more of that fluid through this cool surface and increases the loss of heat in that way. The TEMPEBATURE-REQULATION IK 2 HE BODY. 455 quickened respirations, too, increase the evaporation of water in the lungs and, thus, the loss of heat, 3. Warmth directly dilates the skin-vessels and cold con- tracts them. In a warm room the vessels on the surface dilate as shown by its redness, while in a cold atmosphere tliey contract and the skin becomes pale. But the more blood that flows through the skin the greater will be the heat lost from the surface — and vice versa. 4. Heat induces sweating and cold checks it ; the heat appears to act, partly, reflexly in exciting the sweat-centres from which the secretory nerves for the sudoriparous glands arise, and, partly, directly on those centres, which are thrown into activity, at least in health, as soon as the temperature of the blood is raised. In fever of course we may have a high temperature with a dry non-sweating skin. The more sweat there is poured out, the more heat is used up in evaporating it and the more the Body is cooled. 5. Our sensations induce us to add to or diminish the lieat in the Body according to circumstances; as by cold or warm baths, and iced or hot drinks. As regards temperature-regulation by modifying the rate of heat production in the Body the following points may be noted; on the whole such regulation is far less important than that brought about by changes in the rate of loss, since the necessary vital work of the Body always necessitates the continuance of oxidative processes which liberate a tol- erably large quantity of heat. The Body cannot therefore be cooled by diminishing such oxidations ; nor on the other hand can it be safely warmed by largely increasing them. Still, within certain limits, the heat production may b'^ con- trolled m several ways — 1. Cold increases hunger; and increased ingestion of food increases bodily oxidation as shown by the greater amount of carbon dioxide excreted in the hours succeeding a meal. This increase is probably due to the activity into which the digestive organs are throAvn. 2. Cold inclines to voluntary exercise; warmth to muscu- lar idleness; and the more the muscles are worked the more heat is produced in the Body. 456 THE nUMAN BODY. 3. Cold tends to produce involuntary muscular move- ments, and so increased lieat production; as chattering of the teeth and shivering. 4. Cold applied to the skin increases the bodily chemical metamorphoses and so heat production. At least the tem- perature in the armpit rises at first on entering a cold bath, though the heat carried off from the surface soon overbalances its increased production. The phenomenon may, however, be explained in another way, the rise being attributed to a sudden diminution of loss from more exposed parts of the skin, dependent on contraction of the cutaneous arteries. In some cases, however, the temjiorary rise is accompanied by an increased excretion of carbon dioxide, which would indicate that the surface cooling does really increase the oxidations of the Body. 5. Certain drugs, as salicylic acid, and perhaps quinine, diminish the heat production of the Body. Their mode of action is still obscure. On the whole, however, the direct heat-regulating me- chanisms of the Human Body itself are not very efficient, especially as protections against excessive cooling. Man needs to supplement them by the use of clothing, fuel, and exercise. Local Temperatures. Although, by the means above described, a wonderfully uniform bodily temperature is maintained, and by the circulating blood all parts are kept at nearly the same warmth, variations in both respects do occur. The arrangements for equalization are not by any means fully efficient. External parts, as the skin, the lungs (which are really external in the sense of being in contact with the air), the mouth, and the nose chambers, are always cooler than internal; and even all parts of the skin have not the same temperature, such hollows as the armpit being warmer than more exposed regions. On the other hand, a secreting gland or a working muscle becomes warmer, for the time, than the rest of the Body, because more heat is liberated in it than is carried off by the blood flowing through. In such organs the venous blood leaving is warmer than the arterial coming lo ibcm; while the reverse is the TRERMIG NERVES. 457 case with parts, like the skin, in which the blood is cooled. An organ colder than the blood is of course warmed by an increase in its circulation, as seen in the local rise of tem- perature in the skin of the face in blushing. Thermic Nerves. All nerves, such as motor or secre- tory, which can throw working tissues into activity are in a certain sense thermic nerves: since they excite increased oxidation and heat production in the parts under their con- trol. A true, purely thermic nerve would be one which increased the heat production in a tissue without otherwise throwing it into activity; and whether such exist is still undecided. Certain phenomena of disease, however, seem to render their existence probable. If we return for a moment to our former comparison of the working Body to a steam-engine, such nerves might be regarded as agencies increasing its rate of rusting without setting it at work. The oxidation of the iron would develop some heat, but by processes useless to the steam-engine, although such are, in moderation, essential to living cells; the vitality of these even when at rest, seems to necessitate a constant, if small, breaking down of its substance. In an amoeboid cell no doubt such processes occur quite independently of the nervous system; but in more differentiated tissues they may be controlled by it. Just as a muscle does not nor- mally contract unless excited through its nerve, although a white blood corpuscle does, so may the natural nutritive processes of the muscle-fibre in its resting condition be de- pendent on the nerves going to it. If these be abnormally excited the muscle Avill break down its protoplasm faster than it constructs it, and consequently waste; at the same time the increased chemical degradation of its sub- stance will elevate its temperature. Febrile conditions, in which many tissues waste, without any unusual manifesta- tion of their normal physiological activity, would thus be readily accounted for as due to superexcitation of the thermic nerves. Moreover, it is found that lesions or sec- tions of the spinal cord are followed by a rise in the tem- perature of those parts of the Body sujjplied with nerves arising below the diseased or divided portion. Now 458 THE HUMAN BODY. division of the spinal cord in two ways tends to lower the temperature of parts below the injury: in the first place, the muscles are paralyzed and so a great source of heat is cut otf ; and in the second, the yaso-motor nerves traveling down from the medulla-«-y centre are cut, and hence the skin arteries behind the section dilate and carry more blood to the surface to be cooled. To explain the rise of tem- perature it has therefore been concluded that there are true thermic centres in the spinal cord, which centres, like others in that organ (Chap. XXXV.), are held in check or inliibitecl by brain-centres; when the controlling influence of the latter is removed the former may excite excessive oxida- tions in the tissues to which they are distributed, and so produce the rise of temperature. The proof, however, is not complete; for the raised temperature may, after all, be due merely to an excessive supply of blood, warmed else- where m the Body, to the dilated skin-vessels. Clothing. To man, as social animal, endowed with moral feelings, clothing has certain uses in the interests of morality; but for such purposes the amount necessary is not great, as we find in many tribes living in warm climates. Except in tropical regions, liowever, clothing has in addi- tion an important physiological use in regulating the bodily temperature. While the majority of other warm-blooded animals have coats of their own, formed of hairs or feathers, over most of man's Body his capillary coating is merely rudi- mentary and has lost all i)hysiological importance; and so lie has to protect himself by artificial garments, which Ills gesthetic sense has led him to utilize also for purposes of ;idornment. Here, however, we must confine ourselves to clothes from a physiological point of view. In civilized societies every one is required to cover most of his Body with something, and the question is what is the best covering; the answer will vary, of course, with the climatic conditions of the country dwelt in. In warm I'egions, clothing, in general terms, should allow free radiation or conduction of heat from the surface; in cold it should do the reverse; and in temperate climates, with varying tem- peratures, it should vary with the season. If the surface CLOTHING. 459 of the Body be exposed so tliat ciirrents of air can freely traverse it much more lieat will be carried off (under those usual conditions in which the air is cooler than the skin) than if a stationary layer of air be maintained in con- tact with the surface. As every one knows, a "draught" cools much faster thaa air of the same temperature not in motion. All clothing, tlierefore, tends to keep up the temperature of the Body by checking the renewal of the layer of air in contact with it. Apart from this, however, clothes fall into two great groups; those which are good, and those which are bad, conductors of heat. The former allow changes in the external temperature to cool or heat rapidly the air stratum in actual contact with the Body, while the latter only permit these changes to act more slowly. Of the materials used for clothes, linen is a good conductor; calico not quite so good; and silk, wool, and fur are bad conductors. Whenever the surface of the Body is suddenly chilled the skin-vessels are contracted and those of internal parts reflexly dilated ; hence internal organs tend to become congested, a condition which readily jxasses into the diseased state known as inflatmnation. When hot, therefore, the most unadvisable thing to do, is to sit in a draught, throw off the clothing, or in other ways to strive to get sud- denly cooled. Moreover, while in the American summer it is tolerably safe to wear good-conducting garments, and few people take cold then, this is by no means safe in the spring or autumn, when the temperature of the air is apt to vary considerably within the course of a day. A person going out, clad only for a warm morning, may have to re- turn in a very much colder evening; and if his clothes be not such as to prevent a sudden surface chill, will get off lightly if he only " take" one of the colds so prevalent at those seasons. In the great majority of cases, no doubt, he suffers nothing worse, but many persons, especially of the female sex, often acquire far more serious diseases. When sudden changes of temperature are at all probable, even if the pro- vailing weather be warm, the trunk of the Body should be always protected by some tolerably close-fitting garment 460 THE HUMAN BODY. of non-conducting material. Those whose skins are irri- tated by anything but linen should wear immediately out- side the under-garments a jacket of silken or woolen material. In midwinter comparatively few joeople take cold, because all then wear thick and nonconducting cloth- ins: of some kind. CHAPTER XXX. SENSATION AND SENSE-ORGANS. The Subjective Functions of the Nervous System. Changes in many parts of our Bodies are accompanied or followed by those states of consciousness which we call sew- sations. All such sensitive parts are in connection, direct or indirect, with the brain, by certain afferent nerve-fibres called sensory. Since all feeling is lost in any region of the Body when this connecting path is severed, it is clear that all sensations, whatever their primary exciting cause, are finally dependent on conditions of the central nervous sys- tem. Hitherto we have studied this as its activities are revealed through movements which it excites or prevents; we have seen it, directly or reflexly, cause muscles to con- tract, glands to secrete, or the pulsations of the heart to cease; we have viewed it ohjedively, as a motion-regulating apparatus. Now we have to turn to another side and con- sider it (or parts of it) as influencing the states of conscious- ness of its possessor: this study of the subjective activities of the nervous system is one of much gi'eater difficulty. It may be objected that considerations concerning states of feeling h-ave no proper place in a treatise on Anatomy and Physiology; that, since we cannot form the beginning' of a conception how a certain state of the nervous system causing the feeling redness, another the feeling blueness, and a third the emotion anger, all examination of mental phe- nomena should be excluded from the sciences dealing with the structure and properties of living things. But, although we cannot imagine how a nervous state {neurosis) gives rise to a conscious state (psychosis), we do know this, that dis- tinct phenomena of consciousness never come under our 462 THE HUMA^' BODY. observation apart from a nervous system, and so are pre- sumably, in some way, endowments of it; we are, therefore, justilied in calling them 2)ro2)erties of the nervous system; and their examination, especially with respect to what nerve-parts are concerned with different mental states, and what changes in the former are associated with given phe- nomena in the latter, forms properly a part of Physiology. Whether masses of protoplasm, before the differentiation of definite nerve-tissues, possess some ill-defined sort of con- sciousness, as they possess an indefinite contractility before they have been modified into muscular fibres, may for the present be left undecided: though those who accept the doc- trine of evolution will be inclined to assent to the proposition. While, however, the Physiologist has a right to be heard on questions relating to our mental faculties, it is never- theless true that many laws of thought have been esta- blished, concerning which our present knowledge of the laws of the nervous system gives us no clue; the science of Psychology has thus a well-founded claim to an independent existence. But, in so far as its results are confined merely to the successions and connections of mental states, as estali- lished by observation, they are merely descriptions, and noi explanations in a scientific sense: we know that so many mental phenomena have necessary material antecedents and concomitants in nervous changes, that we are justified in believing that all have such, and in continuing to seek for tliem. We do not know at all how an electric current sent round a bar of soft iron makes it magnetic; we only know that the one change is accompanied by the other; h\xi we say we have explained the magnetism of a piece of iron if we have found an electric current circulating around it. Similarly, we do not know how a nervous change causes a mental state, but we have not explained the mental state until we have found the nervous state associated with it and how that nervous state was produced. As yet it is only with respect to some of the simplest riates of consciousness that we know much of the necessary physiological antecedents, and among these our sensations are the best investigated. As regards siich mental pheno- sensatto:n' and organs of special sense. 4G3 meua a,s the Association of Ideas, and Memory, physio- logy can give ns some light; but so far as others, such as the Will and the Emotions, are concerned, it has at present little to offer. The phenomena of Sensation, therefore, occupy at present a much larger portion of physiological works than all other mental facts put together. Common Sensation and Organs of Special Sense. A sensory nerve is one which, when stimulated, arouses, or may arouse, a sensation in its possessor. The stimulant is in all cases some form of motion, molar [e.g. mechanical pressure) or molecular (as ethereal vibrations or chemical changes). Since all our nerves lie within our Bodies as circumscribed by the skin, and are excited within them, one might, a priori be inclined to suppose that the cause of all sensations would appear to be within our Bodies themselves; that the tlwwg felt would be a modified i)ortiou of fhe feeler. This is the case with regard to many sensations; a head- ache, toothache, or earache gives us no idea of any external object; it merely suggests to each of us a particuhir state of a sensitive portion of myself. As regards many sensations, however, this is not so; they suggest to us ex- ternal causes, to properties of which and not to states of our Bodies, we ascribe them ; and so they lead us to the concejition of an external universe. A knife laid on the skin produces changes in it which lead us to think not of a state of our skin, but of states of some object outside the skin; we believe Ave feel a cold heavy hard thing in contact with it. Nevertheless we have no sensory nerves going into the knife and informing us directly of its condition; what we really feel are the modifications of our Body produced by it, although we irresistibly think of them as properties of the knife — of some object that is no part of our Body, and not as states of the latter itself. Let now the knife cut through the skin; we feel no more koiife, but experience pain, which we thmk of as a condition of ourselves. We do not say the knife is painful, but that our fiiiger is, and yet we have, so far as sensation goes, as much reason to call the knife painful as cold. Applied one way it produced local changes arousing a sensation of cold, and in another local 464 THE HUMAN BODY. changes causing a sensation of pain. Nevertheless in the one case we speak of the cold as being in the knife, and in the other of the pain as being in the finger. Sensitive parts, such as the surface of the skin, through which we get, or believe Ave get, information about outer things, are of far more intellectual value to us than sensitive parts, such as the subcutaneous tissue into which the knife may cut, which give us only sensations referred to conditions of our Bodies. The former are called Sense-organs proper, or Organs of Special Sense; the latter are sensitive paris, or Organs of Common Sensation. The Peripheral Reference of our Sensations. The fact that we refer certain sensations to external causes is only one case of a more general law, in accordance with which we do not ascribe our sensations, as regards their lo- cality, to the brain, where the neurosis is accomi^anied by the sensation, but to a peripheral part. With respect to the brain, other parts of the Body are external ol)jects as much as the rest of the material universe, yet the majority of our common sensations are felt at the places where the sensory nerves concerned are irritated, and not in the brain. Even if a nerve-trunk be stimulated in the middle of its course, we refer the resulting sensation to its outer endings. A blow on the inside of the elbow-joint, injuring the ulnar nerve, produces not only a local pain, but a sense of ting- ling ascribed to the fingers to which the ends of the fibres go. Persons with amputated limbs have feelings in their fingers and toes long after they have been lost, if the nerve- trunks in the stump be irritated. To explain such facts we must trench on the ground of Psychology, and so they can- not be fully discussed here; but they are commonly ascribed to the results of experience. The events of life have taught us that in the great majority of instances the sensory im- ])ulses which excite a given tactile sensation, for example, have acted upon the tip of a finger. The sensation goes when tbe finger is removed, and returns when it is rejilaced; and the eye confirms the contact of the external object with the tinger-tip wlien we get tlie tactile sensation in question. Wc thus come firmly to assocuite a particular region of the QUALITIES OF SENSATIONS. 465 skin with a given sensation, and whenever afterwards the nerve-fibres coming from the finger are stimulated, no mat- ter where, we ascribe the origin of the sensation to some- thing acting on the finger-tip. The Differences between Sensations. In both groups of sensations, those derived through organs of special sense and those due to organs of common sensation, we dis- tinguish kinds which are absolutely distinct for our con- sciousness, and not comparable mentally. We can never get confused between a sight, a sound, and a touch, nor be- tween pain, hunger, and nausea; nor can we compare them with one another; eacli is sai generis. The fundamental difference which_thvis separates one sensation from another is its modality. Sensations of the same modality may differ; but they shade imperceptibly into one another, and are com- parable between themselves in two ways. First, as regards qualityj while a high and a low pitched note are both auditory sensations, they are nevertheless different and yet intelligibly comparable; and so are blue and red objects. In the second place, sensations of the same modality are distinguishable and comparable as to amount or intensitij: we readily recognize and compare a loud and a weak sound of the same pitch; a bright and feeble light of the same color; an acute and a slight pain of the same general char- acter. Our sensations thus differ in the three aspects of onodality, quality untJtin the same modality, and intensity. Certain sensations also differ in what is known as the *' local sign," a difference by which we tell a touch on one part of the skin from a similar touch on another; or an ob- ject exciting one part of the eye from an object like it, but in a different location in sjoace and exciting another part of the visual surface. As regards modality, we commonly distinguish five senses, those of sight, sound, touch, taste, and smell; it is doubtful whether temperature should not be added. The varieties of common sensation are also several; for example, pain, hunger, satiety, thirst, nausea, malaise, hien-etre (feel- ing "good"), fatigue. The muscular sense stands on the intermediate line between special and common sensations; 466 THE HUMAN BODY. we gather by it how much our various muscles are con- tracted, and so learn the position of various parts of the Body, on the one hand, and the resistance opposed to bodily movement by external objects, on the other. In fact, we cannot draw a sharp line between the special senses and common sensations: all the Body, we conclude from ob- servations on the lower animals, is, at an early stage of its development, sensitive; very soon its cells sej^arate them- selves into an outer layer exposed to the action of external forces and an inner layer protected from them: and some of the former cells become especially sensitive. From them, as development proceeds, some are sej)arated and buried be- neath the surface to become the brain and spinal cord; of those which remain superficial, some are modified so that they (in the eye) become especially excited by ethereal vi- brations; others (in the ear) become especially responsive to sound vibrations; others to slight chemical changes (in mouth and nose), and others (in the skin) to variations in pressure or temperature. All our sensations are thus modifications of one common primary sensibility, represented by that of the skin, or rather by the primitive rej^resentative of the skin in such an animal as the Hydra (see Zoology). The cutaneous sen- sations, being less differentiated, shade off more readily into the common sensibility of the other living tissues than do the activities of the highly differentiated cells in the e3'e and ear. We find, accordingly, that while a powerful pres- sure or a high temperature acting on the skin readily arouses a sensation of pain, that this is not the case with the more specialized visual and auditory organs. Their super-ex- citement maybe disagreeable, but never passes into j^ai'w, in the ordinary sense of the word. Similarly the special skin sensations, touch and temperature, may sometimes be con- founded, while a sound and a sight cannot be: the mo- dality of the less modified skin-senses is less comj)lete. The study of comparative anatomy and development thus shows that the irritable parts of our sense-organs are but special dif- ferentiations of the primary external layer of cells covering the Body when it is very young. Some of these become nerve STRUCTURE OF SENSE-ORGANS. -iG? end organs in the eye, others end organs in the ear, and so on; while others, less changed, remain in the skin as organs of touch and temperature; and so, frona a general exterior surface responding equally readily to many external natural forces, we get a surface modified so that its various parts resj^oud with different degrees of readiness to different ex- ternal forces; and these modified parts constitute the essen- tial portions of our organs of special sense. Every sense organ thus comes to have a special relationship to some one natural force or form of energy — is a specially irritable mechanism by which such a force is enabled to excite sen- sory nerves; and is, moreover, commonly supplemented by arrangements which, in the ordinary circumstances of life, prevent other forces from stimulating the nerves connected with it. Not all natural forces have sense-organs with ref- erence to them developed in the Human Body; for exam- ple, we have no organ standing to electrical changes in the same relation that the eye does to light or the ear to sound. The Essential Structure of a Sense-Organ. In every sense-organ the fundamental part is thus one or more end organs^ which are highly irritable tissues (p. 31), so con- structed and so placed as to be normally acted on by some one of the modes of motion met with in the external world. A sensory apparatus requires in addition at least a brain-cen- tre, and a sensory nerve-fibre connecting this with the ter- minal ap2)aratus; but one commonly finds accessory parts added. In the eye, e.g., we have arrangements for bringing to a focus the light rays which are to act on the end organs of the uerve-fibres; and in the ear are found similar subsidiary parts, to conduct sonorous vibrations to the end apparatus of the auditory nerve. Seeing and hearing are the two most specialized senses; the stimuli usually arousing tliem are peculiar and quite distinct from the group of general nerve stimuli (see p. 188), while those most frequently, or naturally, acting upon our other sense-organs are not so peculiar; they are forces wJaich act as general nerve stmiuli when directly applied to nerve- fibres. The end organs, however, as already pointed out 468 THE HUM AX BODY. (p, 100), so increase the sensitiveness of the parts contain- ing them that degrees of change in the exciting forces, which would be totally unable to stimulate neiTe-fibres themselves are appreciated. These terminal apparatuses are therefore as truly mechanisms enabling changes, which would not otherwise stimulate nerves, to excite them, as are the end organs in the eye or ear. The Cause of the Modality of our Sensations. Seeing that the external forces usually exciting our difl'erent sen- sations differ, and that the sensations do also, we might at first be inclined to believe that the latter difference de- pended on the former: that brightness differed from loud- ness because light was different from sound. In other words, we are apt to think that each sensation derives its specific character from some property of its external physi- cal antecedent, and that our sensations answer in some way to, and represent more or less accurately, properties of the forms of energy arousing them. It is, however, quite easy to show that we have no sufficient logical warrant for such a belief. Light falling into the eye causes a sensation of luminosity, a feeling belonging to the visual group or modality; and, since usually nothing else excites such feelings and light entering the healthy eye always does, we come to believe that the physical agent light is some- thing like our sensation of luminosity. But, as we have already seen (p. 191), no matter how we stimulate the optic nerve we still get visual sensations; close the eyes and press with a finger-nail on one eyelid; a sensation of touch is aroused where the finger meets the skin; but the pressure on the eyeball distorts it and stimulates the optic nerve- fibres in it also, and the result is a luminous patch seen in front of the eye in such a position as a bright body must occupy in space to radiate light to that part of the expan- sion of the optic nerve. Finding, then, the same kind of sensation, a visual one, produced by the totally different causes, pressure and light, we are led to doubt if the dif- ferences of modality in our sensations depend upon the dif- ferences of the natural forces arousing them; and this doubt is strengthened when we find still other forces (p. 191) WHY OUR SEirSATIONS DIFFEB. 469 giving rise to visual sensations. But then, since light and pressure, electricity and cutting, all cause visual sensations, we have no valid reason for supposing that light, more than either of the others, is really in any way like our sensation of light: or that sight-feeling differs from sound-feeling because objectively light differs from sound. The eye is an organ specially set apart to be excited by light, and accord- ingly so fixed as to have its nerve-fibres far more often ex- cited by that form of force than by any other; but the fact that light sensations can be otherwise aroused shows plainly that their kind or character has nothing directly to do witli any property of light. Just as by pinching or heating or galvanizing a motor nerve we can make the muscles attached to it contract, and the contraction has nothing in common with the excitant, so the visual sensation, as such, is inde- pendent of the stimulus arousing it and, of itself, tells us nothing concerning the kind of stimulus which has operated. Differences in kind between external forces being thus eliminated as possible causes of the modalities of our sen- sations, we next naturally fall back upon differences in the sense-organs themselves. They do undoubtedly differ both in gross and microscopic structure, and the fact that pres- sure on the closed eye arouses a touch-feeling where the skin is compressed, and a sight-feeling where the optic nerve is, might well be due to the fact that a peripheral touch- organ was different from a peripheral sight-organ, and the same force might therefore produce totally different effects on them and so cause different kinds of feelings. However, here also closer examination shows that we must seek far- ther. Sensation is not produced in a sense-organ, but far away from it in the brain; the organ is merely an apparatus for generating nervous impulses. If the optic nerves be divided, no matter how perfect the eyeballs, no amount of light will arouse visual sensations; if the spinal cord be cut in the middle of the back no pressure on the feet will cause a tactile or other feeling; though the skin, and its nerves and the lower half of the spinal cord be all intact. In all cases we find that if the nerve-paths between a sense-organ 470 THE HVMAN BODY. and the brain be severed no stimidation of the organ will call forth a sensation. The final production of this clearly depends, then, on something occurring in the brain, and so tliekind of a sensation is presumably dependent upon brain events rather than on occurrences in sense-organs. Still it might be that something in the sense-organ caused one sen- sation to differ from another. Each organ might excite the brain in a different way and cause a different sensation, and so our sensations differ because our sense-organs did. Such a view is, however, negatived by observations which show that perfectly characteristic sensations can be felt in the absence of the sense-organs through which they are normally ex- cited. Persons whose eyeballs have been removed by the surgeon, or completely destroyed by disease, have frequently afterwards definite and unmistakable visual sensations, quite as characteristic as those which they had while still possess- ing the visual end organs. The tactile sensations felt in amputated limbs, referred to above, afford another example of the same fact. The persons still feel things touching their legs or lying between their long-lost toes; and the sen- sations are distinctly tactile and not in any way less different from visual or auditory sensations than are the touch-feel- ings following stimulation of those parts of the skin which are still possessed. It is, then, clear that the modality of our sensations is to be sought deeper than in properties of the end organs of the nerves of each sense. Properties of external forces and properties of periphe- ral nerve-organs being excluded as causes of differences in kind of sensation, we come next to the sensory nerve-fibres themselves. Is it because optic nerve-fibres are different from auditory nerve-fibres that luminous sensations aie different from sonorous? This question must be answered in the negative, for we have already (p. 193) seen reason to believe that all nerve-fibres are alike in essential structure and that their properties are everywhere the same; that all they do is to transmit "nervous impulses" when excited, and that, no matter what the c::ritant, these impulses are molecular movements, always alike in kind, though they may differ in amount and in rate of succession. Since, WHY OUR 8EXSATI0N8 DIFFER. 471 then, all that the optic nerve does is to send nervous impulses to the brain, and all that the auditory and gustatory and touch and olfactory nerve-fibres do is the same, and these impulses are all alike in kind, we cannot explain the differ- ence in quality of visual and other sensations by any dif- ferences in property of the nerve-trunks concerned, any more than we could attempt to explain the facts that, in one case, an electric current sent through a thin platinum wire heats it, and, in another, sent through a solution of a salt decom- poses it, by assuming that the different results depend on differences in the conducting copper wires, which may be absolutely the same in the two cases. We are thus driven to conclude that our sensations primarily differ because different central nerve-organs in the brain are concerned in their production. That just as an efferent nerve-fibre will, when stimulated, cause a secre- tion if it go to a gland-cell, and a contraction of it to go to a muscle-fibre, so an optic nerve-fibre, carrying impulses to one brain apparatus and exciting it, will cause a visual sensation, and a gustatory nerve-fibre, connected with another brain-centre, a taste sensation. In other words, our kinds of sensation depend fundamentally on the proper- ties of our own cerebral nervous system. For each special sense we have a nervous apparatus with its peripheral terminal organs, nerve-fibres, and brain-centres; and the excitement of this apparatus, no matter in what way, causes a sensation of a given modality, determined by the proper- ties of its central portion. Usually the apparatus is excited by one particular force acting first on its peripheral organs, but it may be aroused by stimulating its nerve-fibres directly or, as in certain diseased states (delirium), or under the action of certain drugs, by direct excitation of the centres. The sensations of dreams, frequently so vivid, and halluci- nations, a,re also probably in many cases due to direct excitation of the central organs of sensory apparatuses, though no doubt also often due to peripheral stimulation. But no matter how or where the apparatus is excited, pro- vided a sensation is produced it is always of the modality of that sense apparatus. 472 THE HUM AN BODY. It is possible that the general statement in the last para- graph needs some limitation, Wliile in senses of distinct modality, this character can be ascribed only to brain projjer- ties (so that Ave may be pretty sure that a man, the inner end of whose optic nerve was in physiological continuity with the outer end of his auditory, and the inner end of his auditory with the outer end of his optic, might hear a pic- ture and see a symphony), yet, perhaps, differences in the rhythm or intensity of afferent nervous impulses may cause differences in modality in less differentiated senses. Thus contact with a cold soft object may be felt as heat, thought to be due to the approach of a warm body; and from such cases we must perhaps conclude that touch and temperature depend on excitations in different ways of one and the same brain-centre; impulses of a certain rhythm producing a sensation of heat, and those of another (deter- mined by the different heat and touch end organs) causing a tactile sensation. If this be so, however, heat and touch would be but extreme varieties of one kind of sensation, and comparable to yellow and blue. Again, a heavy pres- sure, gradually increased, arouses sensations which pass im- perceptibly from touch to pain, and the result may be due to the fact that regular and orderly afferent impulses, determined through tactile r erve-endings, excite the centre in one Avay; while irregular, disorderly, and violent, excited Avhen the nerve-trunks beneath the skin are directly stimu- lated, may cause a different sensation; much as the same musical notes combined in one order cause pleasure but in another are disagreeable, causing a sort of pain, although the same brain-centres are stimulated in the two cases. The pain from a heavy weight may, however, be merely due to the fact that it excites the nerves very powerfully and gives rise to impulses which radiate farther in the brain than those causing touch sensations, and so excite new centres, the modality of Avliich is a pain sensation. However differences in nervous rhythm may account for minor differences in sensation, it remains clear that the characters of our sensations are creations of our own organ- FECHNER8 LAW. 473 ism; they depend on properties of our Bodies and not on properties of external things, except in so far as these may or may not be adajjted to arouse our different sensory apparatuses to activity. From the kind of the sensation we cannot, therefore, argue as to the nature of the excitant: we have no more warrant for supposing that hght is like our sensation of light than that the knife that cuts us is like our sensation of pain. All that we know with certainty is states of our own consciousness, and although from these we form working hypotheses as to an external universe, yet, granting it, we have no means of acquiring any real knowl- edge as to the properties of things about us. What we want to know, however, for the practical purposes of life is, not what things are, but how to use them for our advan- tage, or to prevent them from acting to our disadvantage; and our senses enable us to do this sufficiently well. The Psycho-Physical Law. Although our sensations are, in modality or kind, independent of the force exciting them, they are not so in degree or intensity, at least within cer- tain limits. We cannot measure the amount of a sensation and express it in foot-pounds or calories, but we can get a sort of unit by determining how small a difference in sen- sation can be perceived. Supposing this smallest perceptible difference to be constant within the range of the same sense, (which is not proved,) it is found that it is produced by dif- ferent amounts of stimuli, measured objectively as forces; and that there exists in some cases a relation between the two which can be expressed in numbers. The increase of stim- ulus necessary to produce the smallest perceptible change in a sensation is proportional to the strength of the stimulus already acting; for example, the heavier a pressure already acting on the skin the more must it be increased or dimin- ished in order that the increase or diminution may be felt. Expressed in another way the facts may be put thus: sup- pose three degrees of stimulation to bear to one another ob- jectively the ratios 10, 100, 1000, then their subjective ef- fects, or the amounts of sensation aroused by them, will be respectively as 1, 2, 3; in other words, the sensation in- 474 THE HVMAX BODY. creases proportionately to the logaritlim of tlie strength of the stimulus. Examples of this, which is known as "■ Weber's " or "Fechner's psycho-physical Icciv" will be hereafter pointed out, and are readily observable in daily life; wo have, for example, a luminous sensation of certain intensity when a lighted candle is brought into a darkroom; this sen- sation is not doubled when a second candle is brought in; and is hardly affected at all by a third. The law is only true, however (and then but api^roximately), for sensations of medium intensity; it is applicable, for example, to light sensations of all degrees between those aroused by the light of a caudle and ordinary clear daylight: but it is not ti'ue for luminosities so feeble as only to be seen at all with diffi- culty, or so bright as to be dazzling. Besides their variations in intensity, dependent on varia- tions in the strength of the stimulus, our sensations also vary with the irritability of the sensory apparatus itself; which is not constant from time to time or from person to person. In the above statements the condition of the sense- organ and its nervous connections is presumed to remain the same throughout. Perceptions. In every sensation we have to carefully distinguish between the pure sensation and certain judg- ments founded upon it; we have to distinguish between what we really feel and what we think we feel; and very often firmly believe we do feel when we do not. The most important of these judgments is that which leads us to ascribe certain sensations, those aroused through organs of special sense, to external objects — that outer reference of our sensations which leads us to form ideas concerning the existence, form, position, and properties of external things. Such representations as these, founded on our senses, are called pierceptions. Since these always imply some mental activity in addition to a mere feeling, their f nil discussion belongs to the domain of Psychology. Phy- siology, however, is concerned with them so far as it can determine the conditions of stimulation and neurosis under which a given mental representation concerning a sensation is made. It is quite certain that we can feel PERCEPTIONS. 475 nothing biTt states of ourselves, but, as already pointed out, we have no hesitation in saving we feel a hard or a cold, a rough or smooth body. When we look at a distant object we usually make no demur to saying that we perceive it. What we really feel is, however, the change produced by it in our eyes. There are no parts of our Bodies reaching to a tree or a house a mile off — and yet we seem to feel all the while that we are looking at the tree or the house and feeling them, and not merely experiencing modifications of oar own eyes or brains. When reading we feel that what we really see is the book; and yet the existence of the book is a judgment founded on a state of our Body, which alone is what we truly feel. We have the same experience in other cases, for example with regard to touch. Hairs are quite insensible, but are imbedded in the sen- sitive skin, which is excited when they are moved. But if the tip of a hair be touched by some external object we be- lieve we feel the contact at its insensible end, and not in the sensitive skin at its root. So, the hard parts of the teeth are insensible; yet when we rub them together we refer the seat of the sensation aroused to the points where they touch one another, and not to the sensitive parts around the sockets where the sensory nerve impulse is really started. Still more, we may refer tactile sensations, not merelv to the distal ends of insensible bodies implanted in the skin, but to the far ends of things which are not parts of our Bodies at all; for instance, the distant end of a rod held between the finger and a table. We then believe we feel touch or pressure in two places; one where the rod touches our finger, and the other where it comes in contact witli the table. We have, simultaneously, sensations at two places separated by the length of the rod. If we hold the rod immovably on the table we feel only its end next the fin- ger. If we could fix it immovably on the finger while the other end was movable on the talkie, we would lose the sen- sation at the finger and only belie-tte we felt the pressure where the rod touched the table. When a tooth is touched with a rod we only feel the contact at its end, unless it is 476 TEE HUMAN BOD Y. loose in its socket; and then we get two sensations on touching its free end with a foreign body. This irresistible mental tendency to refer certain of our states of feeling to causes outside of our Bodies, and either in contact with them or separated from them by a certain space, is known as the phenomenon of the extrmsic refer- ence of our sensations. The discussion of its origin belongs jjroperly to Psychol- ogy, and it will suffice here to point out that it seems largely to depend on the fact that the sensations extrinsically referred can be modified by movements of our Bodies. Hunger, thirst, and toothache all remain the same whether we turn to the right or left, or move away from the place we are standing in. But a sound is altered. We find that in a certain position of the head it is heard more by the right ear than the left; but on turning round the reverse is the case; and half way round the loudness in each ear is the same. Hence we are led, by mental laws outside of the physiological domain, to suspect that its cause is not in our Body, but outside of it; and depends not on a condition of the Body but on something else. And this is confirmed when going in one direction we find the sound inci'eased, and in the other that it is diminished. This implies that we have a knowledge of our movements, and this we gain through the muscular sense. It constitutes the reactive side of our sensory life, associated with the changes,we produce in external things; and is correlated and contrasted with the passive side, in which other things produce sensations by acting upon us. As regards our common sensations we find something of the same kind. The more readily they can be modified by movement the more definitely do we localize them in space, though in this case within the Body instead of outside it. Hunger and nausea can be altered by jiressure on the pit of the stomach; thirst by moistening the throat with water; the desire for oxygen (respiration-hunger) by movements of the chest; and so we more or less definitely ascribe these sensations to conditions of those parts of the Body. Other general sensations, as depression, anxiety, and so on, are not SENSORY ILLUSIONS. 477 modifiable by any particular movement, and so appear to us rather as mental states, pure and simple, than bodily sensations. Sensory Illusions. "I must believe my own eyes" and "we can't always believe our senses" are two expres- sions frequently heard, a^id, each expressing a truth. No doubt a sensation in itself is an absolute incontrovertible fact: if I feel redness or hotness I do feel it and that is an end of the m'atter: but if I go beyond the fact of my having a certain sensation and conclude from it as to j)roperties of something else — if I form adjudgment from my sensatio7i — I may be totally wrong; and in so far be unable to believe my eyes or skin. Such judgments are almost inextricably woven up with many of our sensations, and so closely that we cannot readily separate the two; not even when we know that the judgment is erroneous. For example, the moon when rising or setting, appears bigger than when high in the heavens — we seem to feel directly that it arouses more sensation, and yet we know certainly that it does not. With a body of a given brightness the amount of change produced in the end organs of the eye will depend on the size of the image formed in the eye, provided the same part of its sensory surface is acted U23on. Now the size of this image depends on the distance of the object; it is smaller the farther off it is and bigger the nearer, and measurements show that the area of the sensitive surface affected by the image of the rising moon is no bigger than that affected by it when overhead. Why then do we, even after we know this, see it bigger ? The reason is that when the moon is near the horizon we imagine, unconsciously and irresistibly, that it is farther off; even astronomers who know perfectly that it is not, cannot help forming this unconscious and erroneous judgment — and to them the moon appears in consequence larger when near the horizon, just as it does to less well-informed mortals. In fact we have a conception of the sky over which the moon trav- els, not as a half sphere but as somewhat flattened, and hence when the moon is at the horizon we unconsciously judge that it is farther off than when overhead. But any 478 THE HUMAN BODY. body which excites the same extent of the sensitive surface of the eye at a great distance that another does at less, must be larger than the latter: and so we conclude that the moon at tlie horizon is larger than the moon in the zenith, and are ready to declare that we see it so. So, again, a small bit of light gray paper on a white sheet looks gray: but j^laced. on a large bright green surface it looks purple; and on a bright red surface looks blue-green. As the same bit of gray paper is shifted from one to the other we see it change its color: it arouses in us different feelings, or feelings which we interpret differently, although objectively the light reflected from it remains the same. Similarly a medium-sized man alongside of a very tall one appears short, but when walking with a very short one, tall. Such erroneous perceptions as these are known as sensory illusions; and we ought to be constantly on guard against rhem. CHAPTER XXXI. THE EYE AS AN OPTICAL INSTRUMENT. The Essential Structure of an Eye. Every visual organ consists primarily of a nervous expansion, provided with end organs by means of which light is enabled to excite nervous impulses, and exposed to the access of objective light; such an expansion is called a retina. By itself, however, a retina would give no visual sensations referable to distinctly limited external objects; it would enable its possessor to tell light from darkness, more light from less light, and (at least in its highly developed forms) light of one color from light of another color; but that would be all. Were our eyes merely retinas we could only tell a printed page from a blank one by the fact that, being jjartly covered with black letters, (which reflect less light,) it would excite our visual organ less powerfully than the spotless white page would. In order that distinct objects and not merely degrees of luminosity may be seen, some arrangement is needed which shall bring all light entering the eye from one point of a luminous surface to a fomis again on one point of the sensitive surface. If A and B (Fig. 121) be two red spots on a black surface, K, and rr be a retina, then rays of light diverging from A would fall equally on all parts of the retina and excite it all a little; so with rays starting from B. The sensation aroused, supposing the retina in connection with the rest of the nervous visual apparatus, would be one of a certain amount of red light reachmg the eye; the red spots, as definite objects, would be mdislinguishable. If, however, a convex glass lens L (Fig. 122) be put in front of the retina, it will cause to converge again to a single point all the rays from A falling 480 THE HUMAN BODY. upon it; so, too, with the raj^s from B: and if the focal dis- tance (see Physics) of the lens be properly adjusted these points of convergence will both lie on the retina^ that for Fig. 121.— Diagram illustrating the indistinctness of vision with a retina alone. K, a surface on which are two spots, A and B; r r, tlie retina. The diverging lines represent rays of light spread uniformly over the retina from each spot. rays from A at a, and that for rays from B at b. The sensitive surface could then only be excited at two limited and separated points by the red light emanating from the spots; consequently only some of its end organs and nerve- FiG. 122.— Illustrating the use of a lens in giving definite retinal images. A, B, K, r r, as in Fig. l~'l. L, a biconvex lens so placed that it brings to a focus on the points a and 6 of the retina, rays of light diverging from A and B re- spectively. fibres would be stimulated and the result would be the recognition of two separate red objects. In our eyes are found certain refracting media which lie in front of the retina and take the place of the lens L in Fig 122. That portion of physiology which treats of the physical action of these media, or in other words of the eye as an optical in- strument, is known as the dioptrics of the eye. The Appendages of the Eye. The eyeball itself con- sists of the retina and refracting media, together with supporting and nutritive structures and other accessory THE EYELIDS. 481 apparatuses, as, for example, some controlling the light- converging power of the media, and others regulating the size of the aperture (jjupil) by which light enters. Out- side the hall lie muscles which bring about its movements, and other parts serving to protect it. Each orbit is a pyramidal cavity occupied by connective tissue, muscles, blood-vessels and nerves, and in great part by fat, which forms a soft cushion on which the back of the eyeball lies and rolls during its movements. The contents of the orbit being for the most part incompressible, the eye cannot be drawn into its socket. It simply rotates there, as the head of the femur does in the acetabulum. When the blood-vessels are gorged, however, the eyeballs may be caused to protrude (as in strangulation), and when the ves- sels empty it recedes somewhat, as is commonly seen after death. The front of the eye is exposed for the purpose of allowing light to reach it, but can be covered up by the eyelids, which are folds of integument, movable by muscles and strengthened by plates of fibro-cartilage. At the edge of each eyelid the skin which covers its outside is turned in, and becomes continuous with a mucous membrane, the conjunctiva, which lines the inside of each lid, and also covers all the front of the eyeball as a closely adherent layer. The upper eyelid is larger and more mobile than the lower, and when the eye is closed covers all its transparent part. It has a special muscle to raise it, the levator palpe- hrcB superioris. The eyes are closed by a flat circular mus- cle, the orbicularis palpebrarum, which, lying on and around the lids, immediately beneath the skin, surrounds the ajjer- ture between them. At their outer and inner angles {can- f/ii) the eyelids are united, and the apparent size of the eye depends upon the interval between the canthi, the eyeball itself being nearly of the same size in all j^ersons. Near the inner canthus the line of the edge of each eyelid changes its direction and becomes more horizontal. At this point is found a small eminence, the lacJirpnial pajnlla, on each lid. For most of their extert the i?iner surfaces of the eyelids are in contact with the outside of the eye- 482 THE HUMAN BODY. biill but, near their inner ends, a red vertical fold of con- junctiva, the semilunar fold {plica semilunaris) intervenes. This is a remnant of the third eyelid, or nictitating membrane, found largely developed in many animals, as birds, in wliich it can bo drawn all over the exposed part of the eyeball. Quite in the inner corner is a reddish ele- vation, the caruncula laclirymnlis, caused by a collection of sebaceous glands imbedded in the semilunar fold. Opening along the edge of each eyelid are from twenty to thirty minute compound sebaceous glands, called the Meibomian follicles. Their secretion is sometimes abnormally abun- dant, and then appears as a yellowish matter along the edges of the e3'elids, which often dries in the night and causes the lids to be glued together in the morning. The eyelaslies are short curved hairs, arranged in one or two rows along each lid where the skin joins the conjunctiva. The Lachrymal Apparatus consists of the tear-gland in each orbit, the ducts which carry its secretion to the upper eyelid, and the canals by which this, unless when excessive, is carried off fi'om the front of the eye without running down over the face. The lachrymal or tear gland, about the size of an almond, lies in the upper and outer part of the orbit, near the front end. It is a compound racemose gland, from which twelve or fourteen ducts run and open in a row at the outer corner of the upper eyelid. The se- cretion there poured out is spread evenly over the exposed part of the eye by the movements of winking, and keeps it moist; finally it is drained off by two lachrymal cancds, one of which oj^ens by a small pore {punctum lachrymalis) on each lachrymal papilla. The aperture of the lower canal can be readily seen l)y examining the corresponding papilla in front of a looking-glass. The canals run inwards and ojoen into the lachrymal sac, which lies just outside the nose, in a holloAV where the lachrymal and superior maxillary bones {L and M.v, Fig. 26)* meet. From the sac the nasal duct proceeds to open into the nose-chamber below the in- ferior turbinate bone {q. Fig. 89, p. 309). Tears are constantly being secreted, but ordinarily in * Page 74 MUSCLES OF THE EYEBALL. 483 such quantity as to be drained off into tlie nose, from which they flow into the pharynx and are swallowed. When the lachrymal ducts are stopped up, however, their continual presence makes itself unpleasantly felt, and may need the aid of a surgeon to clear the passage. In iveeping the secretion is increased, and then not only more of it en- ters the nose, but some flows down the cheeks. The fre- quent swallowing movements of a crying child, sometimes spoken of as ''gulping down his passion," are due to the need of swallowing the extra tears which reach the pharynx. Fig. 123.— The eyeballs and their muscles as seen when the roof of the orbit has been removed and the fat in the cavity has been partly cleared away. On the right side the superior rectus muscle has been cut awaj'. a, external rec- tus; s, superior rectus; i, internal rectus; t, superior oblique. The Muscles of the Eye (Fig. V2o). The eyeball is spheroidal in form and attached behind to the optic nerve, 11, somewhat as a cherry might be to a thick stalk. On its outside are inserted the tendons of six muscles, ionv straight and two oblique. The straight muscles lie, one {superior rectns), s, above, one {inferior rectus) below, one {external rectus), a, outside, and one {internal rectus), i, inside the eyeball. Each ai'ises behind from the bony margin of the foramen through which the optic nerve enters the orbit. In the figure. 484 THE HUMAN BODY. which represents tlie orbits opened from ahove, the superior rectus of tlie right side has been removed. The superior oblique or pulley {trochlear) muscle, t, arises behind near the straight muscles and forms anteriorly a tendon, ■u, which passes through a fibro-cartilaginous ring, or pulley, placed at the notch in the frontal bone, Avhere it bounds superiorly the front end of the orbit. The tendon then turns back and is inserted into the eyeball between the ujiper and outer recti mu.-cles. The inferior oblique muscle does not arise, like the rest, at the back of the orbit, but near its front at the inner side, close to the lachiymal sac. It passes thence outwards and backwards beneath the eyeball to be inserted into its outer and j^osterior part. The-inner, upper, and lower straight muscles, the inferior oblique, and the elevator of the upper lid are supplied by branches of the third cranial nerve (see p. 168). The sixth cranial nerve goes to the outer rectus; and the fourth to the superior oblique. The eye may be moved from side to side; up or down; obliquely, that is neither truly vertically nor horizontally, but partly both; or, finally, it may be rotated on its antero- posterior axis. The oblique movements are always accom- panied by a slight amount of rotation. When the glance is turned to the left, the left external rectus and the right in- ternal contract, and vice versa; when up, both superior recti; when down, both the inferior. The superior oblique muscle acting alone will roll the front of the eye downwards and outwards with a certain amount of rotation; the inferior oblique does the reverse. In oblique movements two of the recti are concerned, an upper or lower with an inner or outer; at the same time one of the oblique also always contracts. Movements of rotation rarely, if ever, occur alone. The natural combined movements of the eyes by which both are directed simultaneously towards the same point depends on the accurate adjustment of all its nervo-muscu- lar apparatus. When the co-ordination is deficient the person is said to squint. A left external squint would be caused by paralysis of the inner rectus of that eye, for then, AJ!^ATO^rY OF EYEBALL. 485 when the eyeball had been turned out by the external rec- tus, it would not be brought back again to its median position. A left internal squint would be caused, similarly, by pai'alysis of the left external rectus; and probably by disease of the sixth cranial nerve or its brain-centres. Dropping of the upper eyelid (ptosis) indicates paralysis of Fig. 134. — The left eyeball iu horizontal section from before back. 1, sclerotic; 2, junction of sclerotic and cornea: 3, cornea; 4, 5. conjunctiva: 6 posterior elastic layer of cornea: 7, ciliary muscle: 10. choroid; 11. 13 ciliary processes; 14, iris; 15, retina; 16, optic nerve; 17, artery entering retina m optic nerve; 18, fovea centralis: 10. region where sensory part of retina ends; •^•^, suspensory ligament; 23 is placed in the canal of Petit and the line from 25 points to it; 24, the anterior part of the hyaloid membrane: 26, 27, 28 are placed on the lens; 28 points to the line of attachment aroimd it of the suspensory ligament: 29, vitreous himior; 30, anterior chamber of aqueous humor; 31, posterior chamber of aqueous humor. its elevator muscle (p. 481), and is often a serious syinjDtom, as pointing to disease of the brain-parts from which it is innervated. The Globe of the Eye is on the whole spheroidal, but consists of segments of two spheres (see Fig. 124). a portion of a sphere of smaller radius forming its anterior transparent part and being set on to the front of its posterior segment, which is part of a larger sphere. From before back it 486 THE HUMAN BODY. measures about 22.5 milliTnetens (^ inch), and from side to side about 25 millimeters (1 inch). Except when looking a-t near objects, the antero-posterior axes of the eyeballs are nearly parallel, but the optic nerves diverge considerably (Fig. 123); each joins its eyeball, not at the centre, but about 2.5 mm. (^V inch) on the nasal side of the posterior end of its antero-posterior axis. In general terms the eyeball may be described as consisting of three coats and three refrad- irifj media. The outer coat, 1 and 3. Fig. 124, consists of the sderofic and the cornea, the latter being transparent and situated in front; the former ie opaque and white and covers the back and sides of the globe and part of the front, where it is seen between the eyelids as the white of the eye. Both are tough and strong, being composed of dense connective tis- sue. The white of the eye and the cornea are also covered over by a thin layer of the conjunctiva, 4 and 5. Behind the proper connective-tissue layer, 3, of the cornea is a thin structureless membrane, 6, lined inside by a single layer of epithelial cells; it is called the membrane of Descemst, or posterior elastic layer. The second coat consists of the clioroid, 9, 10, the ciliary processes, 11, 13, and the iris, 14. The choroid consists mainly of blood-vessels supported by loose connective tissue containing numerous corpuscles, which in its inner layers are richly filled with dark brown or black pigment granules. Towards the front of the eyeball, where it begins to dimin- isli in diameter, the choroid is thrown into plaits, the ciliary processes, 11, 12, 13. Beyond these it continues as the iris, which forms the colored part of the eye which is seen through the cornea; and in the centre of it this a circular aperture, the pupil: so the second coat does not, like the outer one, comj)letely envelop the ball. In the iris are two sets of plain muscular fibres; a circular around the margin of the i)upil and narrowing it when they contract; the other set radiate from the inner to the outer margin of the iris and by their contraction dilate the pupil. The pigment in the iris is yellow, or of lighter or darker brown, according to the color of the eve, and more or less HISTOLOGY OF RETINA. 487 abundant according as the eye is black, brown, or gray. In blue eyes tlie pigment i;3 confined to the deeper layers and modified in tint by light absorption in the anterior color- less strata through which the light passes. The third coat of the eye, the retina, 15, is its essential portion, being the part m which the light produces those changes that give rise to impulses in the optic nerve. It is a still less complete envelope than the second tunic, ex- tending forwards only as far as the commencement of the ciliary processes, at least in its typical form. It is extremely soft and delicate and, when fresh, transparent. Usually when an eye is opened it looks colorless; but by taking proper precautions the natural purjDle color of some of its outer layers can be seen. Its most external layer, more- over, is composed of black pigment cells. On its inner surface two parts, different from the rest, can be seen in a fresh eye. One is the point of entry of the optic nerve, 16, the fibres of which, penetrating the sclerotic and choroid, spread out in the retina. At this jolace the retina is whiter than elsewhere and presents an elevation, the 02)tic mound. The other peculiar region is the yellow sjjot {macula lutea), 18, which lies nearly at the posterior end of the axis of the eyeball and therefore outside the optic mound; m its centre the retma is thinner than elsewhere and so a •!^\i{ fovea centralis), 18, is formed. This appears black, the thinned retina there allowing the choroid to be seen through it more clearly than elsewhere. In Fig. 125 is represented the left retina as seen from the front, the elliptical darker patch about the centre being the yellow spot, and the white circle on one side, the optic mound. The vessels of the retina arise from an artery (17, Fig. 124) which runs m with the optic nerve and from which branches diverge as shown in Fig. 125. The Microscopic Structure of the Retina. A simplified stratum, continuous with the proper retina, and formed of a layer of nucleated columnar cells is continued over the ciliary processes; elsewhere the membrane has a very com^ plex structure and a section taken, except at tlie yellow spot or the optic mound, shoM's ten layers, partly sensory 488 THE HUMAN BODY apparatuses and nerve-tissues, and partly accessory struc- tures. Beginning (Fig. 12G) on the inner side we find, first, the itUernal llmUuig monhrane, 1, a thin structureless layer. Next comes the nerve-fibre layer, 2, formed by radiating fibres of the optic nerve; third, the nerve-cell layer, 3; Fig. 135.— The left retina as it would be seen if the front part of the eyeball with the lens and vitreous humor were removed. fourth, the i7iner molecular layer, 4, consisting partly of very fine nerve-fibrils, and largely of connective tissue; fifth, the inner granular layer, 5, composed of nucleated cells, with a small amount of protoplasm at each end, and a nucleolus. These granules, or at any rate the majority of tliem, have an tjiner process running to the inner molecular layer and an outer running to, 6, the outer molecular layer, which is thinner than the inner. Then comes, seventh, the rod and cone fibre layer, 7, or outer granular layer; composed of thick and thin fibres on each of which is a conspicuous nucleus witli a nucleolus. Next is the thin external limiting membrane, 8, perforated by apertures through which the rods and. cones, 9, of the ninth layer join the fibres of the seventh. Outside of all, next tlie choroid, is the pigmentary layer, 10. In addition, cer- HISTOLOGY OF RETINA. 489 taiu libres rim vertically through the retina from the inner to the outer limiting membrane; they are known as the radial fibres of Miiller and give off lateral branches, which are especially numerous in the molecular layers. Fig. 136. — a section througlti the retina from its anterior or inner surface, 1, in contact witli the hyaloid memh ane, to its outer. 10, in contact witli the choroid. 1, internal limitina: membrane; 2, nerve fibre layer: 3, nerve -eel I layer; 4, inner molecular layer; 5, inner granular layer; 6, outer molecular layer; 7, outer granular layer; 8, external luniting membrane; 9, rod and cone layer; 10, pigment-cell layer. On account of the way in which the supporting and essen- tial parts are interwoven in the retina it is not easy to track 490 THE UUMAN BODY. the latter through it. We shall find, however (Cha]). XXXri.). tliat light ih-st acts u])on the rod and cone layer, traversing all the thickness of inner strata of tlie retina to reach this, before it can start those changes which result in visual sensations; and it is thereiore probable that the rods and cones are m direct continuity with the optic nervc- tibres. The limiting membranes, with the fibres of Miiller and their branches, are undoubtedly accessory. Each rod and cone consists of an outer and an i7iner segment. The outer segments of both tend to split up trans- versely into disks and are very similar, except that those of the rods are longer than those of the cones and do not taper as the latter do. The inner segments of the cones are swollen, while those of the rods are narrow and nearly cy- Imdrical. Over most of the retina the rods are longer and much more numerous than the cones, but near the ciliary processes they cease before the cones do, and in the yellow spot elongated cones alone are found. In this region the whole retina is much modified; at its margin all the layers are thickened but especially the nerve-cell layer, Avliich is here six or seven thick, while elscAvhere the cells are found m but one or two strata. All the fibres also are oblique, reaching in to become continuous with the cones of the central pit, Avhich are long, slender and very closely packed. In the fovea itself all the layers, except that of the cones, thin away, and so the depression is produced. The fovea is the seat of most acute vision; when we look at an object we always turn our eyes so that the light proceeding fi'om it shall be focussed on this spot. Where the optic neiwe enters, all the layers but the nerve-fibre layer, which is very thick, and the internal limiting membrane, are absent. The blood-vessels of the retina lie in the nerve-fibre and nerve-cell layers. The Refracting Media of the Ejne are, in succession from before back, the cornea, the aqueous humor, the crystalline lens, and the vitreous humor. The aqueous humor fills the space between the front of the lens, 28, and the back of the cornea. This space is in- completely divided by the iris into an anterior chamber. THE REFR ACTING MEDIA OF THE EYE. 491 30. and a posterior, 31 (Fig. 124). Chemicalh', the aque- ous humor consists of water holding in solution a small amount' of solid matters, mainly common salt. The crystalline lens (?-S, 26, 27) is colorless, transparent, and biconvex, with its anterior surface less curved than the posterior. It is surrounded by a capsule, and the inner edge of the iris lies in contact with it in front. In consist- ence it is soft, but its central layers are rather more dense than the outer. The vitreous liumor is a soft jelly, enveloped in a thin cai)sule, the hyaloid memhrane. In front, this membrane splits into two layers, one of which, 22, passes on to be fixed to the lens a little in front of its edge. This layer is known as the susjjcnsory ligament of the lens; its line of at- tachment around that organ is not straight but sinuous as represented by the curved line between 28 and 26 in Fig. 124. The space between the two layers into which the hyaloid splits is the canal of Petit. The vitreous humor consists mainly of water and contains some salts, a little albumin, and some mucin. It is divided up, by delicate membranes, into compartments in which its more liquid portions are imprisoned. The Ciliary Muscle. Eunning around the eyeball where the cornea joins the sclerotic is a little vein called the canal of Schlenim; it is seen in section at 8 in Fig. 124. Lying on the inner side of this canal, just where the iris and the ciliary processes meet, there is some 2:ilain muscular tissue, imbedded mainly in the middle coat of the eyeball and forming the ciliary muscle, which consists of a radial and a circular portion. The radial part is much the larger, and arises m front from the inner surface of the sclerotic; the fibres pass back, spreading out as they go, and are inserted into the front of the choroid opposite the ciliary processes. The circular part of the muscle lies around the outer rim of t he iris. The contraction of the ciliary muscle tends to pull forward (radial fibres) and press inward (circular fibres) the front part of the choroid, to which the back part of the sus- pensory ligament of the lens is closely attached. In this way the tension exerted on the lens bv its ligament is diminished. 492 THE HUMAN BODY. The Properties of Light. Before proceeding to the study of the eye as tin opticul instrument, it is necessary to recall brieiiy certain properties of liglit. Light is considered as a form of movement of the particles of an hypothetical medium, or ether, the vibrations being in planes at right angles to the line of propagation of the light. When a stone is thrown into a pond a series of circular waves travel from that point in a horizontal direc- tion over the water, while the particles of water themselves move up and down, and cause the surface inequalities which we see as the waves. Somewhat similarly, light-Avaves spread out from a luminous point, but in the same medium travel equally in all directions so that the point is surrounded by shells of spherical waves, instead of rings of circular waves traveling in one plane only, as those on the surface of the water. Starting from a luminous point light would travel in all directions along the radii of a sphere of which the point is the centre; the light propagated along one such radius is called a ray, and in each ray the ethereal particles swing from side to side in a jilane perpendicular to the direction of the ray. Taking a particle on any ray it would swing aside a certain distance from it, then back to it again, and across for a certain distance on the other side; and then back to its original position on the line of the ray. Such a movement is an oscillation, and takes a certain time; in lights of certain kinds the periods of oscilla- tion are all the same, no matter how great the extent or arrqjlitude of the oscillation; just as a given pendulum will always comj)lete its swing in the same time no matter whether its swings be great or small. Light composed of rays in which the periods of oscillation are all equal is called monochromatic or simple light, while light made of a mixture of oscillations of different periods is called mixed or compound light. If monochromatic light is steadily emitted from a point, then, at certain distances along a ray, we come to particles in the s,2imQ phase ot oscillation, say at their greatest dis- tance from their position of rest; just as in" the concentric waves seen on the water after thi'owing in a stone we would PROPERTIES OF LIGHT. 493 along uny radius meet, at intervals, with water raised most above its horizontal ])lane as the crest of a wave, or depressed most below it as the holloiv of a wave. The distance along the ray from crest to crest is called a ivave-lengtli and is always the same in any given simple light; but differs in different-colored lights; the briefer the time of an oscillation the less the wave-length. When light falls on a polished surface separating two transparent media, as air and glass, part of it is reflected or turned back into the first medium; part goes on into tlie second medium, and is commonly deviated from its original course or refracted. The original ray falling in the surface is the incident ray. Let A B (Fig. 127) be the surface of separation ; a x the incident ray; and CD the perpendicular or normal to the surface at the point of incidence: ax Cwill then be the angle of incidence. Then the reflected ray makes an angle of reflection with the normal which is equal to the angle of incidence: and the reflected ray lies in the same plane as the inci- dent ray and the normal to the surface at x. The re- fracted ray lies also in the same plane as the normal and the incident ray, but does not continue in its original direc- tion, xf; if the medium below A B be denser than that above it, the refracted ray is bent in the direction xd nearer the normal, and making with it an angle of refrac- tion, Dxd, smaller than the angle of incidence, ax C. If, on the contrary, the second medium be less dense than the first, the refracted ray xg is bent away from the normal, and makes an angle of refraction, Dxg, greater than the angle of incidence. The ratio of the sine of the angle of Fio. 127.— Diagram illustrating the refraction of light. A B. surface of separation between two transparent media; CD. the perpendicular to the surf'ace at the point of incidence, .c; a X. incident ray ; .r d. refracted ray. if the second medium be denser than the first; xg, refracted ray. if the second medium is less refractive than the first. The reflected ray is not repre- sented, but would make an angle with Cx equal to the angle a x C. 494 THE HUMAN BODY. incidence to that of the angle of refraction is always the same for the same two media with light of the same wave- length. When the first medinm is air the ratio of the sine of the angle of refraction to that of the angle of incidence is called the refractive index of the second medium. The greater this refractive index the more is the refracted r;iy deviated from its original course. Rays which fall perpen- dicularly on the surface of separation of two media pass on without refraction. The shorter the oscillation periods of light-rays the mxore they are deviated by refraction. Hence mixed light when Fig. 128.— Diagram illustrating the dispersion of mixed light by a prism. sent through a prism is spread out, and decomposed into its simple constituents. For let a x (Fig. 128) be a ray of mixed light composed of a set of short and a set of long ethereal waves. When it falls on the surface A B of the prism, that portion which enters will be refracted towards the normal ED, but the short waves more than the longer. Hence the former will take the direction x y, and the latter the direction x z. On emerging from the prism l)oth rays will again be refracted, bat now from the no»-- mals Fy and G z, since the light is passing from a denser to a rarer medium. But again the ray x y, made up of shorter waves, would be most deviated, as in the direction y v, and REFRACTION BY LENSES. 495 the long waves less, in the direction z r. If a screen were pnt at S S, we would receive on it at separate points, v and r, the two simple lights which were mixed together in the compound incident ray a x. Such a separation of light-rays is called disjjersion. Ordinary white light, such as that of the sun, is com- posed of ethereal vibrations of all possible lengths. Hence when such light is sent through a prism it gives a contin- uous band of light-rays, known as the solar spectrum, reaching from the least refracted to the most refracted and shortest. The exceptions to this statement due to Frauen- hofer's lines (see Physics) are unessential for our present purpose. All of the simple lights into which the compound solar light is thus separated do not, however, excite in us visual sensations when they fall into the eye, but only cer- tain middle ones. If solar light were used with the prism, Fig. 125, certain least refracted rays between r and 8 would not be seen, nor the most refracted between v and S; while between v and r would stretch a luminous band exciting in us the series of colors red (due to the least refracted visible rays), successively through orange, yellow, green, bright blue, and indigo, to violet, which latter is the sensation aroused by the most refrangible visible rays. The still shorter waves beyond the violet, are known mainly by their chemical eifects and make up what are called the actinic rays; the longer invisible waves, beyond the red, exert a powerful heating influence and compose the thermal or darh heat rays. The eye, as an organ for making known to us the existence of ethereal vibrations, has, therefore, only a limited range. Refraction of Light by Lenses. In the eye the refract- ing media have the form of lenses thicker in the centre tlian towards the periphery; and we may here confine our- selves therefore to such converging lenses. If simple light from a point A, Fig. 122, fall on such a lens its rays, emerging on the other side, will take new directions after refraction and meet anew at a point, a, after which they again diverge. If a screen, r r, be held at a it will therefore receive an image of the luminous point A. For every con- 496 THE HUMAN BODY. Fig. 129.— Diagrram illustrating the formation of an image by a converging lens. verging lens there is such a point behind it at which the rays from a given point in front of it meet: the point of meeting is called the conjugate focus of the point from which the rays start. If instead of a luminous point a luminous object be placed in front of the lens an image of the object will be formed at a certain distance behind it, for all rays proceeding from one point of the object will meet m the conjugate focus of that point behind. The image is inverted, as can be readily seen from Fig. 129. All rays from the point A of the object meet at the point a of the image; those from B at l, and those from intermediate points at intermediate positions. If the single lens were replaced by several combined so as to form an ojitical system the general result would be the same, provided the system were thicker in the centre than at its perijohery. The Camera Obscura, as used by photographers, is an instrument which serves to illustrate the formation of images by converging systems of lenses. It consists of a box blackened inside and having on its front face a tuJje containing the lenses; the posterior wall is made of ground glass. If the front of the instrument be directed on ex- terior objects, inverted and diminished images of them will be formed on the ground glass; those images are only well defined, at any one time, which are at such a distance in front of the instrument that the conjugate foci of points on them fall exactly on the glass behind the lens: objects nearer or farther off give confused and indistinct images; but by altering the distance between the lenses and the ground glass, in common language "focusing the instru- ment," either can be made distinct. For near objects the lenses must be farther from the surface on which the image is to be received, and for distant nearer. The reason of this may readily be seen from Fig. 130. If the system of lenses brings the parallel rays a c and h d, pro- ceeding from an infinitely distant object, to a focus at x, REFRACTION IN THE ETE. 497 then the diverging rays/c and fcl, proceeding from a nearer point, will be harder to bend round, so to speak, and will not meet until a point y, farther behind the system than X IS. The more divergent the rays, or what amounts to the same thing, the nearer the points they proceed from, the farther behind the refracting system will y be. F<* Fig. 130,— Diagram illustrating the need of " focusing" in an optical instru nient. The eye is such a system, made up of the four refracting media, cornea, aqueous humor, lens, and vitreous humor. These four media are, however, reduced to three prac- tically, by the fact that the indices of refraction of the cornea and aqueous humor are the same, so that they act together as one converging lens. The surfaces at which refraction occurs are — (1) that between the air and ihe cornea, (2) that between the aqueous humor and the front of the lens, (3) that between the vitreous humor and the back of the lens. The refractive indices of those media are — the air, 1; the aqueous humor, 1.3379; the lens (average), 1.4545; the vitreous humor, 1.3379. From the laws of the refraction of light it therefore follows that (Fig. 131) the rays c d will at the corneal surface be refracted towards the normals N, N, and take the course cl e. At the front of lenses they will again be refracted towards the normals to that surface and take the course e/; at the back of the lens, ])assing from a more refracting to a less refracting medium, they will be bent from the normals N" and take the course / g. If the retina be there, these jjarallel rays will therefore be brought to a focus on it. In the resting condition of the natural eye that is what happens to jDarallel rays entering it; and, since distant objects send into the eye rays which are practicallv parallel, such o1)jects are seen distinctly 498 THE HUMAN BOD 7. without any effort; all rays emanating from a point of the object meet again in one point on the retina. Accommodation. Points on near ob ject s send into the eye divei'ging rays: those therefore would not come lo a focus on the retina but behind it, and would not be seen distinctly, did not some change occur in the eye; since we can see them quite plainly if we choose (unless they be very near mdeed), there must exist some means by which the eye is adapted or accommodated for looking at objects at different distances. That some change does occur one can, also, readily prove Fig. 131.— Diagram illustrating the .surfaces at which light is refracted in the eye. by observing that we cannot see distinctly, at the same moment, both near and distant objects. For example standing at a window, behind a lace curtain, we can if we choose look at the threads of the lace or the houses across the street; but when we look at the one we only see the other indistinctly; and if, after looking at the more distant object, we look at the nearer we ex2:)erience a distinct sense of effort. It is clear, then, that something in the eye is different in the two cases. The resting eye, suited for dis- tinctly seeing distant objects, might conceivably be accom- modated for near vision in several ways. The refracting ACCOMMODATION. 499 indices of its media might be increased; that of course does not happen; the physical properties of the media are the same in both cases: or the distance of the retina from the refracting surfaces might be increased, for example by compression of the eyeball by the muscles around it; how- ever, experiment shows that changes of accommodation can be brought about in the fresh excised eyes of animals, in which no such compression is jDossible; we are thus reduced to the third explanation, that the refracting surfaces, or some of them, become more curved, and so bring more diverging rays sooner to a focus; since a lens of smaller curvature is more converging than one of greater curvature composed of the same material. Observation shows that this is what actually happens: the corneal surface remains unchanged when a near object is looked at after a distant one, but the anterior sur- face of the lens becomes considerably more convex and the posterior slightly so. As already pointed out when light meets the separating surface of two media some is reflected and some refracted (p. 493). If, therefore, a person be taken into a dark room and a candle held on one side of his i-I^^'Bhz!^}^^ images ot a candle- eye, while he looks at a distant object an 5^"^^ as seen re- •J ' . _ "> fleeted from the re- observer can see three images of its flame fracting media of in his pupil, due to that part of the light reflected from the surfaces between the media. One (a. Fig. 133) is erect and bright, reflected from the convex mirror formed by the cornea; the next, h, is dimmer and also erect; it comes from the front of the lens. The third, c, is dim and inverted, being reflected from the concave mirror (see Physics) formed by the back of the lens. If now the observed eye looks at a near object in the same line as the distant point previously looked at, it is seen that the image due to corneal reflection remains unchanged; that due to light from the front of the lens becomes smaller and brighter, indicating (see Ph3'sics) a greater convexity of the reflecting surface; the image from the back of the lens also becomes very slightly smaller, indicating a feebly increased curvature. a 6 d 500 THE HUMAN BODY. Accommodation is brouglit about mainly by the ciliavy muscle. In tlie resting eye it is relaxed and the suspensory ligament of the lens is taut, and, pulling on its edge, drags it out laterally a little and flattens its surfaces, especially the anterior, since the ligament is attached a little in front of the edge. To see a nearer object the ciliary muscle is contracted, and according to the degree of its contraction slackens the suspensory ligament (p. 491), and then the elastic lens, relieved from the lateral drag, bulges out a little in the centre. Short Sight and Long Sight. In the eye the range of accommodation is very great, allowing the rays from jioints infinitely distant up to those from points about eight inches in front of the eye to be brought to a focus on the re- tina. In the normal eye par- allel rays meet on the retina when the ciliary muscle is completely relaxed {A, Fig. 133). Such eyes are emme- tropic. In other eyes the eye- ball IS too long from before back; in the resting state par- allel rays meet in front of the retina (B). Persons with such eyes, therefore, cannot see distant objects distinctly without the aid of diverging (concave) spectacles; they are short- sighted or myopic. Or the eyeball may be too short from before back; then, in the resting state, parallel rays are brought to a focus behind the retina (C). To see even infinitely distant objects, such persons must therefore use their accommodating apparatus to increase the converging power of the lens; and when objects are near they cannot, with the greatest effort, bring the divergent rays proceeding from them to a focus soon enough. To get distinct retinal images of near objects they therefore need converging (con- vex) spectacles. Such eyes are called hypermetropic, or in common language long-sighted. Fig. 133— Diagram illustrating the path of parallel ravs after entering an emmetropic {A), a myopic {B), and a hypermetropic (C) eye. HYGIENE OF THE EYE. 501 Hygienic Remarks. Since muscular effort is needed by the normul eye to see near objects, it is clear why the pro- longed contemplation of such is more fatiguing than look- ing at more distant things. If the eye be hypermetropic still more is this apt to be the case, for then the ciliary muscle has no rest when the eye is used, and to read a book at a distance such tliat enough light is reflected from it into the eye in order to enable the letters to be seen at all, requires an extraordinary effort of accommodation. Such persons complain that they can read well enough for a time, but soon fail to be able to see distinctly. This kind of weak sight should always lead to examination of the eyes by an oculist, to see if glasses are needed; otherwise severe neu- ralgic pains about the eyes are apt to come on, and the overstrained organ may be permanently injured. Old per- sons are apt to have such eyes; but young childen frequently also possess tliem, and if so should at once be provided with spectacles. Short-sighted eyes appear to be much more common now than formerly, especially in those. given to literary pursuits. Myopia is rare among those who cannot read or who live mainly out of doors. It is not so apt to lead to per- manent injury of the eye as is the opposite condition, but the effort to see distinctly objects a little distant is apt to produce headaches and other symptoms of nervous exhaustion. If the mj^opia become gradually worse the eyes should be rested for several months. Short-sighted persons are apt to have, or acquire, peculiarities of appear- ance: their eyes are often prominent, indicative of the abnormal length of the eyeball. They also get a habit of '^ screwing" up the eyelids, probably an indication of an effort to compress the eyeball from before back so that distant objects may be better seen. They often stoop, too, from the necessity of getting their eyes near objects they want to see. The acquirement of such habits may be usually prevented by the use of proper glasses. On the other hand ''it is said that myopia even induces peculiari- ties of character, and that myopes are usually unsuspicious and easil}^ pleased; being unable to observe many little 502 THE HUMAN BODY. matters in the demeanor or expression of those with whom they converse, which, being noticed by those of quiclvcr siglit, might induce feelings of distrust or annoy- ance." In old age the eyeball tends to become flattened; hence cmmetro])ic eyes become hypermetropic and old persons are Ui^ually " long-sighted " and need convex glasses. 8nch a flattening of the eyeball is of course a relief to the myopic eye; and so short-sighted persons can frequently, when old, still read without glasses. But this is poor com- pensation for the mistiness with which everything around them, except very near objects, has been seen throughout their previous life. In all forms of deficient accommodation too strong glasses will injure the eyes irreparably, increasing the defects they are intended to relieve. Skilled advice should there- foi-e be invariably obtained in their selection, except per- haps in the long-sightedness of old age when the sufferer may tolerably safely select for himself any glasses that allow him to read easily a book about 30 centimeters (12 inches) from the eye. As age advances stronger lenses must of course be obtained. Optical Defects of the Eye. The eye, though it an- swers admirably as a physiological instrument, is by no means perfect optically; not nearly so good, for example, as a good microscope objective. The main defects in it are due to — 1. Chromatic Aberration. As already pointed out the rays at the violet end of the solar spectrum are more re- frangible than those at the red end. Hence they are brought to a focus sooner. The light emanating from a point on a white object does not, therefore, all meet in one point on the retina; but the violet rays come to a focus first, then the indigo, and so on to the red, farthest back of all. If the eye is accommodated so as to bring to a focus on the retina j^arallel red rays, then violet rays from the same source will meet half a millimeter in front of it, and cross- ing and diverging there make a little violet circle of diffu- sion around the red point on the retina. In optical instru- OPTICAL DEFECTS OF THE EYE. 503 nients this defect is remedied h\ combining together lenses made of different kinds of glass; such compound lenses are called achromatic. The general result of chromatic aberration, as may be seen in a bad opera-glass, is to cause colored borders to ap- pear around the edges of the images of objects. In the eye we usually do not notice such borders unless we especially look for them; but if, while a white surface is looked at, the edge of an opaque body be brought in front of the eye so as to cover half the pupil, colorations will be seen at its margin. If accommodation is inexact they appear also when the boundary between a white and a black sur- face is observed. The phenomena due to chromatic aberration are much more easily seen if light containing only red and violet rays be used instead of Avhite light con- taining all the rays of intermediate refrangibility. Ordi- dinary blue glass only lets through these two kinds of rays. If a bit of it be placed over a very small hole in an opaque shutter and the sunlight be suffered to enter through the hole, it will be found that with one accommodation (that for the red rays) a red point is seen with a violet border, and with another (that at which violet rays are brought to a focus on the retina) a violet point is seen with a red aureole. 2. Spherical Aberration. It is not quite correct to state that ordinary lenses bring to a focus in one point behind them rays proceeding from a point in front, even when these are all of the same refrangibility. Convex lenses whose surfaces are segments of spheres, as are those of the eye, bring to a focus sooner the rays which pass through their marginal than those through their central parts. If the rays proceeding from a point and traversing the lateral part of a lens be brought to a focus at any point, then those passing through the centre of the lens will not meet until a little beyond that point. If the retina receive the image formed by the peripheral rays the others Avill form around this a small luminous circle of light — such as would be formed by sections of the cones of converging rays in Fig. 122, taken a little in front of r r. This defect is found in all 504 THE HUMAN BODY. optical instruments, as it is impossible in practice to grind lenses of the non-spherical curvatures necessary to avoid it. In our eyes its effect is to a large extent corrected in the following ways — (a) The opaque iris cuts off many of the external and more strongly refracted rays, preventing them from reaching the retina, (b) The outer layers of the lens are less refracting than its central; lience the rays passing through its peripheral parts are less refracted than those passing nearer its axis. 3. Ii'Tegularities in Curvature. The refracting surfaces of our eyes are not even truly spherical; this is especially the case with the cornea, which is very rarely curved, to the same extent in its vertical and horizontal diameters. Sup- pose the vertical meridian to be tlie most curved; then the rays proceeding from points along a vertical line will be brought to a focus sooner than those from points on a hori- zontal line. If the eye is accommodated to see distinctly the vertical line, it will see indistinctly the horizontal and vice versa. Few people therefore see equally clearly at once two lines crossing one another at right angles. The pheno- menon is most obvious, however, when a series of concentric circles (Fig. 134) is looked at: then when the lines appear sharp along some sectors, they are dim along the rest. When this defect, known as astigmatism, is marked it causes serious troubles of vision and re- quires peculiarly shaped glasses to counteract it. 4. Opaque Bodies in the Refracting Media. In diseased eyes the lens may be opaque {cataract) and need removal; or opacities from ulcers or wounds may exist on the cornea. But even in the best eye there are apt to be small opaque bodies in the vitreous \\Vimox caxi?>mgr)iuscm volitantes; that is, the appearance of Fig. 134. OPTICAL DEFECTS OF THE EYE. 505 minute bodies floating in space outside tlie eye, but chang- ing their position when the position of the eye changes, by which fact their origin in interr al causes may be recog- nized. Many persons never see them until their attention is called to their sight by some weakness of it, and then they think they are new phenomena. Visual phenomena due to causes in the eye itself are called entopficj the most interesting are those due to the retinal blood-vessels (ChajD. XXXII. ). Tears, or bits of the secretion of the Meibo- mian glands, on the front of the eyeball often cause distant luminous objects to look like ill-defined luminous bands or patches of various shape. The cause of such appearances is readily recognized, since they disappear or are changed after winking. CHAPTER XXXII. THE EYE AS A SENSORY APPARATUS. The Excitation of the Visual Apparatus. The excita- ble visual apparatus for each eye consists of the retina, the optic nerve, and the brain-centres connected with the latter; however stimulated, if intact, it causes visual sensations. In the great majority of cases its excitant is objective light, and so we refer all stimulations of it to that cause, unless we have special reason to know the contrary. As already pointed out (p. 468) pressure on the eyeball causes a lumi- nous sensation (phospheite), which suggests itself to us as dependent on a luminous body situated in space where such an object must be in order to excite the same part of the retina. Since all rays of light penetrating the eye, except m the line of its long axis, cross that axis, if Ave press the outer side of the eyeball we get a visual sensa- tion referred to a luminous body on the nasal side; if we press below we see the luminous patch above, and so on. Of course different rays entering the eye take different paths through it, but on general optical principles, which cannot here be detailed, we may trace all oblique rays through the organ by assuming that they meet and leave the optic axis at what are known as the nodal ijomts of the system; these {kh'. Fig. 135) lie near together in the lens. If we want to find where rays of light from A will meet the retina (the eye being properly accommodated for seeing an object at that distance) we draw a line from ^ to ^ (the first nodal point) and then another, parallel to the first, from h' (the second nodal point) to the retina. The nodal points of the eye lie so near together that for practical purposes we may treat them as one {Jc, Fig. 136), POSITION OF RETIXAL IMAGES. 507 placed near the back of the lens. By manifold experience we have learnt that a lumiuous body {A Fig. 136) which we see, always lies on the prolongation of the hue joining the excited part of the retina, a, and the nodal point, k. Hence Fig. 135.— Diagram illustrating the points at which incident rays in the eye meet the retina, x x, optic axis; k. first nodal point; k\ second uodal point; 6. point where the image of B would be formed, were the eye properly accom- modated for it; a, tlie retinal point where the image of A would be formed. any excitation of that part of the retina makes us think of a luminous body somewhere on the line a A, and, similarly, any excitation of h, of a body on the line h B or its pro- longation. It is only other conflicting experiences, as that Fig. 136.— Diagrammatic section through the eyeball, nodal point. a; a-, optic axis; fc, with the eyes closed external bodies do not excite visual sensations, and the constant connection of the ])ressure felt on the eyelid with the visual sensation, that enable us when we press the eyeball to conclude that, in spite of what 508 TEE HUMAN BODY. we seem to see, the luminous sensation is not due to objective light from outside the eye. The Idio-Retinal Light. The eyelids are not by any means perfectly opaque; in ordinary daylight they still allow a considerable quantity of light to penetrate the eye, as any one may observe by passing his hand in front of the closed eyes. But even in a dark room with the eyes completely covered up so that no objective light can enter them, there is still experienced a small amount of visual sensation due to internal causes. The field of vision is not absolutely dark but slightly lummous, with brighter fleeting patches trav- ersing it. These are especially noticeable, for example, in trying to see and grope one's way with the eyes open up a perfectly dark staircase. Then the luminous patches attract special attention because they are apt to be taken for the signs of objective realities; they become very manifest when any sudden jar of the Body, due for example to knocking against something, occurs; and have no doubt given rise to many ghost stories. These visual sensations felt in the absence of all external stimulation of the eyes, may for con- venience be spoken of as due to the idio-retinal light. The Excitation, of the Visual Apparatus by Light. Light only excites the retina when it reaches its nerve end Fig. 137. organs, (he rods and cones. The proofs of this are several. 1. Light does not aronse visval sensations when it falls directly on tl\e fibres of the optic nerve. Where this nerve enters (p. 490) is a retinal part possessing only nerve-fibres, and this part is blind. Close the left eye and look steadily with the right at the cross in Fig. 137, holding the book FUNCTION OF RODS AND CONES. 509 vertically in front of the face, and moving it to and fro. It will be found that at about 25 centimeters (10 inches) off the white circle disappears; but when the page is nearer or farther, it is seen. During the experiment the gaze must be kept fixed on the cross. There is thus in the field of vision a blind spot, and it is easy to show by measurement that it lies where the ojDtic nerve centres. When the right eye is fixed on the cross, it is so directed that rays from this fall on the yellow spot, y, Fig. 138. The rays from the circle then cross the visual axis at the nodal point, n, and meet the retina at o. If the distance of the eye from the paper be /, and that of the nodal point from the retina (which is 15 mm. ) be F, then the dis- tance, on the paper, of the cross from the circle will be to the distance of y from o as / is to F. Measurements made in this way show that the circle disappears when its image is thrown on the entry of the optic nerve, which lies to the nasal side of the yellow spot (p. 487). 3. The above experiment having shown that light does not act directly on the optic nerve-fibres any more than it does on any other nerve-fibres, we have next to see in what part of the retina those changes do first occur which form the link between light and nervous impulses. They occur in tlie outer part of the retina, in the rods and cones. This is jn'oved by what is called Purkinje's experiment. Take a candle into a dark room and look at a surface not covered with any special pattern, say a whitewashed wall or a plain window-shade. Hold the candle to the side of one eye and close to it, but so far back that no light enters the pupil from it; that is so far back that the flame just cannot be seen, but so that a strong light is thrown on the white of the eye as far back as possible. Then move the candle a little to and fro. The surface looked at will appear luminous with reddish-yellow light, and on it will be seen Fig. 138. 510 THE HUMAN BODT. dark branching lines which are the shadows of the retinal vessels. Now in order that these shadows may be seen the parts on which the light acts must be behind them, and therefore in the outer layers of the retina since the vessels lie (p. 490) in its inner strata. If the light is kept steady the vascular shadows soon dis- appear; in order to continue to see them the candle roust I^e kept moving. The explanation of this fact may readily be made clear by fixing the eyes for ten or fifteen seconds on the dot of an " i" somewhere about the middle of thi.s page: at first the distinction between the slightly luminous black letters and the highly luminous white page is very obvious; in other words, the different sensations arising from the strongly and the feebly excited areas of the retina. But if the glance do not be allowed to wander, very soon the letters become indistinct and at last disappear altogether; the whole page looks uniformly grayish. The reason of this is that the powerful stimulation of the retina by the light reflected from the white part of the page soon fatigues the part of the visual apparatus it acts upon; and as this fatigue progresses the stimulus produces less and less effect. The parts of the retina, on the other hand, which receive light only from the black letters are very little stimu- lated and retain their original excitability so that, at last, the feebler excitation acting upon these more irritable parts produces as much sensation as the stronger stimulus acting upon the fatigued parts; and the letters become indistin- guishable. To see them continuously we must keep shift- ing the eyes so that the same parts of the visual apparatus are alternately fatigued and rested, and the general irrita- Ijility of the whole is kept about the same. So, in Purkinje's experiment, if the position of the shadows I'cmain the same, the shaded part of the retina soon be- comes more irritable than the more excited unshaded parts, and its relative increase of irritability makes up for the less light falling on it, so that the shadows cease to be perceived. It is for this same reason that we do not see the vessels under ordinary circumstances. When light, as usual, enters the eye from front through the pupil the PURKiyjE'S EXPERIMENT. 511 Fig. 1S9. .shadows always protect the same jDarts of the retina, and these parts are thus kept sufficiently more excitable than the rest to make up for the less light reaching them through the vessels. To see the latter we must throw the light into the eye in an unusual direction, not through the pupil but laterally through the sclerotic. If v, Fig. 139, be the section of a retinal vessel, ordi- narily its shadow will fall at some point on the line prolonged through it from the centre of the pupil. If a candle be held opposite h it illuminates that part of the sclerotic and from there light radiates and illumines the eye. The sensation we refer to light entering the eye in the usual manner through the pupil, and accordingly see the surface we look at as if it were illuminated. The shadow of v is now cast on an unusual spot c, and we see it as if at the point d on the wall, on the prolongation of the line joining the nodal point, I:, of the eye with c. If the candle be moved so as to illuminate the point b' of the sclerotic, the shadow of V will be cast on c' and will accordingly seem on the wall to move from d to d\ It is clear that if we know how far h is from V , how far the wall is from the eye, and how far the nodal point is from the retina (15 mm. or 0.6 inch), and measure the distance on the wall from d to d' , we can calculate how far c is from c': and then how far the vessel throwing the shadow must be in front of the retinal jiarts perceiving it. In this way it is found that the part seeing the shadow, that is the layer on which light acts, is just about as far behind the retinal vessels as the main vascular trunks of the retina are in front of the rod and cone layer. It is, therefore, in that layer that the light initiates those changes which give rise to nervous impulses; which is further made obvious by the fact that the seat of most acute vision is the fovea centralis, where this layer and the cone-fibres diverging from it alone are found (p. 490). 512 THE HUMAN BODY. When we want to see anything distinctly we always turn our eyes so that its image shall fall on the centre of the yellow spot. The Vision Purple. How light acts in the retina so as to produce nerve stimuli is still uncertain. Recent observa- tions show that it produces chemical changes in the rod and cone layer, and seemed at first to indicate that its action was to produce substances which were chemical excitants of nerve-fibres; but although there can be little doubt that these chemical changes play some important part in vision, what their role may be is at present quite obscure. If a perfectly fresh retina be excised raj)idly, its outer layers will be found of a rich purple color. In daylight this rapidly bleaches, but in the dark persists even when putre- faction has set in. In pure yellow light it also remains unbleached a long time, but in other lights disappears at different rates. If a rabbit's eye be fixed immovably and exposed so that an image of a window is focused on the same part of its retina for some time, and then the eye be rapidly excised in the dark and placed in solution of potash alum, a colorless image of the window is found on the retina, surrounded by the visual jiurple of the rest which is, through the alum, fixed or rendered incapable of change by light. Photographs, or optograms, are thus obtained which differ from the photographer's in that he uses light to produce chemical changes which give rise to colored bodies, while here the reverse is the case. If the eye be not rapidly excised and put in the alum after its exposure, the optogram will disappear; the vision purple being rapidly regenerated at the bleached part. This reproduction of it is due mainly to the cells of the pigmentary layer of the retina. Portions of frogs' retinas raised from this bleach more rapidly than those left in contact Avith it, but become soon purple again if let fall back upon the pigment-cells. It thus seemed as if we had got a clue to the physiological action of light in the eye: but experiments show that animals (frogs) exposed for a long time to a bright light may have their retinas completely bleached and still see very well; they can still unerringly catch flics that come VISION PURPLE. 513 withm their reach; and they can also distinguish colors, or at least some colors, as green. Moreover, the vision purple is only found in the outer segments of the rods; there is none in the cones, and yet these alone exist in the yellow spot of the human eye, which is the seat of most acute vision; and animals, such as snakes, which have only cones in the retina, possess no vision purple and nevertheless see very well. It may be that other bodies exist in the retina which are also chemically changed by light, but the changes of which are not accompanied by alterations in color which we can see; and m the absence of the vision purple seeing might be carried on by means of these, which may be less quickly destroyed by light and so still persist in the bleached retinas of the frogs above mentioned. For the present, however, the question of the part, if any, played in vision by such bodies must be left an open one. The Intensity of Visual Sensations. Light considered as a form of energy may vary in cpiantity; physiologically, also, we distinguish quantitative differences in light as degrees of brightness, but the connection between the in- tensity of the sensation excited and the quantity of energy represented by the stimulating light is jiot a direct one. In the first place some rays excite our visual apparatus more powerfully than others; a given amount of energy in the form of yellow light, for example, causes more powerful yisual sensations than the same quantity of energy in the form of violet light. The ultra-violet rays only become visible, and then very faintly, when all others are suj^pressed; but if they be passed through some fluorescent substance (see Physics), such as an acid solution of quinine sulphate, which, without altering the amount of energy, turns it into ethereal oscillations of a longer period, then the light be- comes readibly perceptible. Even with light-rays of the same oscillation period our sensation is not proportioned to the amount of energy in the light; to the amount of heat, for example, to which it would give rise if all transformed into it. If objective light increase gradually in amount our sensation increases 514 THE HUMAN BODY. also, up to a limit beyond which it does not go, no matter how strong the light becomes; but the increase of sensation takes place far more slowly than that of the light, in accord- ance with the psycho-physical law mentioned on page 473. If we call the amount of light given out by a single candle a, then that emitted by two candles will be 2a; and so on. If the amount of sensation excited by the single candle be A, then that due to two candles will not be %A, and that by three will be far less than ^A. If a white surface, F, Fig. 140, be illuminated by a candle at c and another else- where, and a rod, o, be placed so as to intercept the light from c, we see clearly a shadow, since our eyes recognize the difference in luminosity of this part of the paper, reflect- ^^' ' ing light from one caudle only., from that of the rest which is illuminated by two: that is we tell the sensation due to the stimulus a from that due to the stimulus 2rt. If now a bright lamp be brought in and placed alongside, and its light be physically equal to that of 10 candles, we cease to joerceive the shadow s. That is the sensation aroused by objective light = 12« (due to the lamp and tAVO candles) cannot be told from that due to light = 11«; although the difference of objective light is still la as before. Most persons must have ob- served illustrations of this. Sitting in a room with three lights not un frequently some object so intercepts the light from two as to cast on the Avail two shadoAvs which partly overlap. Where the shadoAvs overlap the Avail gets light only from the third candle; around that,Avhere each shadow is separate, it is illuminated by this and one other candle: and the wall in the neighborhood of the shadows by all three. Objectively, therefore, the difference betAveen the deep shadoAv and half shadoAv is that between the light of one candle and that of two. The difference between the half shadoAvs and the Avail around is that between the light of tAVo and three candles. But as a matter of sensa- tion the difference betAvecn the half shadoAv and the full INTENSITY OF VISUAL SENSATIONS. 515 shadow seems mucli greater than that between the Iialf shadow and the rest of the wall; in other words the differ- ence, a, between a and 2a, is a more efficient stimuhis than the same difference, a, between 2« and So. When the total stimulus increases the same absolute difference is less felt or may be entirely unperceived. An example of this which every one will recognize is afforded by the invisibility of the stars in daytime. On the other hand, as the total stimulus increases or de- creases the same fractional difference of the whole is per- ceived with the same ease; i.e. excites the same amount of sensation. In reading a book by lamplight w^e perceive clearly the difference between the amount of light reflected from the black letters and the white page. If we call the total lamplight reflected by the blank parts lOrt and that by the letters 2a, we may say we perceive with a certain distinctness a luminous difference equal to one fifth of the whole. If we now take the book into the daylight the total light reflected from both the letters and the unprinted part of the page nicreases, but in the same proportion. Say the one now is 50a and the other 10a; although the absolute difference between the two is now 40a instead of 8« we do not see the letters any more plainly than before. The smallest difference in luminous intensity which we can perceive is about -^ of the whole, for all the range of lights we use m carrying on our ordinary occupations. For strong lights the smallest perceptible fraction is con- siderably greater; finally we reach a limit where no increase in brightness is felt. For weak illumination the sensation is more nearly proportioned to the total differences of the objective light. Thus in a dark room an object reflecting all the little light that reaches it appears almost twice as bright as one reflecting only half; which in a stronger light it would not do. Bright objects in general obscurity thus appear unnaturally bright when compared wath things about chem, and indeed often look self-luminous. A cat's eyes, for example, are said to "shine in the dark;" and painters to produce moonlight effects always make the bright parts of a picture relatively brighter, when compared 516 THE HUMAN BODY. with things about them, than would be the case if a sunny scene were to be represented; by an excessive use of white pigment they produce the reLatively great brightness of those things which are seen at all in the general obscurity of a moonlight landscape. The Duration of Luminous Sensations. This is greater than that of the stimulus, a fact taken advantage of in making fireworks: an ascending rocket produces the sen- sation of a trail of light extending far behind the position of the bright part of the rocket itself at the moment, because the sensation aroused by it in a lower part of its course still persists. So, shooting stars appear to have luminous tails behind them. By rotating rajjidly before the eye a disk with alternate white and black sectors we get for each point of the retina on which a part of its image falls, alternating stimulation (due to the passage of white sector) and rest, when a black sector is passing. If the rotation be rapid enough the sensation aroused is that of a uniform gray, such as would be produced if the white and Ijlack were mixed and spread evenly over the disk. In each revolution the eye gets as much light as if that were the case, and is unable to distinguish that this light is made up of separate portions reaching it at intervals: the stimulation due to each lasts until the next begins and so all are fused together. If one turns out suddenly the gas in a room containing no other light, tlie image of the flame persists a short time after the flame itself is extinquished. The Localizing Power of the Retina. As already pointed out a necessary condition of seeing definite ob- jects, as distinguished from the power of recognizing dif- ferences of light and darkness, is that all light entering the eye from one point of an object shall be focused on one point of the retina. This, however, would not be of any use had we not the faculty of distinguishing the stimula- tion of one part of the retina from that of another part. This power the visual apparatus possesses in a very high degree; while with the skin we cannot distinguish from one, two points touching it less than 1 mm. [^ inch) apart, with our eyes we can distinguish two points whose retinal images LOCALIZING POWER OF RETINA. 517 are not more than .004 mm. (.00016 inch) apart. The distance between the retinal images of two points is deter- mined by the "visual angle" under which they are seen; this angle is that included between lines drawn from them to the nodal point of the eye. If a and h (Fig. Ill) are 4 Fig. 141. two points, the image of a will be formed at a' on the pro- longation of the line a n joining a with the node, n. Sim- ilarly the image of b will be formed at h'. If a and h still remaining the same distance apart, be moved nearer the eye to c and (Z,theu the visual angle under which they are seen will be greater and their retinal images will be farther apart, at c and d '. If a and h are the highest and lowest parts of an object, the distance between their retinal images will then depend, clearly, not only on the size of the object, but on its distance from the eye; to know the discriminating power of the retina we must therefore measure the visual angle in each case. In the fovea centralis two objects seen under a visual angle of 50 to 70 seconds can be distinguished from one another: this gives for the distance between the retinal images that above mentioned, and corresponds pretty accurately to the diameter of a cone in that part of the retina. We may conclude, therefore, that when two images fall on the same cone or on two contiguous cones they are not discriminated; but that if one or more un- stimulated cones intervene between the stimulated, the points may be perceived as distinct. The diameter of a rod or cone, in fact, marks the anatomical limit up to which we can by practice raise our acuteness of visual discrimina- tion; and in the yellow spot which we constantly use all our lives in looking at things which we want to see dis- tinctly, we have educated the visual apparatus up to about 518 THE HUMAN BODY. its highest power. Elsewhere on the retina our discrimi- nating power is miicli less and diminishes as the distance from the yellow spot increases. This is partly due, no doubt, to a less sensibility of those retinal regions, such as, by other facts, is proved to exist, but in part no doubt is also due to a want of practice. The more peripheral the retinal region the less we have used it for such purposes. It is probable, therefore, that outlying portions of the retina are capable of education to a higher discriminating power, just as we shall find the skin to be for tactile stimuli. While we can tell the stimulation of an upper part of the retina from a lower, or a right region from a left, it must be borne in mind that we have no direct knowledge of which is upper or lower or right or left in the ocular image. All our visual sensations tell us is that they are aroused at difEerent points, and nothing at all about the actual positions of these on the retina. There is no other eye behind the retina looking at it to see the inversion of the image (p. 496) formed on it. Suppose I am looking at a pane in a second-story window of a distant house : its image will then fall on the fovea centralis; the line joining this with the pane is called the visual axis. The image of the roof will be formed on a part of the retina below the fovea, and that of the front door above it. I distinguish that the images of all these fall on different parts of the retina in certain relative positions, and have learnt, by the experience of all my life, that when the image of anything arouses the sensation due to excitation of part of the retina below the fovea the object is above my visual axis, and vice versa; similarly with right and left. Consequently I interpret the stimulation of lower retinal regions as mean- ing high objects, and of right retinal regions as meaning left objects, and never get confused by the inverted retinal image about which directly I know nothing. A new-born child, even supposing it could use its muscles perfectly, could not seize a reachable object which it saw; it would not yet have learnt that attaining a point exciting that part of the retina above the fovea, meant reaching a position in space below the visual axis; but very soon it learns that things COLOR VIS 10 2^. 519 near its brow, that is up, excite certain visnal sensations, and objects below its eyes others, and learns to interpret retinal stimuli so as to localize accurately the direction, with ref- erence to its eyes, of outer objects, and never thenceforth gets puzzled by retinal inversion. Color Vision. Sunlight reflected from snow gives us a sensation which we call white. The same light sent through a prism and reflected from a white surface excites in us no iviiite sensation but a number of color sensations, graduating insensibly from red to violet, through orange, yellow, green, blue-green, blue, and indigo. The prism separates from one another light-rays of different periods of oscillation (p. 494) and each ray excites in us a colored visual sensation, while all mixed together, as in sunlight, they arouse the entirely different sensation of white. If the light fall on a piece of black velvet we get still another sensation, that of black; in this case the light-rays are so absorbed that but few are reflected to the eye and the vis- ual apparatus is left at rest. Physically black repre- sents nothing: it is a mere zero — the absence of ethereal vibrations; but, in consciousness, it is as definite a sensation as white, red, or any other color. We do not feel blackness or darkness except over the region of the jDossible visual field of our eyes. In a perfectly dark room we only feel the darkness in front of our eyes, and in the light there is no such sensation associated with the back of our heads or the palms of our hands, though through these we get no visual sensa- tions. It is obvious, therefore, that the sensation of blackness is not due to the mere absence of luminous stimuli but to the unexcited state of the retinas, which are alone capable of being excited by such stimuli when present. This fact is a very remarkable one, and is not paralleled in any other sense. Physically, complete stillness is to the ear what darkness is to the eye; but silence impresses itself on us as the absence of sensation, while darkness causes a definite feeling of '^'blackness." Young's Theory of Color Vision. Our color sensations insensibly fade into one another; starting with black we can insensibly pass through lighter and lighter shades of gray 520 THE HUMAN BODY. to white: or bt3ginning willi green through darker and darker shades of it to black or through lighter and lighter to white: or beginning with red we can b}' impercej)tiblc Rtejas i)ass to orange, from that to yellow and so on to the end of the solar spectrum: and from the violet, througli purple and carmine, we may get back again to red. Black and white appear to be fundamental color sensations mixed up with all the rest: we never imagine a color but as light or dark, that is as more or less near white or black; and it is found that as the light thrown on any given colored sur- face weakens, the shade becomes deeper until it joasses into black; and if the illumination is increased, the color becomes ''lighter" until it passes into white. Of all the colors of the spectrum yellow most easily passes into white with strong illumination. Black and white, with the grays which are mixtures of the two, thus seem to stand apart from all the rest as the fundamental visual sensations, and the others alone are in common parlance named "colors." It has even been suggested that the jDower of differentiating them in sensation has only lately been acquired ])y man, and a certain amount of evidence has been adduced from passages in the Iliad to prove that the Greeks in Homer's time confused together colors that are very different to most modern eyes; at any rate there seems to be no doubt that the color sense can begreatly improved by practice; women Avhose mode of dress causes them to pay more attention to tlie matter, have, as a general rule, a more acute color sense than men. Leaving aside black, white, gray, and the various browns (which are only dark tints of other colors), we may enun'- erate our color sensations as red, orange, yellow, green, blue, violet and purple; between each there are, however, numer- ous transition shades, as yellow-green, blue-green, etc., so that the number which shall have definite names given to them is to a large extent arbitraiy. Of the above, all but purple are found in the spectrum given when sunlight is separated by a prism into its rays of different refrangibility; rays of a certain Avave-length or period of oscillation cause in us the feeling red; others yellow, and so on; for convenience COLOR VISION. 521 we may speak of these as red, yellow, blue, etc., rays; all together, iu about equal proportions, they arouse the sen- sation of white. A remarkable fact is that most color feel- ings can be aroused in several ways. White, for example, not only by the above general mixture, but red and blue- green rays, or orange and blue, or yellow and violet, taken together in pairs, cause the sensation of white: such colors are called complermntary to one another. The mixture may be made in several ways; as, for example, by causing the red and blue-green parts of the spectrum to overlap, or by painting red and blue-green sectors on a disk and rotating it rapidly; they cannot be made, however, by mix- ing pigments, since what happens in such cases is a very complex phenomenon. Painters, for example, are accus- tomed to produce green by mixing blue and yellow paints, and some may be inclined to ridicule the statement that yel- low and blue when mixed give white. When, however, we mix the pigments we do not combine the sensations of the same name, which is the matter in hand. Blue paint is blue because it absorbs all the rays of the sunlight except the blue and some of the green; yellow is yellow because it absorbs all but the yellow and some of the green, and when blue and yellow are mixed the blue absorbs all the distinc- tive part of the yellow and the yellow does the same for the blue; and so only the green is left over to reflect light to the eye, and the mixture has that color. Grass-green has no complementary color in the solar spectrum; but with purple, which is made by mixing red and blue, it gives white. Several other colors taken three together, give also the sensation of white. If then we call the light-rays which arouse in us the sensation red, a, those giving us the sensation orange 5, yellow c, and so on, we find that we get the sensation white with a, h, c, d, e, /and g all together; or with 5 and e, or with c and /, or with a, d, and e\ our sensation ivhite has no determinate relation to ethereal oscillations of a given period, and the same is true for several other colors; yellow feeling, for example, may be excited by ethereal vibrations of one given wave-length (spectral yellow), or by mixing red and grass-green, which 522 THE HUMAN BODY. are duo to ethereal vibrations of totally different wave- lengths; in other words a physical light in which there are no waves of the "yellow" length may cause in us the sen- sation yellow, which is only one more instance of the gen- eral fact that our sensations, as such, give us no direct information as to the nature of external forces; they are but signs which we have to interpret. The modern view of specific nerve energies (p. 191) makes it highly improbable that our different color sensations can all be due to different modes of excitation of exactly the same nerve-fibres; a fibre which when excited alone gives us the sensation red will always give us that feeling when so excited. The simplest method of explaining our color sensations would therefore be to assume that for each there exists in the retina a set of nerve-fibres with appropriate terminal oi'gans, each ex- citable by its own proper stimulus. But we can distinguish so innumerable and so finely graded colors, that, on such a supposition, there must be an almost infinite number of different end organs in the retina, and it is more reasonable to suppose that there are a limited number of primary color sensations, and that the rest are due to combinations of these. That a compound color sensation may be very different from its components when these are regarded apart, is clearly shown by the sensation Avhite aroused either by what we may call red and blue-green, or green and purple, stimuli acting together; or of yellow due to grass-green and red. To account for our various color sensations we may, therefore, assume a much smaller number of primary sen- sations than the total number of color sensations we expe- rience; all can in fact be explained by assuming any three jDrimary color sensations which together give white, and regarding all the rest as due to mixtures of these in various proportions; there may be more than three, but three will account for all the phenomena, black being a sensation experienced when all visual stimuli are absent. This is known as Young's theory of color vision, and is that at present most commonly accepted. The selection of the three primary sensations is somewhat arbitrary, but they are usually regarded as red, green, and violet. It is COLOR BLINDNESS. 523 assumed that all kinds of light stimulating the end appa- ratuses give rise to all three sensations, but not necessarily in the same proportion. When all are equally aroused the sensation is -white; when the red and green are tolerably powerfully excited and the violet little, the sensation is yellow; when the green powerfully and the red and violet little, the sensation is green, and so on. In this way we can also explain the fact that all colored surfaces when intensely illuminated pass into white. A red light, for example, excites the primary red sensation most, but green and violet a little; as the light becomes stronger a limit is reached beyond which the red sensation cannot go, but the green and violet go on increasing with the intensity of the light, until they too reach their limits; and all three pri- mary sensations being then equally aroused, the sensation white is produced. Color Blindness. Some jiersons fail to distinguish colors which are to others quite different; when such a de- ficiency is well marked it is known as "color blindness," and, assuming Young's theory to be correct, it may be ex- plained by an absence of one or more of the three primary color sensations; observation of color-blind persons thus helps in deciding which these are. The most common form is red color blindness; persons afflicted with it con- fuse reds and greens. Red to the normal eye is red because it excites red sensation much, green some, and violet less; and a white page white, because it excites red, green, and violet sensations about equally. In a person without ]-ed sensation a red object would arouse only some green and violet sensation and so would be indistinguishable from a bluish green; in practice it is found that many persons confound these colors. Cases of green and violet color blindness are also met with, but they are much rarer than the red color blindness or "Daltonism." The detection of colorblindness is often a matter of con- siderable importance, especially in sailors and railroad officials, since the two colors most commonly confounded, red and green, are those used in maritime and railroad signals. Persons attach such different names to colors that 534 THE HUMAN BODY. a decision as to color blindness cannot be safely arrived at by simply showing a color and asking its name. The best plan is to take a heap of worsted of all tints, select one, say a red, and tell the man to put alongside it all those of the same color, whether of a lighter or a darker shade; if red blind he will select not only the reds but the greens, especially the paler tints. Aliout one man in eight is red blind. The defect is much rarer in women. Fatigue of the Retina. The nervous visual apparatus is easily fatigued. Usually we do not observe this be- cause its restoration is also rapid, and in ordinary life our eyes, when open, are never at rest; we move them to and fro, so that parts of the retina receive light alternately from brighter and darker objects and are alternately excited and rested. How constant and habitual the movement of the eyes is can be readily observed by trying to fix for a short time a small spot without deviating the glance; to do so for even a few seconds is impossible without practice. If any small object is steadily "fixed" for twenty or thirty seconds it will be found that the whole field of vision becomes grayish and obscure, because the parts of the retina receiving most light get fatigued, and arouse no more sensation than those less fatigued and stimulated by light from less illuminated objects. Or look steadily at a black object, say a blot on a white page, for twenty seconds, and then turn the eye on a white wall; the latter will seem dark gray, with a white patch on it; an effect due to the greater excitability of the retinal parts previously rested by the black, when compared with the sensation aroused elsewhere by light from the white wall acting on the previously stimulated parts of the visual surface. All persons will recall many instances of such phenomena, which are especially noticeable soon after rising m the morning. Similar things maybe noticed with colors; after looking at a red patch the eye turned on a white wall sees a blue-green patch ; the elements causing red sensations having been fatigued, the white mixed light from the wall now excites on that region of the retina only the other primary color sensations. The blending of colors so as to COLOR CONTRASTS. 52b secure their greatest effect depends on this fact; red and green go well together because each rests the parts of the visual apparatus most excited by the otlier, and so each appears bright and vivid as the eye wanders to and fro; while red and orange together, each exciting and exhaust- ing mainly the same visual elements, render dull, or in popular phrase "kill," one another. Contrasts. If a well-defined black surface be looked at on a la-i'ger white one the j^arts of the latter close to the black look whiter than the rest, and the parts of the black near the white blacker than the rest; so, also, if a green patch be looked at on a red surface each color is heightened near where they meet. This jihenomenon is largely due to fatigue and deficient fixation: a region of the eye rested by the black or the green is brought by a movement of the organ so as to receive light from the white or red surface; phe- nomena due to this cause are known as those of successive contrast. Even m the case of perfect fixation, however, something of the sanjie kind is seen; black looks blacker near white, and green greener near red when the eye has not moved in the least from one to the other. A small piece of light gray paper jDut on a sheet of red, which latter is then covered accurately with a sheet of semi-transparent tissue-paper, assumes the complementary color of the red, I.e. looks bluish green; and gray on a green sheet under similar circumstances looks pink. Such phenomena are known as those of simultaneous contrast, and are explained on psychological grounds by those who accept Young's theory of color vision. Just as a medium-sized man looks short beside a tall one, so, it is said, a black surface looks blacker near a white one, or a gray (slightly luminous white) surface, vv'hich feebly excites red, green, and violet sensations, looks deficient in red (and so bluish green) near a deeper red surface. There are, however, certain phenomena of simultaneous contrast which cannot be satis- factorily so explained, and these have led to other theories of color vision, the most important of which is that de- scribed in the next paragraph. Hering's Theory of Vision. Contrasts can be seen with 52G THE HUMAN BODY. the eyes closed and covered. If we look a short time at a bright object and then rapidly exclude light from the eye, we see for a moment a posiiive afier-image of the object, e.g. a window Avith its frame and panes after a glance at it and then closing the eyes. In these positive after-images the bright and dark parts of the object which was looked at retain their original relationship; they depend on the persist- ence of retinal excitement after the cessation of the stimulus and usually soon disappear. If an object be looked at steadily for some time, say twenty seconds, and the eyes be then closed a, negative after-image is seen. In this the lights and shades of the object looked at are reversed. Frequently a positive after-image becomes negative before disappearing. The negative images are explained commonly by fatigue; when the eye is closed some light still enters through the lids and excites less those parts of the retina previously exhausted by prolonged looking at the brighter parts of the Held of vision; or, Avhen all light is rigorously excluded, the proper stimulation of the visual apparatus itself, causing the idio-retinal light, affects less the exhausted portions, and so a negative image is produced. If Ave fix steadily for thirty seconds a point between two white squares about 4 mm. {\ inch) apart on a large black sheet, and then close and coA'er our eyes, we get a negative after-image in Avhich are seen two dark squares on a brighter surface; this surface is brighter close around the negative after-image of each square, and brightest of all between them. This luminous bound- ary is called the corona, and is explained usually as an effect of simultaneous contrast; the dark after-image of the square it is said makes us mentally err in judgment and think the clear surface close to it brighter than elsewhere; and it is brightest between the two dark squares, just as a middle- sized man between tAvo tall ones looks shorter than if along- side one only. If, however, the after-image be watched it will often be noticed not only that the light band betAveen the squares is intensely white, mucli more so than the normal idio-retinal light, bat, as the image fades aways, often the tAvo dark after-images of the squares disappear entirely with all of the corona, except that part between them which is BERING'S THEORY OF COLOR VISION. b21t still seen as a bright band on a uniform grayish field. Here there is no contrast to produce the error of judgment, and from this and other experiments Hering concludes that light acting on one j^art of the retina i^roduces inverse changes in all the rest, and that this has an important j^art in producing the phenomena of contrasts. Similar pheno- mena may be observed with colored objects; m their nega- tive af ter-im-ages each tint is reiDresented by its complemen- tary, as black is by white in colorless vision. Endeavoring to exclude such loose general explanations us " errors of judgment," Hering proposes a theory of vision which can only be briefly sketched here. We may put all our colorless sensations in a continuous series, passing through grays from the deepest black to the brightest white; somewhere half-way between will be a neutral gray which is as black as it is white. We may do some- tliing similar with our color sensations; as in gray we see black and white so in purple we see red and blue, and all colors containing red and blue may be put in a series of which one end is pure red, the other pure blue. So with red and yellow, blue and green, yellow and green. If we call to mind the whole solar spectrum from yellow to blue, through the yellow-greens, green, and blue-greens, we get a series in which all but the terminals have this in common that they contain some green. Green itself forms, however, a special point; it differs from all tints on one side of it in contain- ing no yellow, and from all on the other in containing no blue. In ordinary language this is recognized: we give it a definite name of its own and call it green. Its sim- plicity compared with the doubleness of its immediate neighbors entitles it to a distinct place in the color-sensa- tion series. There are three other color sensations which like green are simple and must have specific names of their own; they are red, blue, and yellow. Green may be pure green or yellow green or blue green, but never yellow and bluish at once, or reddish. Eed may be pure or yellowish or bluish, but never greenish. Red and green are thus mutually exclusive; yellow and blue stand in a similar relationship. All other color sensations, as orange 528 THE EUMAN BODY. suggest tAvo of the above, and may be described as mixtures of them; but they themselves stand out as fundamental color sensations. Moreover, it follows from the above, that more than two simple colored sensations are never combined in a compound color sensation. Since red always excludes green, and yellow blue, we may call them anti-colors (the complementary colors of Young's theory), and are led to suspect that in the visual organ there must occur, in the production of each, processes which prevent the simultaneous production of the other, since there is no a priori reason in the nature of things why we should not see red and green simultaneously, as well as red and yellow. Along with our color sensations there is always some colorless from the black-white series; which we recognize m speaking of lighter and darker shades of the same color. Hering assumes, then, in the retina or some part of the nervous visual apparatus, three substances answering to the black-white, red-green, and yellow-blue sensational series, the construction of each substance being attended with one sensation of its pair, and its destruction with the other. Thus, when construction of the black-white substance ex- ceeds destruction, we get a blackish-gray sensation; when the processes are equal the neutral gray; when destruction exceeds construction a light-gray, and so on. In the other color series similar things would occur; when con- struction of red-green substance exceeded destruction in any point of the retina we would get, say, a red feeling; if so, then excess of destruction would give green sensa- tion. The intensity of any given simple sensation would depend on the ratio of the difPerence between the construc- tion and destruction of the corresponding substance, to the sum of all the constructions and destructions of visual su))- stances going on in that part of the visual apparatus. A little thought will show that this can hardly be reconciled with the results expressed in Fechner's law. The intensity of a mixed color sensation would l)e the sum of the intensities of its factors, and its tint and shade dependent on the rela- tive proportion of these factors. When the construction HERING'S THEORY OF COLOR VISION. 529 and destruction of tlie red-greeii substance are equal no color sensation is aroused by it; and we get gray, due to those simultaneously occurriug changes in the black-white substance which are always present, but were i)reviously more or less cloaked by the results of the changes in the red-green substance. Red and green in certain pro- portions cause then a white or gray sensation, not because they supplement one another, as on Young's theory, but because they mutually cancel; and so for other comple- mentary colors. Moreover, according to Hering, destruction of a visual substance going on in one region of the retina promotes construction and accumulation of that substance elsewhere, but especially in the neighborhood of the excited spot. Hence, when a white square on a black ground is looked at, destruction of the black-white substance overbalances construction in the place on which the image of the square falls, but around this construction occurs in a high degree. When the eyes are shut, this latter retinal region, with its great accumulation of decomposable material, is highly irritable and, under the internal stimuli causing the idio- retinal light, breaks down comparatively fast, causing the corona, which may be intensely himinous; for with the closed eye the total constructive and destructive processes in the visual apparatus are small, and so the excess of de- struction in the coronal region bears a large ratio to the sum of the whole jn-ocesses. The student must apply this theory for himself to the other phenomena of contrasts and negative images, as also to the gradual disappearance of differences between light and dark objects when looked at for a time with steady fixation; the general key being the pi'inciple that anything leading to the accumulation of a visual substance increases its decompositions under stimu- lation, and vice versa. The main value of Hering's theory is that it attempts to account physiologically for phenomena previously indefinitely ex[)lained psychologically by such terms as '•'errors of judgment,'' which really leave the whole matter whei'e it was, since if (as we must believe) mind is a function of brain, the errors of judgment have 530 THE HUMAN BODY. still to be accounted for on i^hysiological grounds, as due to conditions of the nervous system. Visual Perceptions. The sensations which light excites in us we interpret as indications of the existence, form, and position of external objects. The conceptions which we arrive at in this way are known as visual perceptions. The full treatment of perceptions belongs to the domain of Psychology, but Physiology is concerned with the condi- tions under which they are produced. The Visual Perception of Distance. With one eye our perception of distance is very imperfect, as illustrated by the commcm trick of holding a ring suspended by a string a short way in front of a person's face, and telling him to shut one eye and pass a rod through the ring. If a pen- holder be held erect before one eye, while the other is closed, and an attempt be made to touch it with a finger moved across towards it, an error will nearly always be made. (If the finger be moved straight out towards the pen it will be touched because with one eye we can estimate direction accurately and have only to go on moving the finger in the proper direction till it meets the object.) In such cases, however, we get some clue from the amount of effort needed to "accommodate" the eye to see the object distinctly. When we use both eyes our perception of dis- tance is much better; when we look at an object with two eyes the visual axes are converged on it, and the nearer the object the greater the convergence. We have a pretty accurate knowledge of the degree of muscular effort required to converge the eyes on all tolerably near points. When objects are farther off, their apparent size, and the modifi- cations their retinal images experience by aerial perspective, come in to help. The relative distance of objects is easiest Tictcrmined by moving the eyes; all stationary objects then a})pear displaced in the opposite direction (as for example when we look out of the window of a railway car) and those nearest most rapidly; from the different ajoparent rates of movement we can tell which are farther and nearer. We •TO inseparably and unconsciously bind up perceptions of dis- tance with the sensations aroused by objects looked at, that VISUAL PERCEPTION OF SIZE. 531 we seem to see distance; it seems at first thought as definite a sensation as color, Tliat it is not is sliown by cases of persons born blind, who have had sight restored later in life by surgical operations. Such persons have at first no visual perceptions of distance: all objects seem spread out on a flat surface in contact with the eyes, and they only learn gradually to interpret their sensations so as to form judgments about distances, as the rest of ns did uncon- sciously in childhood before we thought about such things. The Visual Perception of Size. The dimensions of the retinal image determine primarily the sensations on which conclusions as to its size are based; the larger the visual angle the larger the retinal image: since the visual angle depends on the distance of an object the correct perception of size depends largely upon a correct perception of distance; having formed a judgment, conscious or unconscious, as to that, we conclude as to size from the extent of the retinal region affected. Most people have been surprised now and then to find that what appeared a large bird in the clouds was only a small insect close to the eye; the large a^^parent size being due to the previous incorrect judgment as to the dis- tance of the object. The presence of an object of tolerably well-known height, as a man, also assists in forming con- ceptions (by comparison)as to size; artists for this jDurpose frequently introduce . human figures to assist in giving an idea of the size of other objects represented. The Visual Perception of a Third Dimension of Space. This is very imperfect with one eye; still we can thus arrive at conclusions from the distribution of light and shade on an object, and from that amount of knowledge as to the relative distance of different points which is attainable monocularly; the different visual angles under which objects are seen also assist us in concluding that objects are farther and nearer; and so are not spread out on a plane before the eye, but occupy depth also. Painters depend mainly on devices of these kinds for representing solid bodies, and objects spread over the visual field in the third dimension of space. Single Vision with Two Eyes. When we look at a 532 THE HUMAN BOD T. flat object with both eyes we get a similar retinal image in each. Under ordinary circumstances we see, however, not two objects but one. In the habitual use of the eyes we move them so that the images of the object looked at fall on the two yellow spots. A point to tlie left of this' forms its image on the inner (right) side of the left eye and the outer (riglit) side of tlic right. An object verti- cally above that looked at would form an image straight l)elow the 3'ellow spot of each eye; an object to the left and above, its image to the inner side and below in the left eye and to the outer side and below in the right eye; and so on. We have learnt that similar simultaneous excita- tions of these corresponding pointn mean single objects, and so interpret our sensations. This at least is the theory of the experiential or empirical school of psychologists, thougJi others believe we have a sort of intuition on the subject. When the eyes do not work together, as in the muscular incoordination of one stage of intoxication, then they are not turned so that images of the same objects fall on cor- responding retinal points, and the person sees double. AVhen a squint comes on, as from paralysis of the external rectus of one eye, the sufferer at first sees double for the same reason. If a given object is looked at lines drawn from it through the nodal points reach the fovea centralis in each eye. Lines so drawn at the same time from a more distant object diverge less and meet each retina on the inner side of its fovea; but as above pointed out the corresponding points for each retinal region on the inside of the left eye, are on the outside of the right, and vice versa. Hence the more distant object is seen double. So, also, is a nearer object, be- cause the more diverging lines drawn from it through the nodal jooints lie outside of the fovea in each eye. Most people go through life unobservant of this fact; we only pay attention to what we are looking at, and nearly always this makes its images on the two foveae. That the fact is as above stated may, however, be readily observed. Hold one finger a short way from the face and the other a little farther off; looking at one, observe the other without moving STEREOSCOPIC VISION. 533 the eyes; it will be seen double. For any given position of the eyes there is a surface in space, all objects on which produce images on corresponding points of the two retinas: this surface is called the lioropter for that position of the eyes: all objects in it are seen single; all others in the visual field, double. The Perception of Solidity. When a solid o))ject is looked at the two retinal images arc different. If a trun- cated pyramid be held in front of one eye its image will be that represented at P, Fig. 142. If, however, it be held midway between the eyes, and looked at with both, then the left-eye image will be that in the middle of the figure, and the right-eye image that to the right. The small surface, bdca, in one answers to the large surface, b' d' d a, in the other. This may be readily observed by I ^ \ / \ d / V I p / c \ / ■ a fl. Fig. l-W holding a small cube in front of the face and alternately looking at it with each eye. In such cases, then, the retinal images do not correspond, and yet we combine them so as to see one solid object. This is known as sfereoscojJi'c vision, and the illusion of the common stereoscope depends on it. Two photographs are taken of the same object from two different points of view, one as it appears when seen by the left, and the other Avhen looked at with the right eye. These are then mounted for the stereoscope so that each is seen by its proper eye, and the scene or object is seen in distinct relief, as if, instead of flat pictures, solid objects were looked at. Of course in many stereo- scopic views the distribution of light and shade, etc., assist, but these are quite unessential, as may be readily observed by enlarging the middle and right outline drawings of Fig. 534 THE HUMAN BODY. 14:3 to the ordinary size of a stereoscopic slide, and placing them in the instrument. A solid pyramid standing out into space will be distinctly perceived; if the pictures be reversed the pyramid appears hollow. The pictures must not be too diiferent, or their combination to give the idea of a single solid body will not take place. Many persons, indeed, fail entirely to get the illusion with ordinary stereo- scopic slides. The phenomena of stereoscopic vision mili- tate strongly against the view that there are any pre- arranged corresponding points in the two retinas. The Perception of Shine. When we look at a rip2')led lake in the moonlight, we get the perception of a " shiny" or brilliant surface. The moonlight is reflected from the waves to the eyes in a number of bright points: these are not exactly the same for both eyes, since the lines of light- reflection from the surface of the water to each are different. The perception of brilliancy seems largely to depend on this slight non-agi'eement of the light and dark points on the two retinas. A rapid change of luminous points, to and fro between neighboring points on one retina, seems also to produce it. CHAPTER XXXIV. THE EAR AND HEARING. The External Ear. The auditory organ in man con- sists of three portions, known respectively as the external ear, the middle ear or tympanum, and the internal ear or labyrinth; of these the latter is the essential one, contain- ing the end organs of the auditory nerve. The external ear consists of the expansion seen on the exterior of the head, called the concha, and a passage leading in from it, the external auditory meatus {D to c, Fig. 143). This pas- FiG. 143.— The right ear (excluding the concha) seen from the front. D to c, external auditory meatus ; c c, tympanic membrane ; B, the tympanum with the auditory ossicles in it ; B to E, Eustachian tube ; A, labyrinth. sage is closed at its inner end by tlie tympanic or drum mernbrane. It is lined by a prolongation of the skin, through which numerous small glands, secreting the 2vax of the ear. open. The Tympanum, or drum-chamber of tlie ear (B, Fig. 143), is an irregular cavity in the temporal bone, closed ex- ternally by the drum membrane. From its inner side the Eustachian tube {B to E) proceeds and opens into the 536 THE HUM AX BODY. pharynx (//, Fig. 80)*, and the mncons membrane of that cav- ity is continued up the tube to line the tympanum; between this inside, and tlie skin of the external auditory meatus outside, is the proper tympanic membrane composed of connective tissue. The inner wall of the tympanum is bony except for two small ai)ortures, the oval and round foramens, which lead into the labyrinth. Daring life the round aperture is closed by the lining mucous membrane, and the oval in another way, to be described presently. The ti/mpanic memirane, c c, stretched like a drum-head across the outer side of the tympanum, forms a shallow funnel with its concavity outwards. If a sheet of indian- rubber be stretched over a ring and pulled down in the centre, its form will be very like that of the membrane in question. It is pressed by the external air on its exterior, and by air entering the tympanic cavity through the Eustachian tube on its inner side. If the tympanum were closed these pressures would not be always equal when barometric pres- sure varied, and the membrane would be bulged in or out according, as the external or internal pressure on it were the greater. On the other hand, were the Eustachian tube always open the sounds of our own voices would be extremely loud and disconcerting, so it is usually closed; but every time we swallow it is opened, and thus the air- pressure in the cavity is kept equal to that in the external auditory meatus. By holding the nose, keeping the mouth shut, and forcibly expiring, air may be forced under j)res- sure into the tympanum, and will be held in part impris- oned there until the next act of swallowing. On making a balloon ascent or going rapidly down a deep mine, the sudden and great change of aerial pressure outside frequently causes painful tension of the drum membrane, which may be greatly alleviated by frequent swallowing. The Auditory Ossicles. Three small bones lie in the tympanum forming a chain (Fig. 144) from the drum membrane to the oval foramen. The external bone is the malleus OY hammer ; the middle one, ihe incus or an nl; and the internal, the stapes or stirrup. The malleus, M, has * Paire 309. TYMPANIC BONES. 537 an upper enlargement or head, which carries on its inner side ail articular surface for the incus; below the head is a constriction, the neck, and below this two processes complete the bone; one, the long or slendei' process, is imbedded in a ligament which reaches from it to the front wall of the tympanum; the other process, the handle, reaches down between the mucous membrane lining the inside of the drum membrane and the membrane proper, and is firmly attached to the lat- ter near its centre and keeps t h e membrane dragged in there so as to give it its peculiar concave form, as seen from the out- side. The incus has a body and two processes and is much like a molar tooth with two fangs. On its body is an articular hollow to receive the head of the malleus; its short process {Jh) is attached by ligament to the back wall of the tym- panum; the long jirocess {Jl) is directed inwards to the stapes; on the tip of this process is a little knob, which represents a bone {osorhiculare), distinct in early life. The stapes {S) is extremely like a stirrup, and its base (the foot- l)iece of the stirrup) fits into the oval foramen, to the mar- ghi of which its edge is united by a fibrous membrane, allowing of a little play in and out. From the posterior side of the neck of the malleus a ligament passes to the back wall of the tympanum: this, with the ligament imbedding the slender process and fixed to the front wall of the tympanum, forms an antero-pos- terior axial ligament, on which the malleus can slightly rotate, so that the handle can be pushed in and the head out and vice versa. If a pin be driven through Fig. 144: just below the neck of the malleus and perpendicular to the Mm Fig. iM.— The auditory ossicles of the right ear, seen from the front. ..V, mal lens; J. incus; -S', stapes: Mcj), head of the malleus; Mc, neck of ditto; Ml, long process; 31m, handle: Jc,b(^d.y, Jb. short, and Jl, long process, of incus; Jpl, os orbiculare: Sep. head of stapes. 538 TUB HUMAN BODY. paper it will very fairly rci)rcsent this axis of rotation. Connected with the malleus is a tiny muscle, called the tensoi' tympani; it is inserted in the handle of the bone below the axis of rotation, and when it contracts pulls the handle in and tightens the drum membrane. Another muscle (the stapedius) is inserted into the outer end of the stapes, and when it contracts fixes the bone so as to limit its range of moment in and out of the fenestra ovalis. The Internal Ear. The labyrinth consists primarily of cham])ers and tubes hollowed out in the temporal bone and inclosed by it on all sides, except for the oval and round Fc vj)a Fig. 145.— Casts of the bony labyrinth. A. left labyrinth seen from the outer side; B, riglit labyrinth from the inner side; C, left labyrinth from above; Fc, round foramen; Fv, "val foramen; A, horizontal semicircular canal; /la, its ampulla; raa, ampulla of anterior vertical semicircular canal; Vj^ju, ampulla of posterior vertical semicircular canal ; vc, conjoined portion of Uie two ver- tical cauals. foramens on its exterior, and certain apertures on its inner side by which blood-vessels and branches of the auditory nerve enter; during life all these are closed water-tight in one way or another. Lying in the lony lahyrinth thus consti- tuted, are membranous parts, of the same general form but smaller, so that between the two a space is left; this is filled with a watery fluid, called the perilymjili; and the mem- hranoiis internal ear is filled by a similar liquid, the endo- lymph. The Bony Labyrinth. The bony labyrinth is described in three portions, the vestibule, the semicircular canals, and the cochlea; casts of its interior are represented from THE INTERNAL EAR. 53'J different aspects in Fig. 145. The vestibule is the central part and has on its exterior the oval foramen {Fv) into which the base of the stirrup-bone fits. Behind the vestibule are three bony semicircular canals, communicating with the back of the vestibule at each end, and dilated near one end to form an ampulla {vpa, vaa, and ha). The horizon- tal canal lies in the plane which its name implies and has its ampulla at the front end. The two other canals lie vertically, the anterior at right angles, and the pos- terior parallel, to the median antero-posterior vertical plane of the head. Their ampullary ends are turned forwards and open close together into the vestibule; their posterior ends unite {vc) and have a common vestibular opening. The bony cochlea is a tube coiled on itself somewhat like a snail's shell, and lying in front of the vestibule. The Membranous Labyrinth. The membranous vesti- bule, lying iu the bony, consists of two sacs communicating by a narrow aperture. The posterior is called the utriculus, and into it the membranous semicircular canals open. The piiste^kir, called the sac cuius, communicates by a tube with the mcmbranoits cochlea. The mem- branous semicircular canals much resemble the bony, and each has an ampulla; in most of their extent they are only united by a few irregular connective-tissue bands with the periosteum lining the bony canals; but in the amj^ulla one side of the mem- branous tube is closely adherent to its bony protector; at this point nerves enter the former. The relations of the membranous to the bony cochlea are more complicated. A section through this part of the auditory apparatus (Fig. 146) shows that its osseous portion consists of a tube Fig. 146.— a section through the cochlea iu tlie line of its axis. 540 THE ITU MAN BODY. woumi two and a half tiiuus (from left to right in the right ear and vice versa) around a central bony axis, the modiolus. From the axis a shelf, the lamina spiralis, projects and par- tially subdivides the tube, extending farthest across in its lower coils. Attached to the outer end of this bony plate is the membranous cochlea [scala media), a tube triangular' in cross-section and attached by its base to the outer side of the bony cochlear spiral. The spiral lamina and the membranous cochlea thus subdivide the cavity of the bony tu^e (Fig, 147) into an upper portion, the scala vestibicli, S V, and a lo\^•cr, the scala tympani, ST. Between these lie the lamina spiralis {ho) and the mem- FiG. 147.— Section of one coil of the cochlea, niaj?nified. SV, scala vestibuli' B, membrane of Reissner; CC, membranous cochlea (scato media); Us, limbus tamince spiralis, t, tectorial membrane; ST, scala tympani', Iso, spiral lamina ; Co. rods of Corti; b, basilar membrane. branous cochlea (CC), the latter being bounded above by the membrane of Reissner {R) and below by the basilar membrane {b). The inner edge of the lamina spiralis is thickened and covered with connective tissue which is hol- lowed out so as to form a spiral groove (the sulcus spiralis, ss) along the whole length of the membranous cochlea. The latter does not extend to the tip of the bony cochlea; above its apex the scala vestibuli and scala tympani com- municate; both are filled with perilymph, and the former communicates below with the perilymph cavity of the ves- tibule, while the scala tympani abuts below on the round foramen, which, as has already been pointed out, is closed by a membrane. The membranous cochlea contains cer- ORGAN OF CORTI. 541 tain solid structures seated on the basilar membrane and forming the organ of Corti; the rest of its cavity is filled with endolymph, which communicates with that in the sacculus. The Organ of Corti. This contains the end organs of the cochlear nerves. Lining the sulcus spiralis are cuboi- dal cells; on the inner edge of the basilar membrane they become cclumnar, and these are succeeded by a row which bear on their upper ends a set of short stiff hairs, and con- stitute the inner hair-cells, which are fixed below by a narrow aiiex to the basilar membrane; nerve-fibres enter them. To the inner hair-cells succeed the rods of Corti A B Fig. 148. — The rods of Corti. A, a pair of rods separated from the rest; B.a bit of the basilar membrane with several rods on it, showing: how they cover iu the tunnel of Corti; i, inner, and e, outer rods; b, basilar membrane; r, reti- cular membrane. {Co, Fig. 147), which are represented highly magnified in Fig. 148. These rods are stiff and arranged side by side in two rows, leaned against one another by their upper ends so as to cover in a tunnel; they are known respectively as the inner and outer rods, the former being nearer the lamina spiralis. Each rod has a somewhat dilated base, firmly fixed to the basilar membrane; an expanded head where it meets its felloAV (the inner rod presenting there a concavity into which the rounded head of the outer fits) ; and a slender shaft uniting the two, slightly curved like an italic S. The inner rods are more slender and more numerous than the outer, their numbers being about 6000 and 4500 respectively. Attached to the external sides of the heads of the outer rods is the reticular vienibrane [r, i42 THE HUMAN BODY. Fig. 148), which is stiff and perforated by holes. External to the outer rods come four rows of oiiter hair-cells, con- nected like the inner row with nerve- fibres; their bristles project into the holes of the reticular membrane. Be- yond the outer hair-cells is ordinary columnar epithelium, which passes gradually into cuboidal cells lining most of the membranous cochlea. The upper lip of the sulcus spiralis is uncovered by epithelium, and is known as the limbus lamince spi- ralis; from it projects the tectorial membrane {t, Fig. 147) which extends over the rods of Corti and the hair- cells. Nerve-Endings in the Semicir- cular Canals and the Vestibule. Nerves reach the ampulla of each semicii'cular canal, and, perforating its Avail, enter the epithelium lining it which is there several layers thick (Fig. 149). Some of the cells {sjj) are fusiform and have large nuclei; a slender external 2Jrocess runs from each to the cavity of the ampulla and is then continued as slender stiff hair [li), which projects into the endo- lymph. The deeper ends of these cells have been described as joining the terminal branches of nerve- fibres, so that they must be regarded as efid organs. In the utricle and saccule are somewhat similar structures; but collected among the hairs are minute calcareous particles, the ear-stones or otoliths. Fig. 149— The epithelium at the point where tlie nerves enter an ampulla. n, nerves; c, superficial columnar epithelium: sp, hair -ceils; /i, hairs; 6, basal cells. The Loudness, Pitch, and Timbre of Sounds. Sounds, as sensations, fall into two groups— ^zo/es and noises. Physi- PROPERTIES OF SOUND. 543 cally, sounds consist of vibrations, and these, under most circumstances, when they first reach our auditory organs, are alternating rarefactions and condensations of the air, or aerial waves. When the waves follow one another uni- formly, or periodicalli/, the resulting sensation (if any) is a note; when the vibrations are aperiodic it is a noise. In notes we recognize (1) loudness or intensity; (2) pitch; (3) quality or timbre, or, as it has been called, to7ie color; a note of a given loudness and pitch produced by a trum- pet and by a violin has a different character or individu- ality in each case; this cpiality is its timbre. Before un- derstanding the working of the auditory mechanism we must get some idea of the physical qualities in objective sound which the subjective differences of auditory sensa- tions are signs of. The loudness of a sound depends on the force of the aerial waves; the greater the intensity of the alternating condensations and rarefactions of these in the external auditory meatus, the louder the sound. The ^jitch of a note depends on the length of the waves, that is the dis- tance from one point of greatest condensation to the next, or (what amounts to the same thing) on the numljer of waves reaching the ear in given time, say a second. The shorter the waves the more rapidly they follow one another, and the higher the pitch of the note. When audible vibrations bear the ratio 1 : 2 to one another, we hear the musical interval called an octave. The note c on the un- accented octave is due to 132 vibrations in a second. The note c', the next higher octave of this, is produced by 264 vibrations in a second; the next lower octave (great octave, C), by 60; and so on. Sound vibrations may be too rapid or too slow in succession to produce sonorous sensations, just as the ultra-violet and ultra-red rays of the solar spectrum fail to excite the retina. The highest-pitched audible note answers to about 38,016 vibrations in a second, but it differs in individuals; many persons cannot hear the cry of a bat nor the chirp of a cricket, which lie near this upper audible limit. On the other hand, sounds of 544 THE HUMAN BODY. vibrational rate about 40 per second are not well heard, and a little below this become inaudible. The highest note used in orchestras is the tV of the fifth accented octave, produced by the piccolo flute, due to 4753 vibra- tions in a second; and the lowest-pitched is the E, of the contra octave, jiroduced by the double bass. Modern grand pianos and organs go down to C,, in the contra octave (33 vibrations per second) or even A,„ \ll\), but the musical quality of such notes is imperfect; they produce rather a ''buzz" than a true tone sensation, and are only used along with notes of higher octaves to which they give a character of greater depth. Pendular Vibrations. Since the loudness of a tone de- pends on the vibrational amplitude of its physical antece- dent, and its pitch on the vibrational rate, we have still to seek the cause of timbre; the quality by which we recog- nize the human voice, the violin, the piano, and the flute, even when all sound the same note and of the same loudness. The only quality of periodic vibrations left to account for this, is what we may call wave-fomi. Think of the movement of a pendulum; starting slowly from its highest point, it sweeps faster and faster to its lowest, and then slower and slower to its highest point on the opposite side; and then repeats the movements in the reverse direc- tion. Graphically we may represent such vibrations by the outer continuous curved line in Fig. 150. Suj^pose the lower end of the pendulum to bear a writing point which marked on a sheet of paper traveling down uniformly behind it, and at such a rate as to travel the distance 0-1- in two seconds. If the pendulum were at rest the straight vertical line would be draAvn. But if the pendulum were swinging we would get a curved line, compounded of the vertical movement of the jiapcr and the to-and-fro move- ment of the pendulum, writing sometimes on one side of the line 0-1-2 and sometimes on the other: Starting at a moment when the pendulum crosses the middle, 0, we would get described the curve 0 a^ a^, at first sejDaratingfast from the vertical line, then slower, then returning, at first PENDULAR VIBRATIONS. 545 gradually then faster, until it met the vertical again, at the end of 1" and commenced an exactly similar excursion on its other side, at the end of which it would be back at 1, and in just the same position, and ready to repeat exactly the swing, with which we commenced. A pendulum thus executes similar movements in equal periods of time, or its vibrations are periodic. A full swing on each side of the position of rest constitutes a complete vibration, so the vibrational period of a second's pendulum is two seconds: at the end of that time it is precisely where it was two seconds before, and moving in the same direction and at the same rate. It is clear that by examining such a curve we could tell exactly how the pendulum moved, and also in what period if we knew the rate at which the paper on which its point wrote was moving. The vertical line 0-1-2 is called the abscissa; per- pendiculars drawn from it and meeting the curve are ordinates: equal lengths on the abscissa represent equal times; where an ordinate from a given point of the abscissa meets the curve, there the writing point was at that moment; where succes- sive ordinates increase or decrease rapidly the pendulum moved fast from or towards its position of rest, and vice versa. Simi- Jarly, any other periodic movement may be perfectly represented by curves ; and since the form of the curve tells us all about the movement, it iscommon to speak of the "form of a vibration," meaning the form of the curve which indicates its charac- ters. Periodic vibrations like those in Fig. 150, where the ordinates at first grow fast, then more slowly, then dimin- ish slowly and then faster, and represented by a symme- trical curve on one side the abscissa, Avhich is repeated Fig. 150. 546 THE HUMAN BODY. exactly on the other side of the abscissa, are known as pendular vibrations. The Composition of Vibrations. The vibrations of a second's ponduluni set tlie air-particles in contact with it in similar movement, bnt the aerial waves succeed one another too slowly to produce in us the sensation of a musical note. If for the pendulum we substitute a tuning- fork (the prongs of which move in a like way), and the fork vibrates 132 times per 1", then 132 aerial waves will fall on the tympanic membrane in that time, and we will hear the note c of the unaccented octave. If the larger con- tinuous curve in Fig. 150 represent the aerial vibrations in this case, the distance 0 to 1 on the abscissa will represent Y^ of a second. Let, simultaneously, the air be set in movement by a fork of the next higher octave, c', making 264 vibrations per 1"; under the influence of this second fork alone, the aerial particles would move as represented by the smaller continuous curved line, the waves being half as long and cutting the abscissa twice as often. But when both forks act together the aerial movement will be the algel)raic sum of the movements due to each fork; when both drive the air one way they will reinforce one another, and vice verso; the result will be the movement represented by the dotted line, which is still periodic, repeating itself at equal intervals of time, but no longer pendular, since it is not alike on the ascending and descending limbs of the curves. We thus get at the fact that non-pendular vibrations may be pro- duced by the fusion of pendular, or, in technical phrase, by their compositioii. Suppose several musical instruments, as those of an or- chestra, to be sounded together. Each produces its own effect on the air-particles, whose movements, being the algebraical sum of those due to all, must at any given in- stant be very complex; yet the ear can pick out at will and follow the tones of any one instrument. From the com- plex aerial movement it can select that fraction of it which one vibrating body produces. The air in the extei'nal auditory meatus at any given moment can only be in one state of rarefaction or condensation and at one rate ANALYSIS OF VIBRATIONS. 547 and in one direction of movement, this being the resultant of all the forces acting upon it; all clashing, and some push- ing one way and others another. If the resultant move- ment be not periodic it will be recognized as due to noises or to several simultaneous inharmonic musical tones; this is commonly the case when musical tones are not united designedly, and the ear thus get one criterion for distin- guishing movements of the air due to several simultaneous musical tones. However, a composite set of tones will give rise to periodic vibrations when all are due to vibrations of rates which are multiples of the same whole number. In such cases the movement of the air in the auditory meatus has no property except vibrational form by which the ear could distinguish it from a simple tone; when the two tuning-forks giving the forms of vibration (with rates as 1 to 3), represented in Fig. 150 by continuous lines, are sounded together, we get the new form of vibration repre- sented by the dotted line, and this has the same period as that of the lower-pitched fork; yet the ear can clearly dis- tinguish the resultant sound from that of this fork alone, as a note of the same pitch but of different timbre; and with practice can recognize exactly what simple vibrations go to make it up. The Analysis of Non-Pendular Vibrations. If a per- son Avith a trained ear listens attentively to any ordinary musical tone, such as that of a piano, he hears, not only the note whose vibrational rate determines the pitch of the tone as a whole, but a whole series of higher notes, in harmony with the general or fundamental tone; this latter is the 'primary partial tone, and the others are secondary pai'tial tones; nearly all tones used in music contain both. If the prime tone be due to 132 vibrations a second (c), its first upper partial is c' (= 264 vibrations j^er second); the next is the fifth of this octave {g' = 396 = 132 X 3 vibrations per 1'); the next is the second octave, c" (132 X 4 = 528 vibrations per 1'); the next is the major third of the c" (=132 X 5 = 660 vibrations per second = e"), and so on. The only form of vibration which gives no upper partial tones is the pendular; we may coll notes due to such vibra- 548 THE HUMAN BODY. tions simple tones; and we, consequently, recognize in music tones which arc simple (such as those of tuning-forks) and those which are compound; these latter are non-pendular in form. We find, then, that the form of aerial vibrations deter- mines in our sensations the occurrence or non-occurrence of upper partial tones. It also, as we have seen, deter- mines the quality or tinihre of the tone, since vibrational amplitude and rate are otherwise accounted for in sensa- tion by loudness and pitch. It can be proved, by the employment of the higher mathematics, that every periodic non-pendular movement can be analyzed (as the dotted curve of Fig. 150 may be) into a given number of pendular vibrations, that is, every compound vibration into a set of simple ones; and that every periodic non-pendular vibration can be made by the combination of pendular. Moreover, any given comj)ound vibration can be analyzed into but one set of simple ones; no other combination will produce it. Consequently a vibra- tional movement of the air in the external auditory passage, producing a compound musical tone sensation, can be ex- hibited always, but only in one way, as the sum of a num- ber of simple vibrations, whose rates are multiples of that which determines the pitch of the tone. Xow when the trained ear listens to a tone with the object of detecting upper partials if present, it hears them only when the vibrations are non-pendular {i.e. when theoretically they ought to be present), and those it hears are exactly those demanded by theory. By the help of certain instruments their detection is made easy even to the untrained ear. But in ordinary circumstances we do not heed secondary partial tones; we hear a note of the pitch of the primary partial and of a certain timbre; and whenever the upper partials present are different, or of different relative intensities, the timbre of the note varies. Hence it becomes probable that, just as the ear can at will follow any instrument in an orchestra, analyzing the aerial movement so as to select and follow the fraction of the whole due to that one, so it cm and does analyze compound SYMPATHETIC RESONANCE. 549 tones when proceeding from one instrument, and that the npper partials, not rising into consciousness as definite tones but present as subdued sensations, give its char- acter to the whole tone and determine its timbre. It might be, however, that the composition of non-pendular vibrations from pendular was a mere mathematical fiction, liaving no real existence in nature; before we can accept the above explanation of timbre, we must see if there is any evidence that, as a matter of fact, non-pendular vibra- tions, not only may be, but are made up by the combination of pendular. Sympathetic Besonance. Imagine slight taps to be given to a jiendulum; if these be repeated at such intervals of time as to always help the swing and never to retard it, the pendulum will soon be set in powerful movement. If the taps are irregular, or when regular come at such intervals as sometimes to promote and sometimes retard the move- ment, no great swing will be produced; but if they always push the pendulum in the way it is going at that instant, they need not come every swing in order to set up a power- ful vibration; once in two, three, or four swings will do. A stretched string, such as that of a piano, is in so far like a given pendulum that it tends to vibrate at one rate and no other; if aerial waves hit it at exactly the right times they soon set it in sufficiently powerful vibrations to cause it to emit an audible note. By using such strings we might hope to detect the separate pendular vibrations in any non-pendular aerial jDcriodic movement if such really existed; certain strings would pick out the pendular com- ponent agreeing in rate with their own vibrational period and be soon set in powerful movement; while those not ^■ibrating in the same period as any of the pendular compo- nents, would remain jjractically at rest, like the pendulum getting taps which sometimes helped and sometimes impeded its swing. If the dampers of a piano be raised and a note be sung to it, it will be found that several strings are set in vibration, such vibrations being called sympathetic. The human voice emits compound tones which can be mathe- matically analyzed into simple vibrations, and if the piano 550 THE HUMAN BODY. strings set in movement l)y it be examined, they will be found to be exactly those which answer to these pendular vil)rations and to no others. We tims get ex}ierimental grouiul.s for believing that compound tones are really made up of a number of sim})le vibrations, and get an additional justification for the supi)osition that in the ear each note is analyzed into its pendular comjioiients; and that the differ- ence of sensation which we call timbre is due to the effect of the secondary j)artial tones thus perceived. If so, the ear must have in it an apparatus adapted for sympathetic resonance. It may be asked why, if the car analyzes yibrations in this way, do we not commonly perceive it? How is it that what we ordinarily hear is the timbre of a given tone and not the separate upper partials which give it this character? The explanation is more jisychological than physiological, and belongs to the same series as the reason why we do not ordinarily notice the blind spot in the eye, or the double- ness of objects out of the horopter, or the duplicity of stereoscopic images. We only use our senses in daily life when they can tell us something that may be useful to us, and we neglect so habitually all sensations which would be useless or confusing, that at last it needs special attention to observe them at all. The way in which tones are com- bined to give timbre to a note is a matter of no importance in the daily use of them, and so we fail entirely to observe the components and note only the resultant, until we make a careful and scientific examination of our sensations. The Functions of the Tympanic Membrane. If a stretched membrane, such as a drum-head, be struck, it will be thrown into periodic vibration and emit for a time a note of a determined pitch. The smaller the membrane and the tighter it is stretcbcd tlie higher the pitch of its note; every stretched membrane tlins has a rate of its own at which it tends to vibrate, just as a piano or violin string has. When a note is sounded in the air near such a membrane, the alternating waves of aerial condensation and rarefaction will move it; and if the waves succeed at the vibrational rate of the membrane the latter will be set in powerful sympafJietic USES OF DRUM-MEMBRANE. 651 vibration; if tlioy do not push the membrane at the proper times, their eiiects will neutralize one another: hence such membranes respond well to only one note. The tym- panic membrane, however, responds equally well to a large number of notes; at the least for those due to aerial vibra- tions of rates from 60 to 4000 per second, running over eight octaves and constituting those commonly used in music. This faculty depends on two things; (1) the mem- brane is comparatively loosely and not uniformly stretched; (2) it is loaded by the tympanic bones. The drum-membrane is (p. 536) in the form of a shallow funnel with its sides convex towards its cavity; in such a membrane the tension is not uniform but increases towards the centre, and it has accordingly no proper note of its own. Further, whatever tendency such a membrane may have to vibrate rather at one rate than another, is almost com- pletely removed by ''damping" it; i. e. placing in contact with it something comparatively heavy and which yet has to be moved when the membrane does. This is effected by the tympanic bones, fixed to the drum-membrane by the handle of the malleus. Another advantage is gained by the damping; once a stretched membrane is set vibrating it continues so doing for some time; but if loaded its move- ments cease almost as soon as the moving impulses. The dampers of a piano are for this purpose; and violin- players have to "damp" with the fingers the strings they have used when they wish the note to cease. When the aerial waves cease the loaded drum-membrane comes to rest almost immediately, and is ready to respond to the next set of waves reaching it. Functions of the Auditory Ossicles. When the air in the external auditory meatus is condensed it pushes in the handle of the malleus. This bone then slightly rotates on the axial ligament (p. 537) and drags out the incus and with it the base of the stapes; the reverse occurs when air in the auditory passage is rarefied. Aerial vibrations thus set the chain of bones swinging, and pull out and push in the base of the stapes, which sets up waves in the perilymph of the labyrinth, and these are transmitted through the mem- 552 THE HUMAN BODY. brauoiis labyrinth to the endolymi)h. These liquids being chiefly water, and practically incompressible, the end of the stapes could not work in and out- at the oval foramen, were the labyrinth elsewhere completely surrounded by bone: but the membrane covering the round foramen bulges out when the base of the stapes is pushed in, and vice versa; and so allows of waves being set up in the labyrinthic ]i(|uids. These correspond in period and form to those in the auditory meatus; their amplitude is determined by the extent of the vibrations of the drum-membrane. The form of the tympanic membrane causes it to trans- mit to its centre, where the malleus is attached, vibrations of its lateral parts in diminished amplitude but increased power; so that the tympanic bones are pushed only a little way but with considerable force. Its area, too, is about twenty times as great as that of the oval foramen, so that force collected on the larger area is, by pushing the tym- panic bones, all concentrated on the smaller. The ossicles also form a bent lever (Fig, 144) of which the fulcrum is at the axial ligament and the effective outer arm of this lever is about half as long again as the inner, and so the move- ments transmitted by the drum-membrane to the handle of the malleus are communicated with diminished range, but increased j^iower, to the base of the stapes. Ordinarily sound-waves reach the labyrinth in this way through the tympanum, but they may also be transmitted through the bones of the head in general; if the handle of a vibrating tuning-fork be placed on the vertex, for exam- ample. Such sounds seem to have their origin in the head itself. Similarly, when a vibrating body is held between the teeth, sound reaches the end organs of the auditory nerve through the skull-bones; and persons who are deaf from disease or injury of the tympanum can thus be made to hear, as with the audiplione. Of course if deafness be due to disease of the proper nervous auditory apparatus no device can make the ])erson hear. Function of the Cochlea. We have already seen reason to believe that in the ear there is an apparatus adapted for sympathetic resonance, by which we recognize different FUNCTIONS OF COCHLEA. 553 musical tone-colors; the minute structure of the membra- nous cochlea is such as to lead us to look for it there. An old view was that the rods of Corti, which yary in length, were like so many piano-strings, each tending to vibrate at a given rate and picking out and responding to peudular aerial vibrations of its own period, and exciting a nerve Avhich gave rise to a particular tone sensation. "When the labyrinthic fluids were set in non-pendular vibrations, the rods of Oorti were thought to analyze these into their peudu- lar components, all rods of the vibrational rate of these be- ing set in sympathetic movement, but that rod most wh-ose period was that of the primary partial tone; this would determine the pitch of the note, and the less-marked sen- sations due to the others affected would give it its timbre. The rods, however, do not differ in size sufficiently to account for the range of notes which we hear, and they are absent in birds, which undoubtedly distinguish different musical notes; and the nerve-fibres of the cochlea are not connected with them but with the hair-cells. On the whole it seems probable that the basilar mem- brane is to be looked upon as the primary arrangement for sympathetic resonance in the ear. It increases in breadth twelve times from the base of the cochlea to its tip (the less width of the lamina spiralis at the apex more than compensating for the less size of the bony tube there) and is stretched tight across, but loosely in the other direc- tion. A membrane so stretched behaves as a set of separate strings placed side by side, somewhat as those of a harp but much closer together; and each string would vibrate at its own period without influencing much those on each side of it. Probably, then, each transverse band vibrates to sim- ple tones of its own jieriod, and excites the hair-cells which lie on it, and through them the nerve-fibres. Perhaps the rods of Corti, being stiff, and carrying the reticular mem- brane, rub that against the upper ends of the hair-cells which project into its apertures and so help in a subsidiary way, each pair of rods being especially moved when the band of basilar membrane carrying it is set in vibration. The tec- torial membrane is probably a ''damper;" it is soft and 554 THE HUMAN BODY. inelastic, and suppresses the vibrations as soon as the mov- ing force ceases. Function of the Vestibule and Semicircular Canals. Many noises are merely spoiled music; they are due to tones so combined as not to give rise to periodic vibrations; these are probably heard by the cochlea. If a single violent air-wave ever cause a sound sensation (which is doubtful since any violent push of an elastic substance, such as the air, will cause it to make several rebounds before coming to rest) we j)erhapshear it by the vestibule; the otoliths, there in contact with the auditory hairs, are imbedded in a tenacious gummy mass quite distinct from the projier endolymph, and are not adapted for executing regular vibrations, but they might yield to a single powerful impulse and transmit it to the hair -cells, and through them stimulate the nerves. There is reason to believe that the semicircular canals have nothing to do with hearing; their supposed function is described in Chapter XXXV. Auditory Perceptions. Sounds, as a general rule, do not seem to us to originate within the auditory apparatus; we refer them to an exteriud source, and to a certain extent can judge the distance and direction of this. As already mer.tioned, the extrinsic reference of sounds which reach the labyrinth through the general skull-bones instead of through the tympanic chain is imperfect or absent. The recogni- tion of the distance of a sounding body is j^ossible only when the sound is well known, and then not very accurately; from its faintness or loudness we may make in some cases a pretty good guess. Judgments as to the direction of a sound are also liable to be grossly wrong, as most persons have experienced. However, when a sound is heard louder by the left than the right ear we can recognize that its source is on the left; when equally with both ears, that it is straight in front or behind; and so on. The concha has per- haps something to do with enabling us to detect whether a sound originates before or behind the ear, since it collects, and turns with more intensity into the external auditory meatus, sound-waves coming from the front. By turning the head and noting the accompanying changes of sensation AUDITORY PERCEPTIONS. 555 in each ear we can localize sounds better than if the head be kept motionless. The large movable concha of many ani- mals, as a rabbit or a horse, which can be turned in several directions, is probably an important aid to them in de- tecting the position of the source of a sound. That the recognition of the direction of sounds is not a true sensation, buta judgment, founded on experience, is illustrated by the fact that we can estimate much more accurately the direc- tion of the human voice, which we hear and heed most, than that of any other sound. CHAPTER XXXIV. TOUCH, THE TEMPERATURE SENSE, THE MUSCULAR SENSE, COMMON SENSA- TION, SMELL, AND TASTE. Nerve-Endings in the Skin. Many of the afEerent skin-nerves end in connection with hair-bulbs; the fine hairs over most of the cutaneous surface, projecting from the skin, transmit any movement impressed on them, with increased force, to the nerve- fibres at their fixed ends. In many animals, as cats, large, specially tactile, hairs are de- veloped on the face, and these have a very rich nerve- supply. Fine branches of axis cylinders have also been described as penetrating be- tween epidermic cells and ending there without termi- nal organs. In or immedi- ately beneath the skin several peculiar forms of nerve end organs have also been de- scribed; they are known as (1) Touch-cells; (2) Pacinian corpuscles; (3) Tactile co7'- jmscles; (4) End-hulbs. The Pacinian corpuscles (Fig. 151) lie in the subcutane- ous tissue of the hand and foot, and about the knee-joint; but also away from the skin on branches of the solar plexus (p. 172), so that it is doubtful if they are touch-organs. Fig. 151. — A Pacinian corpuscle, magnifled. NERVE END ORGANS IN SKIN 557 They are oval, from 1.5 to 3.5 mm. [^ to ^ inch) long, by about half that width, and have a whitish translucent appearance, with a more opaque centre. When magnified each is found to consist of a core, surrounded by many concentric capsules, h. A nerve-fibre, a, enters at one end, and its axis cylinder, c, runs along the core to the other, where it terminates in one or two little knobs, or a number of fine branches. The tactile corpuscles lie in papillae of the dermis, aiid are oval and about .08 mm. (-j-J-g- inch) in length. They contain a soft core, enveloped by a connective-tissue cap- ' ''1 Fig. 152.— Dermic papillae with tactile corpuscles, A. a corpuscle with four nerve-fibres; «, corpuscle; 6 and c. nerve-fibres, i?, papilla made transparent with acetic acid to show tactile eorpvisele within ; o. proper tissue of the papilla ; 6, tactile corpuscle; c, entering nerve, d, e, nerve-fibres twining round the cor- puscle. C, a papilla, containiug a tactile corpuscle, seen in optical transverse section. sule, and separated into several masses. Two, three, or more nerve-fibres go to each corpuscle and appear to end in plates lying between each of the segments of the core. Tactile corpuscles are numerous in the skin of the hand and foot, but are rare elsewhere. This limited distribution over the surface militated against the belief that they were tactile end organs; but it has lately been found that simpler bodies, the touch-cells, of the same essential structure but receiving only one nerve-fibre each, are distributed all over the skin; the more complex, and probably more irritable, form being found where the epidermis is especially thick. 558 THE HUMAN BODY. The end-hulbs are spheroidal and about .04 mm. (^5-^^ inch) in diameter. Each consists of a core, with a connective- tissue capsule, to which two or three nerve-fibres run; the axis cylinders penetrate the core. End-bulbs are found on one or two regions of the skin, as that on the red part of the lips, in the conjunctiva, and the mucous membrane covering the soft palate, and the tongue. Touch, or the Pressure Sense. Through the skin we get several kinds of sensation; touch proper, heat and cold, and pain; and we can with more or less accuracy localize them on the surface of the Body. The interior of the mouth jDossesses also these sensibilities. Through touch proper we recognize pressure or traction exerted on the skin, and the force of the pressure; the softness or hard- ness, roughness or smoothness, of the body producing it; and the form of this, when not too large to be felt all over. In the latter case, as when we move the hand over an object to study its shape, muscular sensations are com- bined with proper tactile, and such a combination of the two sensations is frequent; moreover, we rarely touch any- thing without at the same time getting temperature sen- sations; so that pure tactile feelings are rare. From an evolution point of view, touch is probably the first distinctly differentiated sensation, and this primary position it still largely holds in our mental life; we mainly think of the things about us as objects which would give us certain tac- tile sensations if we were in contact with them. Though the eye tells us much quicker, and at a greater range, what are the shapes of objects and whether they are smooth, rough, and so on, our real conceptions of round and square and rough bodies are derived through touch, and we trans- late unconsciously the teachings of the eye into mental terms of the tactile sense. A person who saw but had no touch-sense would conceive solid objects very differently from the rest of mankind. The delicacy of the tactile sense varies on different parts of the skin; it is greatest on the forehead, temples, and TOUCH. 559 buck of the forearm, where a weight of 2 milligr. (.03 grain) j^ressing on an area of 9 sq. millim. (.0139 sq. inch) can be felt. On the front of the forearm 3 milHgr, (.036 grain) can be simihirly felt, and on the front of the fore- finger 5 to 15 milligr. (.07-0.23 grain). In order that the sense of touch may be excited neigh- boring skin areas must be differently pressed; when we lay the hand on a table this is secured by the inequalities of the skin, which prevent end organs, lying near together, from being equally compressed. When, however, the hand is immersed in a liquid, as mercury, which fits into all its inequalities and presses with practically the same weight on all neighboring immersed areas, the sense of pressure is only felt at a line along the surface, where the immersed and non-immersed parts of the skin meet. It was in connection with the tactile sense that the facts on which so-called iisycho-physical law (p. 473) is based, were first observed. The smallest perceptible difference of pressure recognizable when touch alone is used, is about ^; I.e. we can just tell a weight of 20 grams (310 grains) from one of 30 (465 grains) or of 40 grams (620 grains) from one of 60 (930 grains); the change which can just be recognized being thus the same fraction of that already act- ing as a stimulus. The ratio only holds good, however, for a certain mean range of pressures; and its existence for any has lately been denied. The experimental difficulties in determining the question are considerable; muscular sensa- tions must be rigidly excluded; the time elapsing between laying the different weights on the skin must always be equal; the same region and area of the skin must be used; the weights must have the same temperature; and fatigue of the organs must be eliminated. Considerable individual variations are also observed, the least perceptible difference not being the same in all persons. The Localizing Power of the Skin. When the eyes are closed and a jioint of the skin is touched we can Avitli some accuracy indicate the region stimulated; although tactile feelings are in general characters alike, they differ in some- thing [loccd sifffi) besides intensity by which we can distin- 500 TUE nUMAN BODY. guisli tlieni; some sub-sensation quality not rising definitely into i)roniinence in consciousness must be present, com- parable to the upper joartials determining the timbre of a tone. Tlie accuracy of the localizing power A'aries widely in different skin regions and is measured by observing the least distance which must separate two objects (as the blunted points of a joair of compasses) in order that they may be felt as two. The following table illustrates some of the differences observed — Tongue-tip 1.1 mm. (.04 inch) Palm side of Itist phaluux of linger 2.2 mm. (.08 inch) Red part of lips 4.4 mm. (.16 inch) Tip of nose 6.G mm. (.24 inch) Back of second pU;i];inx of linger 11.0 mm. (.44 inch) Heel ;>2.0 mm. (.88 inch) Buck of hand 30.8 mm. (1.23 inches) Forearm 39.6 mm. (1.58 inches) Sternum •. 44.0 mm. (1.76 inches) Back of neck 52.8 mm. (2.11 inclic.'.) Middle of hack 66.0 mm. (2.64 inches) The localizing power is a little moi'e acute across the long axis of a limb than in it; and is better when the pressure is only strong enough to just cause a distinct tactile sensa- tion, tluiu when it is more power- '^"""/fiW^'N W?*^ ^'^^^' ^^ ^^ ^^^'^ ^^^'^ readily and nnW rapidly improvable by j^ractice. fW|YiT^^ ^^ might be thought that this \UJwU-Wt¥w localizing power dej^ended directly '^ — 'uSuuXuuuYv ^^^ nerve distribution; that each ; ulUM touch-nerve had connection with I \}JfJ^}'}^M>{ ^ special brain-centre on the one \ \ti\jVW/ I hand (the excitation of which ) j caused a sensation with a charac- I / teristic local sign), and at the other \ /' end was distributed over a certain ^^ •'•' skin area, and that the larger this area the farther apart might two points be and still give rise to only one sensation. If tliis were so, however, the peripheral tactile areas (each being determined by the anatomical distribution of a nerve-fibre) LOCALIZATION OF TACTILE SENSATIONS. 561 must have definite unchangeable limits, which experiment shows that they do not possess. Suppose the small areas in Fig. 153 to each represent a peripheral area of nerve distribu- tion. If any two points in c were touched we would accord- ing to the theory get but a single sensation; but if, while the compass points remained the same distance apart, or were even approximated, one were placed in c and the other on a contiguous area, two fibres would be stimulated and we ought to get two sensations; but such is not the case; on the same skin region the points must be always the same distance apart, no matter how they be shifted, in order to give rise to two just distinguishable sensations. It is probable that the nerve areas are much smaller than the tactile; and that several unstimulated must intervene between the excited, in order to produce sensations which shall be distinct. If we suppose twelve unexcited nerve areas must intervene, then, in Fig. 153, a and b will be just on the limits of a single tactile area; and no matter how the pomts are moved, so long as eleven, or fewer, unexcited areas come between, we would get a single tactile sensation; In this way we can explain the fact that tactile areas have no fixed boundaries in the skin, although the nerve distri- bution in any part must be constant. We also see why the back of a knife laid on the surface causes a continuous linear sensation, although it touches many distinct nerve areas; if we could discriminate the excitations of each of these from that of its immediate neighbors we would get the sensation of a series of points touching us, one for each nerve region excited; but in the absence of intervening unexcited nerve areas the sensations are fused together. The ultimate differentiation of tactile areas takes place in the central organs, as will be more fully pointed out in the next chapter. Afferent nerve impulses reaching the spinal cord from a finger-tip enter the gray matter and tend to radiate some way in it; from the gray region through which they spread, impulses are sent on to perceptive tactile centres in the brain; if two skin-points are so close that their regions of irradiation in the cord overlap, then the two points touched cannot be discriminated in con- 6fi2 THE HUMAN BODY. sciousness, since the stime brain regions are excited. Tlie more poAverful tlie stimulus the wider the irradiation in the cord, and hence the less accurate the discriminating power. The more often an impulse has traveled, the more does it tend to keep its own proper tract through the gray matter of the cord, and get on to its own proper brain- centre alone, hence the increase of tactile discrimination with practice, for we cannot suppose it to be due to a growth of more nerve-fibres down to the skin, and a rearrangement of the old, with smaller areas of anatomical distribution. As a general rule, more movable parts have smaller tactile areas; this probably depends on practice, since they are the parts which get the greatest number of different tactile stimulations. The Temperature Sense. By this we mean our faculty of perceiving cold and warmth; and, with the help of these sensations, of perceiving temperature differences in external objects. Its organ is the whole skin, the mucous membrane of mouth and fauces, pharynx and gullet, and the entry of the nares. Direct heating or cooling of a sensory nerve may stimulate it and cause pain, but not a true temperature sensation; and the degree of heat and cold requisite is much greater than that necessary when a temperature-perceiving surface is acted upon; hence we must assume the presence of temperature end organs. In a comfortable room we feel at no part of the Body either heat or cold, although different parts of its surface are at different temperatures; the fingers and nose being cooler than the trunk which is covered by clothes, and this, in turn, cooler than the interior of the mouth. The tem- perature which a given region of the temperature organ has (as measured by a thermometer) when it feels neither heat nor cold is its temjx'r (dure- sensation zero, and is not associated Avitli any one objective temperature; for not only, as we have just seen, does it vary in different parts of the organ, but also on the same part from time to time. Whenever a skin region has a temperature above its sensa- sation zero we feel warmth and vice versa; the sensation is more marked the greater the difference, and the more TEMPERATURE SENSATIONS. 563 suddenly it is produced; touching a metallic body, which conducts heat rapidly to or from the skin, causes a more marked hot or cold sensation than touching a worse con- ductor, as a piece of wood, at the same temperature. The change of temperature in the organ may be brought about by changes in the circulatory apparatus (more blood flowing through the skin warms it and less leads to its cool- ing), or by temperature changes in gases, liquids, or solids in contact with it. Sometimes we fail to distinguish clearly whether the cause is external or internal; a person coming in from a windy walk often feels a room uncomfortably warm which is not really so; the exercise has accelerated his circula- tion and tended to warm his skin, but the moving outer air has rapidly conducted off the extra heat; on entering tlie house the stationary air there does this less qiiickly, the skin gets hot, and the cause is supposed to be oppressive heat of the room. Hence, frequently, opening of win- dows and sitting in a draught, with its concomitant risks; whereas keeping quiet for five or ten minutes, until the circulation had returned to its normal rate, would attain the same end without danger. The acuteness of the temperature sense is greatest at temperatures within a few degrees of 30° 0. (86° F.); at these differences of less than .1°C. can be discriminated. As a means of measuring absolute temperatures, however, the skin is very unreliable, on account of the changeability of its sensation zero. We can localize temperature sensa- tions much as tactile, but not so accurately. Are Toucli and Temperature Sensations of Different Modality ? Tactile and temperature feelings are ordina- rily so very diffei'ent that we can no more compare them than luminous and auditory; and if we accept the modern modified form of the doctrine of specific nerve energies (p. 191), in accordance with which the same sensory fibre when excited always arouses a sensation of the same quality because it excites the same brain-centre, it is hard to con- ceive how the same fibre could at one time arouse a tactile, and at another a temperature sensation. It has, however, been maintained that touch and temperature feelings 564 THE HUMAN BODY. sometimes pass into one another insensibly, and that they are really extreme varieties of one kind of sensation, comparable to yellow and blue in the visual series. If a half dollar cooled to 5° 0. (41° F.) be placed on a person's brow, and ihen two (one on the other) warmed to 37° C. (98.5° F.), he commonly thinks the weight in the two cases is equal; i.e. the temperature difference leads to errors in his pressure perceptions, the cold body seeming heavier. But this does not prove an essential identity in the sensa- tions; it is conceivable, e.g., that the cold half dollar pro- duces contraction of the cutaneous tissues, leading to com- pression of the tactile end organs, which is mistaken, in mental interpretation, for a heavier external object. When sensations are combined in other cases, as red and blue- green to produce white, or partial tones to form a com- pound, we either cannot at all, or only with difficulty, recognize the components; while in this case the person feels both the cold and pressure distinctly when the half dollar is laid on him. It has also been shown that in certain cases a person mis- takes the contact of a piece of raw cotton with a sm^all ex- posed area of his skin, for the approach of a warm object; and this has been taken to prove that touch and temperature feelings may gradate into one another. However, the feeble touch of the raw cotton might well be less felt than the increased temperature of the skin, due to diminished radia- tion when it was covered by this non-conducting substance; and the constancy with which, in the ordinary circumstances of life, we feel and discriminate clearly, on the same skin region, both temperature and touch sensations, is very strongly against any transitional passage of one into the other. In favor of the view that touch and temperature are sen- sations of distinct modality, with different end organs, afferent nerve-fibres, and brain-centres, are several weighty facts. The regions of most acute discrimination for each sensation are different; and cases of disease are recorded in which persons have been extremely sensitive to variations in temperature, while their tactile sensibility was unaltered ; MUSCULAR FEELINGS. 565 and, conversely, cases in which the patient could feel that he had been touched but was unable to say whether with a hot or cold object. The Muscular Sense. In connection with our muscles we have sensations of great importance, although they do not often become so obtrusive in consciousness as to arouse our attention until we look for them. Certain of these feelings {muscle sensations proijer) are due to the excitation of sensory nerves ending within the muscles themselves: the others {innervation sensations) have probably a central origin and accompany the starting of volitional impulses from brain-cells; they are only felt in connection with the voluntary skeletal muscles. The proper muscle sensations only become marked on powerful or long-continued muscular effort (cramp, fatigue), but a lower grade of them, not distinctly perceived, proba- bly accompanies all muscular activity. The innervation feelings are of far more consequence. They accompany the slightest movement of a skeletal mus- cle, and we derive from them means of determining with great accuracy the force and extent of the contraction willed. The belief that their origin is central mainly rests on the fact that we have sensations, not merely of executed but of intended movements. The actual nature of the movement performed is probably characterized by other contemporary sensations, as of the muscle sense proper, from pressure and folding of the skin, and so on. The innervation feelings thus stand apart and opposed to all our others as primary factors in our mental life; they represent the reactive work of the organism with respect to its environment. Some distinguished physiologists, however, deny their existence, ascribing them all to a peripheral origin, cither in sensory muscle-nerves, or in skin-nerves affected when a part of the Body is moved. As, however, we can determine more accurately the differ- ence between two weights when we lift them than we can when they are merely laid on a hand supported by a table (see below), there are undoubtedly (apart from cramp and fatigue) true muscular sensations distinct from tactile; and 566 THE HUMAN BODY. tliut these, or some of tlium, are central in origin seems proved by certain phenomena observed in disease. Persons suffering ivom. jjarcsis, i.e. muscuhir weakness not amount- ing to complete paralysis, make (until they have learnt to interpret correctly their sensations under the new condi- tions) entirely wrong judgments as to the extent of their movements; they think they have contracted their muscles much more than is really the case. If the muscular sense depended on stimulation of sensory nerves in the mus- cles this error could not arise, for the intensity of the sen- sation would be determined by the amount of contraction which actually occurred. Under the circumstances, how- ever, a stronger volition than previously is needed to cause a movement of given extent or power; and we can readily explain the mistakes if we suppose the sensation is of cen- tral origin, and determined in intensity by the will-iiower exerted. We have already seen how closely muscular sensations are combined with visual in enabling us to form judgments as to the distance, size, and movements of objects. They are as closely combined in ordinary life with tactile sensa- tions; in the dark, when an object is of such size and form that it cannot be felt all over by any one region of the skin, we deduce its shape and extent by combining the tactile feelings it gives rise to, with the muscular feelings accom- panying the movements of the hands over it. Even when the eyes are used the sensations attained through them mainly serve as short-cuts which we have learned by experience to interpret, as telling us what tactile and muscular feelings the object seen would give us if felt; and, in regard to distant points, although we have learnt to apply arbitrarily selected standards of neasurement, it is probable that distance, in relation to perception, is primarily a judgment as to how much muscular effort would be needed to come into con- tact with the thing looked at. "When w^e wish to estimate accurately the weight of an object we always, when possible, lift it, and so combine mus- cular with tactile sensations. By this means we can form much better judgments. While with touch alone just percej)- CO MM ox SENSATIONS. 56? tibly different pressures have the ratio 1 : 3, with the mus- cular sense differences of y^ can be perceived. Common Sensations. Under tliis name are included the sensations Avluch we do not mentally attribute to the prop- erties of external objects, but to conditions of our own Bodies; of them we may here consider pain, hunger, and thirst. Pain arises when powerful mechanical or thermal stimuli act on the skin, and when a sensory nerve-trunk (except the optic, auditory, or olfactory) is directly excited. Most commonly we derive the feeling through cutaneous or sub- cutaneous nerves, and hence it has been supposed that pain is not a sensation of distinct modality due to the excitation of special nerve-fibres, but is dependent on excessive excite- ment of ordinary tactile fibres; and pressure or temperature sensations do undoubtedly gradate into painful as the stimuli increase. If this be so, pain is a sort of incoordi- nate or " convulsive" sensation. We shall see in Chapter XXXV. that, when a sensory nerve is normally excited in a decapitated animal, regular purpose-like reflex movements result: but if the stimulus be very powerful, or a nerve- trunk be directly excited, then inco-ordinate convulsions occur- the afferent impulses radiate farther in the centre and produce a new and useless result. We may supj^ose something similar to occur in the cutaneous nerves of an animal still possessing sensory brain-centres, if the stimuli acting on the skin are such as to excite the end organs very powerfully, or the sensory fibres directly without the inter- mediation of end organs; that a new sensation should be thus aroused, different from tactile though gradually shad- ing off into them, is a phenomenon comparable with the production of new color sensations by combinations of the fundamental ones. In such case, too, we could understand the difference of kinds of "pain" in a more general sense of the word; muscular cramp, dazzling, and disagreeably shrill or inharmouiously combined tones, might all be looked upon as inco-ordinate sensations, each with a charac- acter of its own determined by the central apparatus ex- cited. 5 (J 8 THE HUMAN BODY. The niiilter cuniiot, however, bout present decided. The skin seems indubitably to contain many nerve-fibres which terminate in free axis cylinders without end organs; and these may well be true pain-fibres, fitted to respond to stronger stimuli than those which excite the tactile and thermic end organs. Certain pathological and experi- mental phenomena tend also to prove that the brain-centres concerned in the production of tactile and painful sensa- tions arc different. Persons sometimes lose pain sensations and keej) tactile; a gentle touch is felt as well as usual, but a powerful pinch causes no pain: in one stage of ether and chloroform narcosis the same thing is observed; the sur- geon's hand and knife are felt where they touch the skin, but cutting deeper into the tissues produces no pain. In animals a similar state of things may be produced by divid- ing the gray matter of the cord, leaving the posterior white columns intact; while, if the latter be divided and the gray substance loft uninjured, there is increased sensitiveness to l)ain, and probably touch proper is lost, though that is im- possible to say with certainty. Such experiments make it pretty certain that when sensory afferent impulses reach the cord at any level and there enter its gray matter with the posterior i-oot-fibres, they travel on in different tracks to conscious centres; the tactile coming soon out of the gray network and coursing on in a readily conducting white fibre, while the painful first travel on farther in the gray sub- stance. It is still uncertain if both impulses reach the cord in the same fibres. The gray network conducts nerve im- pulses, but not easily; they tend soon to be blocked in it. A feeble (tactile) impulse reaching it by an afferent fibre might only spread a short way and then pass out into a single good conducting fibre in a white column, and ])roceed to the brain; while a stronger (painful) impulse would radiate farther in the gray matter, and perhaps break out of it by many fibres leading to the brain through the white columns, and so give rise to an inco-ordinate and ill localized sensation. That pains are badly localized, and worse the more intense they are, is a well-known fact, which would thus receive an explanation (see p. 579). SMELL. 569 Hunger and Thirst. These sensations, which regulate the taking of food, are peripherally localized in conscious- ness, the former in the stomach and the latter in the throat, and local conditions no doubt i^lay a part in their produc- tion; though general states of the Body are also concerned. Hunger in its first stages is probably due to a condition of the gastric mucous membrane which comes on when the stomach has been empty some time, and may be temporarily stilled by filling the organ with indigestible substances. But soon the feeling comes back intensified and can only be allayed by the ingestion of nutritive substances; pro- vided these are absorbed and reach the blood, their mode of entry is unessential; the hunger may be stayed by injec- tions of food into the rectum as well as by putting it into the stomach. Similarly, thirst may be temporarily relieved by moisten- ing the throat without swallowing, but then soon returns; while it may be permanently relieved by water injections into the veins, without wetting the throat at all. While both sensations thus depend in part on local peripheral conditions of afferent nerves (pneumogastric and glossopharyngeal), they may be also, and more power- fully, excited by poverty of the blood in foods and water; this probably directly stimulates the hunger and thirst brain-centres. Smell. The olfactory organ consists of the upper por- tions of the two nasal cavities, over which the endings of the olfactory nerves are spread and where the mucous membrane has a brownish - yellow color. This region {regto oJfactoria) covers the upper and lower turbinate bones {o, p, Fig. 89*), which are expansions of the ethmoid (p. 75) on the outer wall of the nostril chamber, the oppo- site part of the iiartition between the nares, and the part of the roof of the nose {n, Fig. 89) separating it from the cranial cavity. The epithelium covering the mucous mem- brane contains two varieties of cells arranged in several layers (2, Fig. 154). Some cells are much like ordinary columnar epithelium but with long branched processes attached to their deeper ends; the others have a large * P. 309. 570 THE HUMAN BODY. nucleus surrounded by a little protoplasm; a slender exter- nal process reaching to the surface; and a very fine deep one. The latter cells have been supj^oscd to be the proper olfac- tory end organs, and to be connected with the fibres of the olfactory nerve, which enter the deeper strata of the epithe- lium and there divide; but it is doubtful whether both kinds of cells are not so connected. Odorous substances, the stimuli of the olfactory appa- ratus, are always gaseous and frequently act powerfully Avhen present in very small amount. We cannot, how- ever, classify them by the sen- sations they arouse, or arrange them in series; and smells are but minor sensory factors in our mental life. We commonly refer them to external objects since we find that the sensa- tion is intensified by " sniff- ing" air powerfully into the nose, and ceases when the nostrils are closed. Their peripheral localization is, how- ever, imperfect, for we con- found many smells with tastes (see below); nor can we judge well of the direction of an odorous body through the olfactory sensations which it arouses. nerves of common sensation, such substances as ammonia Fig. 15i.— Cells from the olfactory epithelium. 1, from the frog; 2. from man ; a, columnar cell, with its bi-anched deep process; 6, so-called olfactory cell; c, its narrow outer p:-ocess; d, its slender central pro- cess. 3, gray nerve-fibres of the ol- factory nerve, seen dividing into fine peripheral branches at a. also by The nose possesses which are stimulated vapor. Taste. The organ of taste is the mucous membrane on the dorsum of the tongue and possibly other parts of the boundary of the mouth cavity. The nerves concerned are the glossopharyngeals, distributed over the hind part of the TASTE. 571 tongue, and the lingual branches of the inferior maxillary division (p. 170) of the trigeminals on its anterior two thirds. On the tongue the nerves run to papillae; the circumval- late (p. 313) have the richest supply, and on these are cer- tain peculiar end organs (Fig. 155) known as taste-buds, which are oval and imbedded in the epidermis covering the side of the papilla. Each consists, externally, of a number of flat, fusiform, nucleated cells and, internally, of six or eight so-called taste-cells. The latter are much like the Olfactory cells of the nose, and are probably connected with nerve-fibres at their deeper ends. The capsule formed by the enveloping cells has a small opening on the surface; Fig. 155.— Taste-buds. each taste-cell terminates in a very fine thread which pro- trudes there. Taste-buds are also found on some of the fungiform papillae, and it is possible that simpler struc- tures, not yet recognized, consisting of single taste-cells are widely spread on the tongue, since the sense of taste exists where no taste-buds can be found. The filiform papillae are probably tactile. In order that substances be tasted they must be in solu- tion: wipe the tongue dry and put a crystal of sugar on it; no taste will be felt until exuding moisture has dissolved some of the crystal. Tastes proper may be divided into sweet, bitter, acid, and saline. Intellectually they are, like smells, of small value; the perceptions we attain through them as to qualities of external objects being of little use. 572 THE HUMAN BODY. except as aiding in the selection of food, and for that pur- pose they are not by any means safe guides at all times. Many so-called tastes (flavors) are really smells; odorif- erous particles of substances which are being eaten reach the olfactory region through the posterior nares and arouse sen- sations which, since they accompany the presence of objects in the mouth, we take for tastes. Such is the case, e.g., with most spices; when the nasal chambers are blocked or inflamed by a cold in the head, or closed by compressing the nose, the so-called taste of spices is not perceived when they are eaten; all that is felt, when cinnamon, e.g., is chewed under such circumstances is a certain pungency due to its stimulating nerves of common sensation in the tongue. This fact is sometimes taken advantage of in the jiractice of domestic medicine when a nauseous dose, as rhubarb, is to be given to a child. Tactile sensations play also a part in many so-called tastes. Most persons taste bitters best with the back of the tongue and sweets towards the tip; but this is not constant. The curious interference of tastes which takes place when the acidity of a sour body is covered by adding a sweet one, which does not in any way chemically neutralize the acid (when sugar is put on a lemon for example), needs explana- tion. CHAPTER XXXV. THE FUNCTIONS OF THE BKAm AND SPINAL CORD. The Special Physiology of Nerve-Centres. We have already studied the general physiological properties of nerves and nerve-centres (Chaj^. XIII.) and found that while the former are mere transmitters of nervous impulses, the latter do much more. In some cases the centres are auto- matic; they originate nerve impulses, as illustrated by tlie beat of the heart under the influence of its ganglia. In other cases a feeble impulse reaching the centre gives rise to a great discharge of energy from it (as when an unex- pected noise produces a violent start, due to many impulses sent out from the excited centre to numerous muscles), so that certain centres are irritable; they contain a store of energy-liberating material which only needs a slight dis- turbance to upset its equilibrium and produce many efferent impulses as the result of one afferent. Further, the im- pulses thus liberated are often co-ordinated. When mucus in the windpipe tickles the throat and excites afferent nerve impulses, these, reaching a centre, cause discharges along many efferent fibres, so combined in sequence and power as to produce, not a mere aimless spasm, but a cough which clears the passage. In still other cases the excita- tion of centres, with or without at the same time some of the above phenomena, is associated with sensations or other states of consciousness. We have now to study which of these powers are manifested by different parts of the cen- tral cerebro-spinal neiwous system, and under what circum- stances and m what degree: what is known of the general functions of the sympathetic and sporadic ganglia has al- ready been stated (p. 183). 574 THE nmr AN BODY. The Spinal Cord as a Centre. The spinal cord, forming (except tlio slender sympathetic) the only direct communi- cation between the brain and most of the nerves of the Body, was considered by the older physiologists as merely a huge nerve-trunk, into which the various spinal nerves were collected on their way to the encephalon. It does, it is true, contain the paths for the conduction of all those im- pulses which, originating in the cerebrum, give rise to voluntary movements of the trunk and limbs; also for all the centrally traveling impulses which give rise to sensa- tions ascribed to those parts; and it is also the path for cer- tain impulses giving rise to involuntary movements as, for example, those which, originating in the respiratory centre, travel to the phrenic and intercostal nerves. If, however, the cord were merely collected and con- tinued nerve-roots it ought to increase considerably in bulk as it approached the skull, and this it does not do in any- thing like the required proportion; a histological examina- tion, too, shows that few, if any, of the entering fibres pro- ceed directly to the brain; they pass to the central core of gray substance, containing nerve-cells and different from anything in a true nerve-trunk. In the gray core some fibres (anterior roots) end in cells of the anterior horns, and others (posterior roots) in that fine network which pervades the whole gray substance (p. 179); the nerve-cells by their branches are also in continuity with this network and so, ultimately, with all the nerve-fibres entering the cord. From the network fibres arise at different levels, and pass out into the white columns; some place two regions of the gray matter in intimate connection, while others go on to the brain. In order to understand physiological facts we must assume, first, that a nervous impulse entering the gray network at any point may, under certain conditions, travel all through it, and give rise to efferent impulses emerging at any level; and, on the other hand, that there are certain lines or paths of easiest propagation between different points in this network, which the impulses keep to under ordinary conditions. Keflex Actions. When a frog is decapitated it lies down BEFLEX ACTIONS OF THE COBD. 575 squat on its belly instead of assuming the more erect jjosi- tion of the uninjured animal; its respiratory moyements cease (their centre being removed with the medulla); the hind legs at first remain sprawled out in any position into wliich they may happen to fall, but after a time are drawn up into their usual j^osition, with the hip and knee joints flexed; having made this movement the animal, if protected from external stimuli, makes no other by its skeletal muscles; it has lost all spontaneity, and only stirs under the influence of immediate excitation. Nevertheless the heart goes on beating for hours; the muscles and nerves, when examined, are found to still have all their usual i^hysiological proper- ties; and, by suitable irritation, the animal can be made to execute a great variety of complex movements. But it is no longer a creature with a will, doing things which we cannot predict; it is an instrument Avhich can be played upon, giving different responses to different stimuli (as dif- ferent notes are produced when different keys of a piano are struck), and always the same reaction to the same stimulus; so that we can say beforehand what will happen when we touch it. Such actions are called 7'eflex or excito-motor and fall into two groups; (1) orderly or inirpose-like reflexes, which are correlated to the stimulus and are often defensive, tending, for instance, to remove an irritated part from the irritating object; (2) disorderly or convulsive reflexes, not tending to produce any definite result, and affecting either a limited region or all the muscles of the body. In higher animals similar phenomena may be observed. If a rabbit's spinal cord be divided at the bottom of the neck the animal is at first thrown into a flaccid limp condition like the frog, but it soon recovers. Voluntary movements in muscles supplied from the spinal cord behind the section are never seen again; but on pinching the hind foot it is forcibly withdrawn. Men, whose spinal cord has been divided by stabs or disease below the level of the fifth cervical spinal roots (above which the fibres of the phrenic nerve, which are necessary for breathing, jiass out), sometimes live for a time, but can no longer move their legs by any effort of the will, nor do they feel touches, pinches, or hot 576 THE HUMAN BODY. things applied to them; if, however, the soles of the feet he tickled the legs are thrown into vigorous movement. As a rule, however, orderly reflexes are less marked and less numerous in the higher animals; in them the organiza- tion is less machine-like, the sj)inal cord being more the servant of the larger brain, and less capable of working without directions. Such animals, when intact, can to a greater extent control the muscular responses which shall be made to stimuli under various conditions; they have less automatic protection in the ordinary risks of life, but a greater range of possil:)le protection. The human spinal cord, controlled by the Ijrain, can adapt the reactions of the Body, with great nicety, to a vast variety of conditions; the frog's cord by itself does this for a smaller number of possi- ble emergencies without troul)ling at all such brain as the animal has, but is less completely under the control of the higher centres for adajDtation to other conditions. The difference being, however, but one of degree and not of kind, it is best to approach the study of the reflex actions of the human spinal cord through an examination of those cxhiljited by the frog. The Orderly Reflex Movements of a Decapitated Frog. For the occurrence of these the following parts must Ije in- tact; {a) the end organs of sensory nerve-fibres; {h) afferent filu'cs from these to the cord; {(•) efferent fil)rcs from the cord to the muscles; {d) the part of the spinal cord l^etween the afferent and efferent fibres; {e) the muscles concerned in the particular movement. If the animal be suspended vertically after the shock of the operation is over, it makes a few attempts to hold its hind legs in their usual flexed position; these soon cease, the legs hang down, and the creature comes to rest. If one flank be now gently scratched with the point of a ])encil a reflex movement occurs, limited to the muscles of that region; they twitch, some- wdiat as a horse's neck when tickled by flies. If a jiinch be given at the same spot, more muscles on the same side come into play; a harder pinch causes also the hind leg of that side to be raised to push away the offending object; more violent and prolonged irritation causes all the muscles DISORDEELY REFLEX MOVEMENTS. 577 of the body to contract, and the animal is convulsed. Here then we see that a feeble stimulation causes a limited and purpose-like response; stronger causes a wider radiation of efferent impulses from the cord and the contraction of more muscles, but still the movements are co-ordinated to an end; while abnormally powerful stimulation of the sensory nerves throws all the motor fibres arising from the cord into activity, and calls forth inco-ordinate spasmodic action. The orderly movements are very uniform for a given stimu- lation; if the anal region be j)inched, both hind legs are raised to push aAvay the forceps; if a tiny bit of bibulous paper moistened with dilute vinegar be put on the thigh, the lower part of that leg is raised to wipe it off; if on the middle of the back near the head, both feet are wiped over the spot; if on one flank, the leg and foot of that side are used, and so on; in fact, by careful working, the frog's skin can be mapped into many regions, the application of acidu- lated water to each causing one particular movement, due to the co-ordinated contractions of muscles in different combinations, and never, under ordinary circumstances, any but that one movement. The above purpose-like reflex movements may all be characterized as defensive, Init all orderly reflexes are not so. For example, in the breeding season the male frog clasps the female for several days with his fore limbs. If a male at this season be decapitated and left to recover from the shock, it will be found that gently rubbing his sternal region with the finger causes him to clasp it vigorously. Disorderly Reflexes or Reflex Convulsions. These come on when an afferent nerve-trunk is stimulated instead of the tactile end organs in the skin; or when the skin is very powerfully excited; or. with feeble stimuli, in certain diseased states {pafhologlcal tetanus), and under the in- fluence of certain poisons, especially strychnine. If a frog or a warm-blooded animal be given a dose of tlie latter drug, a stimulus, such as normally would excite only limited orderly reflexes, will excite the whole cord, and lead to discharges along all the efferent fil^res so that general con- vulsions result. It has been clearly proved that, in such 578 THE HUMAN BODY. cases, not the skin, or afferent or efferent nerves, or the muscles, but ilie spinal cord itself is affected by the poison (at least iirimarily), unless unnecessarily large doses have been given. The Least-Resistance Hypothesis. In order to com- prehend reflex acts we must assume a manifold union of sensory with efferent nerve-fibres; this is anatomically afforded by the minute plexus of the gray network, which is continuous through the whole cord and in which the fibres of the anterior and posterior nerve-roots, directly or indirect- ly end. The continuity of this network serves to exj^lain general reflex convulsions, and the spread of an afferent impulse, or its results, through the whole cord, Avith the consequent emission of efferent impulses through many or all the anterior roots; but, on the other hand, it renders it difficult to understand limited and orderly reflexes, in which only a few efferent fibres are stimulated. To explain them we have to assume a great resistance to conduction in the gray network, so that a nerve impulse entering it is soon blocked and transmuted into some other form of energy; hence it only reaches efferent fibres originating near the point at which it enters, or fibres placed in siiecially easy communication with that. When the frog's flank is tickled, only muscles innervated from anterior roots on the same side of the body, and springing from the same level of the cord, are made to contract; when the stimulus is more powerful the stronger afferent impulse radiates farther, but mainly in directions determined by lines of conduc- tivity in the cord; e.g., to the origin of the efferent fibres which cause lifting of the hind leg to the irritated spot. These paths of easiest conduction, or of least resistance, in some cases lie in the gray matter itself, in others in inter- central or commissural fibres of the highly conductive medullated kind, which, passing out of the gray substance at one level, run in the white columns to it at another, where the efferent fibres of the muscles called into play originate. A still stronger afferent impulse radiates wider still, and, liberating energy from all the nerve-cells in the gray matter, produces a useless general convulsion. Under CONDUCTIOX IN THE CORD. 579 the influence of stryclmine and in pathological tetanus (as observed, for example, in liydrophobia) the conductivity of the whole gray matter is so increased that all paths through it are easy, and so a feeble afferent impulse spreads in all directions. To account for the phenomena of localized skin sensa- tions (p. 559) and of limited voluntary movements we must make a similar hypothesis. If the nervous impulses enter- ing the gray network when the tip of a finger is touched sjiread all through it irregularly, we could not tell what region of the skin had been stimulated, for the central results of stimulating the most varied peripheral parts would be the same. From each region of the gray network of the cord where a sensory skin-nerve enters there must, therefore, be a special path of conduction to a given brain- region, producing results which differ recognizably in con- sciousness from those following the stimulation of a differ- ent skin region. The acuteness of the localizing power will largely depend on the definiteness of the path of least resis- tance in the gray matter, since while traveling in a medul- lated nerve-fibre from the skin to the cord, or (in the white columns) from the gi'ay matter of the latter to the brain, the nervous imjnilse is confined to a definite track. Hence anything tending to let the afferent impulse radiate when it enters the cord «'ill diminish the accuracy with which its peripheral origin can be located. This we see in violent pains; a whitlow on the finger affects only a few nerve-fibres, but gives rise to so powerful ner-ve impulses that when they reach the cord they spread widely and, break- ing out of the usual track of propagation to the brain, give rise to ill-localized feelings of pain often referred ail the way up the arm to the elbow. So the pain from one diseased tooth is often felt along half a dozen, or all over one side of the head. Such cases are comparable to the transforma- tion of an orderly reflex into a general convulsion when the stimulus increases. As an animal shows no spontaneous movements when its cerebral hemispheres are removed, we conclude that the nerve impulses giving rise to such movements start in those 680 THE HUMAN BOLT. parts of the brain. Thence they travel down the white columns of the cord to its gray matter, which they enter at different levels, each in the neighborhood of a centre for producing a given movement. If they there radiated far and wide no definite movement could result, for all tlie muscles supplied from the cord would be made to contract, and not merely those necessary to bend tlie index finger, for example. We must here again, therefore, assume a path of least resistance for the propagation of nerve impulses from a given fibre coming down from the brain, to the efferent fibres going to a certain muscle or group of muscles. The path between the two is almost certainly not direct; a co-ordinating spinal centre intervenes, and all that the brain has to do is to excite this centre, which then secures the proper muscular co-ordination. If the hand be laid flat on the table and its palm be rolled over, many muscles, in- cluding thousands of muscular fibres, have to contract in definite order and sequence. Persons who have not studied anatomy and who are quite ignorant of the muscles to be used can perform the movement perfectly; and even a sl-cilled anatomist and physiologist, if he knew them all and their actions, could not by conscious effort combine them so well as the cord does without such direct interfer- ence. We have then to look on the cord as containing a host of co-ordinating centres for different muscles. These centres are put in nervous connection, on the one hand, with certain regions of the skin, and, on the other, with regions of the brain, and may be excited from either; in the former case the movement is called reflex; in tlie latter it may be reflex, or may be accompanied by a feeling of " will" and is then called voluntary. The more accurately the required centre, and no other, is excited, the more definite and precise the movement. The Education of the Cord- Much of what is called educating our touch or our muscles is really education of the spinal cord. A person who begins to play the piano finds at first much difficulty in moving his fingers inde- pendently; the nervous impulses from tlio l)rainto the cord radiate from the spinal centres of the muscle which it is THE lyni^TION OF REFLEX ACTIONS. 581 desired to move, to others. But with practice the indepen- dent movements become easy. So, too, the localizing power of the skin can be greatly increased by exercise (p. 5(30) as one sees in blind persons, who often can distinguish two stimuli on parts of the skin which are so near together as to give only one sensation to other peojile. Such phenomena depend on the fact that the more often a nervous impulse has traveled along a given road in the gray matter, the easier does its path become, and the less does it tend to wander from it into others. We may compare the gray matter to a thicket; persons seeking to beat a road through from one point to another would keep the same general direction, determined by the larger obstacles in the way, but all would diverge more or less from the straight path on account of undergrowth, tree trunks, etc., and would meet with cojisid- erable difficulty in their progress. After some hundreds had passed, however, a tolerably beaten track would be marked out, along which travel was easy and all after-comers would take it. If instead of one entry and one exit we imagine thousands of each, and that the paths between certain have been often traveled, others less, and some hardly at all, we get a pretty good mental picture of what happens in the passage of nervous impulses through the gray matter of the cord; the clearing of the more trodden paths answering to the effects of use and practice. The human cord and that of the frog must not, however, be looked upon as pathless thickets at the commencement; each individual inherits certain paths of least resistance determined by the structure of the cord, which is the transmitted material result of the life experience of a long line of ancestry. The Inhibition of Reflexes. Since it is possible, as by strychnine, to diminish the resistance in the gray matter, it is conceivably also possible to increase it, and diminish or pre- vent reflexes. Such is found to be actually the case. We can to a great extent control reflexes by the will; for example, the jerking of the muscles which tends to follow tickling. and it is found that after a frog's brain is removed it is much easier to get reflex actions out of the spmal cord. Certain drugs, as bromide of potassium, also diminish reflex excita- 582 777^ HUMAN BODY.. bility. If a frog's brain be remoyed and the animal's toe be dij^ped into very dilute acid, it will be removed after a few seconds; the time elapsing between the immersion and tlie lifting of the foot is known as the reflex time; any- thing diminishing reflex excitability increases this, as the stimulus (which has a cumulative effect on the centre) has to act longer before it arouses the cord to the discharging point. If the sciatic nerve of the other leg be stimulated while the toe is in the acid the reflex time is increased, or the reflex may fail entirely to apj)ear. This is one case of a general law, that any powerful stimulation of one sen- sory nerve tends to inhibit orderly reflexes due to the ex- citation of another. A common example is the well-known trick of pressing on the upper lip to prevent a sneeze. The whole question of reflex inhibition is at present very obscure. It may be due to the excitation of special fibres which inhibit reflex centres, as pneumogastric fibres do the cardio-motor; or to the fact that one nerve impulse in the cord in some cases blocks or interferes with another; or l^artly to both. Psychical Activities of the Cord. Since we can get quite marked reflex movements in the lower part of the Body of a man whose cord is divided and who cannot voluntarily move his lower limbs, and on questioning him find that he feels nothing and is quite ignorant of his movements unless he sees his legs, it is most probable that the sjiinal cord in all cases is devoid of centres of conscious- ness and volition: this is not certain, however; for tliere might well be a less division of physiological labor between llic cord and brain of a frog, than between those of a man. Still we are entitled to good evidence before we admit that two things so similar as the human cord and the frog's pos- sess different properties. Co-ordinated movements follow- i:iga given stimulus, or cries emitted by an animal, will not suffice to prove that it is conscious, since these we know may occur entirely unconsciously in men, who alone can tell us of their feelings. We must look for something that resembles actions only done by men consciously. In the frog it has been maintained that we have evidence of such. If a bit of PSYCHICAL ACTIVITIES OF TEE SPIHAL CORD 583 acidulated paper be put on a decapitated frog's thigh, the animal will bend its knee and use its leg to brush off the irritant; always using this same leg if the stimulus be not so strong as to produce disorderly reflexes. If now the foot be tied down so that the frog cannot raise it, after a few ineffectual efforts it will move the other leg, and may wipe the paper off with it. This it has been said shows a true psychical activity in the cord; a conscious and volun- tary employment of new procedures under unusual circum- stances. But a close observation of the phenomenon shows that it will hardly bear this interpretation; the movements of the other leg are very irregular and inco-ordinate, and much resemble reflex convulsions stirred up by the pro- longed action of the acid, which goes on stimulating the skin nerves more and more powerfully. Even if new muscles came, in an orderly way, into play under the stronger stimulus, that would not prove a volitional con- scious use of them; we see quite similar phenomena when there is nothing purpose-like in the movement. Many dogs reflexly kick violently the hind leg of the same side when one flank is tickled. If this leg be held and the tickling continued, very frequently the opposite hind leg will take on the movements, which it never does in ordinary circumstances. This is quite comparable to the frog's use of its other leg under the circumstances above described, but here it would be obviously absurd to talk of a volitional source for such a senseless movement. / Putting together, then, the testimony of persons with / injured spinal cords, and the observations made on them 1 and on animals, we may tolerably safely conclude that the \ cord contains no centres of consciousness. There are, how- ' ever, persons who maintain that in such cases the cord itself I feels though the individual does not, whatever that may I mean; if the statement is used merely to imply that the I cord is irritable (just as a muscle is) no one denies it; but it is an unnecessarily confusing method of stating the fact. The Cord as a Transmitter of Nervous Impulses. In the gray substance, as we have seen, there is reason to believe that nervous impulses can pass in all directions, 584 THE HUMAN BODY. though they usually take certain easiest roads. The main tracks between bram and cord, however, lie in the white columns; nerve-roots enter the gray matter, and from each region of the latter nicdullated fibres arise, pass out into a white column, and continue to the brain. Once in these fibres the impulse has to keep to a definitely marked out anatomical path, which leads to a brain centre or centres. These paths lie mainly in the lateral columns, the affe- rent (sensory) impulses tending, however, to spread into the posterior columns, and the efferent (volitional) into the anterior; the main bulk of the posterior and anterior columns seems to be made up of commissural fibres join- ing different levels of the cord. The sensory fibres for the most part cross soon after they enter the gray matter, and proceed onwards mainly on the side opposite to that of the nerve-root which conveyed them to the cord, while the effe- rent cross mainly in the medulla before they reach the cord; the crossing seems in neither case complete. Hemisection of the cord, therefore, causes marked, but not absolute, loss of voluntary movement in muscles of the same side supplied with nerves from a part of the cord below the level of the section; and a considerable, but not entire, loss of sensi- bility on the opposite side. Impulses so powerful as to lead to feelings of pain travel mainly in the gray matter (p. 568), and they also for the most part cross the middle line soon after their entry. The Functions of the Brain in General. The brain, at least in man and the higher animals, is the seat of con- sciousness and intelligence; these disappear when its blood- supply is cut off, as in fainting; pressure on parts of it, as by a tumor or by an effusion of blood in apoplexy, has the same result; inflammation of it causes delirium; and when the cerebral hemispheres are unusually small idiotcy is observed. The brain has, however, many other important functions; it is the seat of many reflex, automatic, and co-ordinating centres, which may act as entirely apart from consciousness as those of the spinal cord; experiment makes it probable that psychical faculties are dependent on the fore-brain, while the rest of the complex mass has other, CEREBRAL FUNCTIONS. 585 non-mental, duties. If the cerebral hemispheres be re- moved from a frog, the animal can still perform every movement as well as before; but it no longer performs any spontaneously; it must be aroused by an immediately acting stimulus, and its response to this is as invariable and predicable as that of a frog with its spinal cord only. The movements which can be educed are, however, far more complex; instead of mere kicks in various directions the animal can walk, leap, swim, get off its back on to its feet, and so on. Similar results are observable in pigeons whose fore-brain has been removed; mammals bear the operation badly, but some, as rats, survive it several hours and then exhibit like phenomena. The creatures can move, but do not unless directly stimulated; all their volitional spontaneity is lost, and, apparently, all perceptions also; they start at a loud noise, but do not run away as if they conceived danger; they follow a light with the eyes, but do not attempt to escape a hand stretched forth to catch them; they can and do swallow food placed in the mouth, but would die of starvation if left alone with plenty of it al^out them, the sight of edible things seeming to arouse no idea or conception. It may be doubted, perhaps, whether the animals have any true sensations; they start at sounds, avoid opaque objects in their road, and cry Avhen pinched; but all these may be unconscious reflex acts: on the whole it seems more probable, however, that they have sensations but not perceptions; they feel redness and blueness, hard- ness and softness, and so on; but sensations, as already pointed out, tell in themselves nothing; they are but signs which have to be mentally interpreted as indications of ex- ternal objects: it is this interpreting power which seems deficient in the animal deprived of its fore brain. Functions of the Medulla Oblongata. This contains the paths of conduction between the parts of the brain in front of it and the spinal cord. It is also the seat of many important reflex and automatic centres, especially those governing the organs immediately concerned in the maintenance of life; as the respirator \% circulatory, and masticatory. It may therefore be called the "nerve cen- 586 THE HUMAN BODY. tral organ of tlie nutritive processes." The physiological action of most of the medullary centres has already been described; the more important are — 1. The respiratory centre (p. 390). 2. The cardio-inhibitory centre (p. 250); (there is also some reason to believe that the centre of the accelerator heart-fibres (p. 252) lies in the medulla). 3. Tlie vaso-motor centre (p. 253). 4. The centre for the dilator muscle-fibres of the pupil (p. 486). 5. The centre for the muscles of chewing and swallowing (p. 337), which are commonly thrown into action reflexly, though they may be made to contract voluntarily. 6. The convulsive centre (p. 400). 7. The diabetic centre (p. 442). 8. The centre reflexly exciting activity in the salivary glands, when sensory nerves in the mouth are stimulated. 9. Certain centres for complex bodily movements; an animal with its medulla oblongata can execute much more complicated reflex acts than one with its spinal cord alone. Functions of the Cerebellum, Pons Varolii, and Mid- Brain. These contain paths of conduction between the fore-brain and parts behind, and many important centres, especially those concerned with the maintenance of the equilibrium of tlie Body in various postures and modes of locomotion. If, as has been above suggested, an animal without its cerebral hemispheres has sensations which remain untranslated into terms of external things, these sensations must have their seats in these brain-regions. The crura cerebri (p. 164) are essentially paths of conduction between the cerebral hemispheres and parts behind. Injury of one produces partial loss of sensibility and incomplete muscular paralysis on the ojoposite side of the Body. The anterior pair of eminences of the corpora quadrigemvna are concerned with sight; stimuli reaching thcra through the optic nerve, there, probably, first cause visual sensations, which it is left to the fore-brain to interjiret. If the latter is removed from an animal liglit brought in front of the eye still causes contraction of the j^upil; direct irritation of the eminences in question has the same effect, and destruction of one of them causes blindness of the oppo- site eye. The functions of the posterior pair of eminences FUNCTIONS OF CEREBELLUM. 587 of the corpora quadrigemina in man are uncertain. The cerebellum is the chief organ of combined muscular move- ments; it is the main seat of what we may call acquired reflexes. Every one has to learn to stand, walk, run, and so on; at first all are difficult, but after a time become easy and are performed unconsciously. In standing or walking very many muscles are concerned, and if the mind had all the time to look directly after them we could do nothing- else at the same time; we have forgotten how we learnt to walk, but in acquiring a new mode of progression in later years, as skating, we find that at first it needs all our atten- tion, bnt when once learnt we have only to start the series of movements and they are almost unconsciously carried on for us. At first we had to learn to contract certain muscle groups when we got particular sensations, either tactile, from the soles, or muscular, from the general jDosition of the limbs, or visual, or others (equilibrium sensations, see below) from the semicircular canals. But the oftener a given group of sensations has been followed by a given muscular contraction the more close becomes the associ- tion of the two; the path of connection between the afferent and efferent fibres becomes easier the more it is traveled, and at last the sensations arouse the proper move- ment without volitional interference at all, and while hardly exciting any consciousness; we can then walk or skate without thinking about it. The will, which had at first to excite the proper muscular nerve-centres in accordance with the felt directing sensations, now has no more trouble iu the matter; the afferent impulses stimulate the proper motor centres in an unconscious and unheeded way. Injury or disease of the cerebellum produces great disturbances of locomotion and insecurity in maintaining various pos- tures. After a time the animals (birds, which bear the operation best) can walk again, and fly, but they soon become fatigued, probably because the movements require close mental attention and direction all the time. Sensations of Equilibrium. The semicircular canals have probably nothing to do with hearing. An old view was that, lying in three planes at right angles to one another. 588 THE HUMAN BODY. they served to distinguish tlic direction of sound-waves reach- ing the ear; but as the direction of oscillation of the tym- panic ossicles is the same, no matter what that of the sound- waves entering the external auditory meatus may be, such an hypothesis has no foundation. The cochlea abundantly accounts for the ap})reciation of notes, and such noises as are due to inharmonically combined tones; while the vestibule will suffice for other noises: and it is found that disease of the semicircular canals does not interfere with hearing, but often causes uncertainty in movements and feelings of giddiness. Experiment shows that cutting a semicircular canal is followed by violent movements of the head in the plane of the canal divided; the animal staggers, also, if made to walk; and, if a pigeon and thrown into the air, cannot fly. All its muscles can contract as before, but they are no longer so co-ordinated as to enable the animal to maintain or regain a position of equilibrium. It is like a creature sufl'ering from giddiness; and similar phenomena follow, m man, electrical stimulation of the regions of the skull in which the semicircular canals lie. Such facts suggest that the semicircular canals are organs in which sensory afilerent impulses, assisting in the pre- servation of bodily equilibrium, arise. The unconscious maintenance of the erect position depends on thQ excitation of many co-ordinating motor centres through tactile, mus- cular, visual, and aural sensations; all acting together in normal combination these enable us to stand without thought; but loss of any one, or its abnormal state, will throw the whole mechanism out of gear. Persons who have lost muscular or tactile sensibility stand and walk with diffi- culty; those who have nystagmus (jerking unconscious movements of the eyeballs which cause the visual field to seem to move in space) do the same and feel giddy; and, as we have Just seen, similar phenomena follow injury of the semicircular canals. How the nerves in the latter are stimulated is not certain; being filled with liquid the pressure of this on any ampulla will be increased when the head is bent so as to place that one below; and this may be the excitant; giving rise to EQUILIBRIUM SENSATIONS. 589 afferent impulses, which change the condition of the co-ordinating locomotor centres, with every position of the liead. Or, movements of the endolymph m relation to the wall of the canal may be the stimulus, the current swaying the projecting hairs (Fig. 149).* Place a few small bits of cork in a tumbler of water, and rotate the tumbler; at first the water does not move with it; then it begins to go m the same direction, but more slowly; and, finally, moves at the same angular velocity as the tumbler. Then stop the tumbler, and the water will go on rotating for some time. Now if the head be turned in a horizontal plane similar phenomena will occur in the endolymi^h of the horizontal canal; if it be bent sidewise in the vertical plane, in the anterior vertical canal; and if nodded, in the posterior verti- cal; the hairs moving with the canal would meet the more stationary water and be pushed and so, possibly, excite the nerves at the deep ends of the cells which bear them, and gen- erate afferent impulses which will cause the general nerve- centres of bodily equilibration to be differently acted upon in each case. Under ordinary circumstances the results of these impulses do not become prominent in consciousness as sensations; but they sometimes may. If one spins round for a time, the endolymph takes up the movement of the canals, as the water m the tumbler does that of the glass; on stopping, the lic[uid still goes on moving and stimulates the hairs which are now stationary; and we feel giddy, from the ears telling us we are rotating and the eyes that we are not; hence difficulty in standing erect or walking straight. A common trick illustrates this very well; make a person place his forehead on the handle of an umbrella, the other end of which is on the floor, and then walk three or four times round it, rise, and try to go out of a door . he will nearly always fail, being unable to combine his muscles properly on account of the conflicting afferent impulses. If a person, with eyes shut, be laid on a hori- zontal table which is turned, he can at first feel and tell the direction of the rotation; as it continues he loses the feelinsr, and when the movement stops feels as if he were being turned * Page 543. 590 THE HUMAN BODY. in the opposite direction. All this becomes readily intelli- gible if we sup})Ose feelings to be excited by relative move- ments of the endolymph a,nd the canals inclosing it. The so-called "auditory sacs" of many Mollnsks (see Zoology) are probably organs for equilibration sensations and comparable to our semicircular canals. Functions of the Fore-Brain. Beyond the broad fact that this part of the nervous system is essentially volitional and intellectual in function, we know very little. It is clearly not the seat of the centres of muscular co-ordination; for, after its removal, an animal can still, if properly stimu- lated, execute j^erfectly all its usual movements. The true motor centres lie farther back; those for the less compli- cated combinations, as bending a limb, in the spinal cord, and those for more complex, as standing, walking, or breathing, in the mid- and hind-brains: and are not auto- matic. They may be excited to activity either, reflexly, by afferent impulses, traveling in from sensory regions and associated or not with consciousness; or, directly, by im- pulses, associated with those states of consciousness which we call volitions, passing back from the fore-brain. The fore -brain, also, frequently inhibits movements which, in its absence, would be caused by discharges, reflex in nature, of the lower centres; the will is as often employed in restraining as in exciting muscular contractions. For instance, after the cerebral hemispheres have been removed from a frog, stroking the animal gently on its back will, each time, cause it to croak; the skin stimulation origi- nates afferent impulses which excite the "croaking centre"' to discharge: but if the creature has its fore-brain it croaks or not as it pleases; it can then allow the co-ordinating mechanism to work freely under the stimulus, or check it; and it can also, independently of any immediate stimulus, excite voluntarily the same motor nerve-centre and croak if it chooses. We constantly meet with similar phenomena in ourselves; afferent impulses are all the time at work, tend- ing to produce one action or another; and a gi-eat part of our mental activity consists in deciding which we shall jn-event and which we shall permit. The restraint thus FUNCTIOXS OF THE FORE-BRAIN. 591 exercised by the fore-brain on the lower centres is at least as important as its power of exciting them; strength of character depends, perhaps, more on gTeat inhibitory power in the fore-brain than on its initiating facnlty. The intellectnal powers seem mainly, if not entirely, de- pendent for their necessary material antecedents or con- comitants on the surface convolutions of the cerebral hemispheres; if these alone be remoyod from an animal its mental condition is much the same as if the whole fore- brain be taken away. Some simple and fundamental perceptions seem, however, to remain, having, perhaps, their seats in the deeper gray masses constituting the optic thalami and corpora striata (p. 167); a dog, from which the greater part of the cerebral surfaces had been re- moved, after a time learnt to w^ilk about, apparently volun- tarily, and to find and eat his food; he even learnt not to take the bones of other dogs after he had several times been severely bitten for so doing. But more complex perceptions were lost; before the operation, for example, he was violently terrified by seeing a man fantastically dressed; but after- wards no such things seemed to arouse in him so complex a conception as that of a strange or dangerous object. Although the fore-brain is the seat of consciousness it is itself insensible to cutting or wounding; and was long sui)posed to be entirely inexcitable by general nerve stimuli. It has, however, been found that tolerably powerful electri- cal currents applied to the convolutions produce, in many cases, definite movements; the nature of the movement depending upon the area stimulated. Hence an attempt has been made to detect the functions of different j)arts of the cerebral hemispheres by observing the results of stimu- lating each; and provisionally we may, perhaps, assume that the brain-centres, from which volitional impulses proceed to the co-ordinating centres for the muscle groups called into play, lie in the cerebral regions whose stimulation is followed by the movement. The animals, however, so often recover the power of executing the movement spontaneously after its supposed volitional centre has been removed that the proper interpretation of the experimental results is still 592 TUE HUMAN BODY. doubtful. Stimulation of muuy regions of tlie brain is followed by no results; and that of others by movements the ])o\ver of voluntarily executing which is not, even tempo- rarily, lost when that brain-part is removed; these cerebral areas have been supposed to be concerned Avith mental faculties other than volition, the stimulation exciting sen- sations, perceptions, or emotions; but this is still very doubtful. Localized disease of regions of the human brain lias, so far, given better results than physiological experi- ment on the lower animals. The power of using words to express ideas seems intimately connected with a small area on the fore part of the left cerebral hemisj)here, and to be lost (producing the condition known as aphasia) when that part IS diseased; and many cases have been recorded in which a wound of the skull, has been followed exactly by loss of the power of voluntarily moving those muscle groups which (accepting the results of electrical stimulation in the lower animals) might be supposed to l)e normally excited, through the will, from the cerebral area injured. Absence of recovery unless the brain injury is cured seems, moreover, to be the rule in man; while as we have seen this is not the case in lower animals; this may, perhaps, indicate a more precise division of physiological labor in the human brain ; which is a priori probable, considering its great superiority as a mental ajiparatus. What the use of two cerebral hemispheres is, cannot at present be said. Injury of one produces its main effects, so far as sensation and motion are concerned, on the oppo- site side of the Body; but other faculties, as that of using speech, seem located on one side only; and in the brains of the higher human races the surface markings on the two sides are not perfectly symmetrical, which may indicate some difference in function. It has been suggested that in many cases we only learn to use one side of the brain, and that the other is in reserve in case of injury or disease; but the evidence is inconclusive: a good deal may, however, be said for the view that a good deal of brain in every one's skull IS never used. It is there untaught but ready to be educated. IMPORTANCE OF EXERCISING THE BRAIN. 593 MoYements which are commonly executed together tend to become so associated that it is difficult to perform one alone; many persons, e.g., cannot close one eye and keep the other open. From frequent use, the paths of con- duction between the co-ordinating centres for both groups of muscles have become so easy that a volitional imjDulse reaching one centre spreads to the other and excites both. This association of movements, dependent on the modifica- tion of brain structure by use, finds an interesting parallel in the psychological phenomenon known as the association of ideas; and all education is largely based on the fact that the more often brain regions have acted together the more readily, until finally almost indissolubly, do they so act. If we always tram up the child to associate feelings of disgust with wrong actions and of approbation with right, when he is old he will find it very hard to do otherwise: such an organic nexus will have been established that the activity of the one set of centres will lead to an excitation of that which habit has always associated with it. The nerve-centres are throughout eminently plastic; every thought leaves its trace for good or ill; and the moral truism that the more often we yield to temptation — the more often an evil solici- tation, sensory or otherwise, has resulted in a wrong act — the harder it is to resist it, has its parallel (and we can hardly doubt its physical antecedent) in the marking out of a path of easier conduction from perceptive to volitional centres in the brain. The knowledge that every weak yielding degrades our brain structure and leaves its trail in that organ through which man is the " paragon of animals," while every resistance makes less close the bond between the thought and the act for all future time, ought surely to "give us pause:" on the other hand, every right action helps to establish a ''path of least resistance," and makes its subsequent performance easier. The brain, like the muscles, is improved and strengthened by exercise and injured by overwork or idleness; and just as a man may specially develop one set of muscles and neglect the rest until they degenerate, so he may do with his brain; developing one set of intellectual faculties and 594 THE HUMAN BODY. leaving the rest to lie fallow until, at last, he almost loses the power of using tliem at all. The fierceness of the battle of life nowadays especially tends to produce such lopsided mental developments; how often does one meet the business man, so absorbed m money-getting that he has lost all })ower of aj)preciating any but the lower sensual pleasures; the intellectual joys of art, science, and literature have no charm for him; he is a mere money-making machine. One, also, not unfrequently meets the scientific man with no appreciation of art or literature; and literary men utterly in- capable of sympathy with science. A good collegiate education in early life, on a broad basis of mathematics, languages, and the natural sciences, is a great security against such imperfect mental growth; one danger in American life is the tendency to put lads in a technical college, or to start them in business before they have attained any broad general education. Another danger, no doubt, is the oppo- site one of making the training too broad; a man who knows one or two literatures fairly well, and who has mastered the elements of mathematics and of one of the observational or experimental sciences, is likely to have a better and more utilizable brain than he who has a smattering of half a dozen languages and a confused idea of all the '^ologies." The habits of mental slovenliness, the illogical thinking, and the incapacity to know when a thing really is mastered and understood, which, one so often finds as the results of such an education, are far worse than the narrowness apt to follow the opposite error, which is often associated with the power of accurate logical thought. Those who are deprived of the advantages of a general collegiate educa- tion may now, more easily than at any jDrevious period, cultivate mental breadth by reading some of the many excellent general reviews and magazines, and the readable but exact popular expositions now available on nearly all subjects, which are such a feature of our age. Associating, o:it of working hours, with those whose special pursuits aie different from our own is almost necessary to those who would avoid such an asymmetrical development as almost amounts to intellectual deformity. CHAPTER XXXVI. VOICE AND SPEECH. ■Voice consists of sounds produced by the vibrations of two elastic bands, the U'lie vocal cords, placed in the larynx, an upper modified portion of the passage which leads from the pharynx to the lungs. When the vocal cords are put in a certain position, air driven past them sets them in periodic vibration, and they emit a musical note; the lungs and respiratory muscles are, therefore, accessory parts of the vocal apparatus: the strength of the blast pro- duced by them determines the loudness of the voice. The larynx itself is the essential voice-organ: its size primarily determines the pitch of the voice, which is lower the longer the vocal cords; and, hence, shrill in children, and usually higher pitched in women than in men; the male larynx grows rapidly at commencing manhood, causing the change com- monly known as the " breaking of the voice." Every voice, while its general pitch is dependent on the length of the vocal cords, has, however, a certain range, within limits which determine whether it shall be soprano, mezzo-soprano, alto, tenor, baritone, or bass. This variety is produced by muscles within the larynx which alter the tension of the vocal cords. Those characters of voice which we express l)y such phrases as harsh, sweet, or sympathetic, depend on the structure of the vocal cords of the individual; cords which in vibrating emit only harmonic partial tones (p. 547) are pleasant; while those in which inharmonic partials are conspicuous are disagreeable. The vocal cords alone would produce but feeble sounds; those that they emit are strengthened by sympathetic re- sonance of the air in the pharynx and mouth, the action of 59G THE HUMAN BODY. wliich may bo compared to that of the sounding-board of a viohn. By movements of throat, soft palate, tongue, cheeks, and lips the sounds emitted from the larynx are altered or supplemented in various ways, and converted into articulate language or speech. The Larynx lies in front of the neck, beneath the hyoid bone and above tbe Avindpipe; in many persons it is prominent, causing the projection known as " Adam's apple." It consists of a framework of cartilages, partly joined by true synovial joints and partly bound together Cs Fig. 156. — The more important cartilages of fhe laryiix from behind. <, th>Toid; Cs, its superior, and Ci, its inferior, honi of tlie rightside; **, cricoid cartilage; t, arytenoid cartilage; Pv, the corner to wliich the posterior end of a vocal cord is attached; Prn, corner on which the muscles which approximate or separate the vocal cords are inserted ; co, cartilage of Santorini. by membranes; muscles are added which move the car- tilages with reference to one another; and the whole is lined by a mucous membrane. The cartilages of the larynx (Fig. 156) are nine in number; three single and median, and three pairs. The largest {t) is called the thyroid, and consists of two halves which meet at an angle in front, but separate behind so as to inclose a V-shaped space, in which most of the remaining cartilages lie. The epiglottis (not represented in the figure) is fixed to the top of the thyroid cartilage and overhangs the entry ANATOMY OF LARYNX. 597 from the pharynx to the larynx {c, Fig. 89);* it may be seen, covered by mucous membrane, projecting at the base of the tongue, if the latter be pushed down while the mouth is held open in front of a glass; and is, similarly covered, represented, as seen from behind, at a in Fig. 15T. The cricoid, the last of the unpaired cartilages, is the shape of a signet-ring; its broad part (^*, Fig. 156) is on the pos- terior side and lies at the lower part of the opening between the halves of the thyroid; in front and on the sides it is narrow, and a space, occupied by the crico-thyroid mem- hrane, intervenes between its upper border and the lower edge of the thyroid cartilage. The angles of the latter are produced above and below into projecting horns {Cs and Ci, Fig. 156), and the lower horn on each side forms a joint with the cricoid. The thyroid can be ro- tated on an axis, passing through the joints on each side, and rolled down so that its lower front edge shall come nearer the cricoid cartilage, the membrane there intervening being folded. The arytenoids {\, Fig. 156) are the largest of the paired cartilages; they are seated on the upper edge of the posterior wide portion of the cricoid, and form true joints with it. Each is pyramidal with a triangular base, and has on its tip a small nodule {co, Fig. 156), the cartilage of Santorini. From the tip of each arytenoid cartilage the aryteno-ejnglottidean fold of mucous membrane (10, Fig. 157) extends to the epiglottis; the cartilage of Santorini causes a projection (8, Fig. 157) in this; and a little farther on (9) is a similar eminence on each side, caused by the remaining pair of cartilages, known as the cuneiform, or cartilages of Wrisierg. The Vocal Cords are bands of elastic tissue which reach from the inner angle {Pv, Fig. 156) of the base of each arytenoid cartilage to the angle on the inside of the thyroid where the sides of the F unite; they thus meet in front but are separated at their other ends. The cords are not, how- ever, bare strings, like those of a harp, but covered over with the lining mucous membrane of the larynx, a slit, called the glottis {c, Fig. 157), being left between them. It is the * Page 309. 598 THE HUMAN BODY. projecting cushions formed h\ them on each side of this slit which are set in vibration during phonation. Above each vocal cord is a depression, tlie ventricle of the larynx^ [b' , Fig. 157); this is bounded above by a somewhat pronii- FiG. 157.— The larynx -viewed from its pharyngeal opening. The back wall of the piiarynx has been divided and its edges (11 ) turned aside. 1, body of hyoid ; 2, its small, and 3. its great, horns: 4, upper and lower horns of thyroid carti- lage; 5, mucous membrane of front of pharynx, covering the back of the cricoid cartilage; 6, upper end of gullet; T, windpipe, lying in front of the gullet; 8, eminence caused by cartilage of Santorini; !t. eminence caused by cartilage of "Wrisberg; both lie in, 10, the ari/tcno-cpir/lottidean /oW of mucous membrane, surrounding the opening (aditn^ /fn//7((//.s) from pharynx to larjnx. o, project- ing tip of epiglottis; c, the glotiis, tlie lines leading from the letter point to the free vibrating edges of the vocal cords. 6, the ventricles of the larynx: their upper edges, marking them off from the eminences b, are the false vocal cords. nent edge, the false vocal cord. Over most of the interior of the larynx its mucous membrane is thick and covered by ciliated epithelium, and has many mucous glands im- bedded in it. Over the vocal cords, however, it is repre- sented only by a thin layer of flat non-ciliated cells, and MOVEMENTS OF LARYNX. 599 contains no glands. In quiet breathing, and after death, the free inner edges of the vocal cords are thick and rounded, and seem very unsuitable for being readily set in vibration. They are also tolerably widely separated behind, the aryten- oid cartilages, to which their posterior ends are attached, being separated. Air under these conditions passes through without producing voice. If they are watched with the laryngoscope during phonation, it is seen that the cords approximate behind so as to narrow the glottis; at the same time they become more tense, and their inner edges project more sharply and form a better-defined margin to the glottis, and their vibrations can be seen. These changes are brought about by the delicately co-ordinated activity of a number of small muscles, which move the cartilages to which the cords are fixed. The Muscles of the Larynx. In describing the direc- tion and action of these it is convenient to use the words front or anterior and back or posterior with reference to the larynx itself (that is as equivalent to ventral and dorsal) and not with reference to the head, as usual. The base of each arytenoid cartilage is triangular and fits on a surface of the cricoid, on which it can slip to and fro to some ex- tent, the ligaments of the joint being lax. One corner of the triangular base is directed inwards and forwards {i.e. towards the thyroid) and is called the vocal process {Pv, Fig. 156). as to it the vocal cords are fixed. The outer posterior angle {Pm, Fig. 15G) has several muscles inserted on it and is called the muscnlar process. If it be pulled back and towards the middle line the arytenoid cartilage will rotate on its vertical axis, and roll its vocal processes forwards and outwards, and so widen the glottis; the re- verse will happen if the muscular process be drawn forwards. The muscle producing the former movement is the posterior crico-arytenoid {Cap, Fig. 1^8); it arises from the back of the cricoid cartilage, and narrows to its insertion into the muscular process of the arytenoid on the same side. The opponent of this muscle is the lateral crico-arytenoid, which arises from the side of the cricoid cartilage, on its inner surface, and passes upwards and backwards to the 600 THE HUMAX JiODT. muscular process. The posterior crico-arytenoids, working alone, pull inwards and downwards the muscular processes, turn upwards and outwards tlie vocal processes, and sepa- rate the posterior ends of the vocal cords. The lateral crico-thyroid, working alone, pulls downwards and forwards the muscular process, and rotates inwards and upwards the vocal process, and narrows the glottis; it is the chief agent in producing the approximation of the cords necessary for Taep Fig. 158.— The larj'nx seen from behind and dissected so as to display some of its muscles. The mucous membrane of the front of the pharynx (5, Fig. 157) has been dissected away, so as to display the laryngeal muscles beneath it. Part of the left half of the thyroid cartilage has been cut away, co, cartilage of Santorini ; cii, cartilage of Wrisberg. the production of voice. When both pairs of muscles act together, however, each neutralizes the tendency of the other to rotate the arytenoid cartilage; the downward part of the pull of each is, thus, alone left, and this causes the arytenoid to slip downwards and outwards, off the eminence on the cricoid with which it articulates, as far as the loose capsu- lar ligament of the joint will aWow. The arytenoid car- tilages are thus moved apart and the glottis greatly widened TE^'SIO^' CHAXGES lA VOCAL CORDS. 601 and brongiit into its state in deej^ quiet breathing. Other muscles approximate the arytenoid cartilages after they have been separated. The most important is the transverse arytenoid {A, Fig. 158), which runs across from one ary- tenoid cartilage to the other. Another is the oUique arij- tenoid {Taep), which runs across the middle line from the base of one arytenoid to the ti}) of the other; thence cer- tain fibres continue in the aryteno-epiglottidean fold (10, Fig. 157) to the base of the epiglottis; this, with its fellou', thus embraces the whole entry to the larynx ; when they contract they bend inwards the tips of the arytenoid car- tilages, approximate the edges of the aryteno-epiglottidean fold, and draw down the epiglottis, and so close the pas- sage from the pharynx to the larynx; this is probably their chief function. When the epiglottis has been removed, food and drink rarely enter the larynx in swallowing, the eds^es of the folds of mucous membrane on its sides being: so brought together as to effectually close the aperture be- tween them. Increased tension of the vocal cords is produced mainly by the crico-tliyroid muscles, one of which lies on each side of the larynx, over the crico-thyroid membrane. Their action may be understood by help of the diagram, Fig. 159, in which t represents the thy- roid cartilage, c the cricoid, a an arytenoid, and vc a vocal cord. The muscle in question passes obliquely backwards and upwalrds from near the front end of c (to the right in the diagram) to t, near the pivot (which represents the joint be- tween the cricoid cartilage and the inferior horn of the thyroid). When the muscle contracts it ^^^" ^^^' [)ulls t down into the position indicated by the dotted lines and stretches the vocal cord, if the arytenoid cartilages be kept, by the muscles behind, from slipping forwards at the . discus pro- ligerus; c, ovimi. with, d, germinal vesicle and. e. geiminal spot. The general cavity of the follicle (in which 9 is printed) is flUed with liquid during life. liquid filling the cavity of the follicle. In the discus proli- gerus the ovum, c, lies, having in it a nucleus or germinal vesicle, d, and a nucleolus or germinal spot, e. The ovum is about 0.2 mm. (y^g inch) in diameter; its structure is better represented at A, Fig. 8*, where it is seen to consist of a thick outer coat or cell-wall, a, called the viteUine membrane or zona pellucida; within Avhich is the granular cell protoplasm, called here the vitellus or yelk; and in that again the germinal vesicle and spot. * Page 26. PUBERTY. 13 Puberty. The condition of tlie reproductive organs of each sex described above is that found in adults; altliough mapped out, and, to a certain extent, developed, before birth and during childhood, these parts grow but slowly and remain functionally incapable during the early years of life: then they comparatively rapidly increase in size and become physiologically active: the boy or girl becomes man or woman. This period of attaniing sexual maturity, known as pu- berty, takes place from the eleventh to the sixteenth year commonly, and is accompanied by changes in many parts of the Body. Hair grows more abundantly on the pubes and genital organs, and in the armpits; in the male also on various parts of the face. The lad's shoulders broaden; his larynx enlarges, and lengthening of the vocal cords causes a fall in the pitch of his voice; all the reproductive organs increase in size; fully formed seminal fluid is secreted, and erections of the penis occur. As these changes are completed spontaneous nocturnal seminal emissions take place from time to time during sleep, being usually associ- ated with voluptuous dreams. Many a young man is alarmed by these; he has been kept in ignorance of the Avhole mat- ter, is too bashful to speak of it, and getting some quack advertisement thrust into his hand in the street is alarmed to learn that his strength is being drained off, and that he is on the high-road to idiocy and impotence unless he place himself in the hands of the advertiser. Lads at this period of life should have been taught that such emissions, when not too frequent and not excited by any voluntary act of their own, are natural and healthy. They may, however, occur too often; if there is any reason to suspect this, the family physician should be consulted, as the healthy activity of the sexual organs varies so much in individuals as to make it impossible to lay down numerical rules on the subject. The best preventives in any case are, however, not drugs, but an avoidance of too warm and soft a bed, plenty of muscular exercise, and keeping out of the way of any thing- likely to excite the sexual instincts. In the woman the pelvis enlarges considerably at puberty, 14 THE HUM AX BOB 7. and, commonly, more snbcntaneous adipopo tissue develops over the Body generally, bnt especially on the breasts and hips; consequently the contours become more rounded. The external generative organs increase in size, and the clitoris and nymphfe become erectile. The uterus grows considerably, the ovaries enlarge, some Graafian follicles ripen, and the events treated of in the succeeding paragrai)hs occur. Ovulation. From puberty, during the whole child- Ijcaring period of life, certain comparatively very large Graafian follicles may nearly always be found either close to the surface of the ovary or projecting on its exterior. These, by accumulation of liquid within them, have become distended to a diameter of about 4 mm. {^ inch); finally, the thinned wall of the follicle gives way and the ovum is dis- charged, surrounded by some cells of the discus proligerus. The emptied follicle becomes filled up with a reddish-yellow mass of cells, and constitutes the corpus hiteu7n, which recedes again to the interior of the ovary and disajipears in three or four weeks, unless pregnancy occurs; then the corpus luteum increases for a time, and persists during the greater part of the gestation period. Menstruation. Ovulation occurs during the sexual life of a healthy woman at intervals of about four Aveeks, and is attended with important changes in other portions of the generative apparatus. The ovaries and Fallopian tubes becomes congested, and the fimbriae of the latter are erected and come into contact with the ovary so as to receive any ova discharged. Whether the fimbriae embrace the ovary and catch the ovum, or merely touch it at various points and the ova are swept along them by their cilia to tlie cavity of the oviduct, is not certain. Having entered the Fallopian tube the egg slowly passes on to the uterus, prol)ably moved by the cilia lining the oviduct. In the woml), also, important changes occur at this time; it be- comes swollen and gorged with blood, especially its lining mucous membrane and its glands. If the ovum be not fertilized it dies and is passed out. At the same time the gVipcrficial part of the uterine mucous membrane is softened, MENSTRUATION. 15 broken down, and discharged along with more or less blood, constituting the menses, or monthly sickness, which com- monly lasts from three to five days. During this time the vaginal secretion is also increased, and, mixed with the blood discharged, more or less alters its color, and usually destroys its coagulating power. Except daring pregnancy and while suckling, menstruation occurs at the above intervals, from puberty up to about the forty-fifth year; the periods then become irregular, and finally the discharges cease; this is an indication that ovulation has come to an end, and tlie sexual life of the woman is completed. This time, the diniucteric or ''tarn of life," is a critical one; various local disorders are apt to supervene, and even mental derangement. Hygiene of Menstruation. During menstruation there IS apt to be more or less general discomfort and nervous irri- tability; the woman is not quite herself, and those respon- sible for her happiness ought to watch and tend her with special solicitude, forbearance, and tenderness, and protect her from, anxiety and agitation. Any strong emotion, especially of a disagreeable character, is apt to check the flow, and this is always liable to be followed by serious con- sequences. A sudden chill often has the same effect; hence a menstruating woman ought always to be warmly clad, and take more than usual care to avoid draughts or getting wet. At these periods, also, the uterus is enlarged and heavy", and being (as may be seen in Fig. 163) but slightly supported, and that near its lower end, it is especially apt to be dis- placed or distorted; it may tilt forwards or sideways (fer- sions of the uterus), or be bent where the neck and body of the organ meet {flexion). Hence violent exercise at this time should be avoided, though there is no reason why a properly clad woman should not take her usual daily walk. Painful menstruation {dysmenorrlicea) may be due to very many causes, but it is only within recent years that physi- cians have come to recognize how often it depends on uterine displacements, and in such cases how readily it may usually be remedied by restoring the organ to its proper position, and supporting it there if necessary. A flexion of the organ 16 THE nUMAN BODY. closes the passage of the cervix and is especially apt to cause pain at the menstrual period. The accumulated blood distends the uterus, which makes violent contractions to expel it; finally, if the resistance is overcome, the blood has probably already clotted and its expulsion causes more suf- fering. All this might usually be soon remedied, but the sufferer, ignorant of what is wrong, often goes on montli after month until her health is undermined, hoping that the trouble will get better of itself, which it never will. To submit to the necessary examination and treatment from one of the other sex is, to a refined woman, hoAvever, apt to be a more severe trial than all the jDhysical pain; and there is no recent social movement more deserving of every en- couragement and support than that whose aim is to provide proj)erly trained female medical attendance for women in the diseases peculiar to their sex; such as may now, fortu- nately, be found in most of our large cities. Few except physicians, and perhaps few physicians, know what an amount of relievable pain women endure in silence rather than run the risk of being forced to consult a male doc- tor. If no skilled person of her own sex is at hand the sufferer, if she do anything, is only too apt to take some of the nostrums advertised in such number for '' female com- plaints," or to consult a half -educated lady quack of some novel "school" with a taking title. The result of doing this, or doing nothing, is often permanent valetudinarianism and a life of uselessness, to those who might be active and happy wives and mothers. The absence of the menstrual flow {amenorrhcBa) is nor- mal during pregnancy and Avhile suckling; and in some rare cases it never occurs throughout life, even in healthy women capable of child-ljcaring. Usually, however, the non- appearance of the menses at the proper periods is a serious symptom, and one which calls for prompt measures. In all such cases it cannot be too strongly impressed upon women that the most dangerous thing to do is to take drugs tending to induce the discharge, except under skilled advice; to excite the floAV, in many cases, as for example occlusion of the OS uteri, or in general debility (when its absence is CONCEPTION. 17 a conservative effort of the system), may be the most dis- astrous thing to do. Impregnation. As the ovum descends the Fallopian tube the changes preliminary to menstruation are taking place in the uterus. Its mucous membrane is thickened, all the generative organs of the woman are more or less congested, and her sexual emotions are commonly more easily aroused. Unless the act of coition occur all this passes off with the menstrual flux, and the organs return to a quiet state until the period of the next ovulation. If sexual congress takes place the vagina, uterus, and oviducts are thrown into reflex peristaltic contractions; and there is frequently an increased secretion by the vaginal mucous membrane. Some of the seminal fluid is received into the uterine cavity, and there, or, more probably, in the Fallopian tube, meets the ovum. The spermatozoa are carried along partly, perhaps, by the contractions of the muscular walls of the female cavities, but mainly by their own activity. From observations made on various lower animals it appears that their movements cease immediately on coming into contact with the ovum, and that one only takes part in fertilization. How this lattei occurs in the mammalian ovum is not cer- tain; observations in other groups make it probable that the male element directly fuses m whole or part with the protoplasmic mass of the ovum, but no opening has been de- tected in the zona pellucidaof the mammalian ovum, which IS so thick and firm that it is hard to imagine a spermato- zoon otherwise penetrating it; some, therefore, are inclined to suppose that material is merely passed by dialysis from the spermatozoon into the egg-cell. The fertilized ovum continues its descent to the uterine cavity, but, instead of lying dormant like the tmfertilized, segments (p. 26), and forms a morula. This, entering the womb, becomes imbedded in the soft, thickened, vascular mucous membrane there, from which it imbibes nourish- ment, and which, instead of being cast off in a menstrual discharge, is now retained and grows during the whole of pregnane^ , having important dtities to discharge in connec- tion with the nutrition of the embryo. 18 THE HUM Ay BODY. Sexual congress is most apt to be followed by impregnation if it occur Just before the menstrual flow is due, while a ripe and receptive ovum is in the Fallopian tube. This time is about four days, that being the period the ovum ajipears to take in its passage from the ovary to the womlj. The spermatozoa, however, retain their vitality in the female cavities for some five or six days, so that coition during the ten or twelve days preceding the date when the next menstruation may be expected, is a23t to be followed by pregnancy. There seems reason to believe, however, that ova are occasionally discharged at other than the regular monthly periods of ovulation, especially under the influence of the erection and congestion of the parts which takes place during the sexual act. There is, therefore, no time during a woman's life from jouberty to the climacteric, except during pregnancy or lactation, when sexual congress may not result in impregnation. The exact parts taken by the male and female elements respectively in the formation of the embryo are not known with certainty; seeing that the ovum is much larger than the spermatozoon it is not uncommon to speak of the latter as a mere stimulant or excitant, arousing the egg-cell to developmental activity; but the definite characteristics of ten inherited by children from the father show that the sperma- tozoon IS much more than that; materials derived from it are no doubt an essential constituent of the compound mass Avhich develops into the new human being. Pregnancy. When the mulberry mass reaches the uterine cavity the mucous membrane lining the latter grows rapidly and forms a new, thick, very vascular lining to the womb, known as the decidua. At one point on this the morula becomes attached, the decidua growing up around it. As pregnancy advances and the embryo gi'ows, it bulges out into the uterine cavity and pushes before it that part of the decidua which has grown over it (the decidna rejiexa); at about the end of the third month this meets the decidua lining the opposite sides of the uterine cavity and the two grow together. That part of the decidua (decidua serotina) against which the morula is first attached subsequently un- GESTATION. 19 dergoes a great developineiit iu connection with the forma- tion of the placenta (see below). Meanwhile the whole uterus enlarges; its muscular coat especially thickens. At first the organ still lies within the pelvis, where there is but Httle room for it; it accordingly presses on the bladder and rectum (see Fig. 163) and the nerves in the neighborhood, frequently causing considerable discomfort or pain; and, reflexly, often exciting nausea or vomiting (the morning siclcness of pregnancy). Later on, the pregnant womb escapes higher into the abdominal cavity, and although then larger, the soft abdominal walls more readily make room for it, and less discomfort is usually felt, though there may be shortness of breath and palpitation of the heart from inter- ference with the diaphragmatic movements. All tight garments should at this time be especially avoided; the woman's breathing is already sufficiently impeded, and the pressure may also injure the developing child. Meanwhile, changes occur elsewhere in the Body. The breasts enlarge and hard masses of developing glandular tissue can be felt in them; and there may be mental symptoms: depression, anxiety, and aii emotional nervoits state. During the whole period of gestation the woman is not merely sttpplying from her blood nutriment for the foetus, but also, through her lungs and kidneys, getting rid of its wastes; the result is a strain on her whole system which, it is true, she is constructed to bear and will carry well if iu good health, but which is severely felt if she be feeble or suffering from disease. Many a wife who might have led a long and happy life is made an invalid or brought to jjre- mature death, through being kept in a chronic state of pregnancy. There is a general agreement that sexual con- tinence is j)ossible and a duty in unmarried men, but the husband rarely considers that he should put any bounds on himself beyond those indicated by his own passions; consid- eration for his wife's health rarely enters his head in this connection. The healthy, married woman who endeavors to evade motherhood because she thinks she will thus pre- serve her personal appearance, or because she dislikes the trouble of a family, deserves but little sympathy; she is 30 THE HUMAN BODY. trying to escape a duty voluutarily undertaken, and owed to her husband, her country, and her race; but she whoso strength is undermined and whose life is made one long discomfort for the sexual gratification of her husband calls for all aid, and it is wrong to keep silent on the subject. The professor of gynajcology in a leading medical school, gives it as his deliberate opinion that the majority of American women must at some periods of their lives choose between freedom from pregnancy or early death. He further says that he does not believe that healthy men are so organized that this matter can be regulated by them; but that the question depends on the tact and prudence of the wife. Men, however, as a rule, are not utterly selfish; they commonly err through ignorance or thoughtlessness, not knowing or realizing what a strain frequent pregnancy is on the strength of a delicate woman. A social custom so deep-rooted and ancient as the usual American and English one of married couples constantly occupying the same bed is not easily changed, yet it probably leads to much harm, and especially in this country. Whatever the reason be, there is no doubt that the physical stamina of the average English woman is considerably greater than that of her American cousin, and she bears and brings up large families with greater safety. For a husband, who has reason to believe that child-bearing will injure his wife's health, to always share her couch is a deliberate walking into temptation. Apart from pregnancy, moreover, a woman's health is often injured by frequent sexual intercourse. A johysician who has unusual opportunities of knowing states that he has reason to believe that not only is the act of sexual con- gress at best, from a physical jDoint of view, a mere nuisance to the majority of women belonging to the more luxurious classes of society after they attain the age of twenty-two or twenty-three, but that a very considerable proportion suf- fer acute pain from it such as, if frequent, breaks down the general health. A loving woman, finding her highest happi- ness in suffering for those dear to her, is very unlikely to let her husband know this, so long as she can bear it; but if the possibility is known it will not, perhaps, need much NUTRITION OF THE EMBRYO. 31 acuteness in him to discover sucli suffering when it exists, nor very much real affection to control himself accordingly. In the class of cases referred to, rest of the over-irritable and congested female organs is above all essential. The cause is frequently removable by simple, but skilled, treat- ment; the desirability of rendering this available to a woman in members of her own sex has already been insisted upon. Even when no pain is caused harm may be done: the presi- dent of the Gynaecological Society, in an address delivered before that body, lately stated that if either party to a coi- tion fails of the orgasm damage is apt to ensue, but espe- cially to the woman if she fail; the organs are congested from the stimulus of the sexual act and the normal final orgasm is required for their healthy relief and return to the resting condition. If this be so, it is clear that coition should be restricted to times when the woman's general state encour- ages the orgasm, and unless she generally experiences it sexual congress should be avoided until her health is restored. The Foetal Appendages. In the earliest stages of life, those occurring in the days imme- diately after fertilization of the ovum, there is little or no growth. The ovum segments into a number of cells, but the morula thus formed is little larger than the original egg-cell itself. At first it is a solid mass (F, Fig. 8, p. 26), but its cells soon recede from the centre and become arranged (Fig, 165) around a central cavity containing mainly some absorbed liquid; at one point, c (the embryonal disk), the layer thickens, and from thence the thickening spreads, by cell growth and multiplication, over the whole sac, which is known as the embryonal vesicle; the membrane thus formed is the blastoderm, and Fig. 165.— The embryonal vesicle. a, thinned and distended zona pel- lucida: this soon after disappears altogether; b, tlie blastoderm; c, the embryonal disk. 22 TEE HUMAN BODY. consists of three cell layers; au outer or epihlast, a middle or mesohlast, and an inner or hypoblast. From this simple sac, presenting no resemblance not merely to a human being but to a vertebrate animal, the foetus is built up by cell division and modification. The general history of intra- uterine development is much too long and too complex to enter upon here, but the formation of certain structures lost at or before birth, and associated with the jn-otection and nourishment of the embryo, may be attempted: they are the yelh sac, the amnion, and the allantois. The developing blastoderm especially thickens in the neighborhood of the embryonal disk and there the outlines of the Body are first laid down. Along the thickening a groove appears, which A B Fig. 166.— a, an early blastoderm with the first traces of the primitive groove; B, the same a little later ; /, primitive groove ; d, thickened region of the blas- toderm which directlj- builds up the embryo. marks out the future longitudinal axis and dorsal side of the bod}^, (A, Fig. IGG). This groove elongates, its edges rise (B), and finally arch over, meet, and fuse together above it. The tube thus closed in is the rudiment of the cerebro-spinal axis; from its lining epiblastic cells the brain and spinal cord are developed, and its cavity remains throughout life- as the central canal of the spinal cord and as the cerebral ven- tricles, excluding the fifth (p. 165). Some way outside this dorsal tube the mesoblast splits into an outer leaf, adherent to the epiblast, and an inner adherent to the hypoblast; the conjoined meso-epiblastic layer is the somatopleure; the meso-hypoblastic the splanchnopleiire. The proximal j^arts of the somatopleure {i.e., the regions nearest the central THE FCETAL APPENDAGES. 23 axis) develoj) into the walls of chest and abdomen; farther out it turns up and arches over the back of the embryo, and its edges, there meeting, grow together and form a bag, the amnion, enveloping the foetus. Into this a considerable quantity of liquid is secreted, m which the fcetus floats. At birth the contractions of the uterus, pressing on the amnion, drive part of it down like a wedge into the neck of the uterus, and through its liquid contents an equable pressure is exerted there, until the os uteri is tolerably dilated; the sac then normally ruptures and the "waters" escape. Some- times, however, an infant is born still enveloped in the amnion, which is then popularly known as a caul. While the amnion is develoi)ing, a semi-cartilaginous rod forms along the axis of the Body beneath the floor of the dorsal tube; this is the notochord; when it appears the young being is marked out distinctly as a vertebrate animal, having a dor- sal neural tube above an axial skeleton, and a ventral hcemal tube (p. 4), formed by the proximal regions of the somato- pleure, beneath it. The ventral tube, however, is still widely open, the points where the amniotic folds turn back being far from meeting in the future middle line of the chest and abdomen. Tlie proximal portions of the splanchnopleure incurve to inclose the alimentary tube, which is at first straight and simple. Beyond the point where it bends in for this purpose the splanchnopleure again diverges, and incloses a small globular bag, the yelk sac, which is, thus, attached to the ventral side of the alimentary canal; it at first projects through the opening where the amniotic folds turn back, but has little importance in the mammalian embryo and is soon absorbed. The aUa)Lfois is primarily an outgrowth from the ali- mentary canal, containing blood-vessels. It passes out from the Body on the ventral side where the somatopleures have not yet met, and reaching the inside of the uterus, its dist:il end expands there to make the main part of the placenta (see below). Its narrow proximal portion forms the umbilical cord, around which the somatojDleures, incurving to in- close the belly, meet at the navel some time before birth. 24 TUE HUMAN BODY. The Intra-Uterine Nutrition of the Embryo. At tirst the embryo is uuuri.slied by tibsor])tion of niateriuls from Llie soft vascular lining of the womb; as it increases in size this is not sufficient, and a new organ, the placenta, is formed for the purpose. The allantois plants itself firmly against the decidua serotina (p. 18), and villi developed on it burrow from its surface into the uterine mucous membrane. In the deeper layer of this latter are large sinuses through which the maternal blood flows. Into the allantoic villi foetal blood-vessels run and form capil- lary networks; the blood flowing through these receives, by dialysis, oxygen and food materials from the maternal blood, and gives up to it carbon dioxide, urea, and other wastes. There is thus no direct intermixture of the two bloods; the embryo is from the first an essentially sei)arate and independent organism. The allantois and decidua serotina becoming closely united together form the placenta, which in the human species is, when fully developed, a round thick mass about the size of a large saucer. Parturition. At the end of fj-om 275 to 280 days from fertilization of the ovum [conception) pregnancy terminates, and the child is expelled by powerful contractions of the uterus, assisted by those of the muscles in the abdominal walls. When the child is born, it has attached to its navel the umbilical cord, through which arteries run to, and veins from, the placenta. Shortly afterwards the latter is detached and expelled (both its allantoic and decidual parts), and where it is torn loose from the uterine wall large blood sinuses are left open and exposed. Hence a certain amount of bleeding occurs, but in normal labor this is speedily checked by firm contraction of the uterus. Should this fail to take place profuse hiBmorrhage occurs {flooding) and the mother may bleed to death in a few min- utes unless prompt measures are adopted. For a few days after delivery there is some discharge (the lochia) from the uterine cavity: the whole decidua being broken down and carried off, to be subsetpiently replaced by new mucous membrane. The muscular fibres developed in the uterine wall in such large quantities during pregnancy. PARTURITION. 25 undergo rapid fatty degeneration and are absorbed, and in a few weeks the organ returns almost to its original size. The parturient woman is especially apt to take infec- tious diseases; and these, should they attack her, are fatal in a very large percentage of cases. Very special care should therefore be taken to keeja all contagion from her. There is a current impression that a pregnancy, once commenced, can be brought to a premature end, especially m its early stages, without any serious risk to the woman. It ought to be widely made known that such a belief is erroneous. Premature delivery, early or late in pregnancy, is always more dangerous than natural labor at the jiroi^er term; the physician has sometimes to induce it, as when a malformed pelvis makes normal parturition impossible, or the general derangement of health accompanying the preg- nancy IS such as to threaten the mother's life; but the occa- sional necessity of deciding whether it is liis duty to pro- cure an abortion is one of the most serious responsibilities he meets with in the course of his professional work. Dr. Storer, an eminent gynaecologist, states emphatically, from extended observation, that "despite aiij)arent and isolated instances to the contrary — 1. A larger proportion of women die during or in con- sequence of an abortion, than during or in consequence of child-bed at the full term of i)regnancy. 3. A very much larger number of women become con- firmed invalids, perhaps for life; and — 3. The tendency to serious and often fatal organic disease, as cancer, is rendered very much greater at the so-called "turn of life,"' by previous artificially induced premature delivery. During pregnancy there is a close connection between the placenta and uterus; nature makes preparation for the safe dissolutioii of this at the end of the normal period, but " its premature rupture is usually attended by pro- fuse haemorrhage, often fatal, often persistent to a greater or less degree for many months after the act is completed, and always attended with more or less shock to the maternal system, even though the full effect of this is not noted for 26 THE HUMAN BODY. years." Tlie sume authority states again: "Any deviation from this process at the full term" {i.e., the process, asso- ciated with lactation, by which the uterus is restored to its small non-gravid dimensions) "lays the foundation of. and causes, a wide range of uterine accidents and disease, dis- [dacements of various kinds; falling of the womb downwards or forwards or backwards, with the long list of neuralgic pains in the l)ack, groin, thighs and elsewhere that they occasion; constant and inordinate leucorrhoea; sympathetic attacks of ovarian irritation, running even into dropsy," etc. etc. There is, thus, abundant reason for bearing most things rather than the risks of an avoidable abortion; among these is one not mentioned above, but more terrible than all, insanity. Lactation. The mammary glands for several years aftei birtli remain small, and alike in both sexes. Towards puberty they begin to enlarge in the female, and when fully developed form in that sex two rounded eminences, tho breasts, placed on the thorax. A little below the centre of each projects a small eminence, the nipple, and the skin around this forms a colored circle, the areola. In virgma the areola? are pink; they darken m tint and enlarge during the first pregnancy and never quite regain their original hue. The mammary glands are constructed on the com pound racemose type (p. 2G2). Each consists of from fifteen to twenty distinct lobes, made up of smaller divisions^ from each main lobe a separate galadophorous duct, made by the union of smaller branches from the lobules, runs towards the nipjole, all converging beneath the areola. There each dilates and forms a small elongated reservoir m which the milk may temporarily collect. Beyond this the ducts narrow again, and each continues to a separate opening on the nipple. Imbedding and enveloping the lobes of the gland is a quantity of firm adipose tissue which gives the whole breast its rounded form. During maidenhood the glandular tissue remains imper- fectly developed and dormant. Early in pregnancy it begins to increase in bulk, and the gland lobes can be felt as hard masses through the superjacent skin and fat. Even at par- turition, however, their functional activity is not fully LACTATION. 27 established. The oil-globixles of the milk are formed by a sort of fatty degeneration of the gland-cells, which finally fall to pieces; the cream is thus set free m the watery and albuminous secretion formed simultaneously, while newly developed gland-cells take the place of those destroyed. In the milk first secreted after accouchment (the colostrum) the cell destruction is incomplete, and many cells still float in the liquid, which has a yellowish color; this first milk acts as a purgative on the infant, and probably thus serves a useful purpose, as a certain amount of substances (bdiary and other), excreted by its organs during development, are found in the intestines at birth. Human milk is undoubtedly the best food for an infant in the early months of life; and to suckle her child is useful to the mother if she be a healthy woman. There is reason to believe that the processes of involution by which the large mass of muscular and other tissues developed in the uterine walls during pregnancy are broken down and absorbed, take place more safely to health if the natural milk secretion is encouraged. Many women refuse to suckle their children from a belief that so doing will injure their personal appear- ance, but skilled medical opinion is to the co itrary effect; the natural course of events is the best for this purpose, unless lactation be too prolonged. Of course in many cases there are justifiable grounds for a mother's not undertaking this part of her duties; a physician is the pioper person to decide. In a healthy woman, not suckling her child, ovulation and menstruation recommence about six weeks after childbirth; a nursing mother usually does not menstruate for ten or twelve months; the infant should then be weaned. When an infant cannot be suckled by its mother or a wet- nurse an important matter is to decide what is the best food to substitute. Grood cow's milk contains rather more fats than that of a woman, and much more casein; the following table gives averages in 1000 parts of milk: Woman. Cow. Casein 28.0 54.0 Butter 8:5 5 43.0 Milk sugar 44.5 42.5 Inorganic matters 4.75 7.75 28 THE HUMA^' BODY. The inorganic matters of human milk yield, on analysis, m 100 parts — calcium carbonate 0.9; calcium phosphate 70.6; sodium chloride D.8; sodium sulphate 7.4; other salts 5.3. The lime salts arc of especial importance to the child, which lias still to build up nearly all its bony skeleton. When undiluted covv's milk is given to infants they rarely bear it well; the too abundant casein is vomited in loose coagula. The milk should therefore be diluted with half or, for very young children, even two thirds its bulk of water. This, however, brings down the percentage of sugar and butter below the proper amount. The sugar is commonly replaced by adding cane sugar; but sugar of milk is readily obtainable and is better for the purpose. If used at all it should, however, be employed from the first; it sweetens much less than cane sugar, and infants used to the latter refuse milk in which milk sugar is substituted. Cream from cow's milk may be added to raise the percentage of fats to the normal, but must be perfectly fresh and only added to the milk immediately before it is given to the child. While milk is standing for the cream to rise it is very apt to turn a little sour; the amount of this sour milk carried off with the cream is itself no harm when mixed with a large bulk of fresh milk; it carries with it, however, some of the fungus whose development causes the souring, and this will rapidly de- velop and sour all the milk it is added to if the mixture be let stand. As the infant grows older less diluted cow's milk may gradually be given; after the seventh or eighth month no addition of water is necessary. In the first weeks after birth it is no use to give an infant starchy foods, as arrowroot. The greater part of the starch passes through the bowels unchanged; apparently because the pancreas has not yet fully developed, and has not commenced to make its starch-converting ferment (p. 311). Later on, starchy substances may be added to tlie diet with advantage, but it should be borne in mind that they cannot form the chief part of the child's food; it needs proteids for the formation of its tissues, and amyloid foods contain none of these. Many infants are, ignorautly, half starved by being fed almost entirely on such things as corn-flour or arrowroot. STAGES OF LIFE. 29 The Stages of Life. Starting from the ovum each human being, apart from accident or disease, runs through a life- cycle which terminates on the average after a course of from 75 to 80 years. The earliest years are marked not only by raj^id growth but by differentiating growth or de- velopment; then comes a more stationary period, and finally one of degeneration. The life of various tissues and of many organs is not, however, coextensive with that of the individual. During life all the formed elements of the Body are constantly being broken down and removed; either molecularly {i.e., bit by bit Avhile the general size and form of the cell or fibre remains unaltered), or in mass, as when hairs and the cuticle are shed. The life of many organs, also, does not extend from birth to death, at least in a functionally active state. At the former jieriod numerous bones are represented mainly by cartilage. The pancreas has not attained its full development; and some of the sense-organs seem to be in the sarne case; at least new-born infants appear to hear very imi^erfectly. Tlie reproductive organs only attain full development at pu- berty, and degenerate and lose all or much of their func- tional importance as years accumulate. Certain organs have even a still shorter range of physiological life; the thymus, for example (p. 833), attains its fullest develop- ment at the end of the second year and then gradually dwindles away, so that m the adult scarcely a trace of it is to be found. The milk-teeth are shed in childhood, and their so-called permanent successors rarely last to ripe old age. During early life the Body increases in mass, at first very rapidly, and then more slowly, till the full size is attained, except that girls make a sudden advance m this respect at puberty. Henceforth the woman's weight (excluding cases of accumulation of non-working adipose tissue) remains about the same until the climacteric. After that there is often an increase of weight for several years; a man's weight usually slowdy increases until forty. As old age comes on a general decline sets in, the rib cartilages become calcified, and lime salts are laid down 30 THE HUMAN BODY. in the ai'terial walls, wliieli thus lose their elasticity; the refracting media of the eye become more or less opaqne; the pln'siological irritability of the sense-organs in general diminishes; and fatty degeneration, diminishing their work- ing power, occurs in many tissues. In the brain we find signs of less plasticity; the youth in whom few lines of least resistance have been firmly established is ready to accept novelties and form new associations of ideas; but the longer he lives, the more difficult does this become to him. A man past middle life may do good, or even his best work, but almost invariably in some line of thought which he has already accepted: it is extremely rare for an old man to take up a new study or change his views, philosophical, scientific, or other. Hence, as we live, we all tend to lag behind the rising generation. Death. After the prime of life the tissues dwindle (or at least the most important ones) as they increased in child- hood; it is conceivable that, without death, this })rocess might occur until the Body was reduced to its original microscopic dimensions. Before any great diminution takes place, however, a breakdown occurs somewhere, the enfeebled community of organs and tissues forming the man is unable to meet the contingencies of life, and death supervenes. "It is as natural to die as to be born," Bacon wrote long since; but though we all know it few realize the fact until the sum- mons comes. To the popular imagination the prospect of dying is often associated with thoughts of extreme suffering; ]iersonifying life people picture a forcible and agonizing rending of it, as an entity, from the bodily frame with which it is associated. As a matter of fact, death is prob- ably rarely associated with any immediate suffering. The sensibilities are gradually dulled as the end approaches; the nervous tissues, with the rest, lose their functional capacity, and, before the heart ceases to beat, the individual has commonly lost consciousness. The actual moment of death is hard to define: that of the Body generally, of the mass as a whole, may be taken to be the moment when the heart makes its last beat: arterial DEATH. 31 pressure then falls irretrievably, the capillary circulation ceases, and the tissues, no longer nourished from the blood, gradually die, not all at one instant, but one after another, according as their individual resjiiratory or other needs are great or little. While death is the natural end of life, it is not its aim — we should not live to die, but live prepared to die. Life has its duties and its legitimate pleasures, and we better play our part rather by attending to the fulfilment of the one and the enjoyment of the other, than by concentrating a morbid and paralyzing attention on the inevitable, with the too frequent result of producing indifference to the work which lies at hand for each. Our organs and faculties arc not talents which we may justifiably leave unem])loyed; each is bound to do his best with them, and so to live that he may most utilize them. An active, vigorous, dutiful, un- selfish life is a good preparation for death; when that time, at which we must pass from the realm controlled by physi- ological laws, approaches, when the hands tremble and the eyes grow dim, when "the grasshopper shall be a burden and desire shall fail," then, surely, the consciousness of having "quitted us like men" in the cm])loyment of our faculties while they were ours to use, will be no mean conso- lation. INDEX TO APPENDIX. Abortion, 25. Allantois, 23. Amenorrhoea, 16. Amnion, 2.3. Blastoderm. 21. Breasts, 26. Budding, 1. Caul, 23. Cervix uteri, 10. Childbirth, 24. Climacteric, 15. Clitoris, 11. Coni vasculosi, 6. Corpora cavernosa, 7. Corpus luteum, 14. Corpus spongiosum, 7. Death, 30. Decidua, 18. Development of embryo, 21. Discontinuous growth, 1. Discus proligerus, 12. Displacement of uterus, 15. Dysmenorrhea, 15. Egg-cell, 4. Emliryonal disk, 21. Embryonal vesicle, 21. Epi])last, 22. Epididymis, 6. Erectile tissues, 7. Fallopian tube, 4, 9. Feeding of infants, 27. Fertilization, 4, 17. Foetal appendages, 21. Galactophorous ducts. 2fl Gemmation, 1. Germinal epithelium, 11. Germinal spot, 12. Germinal vesicle, 12. Gland, maiiunary, 26. Graafian follicles, 11. Hermaphroditism, 4. Hernia, inguinal, 5. Hi.stology of ovary, 11; of testis, 8. Hydrocele, 5. Hygiene of menstruation, 15- of pregnancy, 19. Hymen, 11. Hypoblast, 22. Impregnation, 17. Inguinal hernia, 5. Intra uterine development, 21. Intrauterine nutrition of embryo, 24. Lactation, 26. Lochia, 24. Mammary gland, 26. Membrana grauulo.sa, 12. Menstruation, 14. Mesoblast, 22. Milk, 27. Nervi erigentes, 7. Notochord, 23. Organs of reproduction, 4. Ovary, 4, 8. Oviduct, 4, 9. Ovulation, 14. Ovum, 4, 12. INDEX rO APPENDIX. Parthenogenesis, 4. Parturition, 24. Penis, 7. Placenta, 24. Pregnancy, 18. Prepuce, 8. Prostate gl;incl, (i. Puberty, 1:5. Reproduction in general, 1. Reproductive organs, accessory, 4; male, 4; female, 8. Ruplure, 5. Seminal fluid, 8. Seminiferous tubules, 5. Sexual reproduction, 3. Somatopleure, 22. Spermatozoon, 4, 8. Sperm-cell, 4. Splanchnopleure, 22. Stages of life, 29. Testes, 4, 8. Tunica vaginalis, 4. Umbilical cord, 23. Urethra, 7. Uterus, 4, 9. Utricle, 7. Vagina, 10. Vas deferens, 6. Vasa recta, 6. Vasa efferentia, G. Vesiculoe seminales, 6. Vitelline membrane, 12. Vitellus, 12. Vulva, 10. Yelk, 12. Yelk-sac, 22. Zona pellucida, 12 QV^Co T\^(^ XAo.«^\ \ki. ^^f^ a.^ -^oh \