CORNELL UNIVERSITY LIBRARY FROM The Estate of P.R.flacmillanA TEXT-BOOK GENERAL PHYSIOLOGY AND ANATOMY BY WALTER HOLLIS EDDY Chairman of the Department of Biology in the High School of Commerce, New York NEW YORK CINCINNATI CHICAGO AMERICAN BOOK COMPANY4 PREFACE laboratory course in zoology these chapters should be omitted, except for general reading. This text is not intended to be used as a substitute for laboratory study. In my Experimental Physiology and Anatomy are outlined laboratory exercises which should be first performed by the pupil himself, if possible, before he is assigned a lesson in this text. While written as a supplement to the laboratory guide mentioned, the text is suited for use with any other laboratory manual and, where lack of laboratory facilities renders such pre- liminary work impossible, its description of the laboratory exercises is sufficiently full to permit its use where a text- book alone must serve. The text meets both the optional and required work of the New York State Syllabus and the requirements of the Harvard entrance examinations. Its topical arrangement and the printing of less impor- tant details in finer type make it adaptable to longer or shorter courses of study; and in these structural details the author has endeavored to make the book meet the needs of all classes of schools. I wish to acknowledge the helpful suggestions given me by my colleagues of the High School of Commerce, and by Professor Geis of Columbia University and Dr. Darling of Harvard University. I am especially indebted to my colleague Mr. C. W. Hahn for his very careful read- ing of the proofs and his many helpful suggestions, and to my wife for assistance in the revision of grammatical arrangement and in the mechanical details of the work. W. H. EDDY. The High School nv Commerce, New York.Page 7 12 28 39 51 73 90 107 133 146 159 180 191 212 221 244 252 265 278 294 CON T EN T S What the Study of Physiology Means. . Chemical Composition of Living Matter . . . . The Structural Units of Living Matter . . . Structural Units of Human Tissues — Histology Foods and Nutrition ................. Food Accessories; Alcohol and Tobacco Digestion . . ................... Digestion (continued) Absorption . . Digestion in the Lower Animals . Blood and Lymph . . . Circulation . . Circulation (continued) . . . Circulation in the Lower Animals The Skeleton . . ... The Skeleton (continued) . . Skeletons of the Lower Animals Muscles ....................... Muscles (continued) ....... Movement in Lower Animals . . 56 CONTENTS Page XXI. Respiration 312 XXII. Respiration in Lower Animals........................340 XXIII. Excretion . 352 XXIV. Skin Structure and Excretion in the Lower Animals 374 XXV. The Nervous System . . .... . 383 XXVI. The Nervous System (continued)........................411 XXVII. The Nervous System in the Lower Animals . 429 XXVIII. The Special Senses................. ... 436 XXIX. The Special Senses (continued).....................452 XXX. The Special Senses (concluded) ............ 470 XXXI. Sensation in the Lower Animals ................ 4S0 XXXII. The Voice.......................................... 486 XXXIII. Bacteria and Sanitation ........................... 493 Index 505PHYSIOLOGY AND ANATOMY I. WHAT THE STUDY OE PHYSIOLOGY MEANS. From our earliest days we are familiar with living forms. We know that each living form must perform cer- tain actions every day in order to live; but few of us have a clear idea of why these actions maintain life, or of the part each separate action plays in keeping us alive. For example, if asked why we eat, we answer, “ To keep us alive,” but that answer does not explain how food gives life. Again, we see gardeners watering plants, repotting them in new earth, taking care to place them in sunlight, and we recognize that all these actions are necessary to the life of the plants. We know that dogs and cats, horses and cows, must be fed and cared for to keep them alive and healthy. In fine, we know that to maintain life all living bodies, whether plant or animal, must repeat certain actions continually, or else die ; but we do not know how these actions produce life. It is this question of “ how ” and “ why ” that the present study of Physiology is to answer. Method of study.-—How shall we begin the study? An example will indicate the method. If we see a man moving the handle of a certain machine up and down and see a stream of water issuing from another part of the same machine, we know at once that the movement of the handle in some way causes the flow of water. In 78 WHAT THE STUDY OF PHYSIOLOGY MEANS order to answer the question “ how ” we should first pull our pump or a similar machine to pieces and find what parts it has and how they are placed in relation to one another. We should call this the study of the structure of the pump. Next, we should set the machine in motion and observe the action of the parts and how the move- ment of one affects the other. We should call this the study of function, or use of parts. Combining our knowl- edge of structure and function we might go still further, and, by experimenting, find that certain actions, such as lubricating the parts with oil, improve the action of the machine as a whole. We call this the study of the con- ditions best suited to the action of the machine. Applied to the study of living objects such a method of investigation resolves itself into the following divisions of research. First. The study of the structure of the living bodies. Such a study is called anatomy {ana, up; temnein, to cut) and is dependent upon dissection. Second. The study of the use or functions of the various structures. Such a study is called 'physiology (physiologia, an investigation into the nature of things) and is dependent upon experimenting with living structures. Third. The study of methods that will cause the parts to act to the best advantage. This study is called hygiene {hygeia, health) and like physiology is dependent upon experimentation. Formulating these three directions we have the follow- ing definitions: Anatomy. —The study of the structure of a plant or animal to determine the parts present and the relative position of these parts.METHOD OF STUDY 9 Physiology. —The study of the function or use of the parts of a plant or animal body and their method of action. Hygiene. —The study of the means best adapted to maintain the parts and actions of the plant or animal body in a working or healthful state. Anatomy.— Like a pump, a living body is made up of parts arranged in definite positions. It is manifestly impossible to locate all these parts from the study of the living body. To get at the internal structure it is neces- sary to cut or dissect the dead body. In the case of the human body such dissection is of course impossible for the high school pupil. It is here that a fact of great im- portance comes to our rescue, namely, that the arrange- ment of parts in our bodies is so similar to that in certain other animal forms that by studying these forms we may learn of our own structure. In other words, the dissec- tion of the rat, cat, or rabbit, enables us to know our own structures. Even with this difficulty overcome, however, many parts are so small that the eye cannot make them out. To remedy this there has been devised an instrument called the microscope; and so important is this side of anatomical study that it has received a special name. Histology (histos, a tissue; logos, speech) is the microscopic study of plant and animal parts. Finally, chemists tell us that the parts themselves are composed of simpler substances called chemical elements. This branch of our study is called chemical anatomy. When we have studied a body from these three points of view, its gross structure, its microscopic structure, and its chemical composition, we shall have exhausted all that anatomy has to tell us,10 WHAT THE STUDY OF PHYSIOLOGY MEANS Physiology. —Anatomy tells us only of the structure of the lifeless body. It is physiology that tells us of the action of the parts in the living body. Here the problem of investigation becomes difficult, but again we call to aid the fact that vital processes in plants and animals are similar to those in our own bodies, and the experiments we employ with the former tell us the workings of our own body. These experiments are supplemented by the reports of surgeons of their operations on human bodies, while another branch of science, called physiological chem- istry, has taught us how to reproduce artificially many of the actions which take place normally in our bodies. Hygiene. —The best engineer is the man who knows not only how his engine is made and acts, but also how to keep it in order and to run it to the best advantage. It is this knowledge — how to run our bodies to the best advantage, that is, to keep them healthy — that the study of hygiene gives us. It can be acquired only by taking to ourselves the experience of those most familiar with the running of our bodies. There are, however, certain of these experiences which have been formulated under the laws of sanitation which will give us a better foundation for our apprenticeship. Such laws are those that have been formulated by doctors, physicists, and bacteriologists, and we shall study them in their proper place. Relation to other sciences. —The preceding pages indi- cate in a general way what the scope of our study is to be. AVhat relation does this study bear to other sciences? Biology (bios, life; logos, speech) is the name given to the study of living as distinct from lifeless matter. This subject naturally groups itself into the study of plants, botany, and the study of animals, zoology. From what hasBIOLOGY 11 been said already it will be seen that the terms anatomy, physiology, and hygiene refer to methods of study applied to either plants or animals, for example, plant physiology, animal anatomy, etc. —or they may lie restricted to indi- vidual members of each group, human physiology, horse anatomy, yeast physiology, etc. In short, physiology, anatomy, and hygiene are merely one phase of the study of living matter (biology) and while our present problem is the human body, we must bear in mind that many of the laws that hold true of our bodies are merely special cases of laws which apply to all living organisms. The great problem which all biologists, zoologists, and physiologists are seeking to answer is, “ AVhat is life? ” and in whatever field they may be working, their results arc valuable and throw light upon this main, great problem.II. CHEMICAL COMPOSITION OF LIVING MATTER. Chemical elements. —The dissection of a plant or animal shows it to be made up of many different kinds of materials, some soft, some hard, some elastic. Dis- section, however, can tell us nothing of the composition of these separate masses of material, and to find out what bone, muscle, and fat are made of, we must go to the chemist. As the result of methods which chemists have learned to apply to the problem, they tell us that muscle, fat, bone, wood, and in fact all the other substances of the universe are built up of simple substances which are called elements. By examining all the different kinds of matter, both living and lifeless, to be found in the universe, it has been found possible to reduce the number of these simple substances or elements to some seventy-six, and just as all the words in the language may be built up of the twenty-six letters of the alphabet, so all matter is built up by varying combinations of these seventy-six elements. Some of these elements are very rare, while others are exceedingly abundant. Some are never found in the pure state but always combined with other elements, while others are found both in a free state and in combin- ation. Of the seventy-six known elements, twelve 1 are always found in living matter and six of the remaining sixty-four are sometimes present. 1 The twelve that are always present in living matter are phosphorus, sulphur, carbon, oxygen, hydrogen, nitrogen, chlorine, potassium, sodium, calcium, magnesium, and iron. 12PHOSPHORUS 13 Characteristics of chemical elements. —The distinguish- ing feature of a chemical element is that it cannot be resolved by any known means into a simpler substance. Thus while it is possible to resolve a substance like com- mon salt into two substances known respectively as sodium and chlorine, no one has ever been able to separate sodium or chlorine into anything simpler, and these two sub- stances are, therefore, called elements. Iron, sulphur, gold, silver, are familiar examples of elements. Since the chemical elements are the substances out of which all matter is formed, it will be well for us to examine a few of them to learn their properties and the manner in which they combine to form compounds. We shall select for examination those which are particularly concerned in the composition of our body materials. Phosphorus. — (See Ex. III.) This element forms a large part of our bones; these are, in fact, composed mainly of a compound of this element with calcium and oxygen. In its pure state phosphorus has a yellow color and a waxy consistency. When exposed to the air it gives off white fumes which have a peculiar odor. These fumes are a compound of phosphorus with the oxygen of the air, and to keep phosphorus from combining with oxygen it is necessary to coArer it with water. These fumes teach us of a peculiar property of phos- phorus, namely, its power to combine with another element. Such power on the part of an element is called its chemi- cal affinity. All elements possess this affinity for certain other elements, but the strength of the affinity varies with the element and with the external conditions. To this affinity of elements all chemical compounds are due.14 CHEMICAL COMPOSITION OF LIVING MATTER Chemical compounds distinguished from physical mix- tures. — Most of the substances found in nature are mixtures of several substances. If we examine a piece of wood we can pick out several different kinds of matter which are intimately united in the compound wood. Similarly a piece of flesh shows pieces of fiber, and bits of fat, blood, and muscle, which are all intimately mixed together, but each of which possesses properties peculiar to itself. If we examine a bit of common salt and com- pare it with the elements sodium and chlorine, we find that the salt has properties which are distinct from either the sodium or the chlorine. In short the sodium and chlorine which have combined to form salt have lost their identity and individual properties, and the compound resulting from their union has entirely new properties. These examples illustrate the distinction between a mechanical mixture and a chemical compound. The flesh is a mechanical mixture of fat, blood, etc. The salt is a chemical compound. Whenever two or more elements combine, losing their individual properties and producing a substance with entirely new properties, the combinaton is called a chemi- cal compound. The white fumes which result from the union of phosphorus with the oxygen of the air arc entirely distinct in properties from the phosphorus and oxygen. In their formation, therefore, we have an example of the formation of a chemical compound. If the phosphorus is exposed to the air in a dark room a dull glow of light may be seen. This glow is due to the heat generated in the act of union of the phosphorus and oxygen. A certain amount of heat is always generated when chemical elements combine.SULPHUR 15 Sulphur. — (See Ex. IV.) Another important element in our body compounds is sulphur. Most of us have seen this element in its pure state and recall its yellow color. The sulphur used on sulphur matches is the element in its pure state, and wc may recall the fact that when it burns it gives off suffocating fumes and a blue flame. In its free state, this element is found only around volcanic regions. Combined with other elements (chemical com- pounds of sulphur) it is extremely abundant. Muscle is such a compound of sulphur with other elements. Pure sulphur is odorless and tasteless. When flesh decays it gives off a disagreeable odor. This odor is a property of a gas called hydrogen sulphide, which the sulphur forms as flesh decays, by combining with the element hydrogen. The ordinary hen’s egg con- tains sulphur, and in decay this gas is formed abundantly. The blackening of a silver spoon used in eating an egg is due to a black compound called sulphide of silver, which the sulphur of the egg forms with the element silver. The fumes from binning sulphur (a compound of sul- phur and oxygen), the bad smelling gas (sulphur and hydrogen) and the black compound (sulphur and silver) bring out the fact that a chemical element may have an affinity for more than one element. The fact that it occurs free in the air and yet combines with oxygen to form a compound when heated is an illustration of the way in which external conditions may affect the affinity of an element, while the blue flame which results when sulphur combines with oxygen and which is absent in the forma- tion of the other two compounds, shows that the amount of heat generated by the union of two elements may vary greatly with the elements.16 CHEMICAL COMPOSITION OP LIVING MATTER Carbon. — (Sec Ex. V.) When we partly burn a piece of wood, bone, or flesh, we find it transformed into a black substance which we call charcoal. In nature we find this same substance in a much harder form and we call it coal. When we use our lead pencils the mark we make is due to another form of this substance called graphite. Finally in the diamond we have a substance very different in appearance from charcoal, coal, or graphite, but essen- tially like them in some respects. All of these substances are forms of a very abundant chemical element called carbon. So important a, part of all living matter is this element that the study of its compounds is made a special branch of chemistry under the name Organic Chem- istry. In most of its forms, carbon shows a black color and is without odor or taste. In its free state it has so little affinity for other elements that farmers often char the ends of fence posts before setting them in order to insure against decay. When heated to a certain temperature, however, all forms of carbon burn and form a colorless gas, which may be readily detected by its power to turn clear lime water milky. This gas is a chemical compound of carbon with the oxygen of the air and is called carbon dioxide. It is this gas that is used to charge water in the manufacture of ordinary soda water. Just as in the union of phos- phorus and sulphur with oxygen, the formation of this gas is accompanied by the generation of much heat, and this fact is made use of in our bodies as well as in our stoves and furnaces. If we blow our breath through clear lime water the milky color that results gives unmistakable evidence that this gas is being constantly formed in our bodies and given off with the exhaled air.OXYGEN 17 A study of the element carbon demonstrates three important facts. First. It is present in large amounts as a part of all living matter. Seco7id. Its combination with oxygen to form carbon dioxide is always accompanied by the production of heat, and the formation of this gas in our bodies results in main- taining a constant body temperature. Third. A chemical element in its free state may occur in forms which appear very different to the eye, and must be tested by other means than sight to determine its identity. Oxygen. — (See Exs. VII. and VIII.) This important element occurs free in the air in the form of a colorless, odorless, and tasteless gas. In its combined form it is abundant in both living and non-living matter. It is of great importance to all living processes because it has a great affinity for other elements, and because, in combin- ing with other elements, it sets free a great amount of energy in the form of heat. While oxygen is present in the air in a free state, it is so mixed with other gases that special processes must be resorted to to obtain it pure. The simplest method of obtaining oxygen is to heat a compound which contains it. Some of these compounds when so heated break up and set free the pure gas. Two compounds which are especially favorable to this method are oxide of mercury (a compound of mercury and oxygen), and potassium chlo- rate (potassium, chlorine, and oxygen). If we heat some oxide of mercury in a tube, as shown in Figure 1, we note that it gradually disappears. If, while the heating is going on, we insert the glowing end EDDY. TUYS. —2.18 CHEMICAL COMPOSITION OF LIVING MATTER Fig. 1 — Apparatus for heating mer- curic oxide. of a charred match into the mouth of the tube the pres- ence of the oxygen gas will become evident by the in- creased brilliancy of the glow, due to the rapid union of the charcoal with the liberated oxygen. The presence of the mercury will be seen in the drops which condense on the cooled sides of the tube. Such an experiment not only illustrates the manner in which a compound may be separated into its elements, but also demonstrates an important property of oxygen, namely, that in its pure state its affinity for carbon is much greater than when mixed with other gases in the air. If we heat a mixture of manganese dioxide1 and potas- sium chlorate and collect the liberated gas over water as shown in Figure 2, it is pos- sible to compare the union of sulphur and phosphorus with tlois pure product, with the result when these elements combined with the mixed oxygen of the air. In every case the union is much more vio- lent than when ordinary air is present, and results in more heat being liberated. Oxidation and combustion. — Such unions of elements with oxygen as have been described in the preceding pages, are spoken of as oxidations and the compounds 1 The manganese dioxide undergoes no change, hut merely facili- tates the decompositiou of the potassium chlorate. Fig. 2 — Apparatus arranged for obtain- ing oxygen.OXIDATION AND COMBUSTION 19 formed are called oxides. For example, a chemist would describe the formation of carbon dioxide as the oxidation of carbon to form an oxide. In every case the formation of an oxide is accompanied by the production of heat, but sometimes the union of the elements takes place much more rapidly than at others, and when this occurs the amount of heat produced is large and often accompanied by flames or light. Such rapid unions are called rapid oxidation or combustion, to distinguish them from slower unions or slow oxidation. All burning then may be ex- plained as the rapid oxidation of the substance burned, and it is evident, therefore, that burning cannot take place in the absence of oxygen. Further, since in burn- ing the oxygen combines with the substance to form an oxide, it follows that for a substance to continue burning implies a continuous supply of oxygen until the substance has entered completely into combination. It is for these reasons that we supply our stoves with draughts which will permit the continuous entrance of oxygen. This explanation of the true nature of combustion also explains the methods of extinguishing fires, all such methods aim- ing to separate, through the aid of some substance which does not combine with oxygen, the burning substance from its air or oxygen supply. In short, pouring water on a fire puts an incombustible substance between the burn- ing matter and the air, and thus prevents further oxida- tion. Water is more frequently used than sand or dirt for this purpose because it is easier to handle, but in theory one is as effective as the other. The special interest of oxidation to the physiologist consists in the fact that it furnishes a means for the pro- duction of heat, and we shall learn later how the body20 CHEMICAL COMPOSITION OF LIVING MATTER makes use of oxidation to maintain the body temperature, while the necessity for a continuous oxidation explains the presence of carbon in the body and the continuous entrance of oxygen and exit of carbon dioxide in breathing. Nitrogen. — Experiments with pure oxygen show that combustion takes place more rapidly in it than in air. This implies that air must contain something besides oxygen, which dilutes it and makes oxidation slower. What is this substance? A simple experiment answers this question. (See Ex. IX.). Place a piece of phos- phorus the size of a pea in an evaporating dish that is floating on water. Light the phosphorus and thus raise its temperature to the combining point. Then cover the dish quickly with a bell jar (Fig. 3). The phosphorus Further, since the water rises in the jar to take the place of the consumed oxygen the volume of gas left in the jar subtracted from its original volume will give us the proportion of oxygen present in a given volume of air; roughly one fifth of its volume. What is this gas that is left? If we examine it by Fig. 3 — Apparatus arranged for obtaining nitrogen from air. will at once begin to combine with the oxygen in the in- closed air, and when it stops burning it will be because there is no more oxygen to combine with it to form the white fumes of phosphorus oxide. If we let the fumes settle they will dissolve in the water and there will be left in the jar only what was in the air before the oxygen was taken out.HYDROGEN 21 inserting heated elements as we did oxygen, we find that while it is colorless, odorless, and tasteless like oxygen, it has not the power of that element to combine with these other elements or produce combustion. This residual gas which is so lacking in chemical affinity is an element called nitrogen. Like sulphur, however, its lack of affinity is due rather to lack of the right conditions for combina- tion than to absence of that power. In nature, nitrogen occurs very abundantly in combination with other ele- ments, and some of its compounds form important parts of our body material, notably muscle or flesh. Owing to the weakness of its affinity for most elements its compounds arc apt to break up readily, and on that account are spoken of as un- stable compounds. Most of our explosives, such as gunpowder and nitroglycerine, owe their explosive power to the fact that they are unstable compounds of nitrogen. Fig. 4—^.apparatus for obtaining hydrogen and oxygen with the aid of an electric current. B, method of collecting the gases. Hydrogen. — (See Exs. X. and XI.). If we allow an electric current to pass through water (Fig. 4, A) which has had a little acid added to it to make it a conductor22 CHEMICAL COMPOSITION OF LIVING MATTEB of the current, we find that the current has the power to separate the water into two gases. One of the gases is twice as great in volume as the other and shows very different properties. If we collect these gases (Fig. 4, B) and examine them we find that oxygen is the smaller vol- ume. The other gas is colorless, odorless, and tasteless, like the first, but when we introduce a lighted splinter into it, it takes fire with an explosion and burns with a very hot colorless flame. If, instead of using water and the electric current, we pour sulphuric acid upon zinc (Fig. 5) the zinc will cause the acid to split and set free large quantities of this same substance which we may collect and examine more carefully. Such an exami- nation will show that not only does this substance burn, but that when it does burn it forms water. This peculiar gas is a chem- ical element called hydro- gen and since burning is simply the result of the combining of an element with oxygen, we must conclude that water is not an element but a compound of hydrogen and oxygen. The explosive force with which this combination takes place shows that the affinity of hydrogen for oxygen is very strong. Hydro- gen has a strong affinity for many other elements beside oxygen, especially for the element carbon; and its com- pounds with this element form a very large part of living matter. Aside from being one of the most active of Fig. 5 — Apparatus arranged for obtain- ing hydrogen from zinc and sulphuric acid.ACIDS, BASES, AND SALTS ; NEUTRALIZATION 23 chemical elements, hydrogen is also noteworthy as being the lightest chemical element known. Metallic elements. — All the elements we have con- sidered so far are called by chemists, nonmetals, to distin- guish them from another group of elements called metals. Some of these metallic elements are very familiar to us under the names of iron, zinc, copper, gold, etc. Beside these more familiar metals, there are others that form important parts of our bodies but which very rarely occur uncombined owing to their greater affinity. _ Such are the elements calcium, potassium, sodium, and magnesium. The most important characteristic of metals is that when they combine, it is with a nonmetallic element, and very rarely with another metal. Such compounds of metals and nonmetals fall very readily under two classes, bases, and salts. All chemical elements are divisible into metals and nonmetals. Acids, bases, and salts; neutralization. —When iron is exposed to the air it takes on a reddish coating that we call rust. This action is hastened by the presence of moisture. This rust is merely a compound of the metal iron with the oxygen of the air or water. Similarly other metals such as potassium, sodium, and calcium, etc., are able to unite with the oxygen of air or water and form oxides. If we dissolve some of these latter oxides in water the mixtures have peculiar properties. They have a slippery feel and a soapy taste, and when we put into them a piece of paper dyed red with the vegetable coloring matter, litmus, they change the color of the paper from red to blue. Such mixtures are not solutions but true chemical compounds of the oxides and the water, and are called bases. If we put the paper which was colored blue by these bases24 CHEMICAL COMPOSITION OP LIVING MATTER into vinegar or lemon juice it will turn red again. The reason for this is that vinegar and lemon juice contain certain combinations of nonmetallic elements called acids. The acid present in vinegar is a combination of hydrogen, oxygen, and carbon, called acetic acid. Some other powerful acids are sulphuric acid (hydrogen, sul- phur, and oxygen), nitric acid (hydrogen, oxygen, and nitrogen), and hydrochloric acid (hydrogen and chlorine). All acids have a sour taste, and contain no metals in their composition. If we add drop by drop some of a base to an acid, we shall evidently come to a point when the tendency of the acid to turn litmus red is just counterbalanced by the tendency of the base to turn it blue. When this point is reached the base is said to have neutralized the acid, and the process is called neutralization. Finally, if we drive off the water of such a neutralized mixture by evap- oration, we have left a product which turns litmus neither blue nor red and is called a salt. Analysis of such a prod- uct shows it to be composed partly of the metal which was in the base and partly of the nonmetals which were in the acid. You can make common table salt yourself by mixing hydrochloric acid and the base caustic soda (oxide of sodium) in this way. (See Ex. XII.) Salts, or compounds of metals and nonmetals, form an important part of the composition of our bodies, bones being a salt of calcium, phosphorus, and oxygen. Since most of the minerals in nature, limestone, quartz, etc., are metallic salts, such salts in the living matter are spoken of as mineral salts to distinguish them from compounds of purely nonmetallic elements.MINERALS 25 Chemical Compounds of the Body. The elements previously enumerated very rarely occur in an uncombined state in plant or animal matter; the only important exception to this rule being the element oxygen, and even that is more abundant in combined form than in the free state. The various compounds (acids, oxides, salts, and bases) are so numerous and varied that it would take too long to enumerate them all. Fortunately, they admit of classification into a few char- acteristic groups of life properties. These groups are called, respectively, minerals, gases, and carbon com- pounds. Minerals. —Under this head are grouped water, mineral salts, and all acids that do not contain carbon. The only important acid in our bodies under this group is hydro- chloric, and this is found in small quantities in the gastric juice of the stomach. Water forms more than fifty per cent, by weight, of all living matter. In the human body approximately 59% is water; the bones contain 22%, the liver 69%, the muscles 75%, and the kidneys 82%, while the fluids of the body are still richer in water content. Mineral salts will not burn, and by burning different parts of the body until nothing but the ash or mineral salts remain we cannot only demonstrate their presence in every part of the body, but also determine the form and proportion present. Such determination shows them to be of great variety as to kind. The most common are the chlorides, carbonates, and phosphates of the metals sodium, calcium, and potassium. These names indicate the predominant elements in these compounds. Bone ash also represents nearly 33% by weight of the entire bone,26 CHEMICAL COMPOSITION OF LIVING MATTER and is composed mainly of calcium carbonate and calcium phosphate. Gases. — Oxygen and carbon dioxide occur in all living matter. They may be free, as in the lungs, or absorbed in the fluids of the body. These two are the most impor- tant gases present, but others are present in smaller quantities. Carbon compounds. —The compounds of this element in living matter are so numerous and so important that they have been given the name of organic compounds, or life compounds. They may be classified under two groups, nitrogenous and non-nitrogenous. Nitrogenous carbon compounds. — As the name indi- cates, these compounds contain the element nitrogen in addition to the elements carbon, hydrogen, and oxygen. They are of many forms. Proteid, which contains sulphur, in addition to the above mentioned elements, is the name given to the form which is perhaps most abundant, and which under the name of albumin and fibrin in the blood, myosin in the muscles, and casein in milk, is very im- portant in its relation to vital processes. When proteid burns it loses part of its carbon and the result is another nitrogen compound called urea, which is given off as a waste by the kidneys. Still another form of nitrogen compound is found in the gelatine of the bones and the keratin of the hair and nails. Non-nitrogenous carbon compounds.— In this group are placed all carbon compounds found in living matter, which do not contain nitrogen. This group is divisible into two sub-groups, carbohydrates and fats. Carbohydrates. — Besides carbon these compounds con- tain only hydrogen and oxygen, which are always in theNON—NITROGENOUS CARBON COMPOUNDS 27 proportion in which they occur in water, namely, two parts of hydrogen to one of oxygen. Carbohydrates fall into two main divisions, the sugars and the starches. The most important of the sugars is glucose or grape sugar, found in the blood and in many other parts of the body. Starch occurs in the body in a form called glyco- gen, which is stored in the liver. When a carbohydrate is oxidized it liberates much heat, and the oxidation in the body of these compounds furnishes us much of our warmth. Fat. — This substance is present in the body in varying amounts. Like the carbohydrates it is a compound of carbon, hydrogen, and oxygen, and like them it produces heat when oxidized and thus acts as a body fuel. It occurs largely in a liquid state in the living body, in the muscles, the blood, the bones, and connective tissues. We shall consider the properties of these carbon com- pounds further, in our study of foods.III. THE STRUCTURAL UNITS OF LIVING MATTER. The analyses of the chemist tell merely what elements or combinations of elements enter into the building of living matter. The principal interest in such analyses lies in the proof they give that all living matter, plant or animal, consists of essentially the same chemical elements. When we speak of the struc- tural composition of a given plant or animal we do not refer to its chemical make up, but to the form which these compounds take. Thus the most evident structural forms these com- pounds take in our bodies we call eyes, ears, bones, flesh, etc., while in plants they receive the name of leaves, roots, stems, etc. Gross structure of the body. — Roughly, the body consists of a head, a trunk, and append- ages (arms and legs). All these parts agree in being made up of soft matter supported by a framework of hard matter. Lower animals, like the crayfish and grasshopper, also consist of hard and soft matter, but with them the hard matter, when present, is on the 28 Fig. 6— A, diagram of a human trunk split to show the internal skeleton. It, diagram of a grass- hopper split to show the external skeleton. Hard parts are in black.GEOSS STKUCTUKE OF THE BODY 29 outside and forms what is called an external skeleton, while in man it is internal with the soft parts packed upon it. If we examine a diagram (Fig. 7) of the human body split lengthwise the arrange- ment of these soft and hard parts is made clearer. Such a diagram shows that the head is really a box of bone resting on top of the backbone with an external cover- ing of soft parts, the eyes, ears, cheeks, and scalp, and containing within a mass of soft material called the brain. This latter is continued down the center of the backbone in a long cord called the spinal cord. The trunk is divided into two cavities, one above the other. The upper cavity, the chest, is protected on each side by arches of bone called the ribs, on the front by the breastbone, and is separated from the lower cavity by a thin membrane called the diaphragm. The lower cavity is supported at the base by a broad basket of bones called the pelvis and at the back by the back- bone. The upper cavity is filled with the heart and lungs, while the lower contains a number of soft organs (stomach, intestines, kidneys, etc.). When Fig. 7 — Diagram of a human trunk split lengthwise; a, brain and cord cavity; b, nasal cavity : c, mouth cav- ity ; d, regions of alimentary canal; 0 + o; 2 + 2>3 + 3 = 28. Structure of the teeth.—(See Fig. 27.) The part of a tooth above the gum is called its crown. When a tooth is pulled we find that the crowm is only a small part of the entire tooth. At the point where it enters the gum it narrow's into a part called the neck and extends below in one or more projections called the fangs or roots ivhich anchor it firmly in its bony socket. The incisors and canines have only one fang, the bicuspids may have two, while the molars may have as many as five and as few as two. The internal structure of all human teeth is practically the same. If we split a tooth in two, lengthwise, examina- tion shows that the crown is covered with an external layer of hard matter called enamel. This thins down at the neck and is succeeded by a fang covering called cement. The main bulk of the tooth is formed of a bonelike sub- stance called dentine which surrounds a small central cavity called the pulp cavity. During life this cavity is filled with nerve fiber and blood vessels interlaced in a network of connective tissue, called the pulp. It is lined with a layer of epithelial cells which secrete the mineral matter of which the outer layers are formed. This cavity has a small EDDY. PKYS.- 7 Fig. 27 — Section of a tooth; k, crown; e, enamel; n, neck ; d, dentine ; /, fangs; c, cement; p, puip cavity inclosed by dentine.98 DIGESTION opening at the base of each fang through which the nerves and blood vessels communicate with the rest of the body. The blood supplies the epithelial cells with the material out of which they build the tooth, and the nerves probably direct this process. In some animals the teeth grow throughout life, but in man their growth ceases upon the attainment of a certain size. Digestive action of the teeth. —The teeth grind and Pig. 28 — The tongue. break the food into small particles, and thus prepare it for the solvent action of the digestive juices. If the food is not properly chewed its solution is necessarily delayed justGLANDS AND SECRETIONS 99 as a lump of sugar dissolves less quickly than the same amount of pulverized sugar. The grinding action of the teeth is called mastication. The tongue. —(See Fig. 28.) The tongue is a soft, fleshy mass of matter mainly composed of muscle with a cover- ing of mucous membrane. It is fastened at its base to a bone called the hyoid. Its tip projects forward, and is free and very mobile. It has two functions, one mechanical, the other sensitive in nature. As a sense organ it is the seat of taste. As a mechanical organ it mixes the food with saliva and pushes it into the proper position for chew- ing and swallowing. It also serves as an indicator of the health of the digestive system by becoming coated with a “ fur ” or coat of yellowish mucus whenever the digestion is impaired. Its healthy color is red. Chemistry of Digestion. Both tongue and teeth are merely mechanical agents which prepare the food for the solvent action of a liquid called saliva. This liquid is a typical digestive fluid in that it is produced by a structure called a gland, and exerts a solvent action through chemical change induced by means of a contained ferment or enzyme. We shall understand its action better if first we make clear what a gland and a ferment are. Glands and secretions. — (See Ex. XXXII.) Every cell has the power of taking in food and giving off waste matter. Evidently, then, the character of the food which a cell takes from the blood determines, in a way, the kind of waste which it shall give off. Sometimes the matter so given off is a solid, and the result is the formation of hard100 DIGESTION structures like the bones and teeth. When it is liquid or semiliquid the matter is called a secretion. There are many kinds of secretion given out by the various cells of the body, and some of the secretions have an important part to play in the action of the body. So important are some of these secretions, in fact, that certain cells are set aside whose sole work is their preparation. These special- ized cells often become grouped together in certain regions, and are then spoken of collectively as glands. A gland, then, is merely a collection of cells whose special duty is the giving off of a certain kind of secretion. To this group of secretions belong all the digestive fluids, and the cells or group of cells which secrete them are called the diges- tive glands. In this group are included the salivary glands, the pancreas, the liver, and the mucous membrane. The form of tissue which furnishes most of these speci- alized cells is the epithelial, and the simplest form of a gland would be a layer of these cells so placed as to receive food from the blood and give off their secretion. Such a glandular structure is called a secreting membrane, and an example is the mucous membrane which lines all parts of the alimentary tract. (See Fig. 29.) In so complicated a structure as the body, however, it is necessary to economize space, and it is easy to see that one of the simplest methods of doing this is to arrange a layer in a simple fold or sac. In this form (see Fig. 29) the secretion is poured out into the central cavity or lumen, which, when filled, overflows through the mouth or opening of the gland. Of such a character are the gastric glands of the stomach, and the simple glands {crypts) of the intestine. Evidently, the next step in saving space would be to collect a series of these simpleGLANDS AND SECRETIONS 101 sacs about a common duct, and such arrangements are called tubular, racemose or tubular-racemose according as the separate elements are tubelike or branched or a com- bination of the two. Such glands are called compound 2 0 Fig. 29 — Forms of glands; A and B, simple layers of mucous membrane cells; 1, a simple tubular gland ; 2, a forked tubular gland ; 3, a simple saccular gland with duct, d ; 5, a saccular gland still more divided ; 6, a, compound racemose gland with common duct, d.102 DIGESTION glands, and in this class are the salivary glands, liver, and pancreas. Enzymes. — (See Ex. XXXI.) The secretions of the digestive glands owe their power of dissolving nutrients to the presence of certain complicated chemical compounds called enzymes. The action of these enzymes is little understood at the present time, but their effect is well known. In general, they all agree in causing a chemical change in a nutrient without being themselves consumed in the process. For example, if we add a little malt (see p. 83), which contains the enzyme diastase, to some starch paste, the starch is gradually changed to sugar, but the amount of the diastase which causes the change is neither increased nor diminished by the process. Similarly, yeast added to a sugar solution causes the sugar to break up into two simpler compounds, alcohol and carbon dioxide, while the number of yeast plants is not decreased by the process. The name ferment was long used to describe these com- pounds, and hence the action of yeast or bacteria was described as fermentation. We now know that yeast and bacteria produce their effect through definite chemical compounds, and to all such compounds we give the name enzymes. The salivary glands. —(See Fig. 30.) There are three pairs of these glands in man. They arc called the parotid, the submaxillary, and the sublingual. All these glands are of the compound, racemose type, and resemble bunches of grapes in which the grapes are the sacs of secreting cells, and the connecting stems are the hollow ducts. The parotid glands lie in front of and under the ear on eachACTION OF THE SALIVAEY GLANDS 103 side of the head, and their ducts enter the mouth on the inner side of the cheeks, just about opposite the second upper molar teeth. It is the inflammation and enlarge- ment of these glands that causes the disease known as the mumps. The submaxillary glands lie below the two halves of the lower jaw, and their ducts open into the mouth at points near the middle line of the under side of the tongue. The sublingual glands lie under the mucous membrane below the tongue and floor of the mouth. They open into the floor of the mouth by many small ducts. Action of the salivary glands. —Each cell of a gland obtains its food sup- ply from the blood vessels, and out of this food it manu- factures the secretion known as saliva. Since the amount of saliva needed to di- gest the food we eat varies with the amount of food pres- ent in the mouth, it is evident that the secretion of the cells must be under con- trol. In this respect the glands are like a factory whose output is regulated to meet the conditions of supply and demand. In the case of the cells this control is exercised by nerves whose general arrangement is indi- Fig. 30 — Relation of salivary glands to the mouth cavity.104 DIGESTION cated in the diagram. (See Fig. 31.) The results of this arrangement are as follows: The nerves may be stimu- lated in several ways: by the contact of the food in the mouth with the nerve ends, by the odor of the food which stimulates the nerve ends in the nose, or by the stimula- points. One impulse goes to the blood vessels and causes them to dilate, and thus supply more blood and food. The other impluse is given to the cells, and stimulates them to greater secretive activity. The result of these two directing impulses is an increased flow of saliva. In a similar way, this threefold nerve connection also accounts for the dryness of the mouth when certain con- ditions are present. For example, fright, a bad smell, or unpleasant taste, may start impulses along the nerve which, instead of increasing the flow of blood and secre- tion, may constrict the blood vessel and diminish the secretive activity of the cells. The saliva. —The secretion produced by the salivary glands is called the saliva. The liquid found in the mouth and constituting the spit is not pure saliva, but a mix- ture of this liquid with a secretion of the mouth lining called the mucus, together with a number of other more ox- less solid elements. This mixture is a colorless, cloudy, Impulse Controlling Size Of Blood Tube And Amount Of Blood Supply Fig. 31 — Diagram to illustrate mouth control. Impulse Controlling Secretion common Secreting Cells Gland Duct tions of the nerves at the brain end, as by thought of food. When the nerves are stimulated in any of these three ways the impulse they carry is com- municated to twoACTION OF SALIVA 105 slightly alkaline liquid, with a ropy consistency due to the mucin in it, and is often frothy from the presence of air bubbles. Pure saliva is a clear, viscid, colorless liquid consisting mainly of water and a certain enzyme called ptyalin. It is this enzyme which is responsible for the chemical action of saliva in digestion. Action of saliva. —Of all the nutrients taken into the mouth starch is the only one acted upon by the saliva. If clear saliva is allowed to act upon starch it changes it to sugar in exactly the same way that the diastase of malt changes grain starch to sugar. This sugar is, of course, readily soluble in the watery part of the saliva, and thus the nutrient starch is forced in solution. That this transformation is actually produced may be readily demonstrated by chewing a little dry biscuit for a few moments. The dry, tasteless mass soon becomes sweet to the taste as its starch is changed to sugar by the saliva. The chemical changes involved in this transformation are rather complicated, but for our purpose it is sufficient to state that they have their origin in the action of the enzyme ptyalin, and we can, therefore, say that it is the ptyalin which produces the change. Experiments with starch paste and clear saliva (see Exs. XXXV. a and XXXVI.) show that this transfor- mation takes place best at the normal mouth temperature (98° F.), while high temperatures (above 145° F.), or very low temperatures (below 32° F.), or the presence of strong acids or alkalies are sufficient to stop completely the ptyalin action. Further, cooked starch is much more easily changed to sugar than uncooked, and requires less time for the action of the enzyme. Saliva also aids digestion in a mechanical way by fit-106 DIGESTION ting the food for swallowing. It is impossible to swallow dry food, and the moistening with the saliva makes this ac- tion possible, and at the same time acts as a lubricant and allows it to slip down more easily. In fact, so short is the time that food spends in the mouth in ordinary eating that probably very little starch is changed into sugar there, and the mechanical action, therefore, is the most important. Saliva, as we shall see later, also aids in producing the sensation of taste. Summary op Digestion in the Mouth. 1. The teeth grind the food into small particles and fit it for rapid and thorough mixing with the saliva. (Mastication.) 2. The tongue rolls the food about into position for mastication and mixes it with the saliva. 3. The saliva through the action of the ptyalin enzyme con- verts part of the starch into soluble sugar. 4. The saliva moistens and lubricates the food mass and thus fits it for swallowing. 5. The food leaves the mouth in proper shape for the action of other juices and with part of its starch converted into sugar.VIII. DIGESTION (continued). After the action of the mouth the food is ready to be passed on to the stomach, where the next steps in diges- tion take place. The first step in this process is the act known as swallowing or deglutition. In this action the Fig. 32— Swallowing or deglutition ; a, tongue ; b, pharynx; ct food bolus ; d, soft palate; e, uvula ; fy epiglottis. A shows food being pushed back by the tongue. £ shows position of uvula and epiglottis as food falls into the pharynx. uvula at the back of the mouth is raised automatically to close the nasal passage, and the tongue simply pushes the food backward until it falls or slips into the pharynx cavity. Structure of the pharynx cavity. — (See Fig. 24.) This cavity is a conical bag just back of the mouth cavity and about five inches in depth. It has seven apertures ; two into the nasal cavity at the top, two into the ear (the Eustachian tube openings), one into the mouth, one into the windpipe (glottis), and one into the esophagus or 107108 DIGESTION gullet. The glottis opening is protected by a sort of trapdoor called the epiglottis, which is closed during swallowing but open at other times to permit the entrance and exit of air in breathing. The whole cavity is lined with mucous membrane, whose secretion of mucus moistens and makes flexible its walls. (Excess of this mucus produces catarrh.) When the food is swallowed the epiglottis is closed and the food falls upon the top of the gullet. Sometimes the epiglottis does not close quickly enough, and some food enters the air passage. This produces choking, that is to say, the for- cible expiration of air in the attempt to force out the misplaced food. If properly swal- lowed, the food is forced along the gullet to the stomach. The esophagus. — The esophagus, or gul- let, is the name given to that part of the alimentary tract which begins at the lower end of the pharynx cavity, and passing through chest and diaphragm, enters the stomach. This portion is in the form of a tube of about nine inches in length and is formed by three layers of tissue, (^ee Fig. 33 and Ex. XLI.) The inside layer is a continuation of the mucous membrane, and the secretion of mucus here together with the saliva acts as a lubricator, making the food slip down easily. The outer layer is formed of muscles, while between this and the inside layer is a layer Fig. 33 — Diagrammatic section of the wall ' of the esophagus ; -S'. Mserous membrane ; L. M.y longitudinal muscles ; Cl Mcircu- lar muscles ; M. AT., mucous membrane. Be- tween the circular muscles and the mucous membrane is a thin layer of connective tissue.THE ABDOMINAL CAVITY AND CONTENTS 109 of connective tissue. The muscular layer is actually a double layer, the inner fibers arranged in a circular man- ner, while the outer ones are extended lengthwise of the tube. By alternate contractions and expansions of these muscles the food is forced along the tube to the stomach. In this action the effect produced is similar to what hap- pens when a ring slightly smaller than the tube is forced along a rubber tube; it forces any bodies inclosed in the tube in the same direction that the ring is moving. The contractions of the circular and longitudinal bands of muscles necessary to this action in the gullet have received the name peristalsis. Through peristalsis of the esophagus walls the food is always forced toward the stomach, regardless of the position of the body. The food undergoes no chemical change in either the pharynx cavity or esophagus. The Abdominal Cavity and Contents. This large body cavity (seep. 29, and plate I), incloses the greater part of the alimentary canal. (See Ex. XXXIII.) It is separated from the chest at the top by a muscular partition called the diaphragm, while its base is formed of the basketlike bones of the pelvis. At the back are the spinal column and the lower ribs, while the sides and front are formed of thick sheets of muscle without bone support. This cavity is lined throughout with a very smooth membrane called the peritoneum. This peritoneum is so arranged that it not only lines the cavity but also, by extensions, forms folds which support the various parts of the alimentary canal. These folds are so arranged that the parts of the canal are hung in it like a110 DIGESTION stone in a sling. (See Fig. 34.) The entire mem- brane is lubricated by a watery liquid or serum, and Fig. 34 — Intestine supported by a fold of the peritoneum (mesentery), showing method of entrance of blood vessels. for this reason it is called a serous membrane. It is also richly supplied with blood vessels. The organs sup- Fig. 35 — External view of the stomach, showing covering of peritoneum and muscles ; 1, esophagus ; 2, cardiac end ; 3, pyloric end ; 4, small intestine ; 5, a portion of peritoneum turned back; 6, longitudinal muscle libers ; 7, circular muscle fibers; 8, oblique muscle fibers.THE STOMACH 111 ported by this membrane are the stomach and the many folded intestines. The stomach.— (See Ex. XXXIII.) This organ is a bag- like swelling (see Fig. 35) of the alimentary canal lying just under the diaphragm and across the upper part of the cavity. The larger end is at the left side of the body, and into its upper surface the esophagus opens by an ori- fice called the cardiac orifice. The right end of the stomach tapers into a narrow neck which is comiected with the be- ginning of the small intestine by an opening called the pylorus or pyloric orifice. The whole bag is supported in a fold of the peritoneum, whose two parts come together at the lower edge of the stomach and hang down like an apron over the front of the ab- dominal cavity. This apron is known as the greater omentum, and is fre- quently the seat of a great accumula- tion of fat. The walls (see Ex. XLI.) of the stomach are composed of four coats or layers. (See Fig. 36.) The outermost is a serous membrane (the peritoneum just mentioned). Next inside this is a muscular coat composed of three layers of muscle fibers. Then comes a layer of connective tissue (the submucous coat), and lining the entire stom- ach is the mucous membrane. The blood vessels of the stomach enter between the folds of the peritoneum, and Fig. 36 — Section of stomach w a 11; n, serous membrane (peritoneum) ; l. m.y longitudinal mus- cles ; c. m., circular muscles ; s. m., sub- mucous coat; m.m.j mucous membrane; d, ducts in mucous membrane.112 DIGESTION branch into capillaries in the submucous layer. The mus- cular coat is made up of unstriped muscle fibers, and varies greatly in thickness at different parts. It is thickest at the pyloric end. The outermost layer of fibers is a con- tinuation of the longitudinal fibers of the esophagus and is Fig. 37—Longitudinal section of the stomach; a, esophagus ; b, greater curvature; c, lesser curvature; d, sphincter pylorus; h-i, small intestine. The ridges in the stomach are folds of mucous membrane. thickest over the curvatures of the stomach. The circular muscle fibers of the esophagus are likewise continued, and form the inner layer of muscle over the stomach. This circular layer becomes thickest at the pyloric end. Between these two layers is a very incomplete oblique layer whose fibers radiate from the cardiac orifice. The thick ring of circular fibers at the pyloric end is called the sphincter < pylorus, and controls the entrance of food into the small intestine. 6GASTRIC GLANDS 113 The mucous coat is a pink membrane which is not elastic, and, though smooth when the stomach is fully distended, is thrown into folds when the organ is empty. This mem- brane derives its pink appearance from the blood vessels underlying it and becomes much redder during digestion, as these submucous capillaries become distended with blood. When examined with a lens the mucous membrane is seen to be covered with shallow pits. These pits are the mouths of many simple tubular glands (the gas- tric glands) which secrete the digestive fluid of the stomach. This fluid is called the gastric juice. Between these special glands are the ordinary mucous glands which secrete the lubricating mucus. Gastric glands. —These glands (see Fig. 38) are simple tubes lined with epithelial cells, formed by the folding of the mucous membrane. Between suc- cessive tubes project layers of the sub- mucous capillaries. These capillaries furnish the food out of which the cells manufacture the fluids and ferments used in stomach digestion. The cells of these glands are of two kinds. Those which secrete the enzymes, pepsin and rennin, are called the chief cells to distinguish them from another group called the border ceZlswhich are supposed to secrete hydrochloric acid. The exact function of these border cells is still unknown. (For arrangement of these cells see Fig. 38.) Fig. 38 — A peptic gland, from cardiac end of stomach. Very much magni- fied. Ay central or chief cells, which make pepsin; B, border or parietal cells, which make acid. [From Miller’s Histology.] EDDY. PHYS. — 8114 DIGESTION Digestion in the Stomach. Gastric juice.—The digestive action of the stomach is due mainly to the action of the digestive fluid produced by the gastric glands. This gastric juice is a thin, nearly colorless fluid, consisting mainly of water and holding in solution a small proportion of hydrochloric acid and two enzymes, pepsin and rennin. As a result of the acid contained it gives a red color to blue litmus, and, since ptyalin can act only in an alkaline medium, the saliva swallowed with the food can exert no digestive action upon starch after the food is acidified. The flow of the gastric juice begins with the entrance of food into the stomach, and the flow increases with the amount of food taken in. This flow may also be stimu- lated by certain food accessories, by the alkaline saliva, and by the flavor of the food. Normally, about three quarts of gastric juice is produced by the glands in twenty- four hours. Digestive action of the enzymes.—Pepsin. This enzyme has the power to transform proteids to a soluble form called peptone. It acts only in an acid medium, and what is known as peptic digestion is therefore the result of the combined action of pepsin and hydrochloric acid. Peptic digestion varies with temperature. Low temperatures retard, while very high temperatures may stop completely the action of the pepsin. Pepsin can be obtained in a fairly pure state (see Ex. XXXV., b) and with the product so obtained it is possible to prepare artificial gastric juice whose action upon various foods can be observed with great accuracy. (See Ex. XXXVII.) In such experi- ments the artificial juice is allowed to act upon food prep-DIGESTIVE ACTION OF THE ENZYMES 115 arations contained in glass vessels. The results of such experiments may be summarized as follows: The proteid thus treated first swells and softens and then is slowly transformed into a soluble form of proteid known as peptone. This peptone is the result of the combined action of the pepsin and hydrochloric acid, the food proteids being broken gradually into simpler and simpler forms, until finally the soluble peptone results. Peptic digestion is found to take place best at a tem- perature of from 98° to 102° Fahrenheit. The action of peptic digestion is confined wholly to the proteid parts of the food, and does not directly affect any other nutrient. In fat meats the albuminous pockets are dissolved away in this action, and the result is the liberation of the fat globules inclosed by the pockets, but the pepsin does not change the fat in any way. In similar manner starch grains held in albuminous pockets may be liberated, but the starch itself is not affected. All these actions of the pepsin and acid which are ob- served in experiments with glass vessels take place nor- mally under similar conditions in the stomach. Rennin. (See Ex. XXXVII., b.) Gastric juice con- tains a second enzyme called rennin. This enzyme plays an important part in the digestive action of the gastric juice upon the particular form of proteid found in milk. When milk proteid (caseinogen) comes in contact with gastric juice, the rennin enzyme causes it to coagulate and separate from the rest of the milk as curd or casein. This action is exactly similar to the curdling of milk by acid when it sours. This separation of milk proteid by the rennin prepares it for the action of the pepsin and116 DIGESTION acid, and once separated, peptic digestion takes place as with ordinary proteid. In children whose diet is almost entirely of milk this rennin action becomes very important. Hydrochloric acid. (See Ex. XXXTX.) Aside from the aid which this acid gives in peptic digestion its pres- ence is important in that it destroys the germs of fermen- tation and disease, and probably dissolves some mineral salts. Its action in destroying germs permits the food to be stored in the stomach for some time without under- going decay. In this way the stomach may act as a storehouse of food. While proteids arc practically the only nutrients digested in the stomach, it is also true that some of the preparatory actions upon other nutrients take place here. For example, the heat of the stomach is sufficient to liquefy the fats of the food, and the digestion of the proteid covers of starch and fat particles liberates these nutrients and prepares them for future digestive changes. Sugar and certain mineral salts may also be dissolved here by the water and acid of the gastric juice. The mixture of all these substances together with the digested and undigested particles of food and the gastric juice results in the formation of a milky fluid called chyme. This chyme is from time to time forced out of the stomach into the small intestine. Mechanical action of the stomach. — Once in the stomach the food is entirely cut off from the rest of the alimentary canal by the contraction of the cardiac and pyloric sphincter muscles. It was long thought that dur- ing the period of stomach digestion the food was forced in continuous circulation round and round the stomach until sufficiently ground, and mixed with gastric juice to permit of its discharge into the intestine. Later investi-SUMMARY OF DIGESTION 117 gators showed that while this mixing does take place the method is somewhat different from what was formerly supposed. According to these investigators, the stomach cavity may be divided into two parts, in only one of which does the actual mixing take place. The cardiac end of the stomach or fundus shows almost no movement and acts simply as a storehouse. The contractions start in the middle of the stomach and run toward the pylorus end, and only the portion in this end is actually in process of mixing. These contraction waves are repeated at inter- vals of two minutes or less, and finally reduce the food in this end to a thin liquid. At intervals depending upon the character of the food, the sphincter pylorus relaxes and portions of the liquid mass are spirted into the intestine while new material flows forward from the storage end to take its place. When the food is liquid these relaxations of the sphincter are very frequent, while for solid food they occur at longer intervals. The time required for the stomach to empty itself in this way is from two to five hours after a meal. The exact time, of course, depends upon the amount of food taken into the stomach, and ease with which it may be broken up. After the stomach has been empty for a certain length of time we feel a sense of hunger, and this is the signal for a renewal of the food supply. Meanwhile, the chyme which has entered the intestine is undergoing further digestive changes. Summary of Digestion in the Stomach. 1. The food enters the stomach by the cardiac orifice, and by the peristaltic action of the muscular walls is mixed with the gastric juice. 2. The hydrochloric acid stops the action of the ptyalin and the digestion of starch, and makes possible the action of the pepsin.118 digestion It also tends to kill bacteria and prevent fermentation and aids in the solution of certain mineral salts. 3. The pepsin converts proteid into soluble peptone, part of which is absorbed directly by the blood vessels of the stomach. 4. The rennin coagulates the proteid in milk and prepares it for the action of the pepsin. 5. The partly digested proteids, and starch together with the undigested matter leaves the stomach as an acid mixture called chyme. Fig. 39 — The stomach and intes- tines ; 1, stomach ; 2, duodenum ; 3, small intestine; 4, end of ileum ; 5, caecum; 6, vermiform appendix ; 7, ascending colon ; 8, transverse colon ; 9, descend- ing colon ; 10, sigmoid flexure ; 11, rectum ; 12, anus. part of the canal consists Digestion in the Intestine. While some starch is changed to sugar in the mouth and some proteid to peptone in the stom- ach, the real seat of digestion in the body is the small intes- tine. Here all kinds of nutri- ents are digested, and the final preparation of the food for use in the body takes place. Here, too, the digestible - matter is separated from the indiges- tible. In short, even if there were no preliminary mouth or stomach action, the small in- testine would be capable of performing all the digestive actions necessary to prepare the food for the body. The intestine. — All of the alimentary canal between the stomach and the anus is known as the intestine. This of a tube of varying diameter,THE SMALL INTESTINE 119 and to its different parts have been given appropriate regional names. The part next to the stomach, and con- tinuing for some twenty feet from the pyloric orifice, is all included under the name small intestine. It varies in diameter from two inches at the stomach end to less than an inch where it enters the large intestine. By large intestine we include the five odd feet of tube extending from the point where the small intestine is considered to end to the anus. This portion of the tract has a diam- eter of from two and a half to one and a half inches. The relative position, size, and general arrangement of the two intestines arc shown in Fig. 39. The small intestine. — (See Ex. XXXII.) The entire part of the alimentary canal included under the name of small intestine is held in place by a fold of the peritoneum called the mesentery. This mesentery is about four inches in length at its back edge where it is fastened to the spinal column, and widens to twenty feet at its outer edge. The intestine is hung in this fold like an arm i$ a sling, and the outer edge of the mesentery and the intestine are very much coiled and folded to confine this long tube to the space adapted for it. The different regions of the small intestine are indicated by three names. The twelve inches nearest the stomach receive the name of duodenum, the succeeding two fifths is called the jejunum, and the re- mainder is called the ileum. These names are mainly useful in permitting us to designate special parts of the tube. Like the stomach the small intestine has four coats (see Ex. XLI.) or layers, an external serous layer (peritoneum), next inside a muscular layer, inside of this a submucous coat, and a lining of mucous membrane. (See Fig. 40.)120 DIGESTION The serous coat is formed of the fold of the mesentery. The muscular coat is in two layers, an outer one of longi- tudinal fibers and an inner of circular fibers. The submucous coat of connective tissue binds together the muscle and mucous membrane layers just as in the stomach, and receives the blood vessels which enter in a fold of the mesentery. The mu- cous coat which lines the whole intestine is pink, soft, and filled with minute vessels of the blood and lym- phatic system. In the upper part of the in- testine it is perma- nently folded into ridges known as the FIG. 40-Wall of small intestine; a, b, mucous valvuloe COnniveilteS, membrane with projecting villi; c, d, connec- . tive tissue layers (submucosa); e, circular WlllCll increase tile muscle fibers; e', longitudinal muscle fibers; digestive Surface and /, serous membrane ; <7, spaces between villi; 0 h, opening of crypts or simple intestinal glands ; delay the passage of i. blood vessels in villi. ., r , .. , T the iood particles. In the lower part these ridges disappear, and to the eye it appears smooth. Examined carefully with a lens the entire inner surface of the intestine is seen to be raised in minute projections called villi, like the pile of velvet.THE LIVER 121 (See Fig. 41.) We shall consider the structure and func- tion of these villi more in detail in our study of the absorp- tion of food. (See p. 134.) Between the bases of these villi are the openings of tiny glands called the crypts of Lieberkiihn. Digestive glands of the small intestine. — The glands which se- crete the digestive fluids of three in kind. Two of these Fig. 41—Yilli on surface of intestine (much magnified). the small intestine are are large and lie outside of the intestine, being connected with it by means of a duct. The other kinds of glands (the crypts) are very numerous and embedded in the mucous membrane itself. The two large glands are known as the liver and the pan- creas, and the ducts from these two glands unite outside the intestine and pour their combined secretions into the intestine through a common duct at a point on the duodenum a few inches from its connection with the stomach. The secretions of these two large glands are known as the bile and the pancreatic juice. The relation of these two glands and the ducts is shown in the diagram (see Fig. 42). The secretion of the crypts is called the intestinal juice. The liver. — (See Fig. 46 and Ex. XXXII.) This is the largest digestive gland in the body. It is a dark red mass located in the upper part of the abdominal cavity just under the diaphragm, and partly overlapping the stomach. The entire gland is covered with peritoneum, and is divided by a fissure into two unequal lobes. Of these two lobes,122 DIGESTION the right is much the larger. These two lobes are con- nected on the under side by ducts which emerge from each lobe and fuse into a common duct called the hepatic Fig. 42—Relation of bile duct and pancreatic duct to stomach and intestine. duct. On the under side is also located a small sac (the gall bladder), in which the bile secreted by the liver is stored until needed by the intestine. From this sac another duct (the cystic), leads, and fuses with the hepatic duct to form the common bile duct which opens into the duodenum. Two large blood vessels (the portalTHE LIVER 123 vein and the hepatic artery) carry blood to the liver, entering it on the lower side and breaking up inside the Fig. 43 — The liver. lobes into capillaries. These capillaries unite again, and the blood passes out of the liver by the hepatic veins. Fig. 44 —Two liver globules ; A, section across central hepatic vein ; B, section splitting the central hepatic vein ; p, capillaries of the portal vein which break up among the liver cells and finally unite in the central hepatic vein h.124 DIGESTION The internal structure shows it to be composed of a mass of many sided bodies called lobules. Each lobule (see Fig. 44) consists of a mass of cells (hepatic cells) sur- rounded by a network of capillaries. Branches of the bile ducts also enter these lobules and receive the bile which the hepatic cells secrete from the blood. Each hepatic cell, therefore, is a trqe gland cell, and the combined secre- tion of all these cells is collected in the branched ducts and flows downward until it enters the common hepatic duct. From this point it may take one of two courses. Fig. 45 — The pancreas ; 1, central duct; 2, pancreatic duct; 3, stomach. It either enters the common bile duct and flows directly into the intestine or it backs up and accumulates in the gall bladder. In this way bile is stored in this bladder so that when necessary it may be called on to supply the intestine with increased amounts of bile. In such case, bile flows from the lobes of the liver and from the gall bladder directly into the intestine. The bile is a thick golden-brown liquid of bitter taste. About a quart is produced daily, and performs very important functions in digestion. The pancreas.— (See Fig. 45 and Ex. XXXII.) ThisACTION OF THE PANCREATIC JUICE 125 organ, often called the sweetbread, is an elongated race- mose gland of a pinkish-yellow color lying along the greater curvature of the stomach. The cells of this gland secrete a digestive fluid called the pancreatic juice, which they pour into a central duct. This duct joins the bile duct close to its intestinal opening, and bile and pan- creatic juice mingle and enter the intestine by the com- mon opening. (See Fig. 42.) Pancreatic juice.— This is a watery secretion which contains three very important enzymes. One (trypsin) changes proteid to peptone, one (amylopsin) changes starch to sugar, and the third (steapsin) acts upon fats. Digestion in the Small Intestine. The partly digested food or chyme which is forced from time to time through the pyloric orifice of the stomach is now ready for the action of the various digestive fluids of the small intestine. (See Ex. XXXVIII.) The digestive action which takes place in the intestine is, therefore, due to the combined action of three digestive fluids, pan- creatic juice, bile, ami intestinal juice. Rhythmic peris- taltic movements of the muscle walls of the intestine keep the contents of the intestine in constant motion, and thus mix them with these combined juices. Action of the pancreatic juice.— This juice owes its digestive action to its three enzymes, trypsin, amylopsin, and steapsin. Trypsin. (See Ex. XXXVIII., a.) Trypsin is a more powerful enzyme than pepsin and, unlike that, acts best in an alkaline medium. Like pepsin, its action is affected126 DIGESTION by temperature; it is more active at ordinary body tem- perature (98-102° F.), while it is completely destroyed at a temperature of 172° to 176° F. Its action upon proteid is very similar to that of pepsin in that it converts pro- teid into soluble peptone. It differs from pepsin in that there is no swelling of the proteid under its action. The process is rather an eating away of the proteid part, and in this action the indigestible matter is left in its original shape to be broken into fragments later by the churning action of the intestine. It also reduces some albuminoids to peptone. Most of the proteid nutrient of our food receives its final preparation for absorption in the small intestine under the action of this enzyme. Amylopsin. (See Ex. XXXVIII., b.) Practically all the starch of foods is digested by the action of this enzyme. It is possible that some of the ptyalin of the saliva whose action was stopped by the acidity of the stomach shares in this digestive action upon returning to the alkaline medium of the intestine. It would be impossible to determine this definitely, since amylopsin and ptyalin are apparently identical in structure and action, and in Ger- man texts on physiology the name ptyalin is used for both enzymes. The use of the two names is, however, convenient in indicating their origin. Both convert starch to sugar under similar conditions and thus transform it into a soluble substance. Steapsin. (See Ex. XXXVIII., c.) This enzyme (also called lipase), which is extremely sensitive to temperature, performs two distinct operations upon fats, saponification and emulsification. In saponification the fat is first split into glycerine and fatty acids. These latter unite with the alkalies andACTION OF INTESTINAL JUICE m alkaline salt present, and form soaps which are, of course, soluble. In emulsification the fat is separated into tiny droplets and surrounded by the fatty acid or alkaline material in such a way as to prevent these drops from uniting again. In this form they may be absorbed directly without further action. Whether emulsification precedes saponification or vice versa is still a disputed point. In general, then, the pancreatic juice first transforms the acid chyme of the stomach into an alkaline fluid through the neutralizing action of the alkaline salts in the juice, and then by the action of the three enzymes it converts proteids, albuminoids, starches, and fats into substances fit for absorption. Chyme is neutralized mainly by bile salts. Action of the bile. — (See Ex. XXXVIII., d.) It has been found by experiment that when pancreatic juice is mixed with bile the emulsifying power of the steapsin is greatly increased thereby. The alkaline f ile salts are of aid in the splitting of fats and consequently in the saponification. Its presence is also supposed to prevent, to a certain degree, the putrefaction of foods, though this antiseptic power is disputed by some authorities. Finally, it takes a direct part in destroying the peptic digestive action of the acid chyme. It not only neutralizes the acid but also causes a precipitation or separation of the proteids in such a form that they may be acted upon directly by the trypsin. Action of intestinal juice (succus entericus). — This juice is produced by the simple glands known as the crypts of Lieberkiihn. It is a yellowish liquid with a strong alka- line reaction. This alkaline character it derives from the presence of sodium carbonate, and the presence of this128 DIGESTION alkali is important in aiding the emulsification of fats. It varies in quantity in different regions of the intestine, being scarce in the upper part and very plentiful in the lower or ileac region. Its specific action is due mainly to the presence of two kinds of enzymes. One, similar to amylopsin, aids in converting the starches to sugar. The other (invertase) converts cane sugar and other forms of sugar to grape sugar so that all sugars and starches are finally absorbed as grape sugar. Summary. 1. The food enters the intestine as an acid mixture called chyme. 2. It is mixed in the intestine with three juices, bile, pan- creatic juice, and intestinal juice. 3. The three juices transform the acid chyme into an alkaline mixture called chyle. 4. In this alkaline medium the various enzymes of the three juices transform proteids and albuminoids to peptone; fats to soap and emulsions; starches and sugars, to grape sugar. 5. The action of the enzymes is aided by the mechanical action of the muscle walls. 6. The resulting mixture is then ready for absorption. Digestion of the Large Intestine. That part of the alimentary tract included under the name of the large intestine is subdivided, like the small intestine, into regions. These regions are termed the caecum, the colon, the sigmoid flexure, and the rectum. The caecum is the name applied to the saclike pouch into which the small intestine opens at the top by an orifice, guarded by a valve called the ileocolic valve. AtACTION OF THE LARGE INTESTINE 129 the bottom of this etecum is a wormlike extension a few inches in length called the vermiform appendix. Inflamma- tion of this appendix produces the disease known as appendicitis. The appendix has no known function, and is supposed to represent a remnant left in man’s develop- ment. From the point where the small intestine enters the caecum the intestine extends upward, along the right side of the abdomen, across the cavity just below the stomach and down the other side. These portions of the intestine have the greatest diameter (2J inches) of any part of the intestine and are called the ascending, trans- verse, and descending colons according to position. At the base of the descending colon is an S-shaped bend called the sigmoid flexure, and from this a straight portion (the rectum) proceeds to the external opening or anus. The large intestine shows the same number of coats as the small intestine. The muscular layer, however, is not uniformly developed but arranged in bands separated by spaces where it is wanting. This arrangement gives the tube a puckered appearance. The mucous coat has no valvulae conniventes but is filled with closely set simple glands similar to the crypts of Lieberkiihn. Action of the large intestine.— By contractions of the small intestine its contents are forced through the ileo- colic valve into the large intestine. Before these con- tents reach the large intestine, however, nearly all the nutrient part has been digested and absorbed, and what passes through the ileocolic valve consists mainly of indi- gestible material called feces. These faeces consist mainly of water, cellulose, elastic tissue, and mucin. With these substances may be present some starch and fat. If this EDDY. PHYS. —9130 DIGESTION is the case the digestion and absorption is continued in the large intestine until finally the residue is collected in the rectum and then passed out of the body. There are no distinctive enzymes secreted by the glands of the large intestine, and what little digestion goes on there is due either to secretions which pass through from the small intestine or to bacteria. These bacteria induce fermentation, and cause the putrefaction of certain parts of the faeces. The principal function of the large intestine, therefore, is the removal of indigestible material from the body. In case of over-feeding the faeces will contain more or less nutrient matter which the small intestine was unable to digest and absorb. Alcohol and Digestion. Alcohol, in small amounts, stimulates the flow of gastric juice and saliva, and this increased flow is accompanied by an increase in the enzyme and hydrochloric acid con- tent of these juices. Furthermore, whep taken in small quantities, it is itself very quickly absorbed by the blood vessels. Hence, the amount which remains in the stomach at any given time is entirely insufficient to interfere with the action of the digestive fluid upon the nutrients. From these statements, then, it would appear that digestion is aided by the use of small quantities of alcohol. Wherein lies the danger? Aside from the danger of habit forming, which is great and in itself sufficient reason for avoidance of the use of alcohol, the fact that this flow of juices is due to unnatural stimulation is the dangerous feature of its use. Natural foods stimulate the flow only to the extentALCOHOL AND DIGESTION 131 of a normal production of fluid. By normal production we mean a flow sufficient to use up the materials out of which the juices are made at a rate not faster than the body can naturally supply them for the purpose. Since the supply of this material remains fairly constant in amount it follows that any unnatural stimulation will cause this material to be used up faster than it can be sup- plied, and periods of increased flow must, therefore, be followed by periods of decreased flow. While tire occa- sional use of alcohol, therefore, may, as in cases of sickness, be of advantage, its continued use results ultimately in decrease in supply of fluids. Furthermore, the action of a stimulant is such that the secretive cells soon fail to respond to a slight stimula- tion, and hence the amount taken must be continually increased or fail of effect, thus leading to such increase as seriously to affect other parts of the body, such as the nerve centers. Finally, in large amounts, the effect of alcohol is to retard seriously the action of the fluids upon the foods, and to injure the digestive organs. While, at first, small amounts of alcohol may aid digestion, they soon fail of effect, and tend to encourage habits of excess which retard digestive processes and injure the body. Alcoholic beverages also often contain other substances which counteract the advantage of increased flow. Wines, for example, contain acids, and while the alcohol may stimulate the flow of saliva, this increased flow is of no value since ptyalin cannot act in an acid medium, and simply drains the body of valuable fluid. Since the alcohol is mainly absorbed directly from the stomach its effect upon the intestinal secretives is practically nil, and may be neglected as a factor in digestive processes.TABULAR SUMMARY OF DIGESTION. (See Ex. XL.) to IS' utrient Region of Alimentary- Tract, Where Digested Digestive Fluid Enzyme Digested Product Starch Mouth (very little) Sm. intestine Sm. intestine Saliva Pancreatic juice Succus entericus Ptyalin Amylopsin Amyloptic enzyme Grape sugar Grape sugar Grape sugar Proteid Stomach Sm. intestine Gastric juice Pancreatic juice Pepsin and HCl 1 Trypsin Peptone Peptone Albuminoids Sm. intestine Pancreatic juice Trypsin Peptone Sugars other than Grape Sm. intestine Succus entericus Invertase Grape sugar Pats Sm. intestine Pancreatic juice Steapsin Soap or emulsion Soluble salts Mouth, stomach, and intestine Water of digestive juices Salts in solution Insoluble salts Stomach HCl of gastric juice Soluble salt Water Requires no digestive action 1 HC1 — hydrochloric acid. DIGESTIONIX. ABSORPTION. In the processes of digestion, food is broken up and transformed in various ways into soluble substances, such as peptone, sugar, etc. This food, however, is still in the alimentary canal, and far removed from the cells of the body which it is to repair and build up. In this respect, then, the alimentary canal is like a big factory whose whole action is centered in the preparation of food products. A delivery system is necessary for factory products to be made available for use, and, likewise, in the body, a delivery system is necessary to distribute to all parts of the body the food prepared by the alimentary canal. In the body such a system is found in the blood system, and it is evident that the first step necessary to distribution of food is the transfer of the digested food matter to the blood. In the business world such a trans- fer of goods to a delivery system is called shipping. In the body the processes which are concerned in transfer of digested food to the blood are all included under the name of absorption. The object of shipping and absorp- tion is identical, namely, to get prepared products into circulation. Relation of the blood vessels to the alimentary tract. — The transfer of food to the blood may take place in one of two ways. It may pass directly into the blood vessels which line the walls of the intestine and stomach, or it may enter another set of tubes called the lymphatics. These lymphatics are closed tubes like the blood vessels, 133134 ABSORPTION and the food which they collect from the intestine they convey, by a common duct, to a large vein at the base of the neck, and place in the blood circulation at that point. In either case the ultimate disposal of the food is the same, namely, the putting it into the blood. The knowledge of the relation of these collecting blood vessels and lymphatics to the alimentary canal is important to the understanding of the method of transfer. In the study of the structure of the stomach and intes- tine it was noted that beneath the lining of the mucous membrane and separating it from the muscle layers was a coat of tissue called the submucous layer. Further, it was noted that blood vessels entered this layer through a fold of the peritoneum and within the layer spread out over the mucous lining in a many branched network of fine tubes or capillaries. These tiny, thin walled capillaries, which are thus in contact with the mucous membrane in all parts of the intestine and stomach, are the direct receivers of the digested food of these tracts. These capillaries are all connected and finally fuse into a single large vein called the portal vein. Again, we may recall that the lining of the small intes- tine was not smooth, except in appearance, and that the surface of the mucous lining was raised in tiny projections called villi. (See p. 120.) A lengthwise section of one of these villi (see Fig. 46) shows the following structure: Each villus is seen to be a tiny conical projection of the mucous membrane (one fiftieth to one eighth of an inch in length). The outside, or surface exposed to the digestive fluids, is composed of a single layer of mucous membrane cells, while the core of the villus is a loose network of blood capillaries and connective tissue cells, together withABSORBENT VESSELS 135 one or more larger tubes or lymph capillaries. Like the blood vessels, these latter are branches of larger vessels in the submucous layer; and fluid which once enters the lymph capillaries of the villus flows first into these larger lymph vessels, while these in turn are col- lected and fuse into a single tube about the diameter of a goose quill, called the thoracic duct. This duct passes up in front of the spinal column and finally opens into a large vein in the neck under the left collar bone. Owing to the milky color of the fluid which is in these vessels and the absence of blood they have received the name of the ladeals, and this name also serves to distinguish them from other parts of the lymphatic sys- tem. The blood capilla- ries as already mentioned are collected together and finally pour their contents into the portal vein. From this descrip- tion it must be clear that the digested food of the stomach and intestine in order to enter the blood and lymph must pass through two very thin layers, namely, the single layer of mucous cells, wrhich form the lining of the tracts, and Fig. 46 — Section of a villus; a, layer of mucous membrane cells which absorb food and carry it to the tubes within ; &, an artery ; c, blood capillaries ; d, a lacteal.136 ABSORPTION the walls of the blood and lymph vessels themselves. The manner in which this is accomplished involves a discus- sion of certain peculiar physical processes known as osmosis and dialysis. Osmosis and dialysis.— Tie a thin membrane, such as parchment or the skin which a butcher uses to hold sau- sage meat, tightly about the base of a student lamp chim- ney. Then, into the top of the chimney fit a cork with an eighth of an inch glass tube passing through a hole in its center. Before fitting in the cork and tube, fill the base of the chimney with water in which has been dissolved some colored salt, such as potassium bichromate. Then insert the cork and tube. If the parch- ment is tightly tied there will be no leakage. Finally, suspend this whole apparatus in a jar of clear water so that the liquid in the jar and the chimney is at the same level. If these direc- tions are carefully followed, the result will be an apparatus such as is pictured in Fig. 47, and the liquid in the chimney will have a reddish-yellow color. (See Ex. XXIX.) If we let this apparatus stand for a few hours, at the end of that time we shall Fig. 47—Dialyzer. be able to observe two curious results. First, the level of the liquid in the chimney will be higher than that in the jar; second, the clear water in the jar will have become colored like that in the chimney. Now it is evident that what has happened is not due toOSMOSIS AND DIALYSIS 137 ordinary leakage from the chimney, for while such explana- tion might account for the similarity in color of the two liquids, it would not account for the rise in liquid in the chimney. What has happened is as follows: In some way, the soluble coloring matter in the chimney has passed through the membrane into the jar of clear water, while some of the water in the jar has passed through the membrane and into the chimney as evidenced by the higher level in the chimney. This tendency of two liquids to mix by passage through a separating membrane is called osmosis and the experiment just described is an example of osmosis. If we continue the experiment and substitute for the potassium bichromate solution other solutions, such as grape sugar, white of egg, starch paste, peptone, etc., we shall obtain varying results. When we test the water in the jar we find that while grape sugar, peptone, and salt solutions pass readily through the membrane into the jar, white of egg and starch paste do not pass through it at all. In other words, peptone, grape sugar, and salt solutions are substances which mix with water by osmosis, while starch paste and albumin do not. Soluble substances which mix by osmosis are called crystalloids; those which do not are called colloids. Grape sugar, salt solution, potassium bichromate solution, and peptone are therefore crystalloids; starch paste and albumin are colloids. Fur- thermore, observations of the height to which the water rises in the chimney in each of the above cases shows that the rate of exchange is unequal, the water in each case pass- ing through more rapidly than the crystalloid solution, and experiment has shown that this rate of passage is deter- mined by the character of the crystalloid employed. This138 ABSORPTION relation of the crystalloid solution to the rate of exchange is spoken of as the osmotic pressure of the crystalloid. To say that potassium bichromate has a greater osmotic pres- sure than peptone means simply that the rate of flow of watqr through a membrane toward the potassium bichro- mate solution is greater than it is toward peptone solution. This process of osmosis enables us to separate a crystalloid from a colloid by putting the mixture in a membrane bag and suspending the bag in water. Under those conditions the crystalloid solution will pass through the membrane while the colloid will not. Separation of a crystalloid from a colloid by osmosis is called dialysis. Relation of osmosis to absorption in the body. — If, now, we refer to the conditions in the intestine we shall note that the relation of the digested food to the fluid in the lymph capillaries and blood capillaries fulfils exactly the condi- tions for osmosis and dialysis. In other words, the mucous membrane and capillary walls furnish the thin membrane separating two fluids, and by dialysis and osmosis the solu- ble crystalloid substances in the intestine are separated from the colloids and pass into the blood. In this explanation of the relation of digestion to dial- ysis is seen a new reason for the action of the various enzymes upon foods. In other words, they not only render them soluble but at the same time transform them into crystalloids in order to make possible this action of dialysis. While the ordinary process of osmosis accounts for the passing of crystalloid material from the alimentary tract to the lymph and blood, it must be admitted that it does not fully explain the disposal of proteid matter. For example, if peptone were a simple crystalloid sub-WHERE NUTRIENTS ARE ABSORBED 139 stance like grape sugar we should expect to find it in blood. The fact remains, however, that while we know that peptone leaves the alimentary tract as peptone it is not only never found in the blood as 'peptone, but when injected into the blood it acts as a poison. Now, since the only thing that intervenes between the alimentary tract and the blood and lymph are the cells which com- pose the membranes (mucous membrane and capillary wall cells), it must follow that in passing through these cells the protoplasm in them acts upon the peptone and converts it into another and harmless form of proteid and delivers it in this form to the blood. What the exact nature of this protoplasmic action is, is still unknown. We may, however, sum up the facts known as to transfer of foods as follows: Summary. Digestion by transforming foods to crystal- loids makes possible their transfer into the blood and lymph. This passage may or may not be accompanied by changes in the character of the substances in solution, according as the protoplasm does or does not act upon them in transit. Before leaving this subject, however, it must be stated that since the membranes concerned are made up of separate cells, there may be some seeping of liquid through the spaces between the cells. Such pas- sage of liquids is called filtration, and is not to be con- fused with osmosis. Since these spaces are exceedingly minute, it is safe to say that the osmotic exchange accounts for the transfer of the major part of the food. Where nutrients are absorbed.— Proteid. While some of the proteid is changed to peptone in the stomach, the amount of peptone absorbed by the blood vessels of the stomach is relatively small, as most of the peptone is140 ABSORPTION carried into the small intestine with the chyme. In the small intestine, therefore, is found the main seat of proteid absorption. A little may pass into the large intestine and be absorbed there, but this represents a very small amount of the whole, less in fact than in the stomach under ordi- nary conditions. Wherever it is absorbed it passes into the blood and never into the lymphatics or lacteal system. The reason for this selection is another of the facts not yet explained. In passing through the mucous mem- brane cells, as already mentioned, the protoplasm of those cells breaks down the peptone into a simpler form of proteid and delivers it in that form to the blood. While this action of the protoplasm is not yet fully explained, indications are that the change takes place in the mucous membrane cells, and is due to an enzyme present in the protoplasm. At present we know nothing of what this enzyme is or its composition. In view of the poisonous action of peptone upon the blood and the system, its importance is evident. Starch and sugar. By the time digestion is completed in the small intestine, starches and sugars have all been changed to grape sugar, and this substance passes into the blood without change. While some may be absorbed in the stomach and large intestine, the main seat of absorption is the small intestine, as in the case of proteid. Like proteid, grape sugar enters the blood, and not the lacteals. Fats. Fats are absorbed either as emulsions or soaps, and instead of entering the blood they are the only form of nutrient that passes into the lacteals. In passing through the mucous cells of the villi the soaps are changed back into emulsions so that the fat which enters the lac-WHERE NUTRIENTS ARE ABSORBED 141 teals is all in the form of an emulsion. It is the milky character of this emulsion which accounts for the appear- ance and name of this branch of the lymphatics. The seat of absorption of fats is the villi of the small intestine, and the lacteals, which receive it finally, put it into the blood circulation at the opening of the thoracic duct into the left jugular vein. (See Fig. 4S.) Water and salts. These nutrients are absorbed directly into the blood mainly in the small and large intestine. The Fig. 48 — Diagram of absorption, showing Splits and tllG water pass how food reaches the blood circulation. through by osmosis owing to the osmotic pressure of salts contained in the blood. In the case of water, the long time required by the food to pass through the large intestine permits the great bulk of water to become absorbed in this tract. This is further evidenced by the fact that while the chyle which enters the large intestine is very thin and watery, the matter that is passed out of the rectum is mainly solid. Very little water or salt is absorbed in the stomach; Alcohol. Alcohol is very quickly and completely absorbed by the blood vessels of the stomach, very little ever pass- ing into the intestine. This rapid absorption in the stomach accounts for the quick effect of this substance upon the body. It also has its beneficial side in that, as a solvent of certain drugs, it enables them to be put into circulation quickly.142 ABSORPTION Summary of nutrient absorption. — In general, the seat of greatest digestion is the seat of greatest absorption, namely, the small intestine; the large intestine blood vessels taking up only water and such food as may have escaped absorption in the small intestine. Water, salts, grape sugar, and proteid enter the blood capillaries directly and, except in the case of the proteid, without change. All these nutrients are thus conveyed to a large vein called the portal vein which carries them to the liver. Fats enter the jugular vein by way of the lymph capillaries and thoracic duct (the lacteals) in the form of an emulsion. They do not, therefore, enter the liver like the other nutrients. The nutrients which the portal vein carries to the liver are in the form of water, dissolved salts, blood proteid, and grape sugar. Action of the liver as an absorbent organ.— The nutri- ents brought to the liver by the portal vein undergo still further changes before entering that part of the blood circulation which supplies the tissues with food. The most important of these changes is the transformation of grape sugar to a starch or glycogen (sometimes called liver starch). Another change which takes place in the liver is the elimination of poisonous compounds which may be absorbed with the foods, and which, if permitted to circulate to the tissues, would result in disease. Glycogen.— The cells of the liver take up the grape sugar brought to it by the portal vein, which itself divides in the liver into many tiny capillaries. This grape sugar the liver changes into a carbohydrate of the same compo- sition as vegetable starch, hence the name liver starch. This substance, like vegetable starch, may be changed into grape sugar again by the action of ptyalin and other diastasic enzymes. Acids also transform it directly into grape sugar. It may be distinguished from starch by its action on iodine solutions to which it gives a wine-OTHER SOURCES OE GLYCOGEN 143 red color instead of the blue color of ordinary starch. The manner in which this glycogen is formed from the grape sugar of the portal vein is explained as follows: The protoplasm of the liver cells causes the sugar to lose water, and the result is a chemical change resulting in glycogen. It has been demonstrated that blood which contains over a certain proportion of grape sugar throws off this excess through the kidneys, and it is thus lost to the body as a nutrient. To avoid this loss the liver cells take up this sugar and convert it into glycogen, which they store in the body of the cell and feed out to the tissue- supply system as it is needed. Before they feed it out they reconvert it into grape sugar, and thus the amount of sugar contained in the blood which supplies the tissues never exceeds a certain proportion (0.1% to 0.2%). The liver, therefore, acts as a storehouse of the carbohydrate or energy supplying nutrient of the body. How it changes the glycogen back to sugar is not fully known, but it is supposed that the liver cells contain an enzyme similar to diastase, which performs this action, as needed. When we consider that the carbohydrates furnish the main part of the energy of the body, this regulative action of the liver becomes very important to the activity of the body. Other Sources of Glycogen. Experiments have demonstrated that, to some extent, glycogen may be formed also from proteids. For example, when a person suffering from diabetes is fed upon purely proteid foods, all carbo- hydrates being excluded from his diet, sugar is still formed. This can be explained only on the assumption that it is manu- factured from proteid. This manufacture is believed to take144 ABSORPTION place when the peptone passes through the mucous cells; that there a certain per cent of the proteid is split off as a sugar. If that is the case, it is evident that this sugar may be transformed in the liver to glycogen, and hence proteid is to a certain extent a glycogen former. Some evidence seems to indicate that fat may also be changed to glycogen, but this has' not been satisfactorily proved at the present time. Glycogen is formed in the muscles and in other parts of the body than the liver. In all these cases, it is made from the grape sugar in the blood and acts as a local supply of reserve energy. The great storehouse, however, is the liver. Poisons.— In the intestines are found many forms of bacteria. These in turn may bring about fermentation of the food, and in this way are produced certain injurious substances or poisons which may be absorbed with the digested nutrients. In that case they are brought in the blood of the portal vein to the liver, and one of the important functions of this organ is to destroy these poisons or return them to the intestine with the bile. If, however, any interruption of the bile making of the liver occurs, it loses its power to destroy and give off these poisons, and the result is that they pass into the tissue-supply blood, and poison the body. The results of the entry of these poisons into the system usually manifest themselves in a coated tongue, loss of appetite, headache, and dullness, and such a condition is called biliousness. Certain drugs have the power to stimulate the liver cells and cause a great outpouring of bile which carries with it the poisons, and thus restores the liver to its normal action and sometimes removes the source of trouble. Defaecation.— The removal of the feces from the large intestine results not only in the removal from the body of indigestible material, but also has a vital connection with the poison removing power of the liver. When the bile formed in the liver is poured into the intestine by the gall duct it carries with it the poisons which the liver cells have separated from the blood of the portal vein. If these poi- sons are allowed to remain in the small intestine they willHVGIENE OF DIGESTION AND ABSORPTION 145 again be absorbed, taken to the liver and so on indefinitely, or until the power of removal of the liver is exhausted. The only way in which this can be avoided is for the bile which contains them to be carried into the large intestine and out with the feces. It is this fact that makes regular operations of the bowels necessary and makes constipation dangerous. In severe cases of constipation stimulants which increase peristalsis, and thus induce defecation, are often given, but it must be borne in mind that all such rem- edies are purely stimulants and should be resorted to only under the advice of a physician. Regularity and care in diet will make their use unnecessary in most cases. Hygiene of Digestion and Absorption. The body requires the digestion and absorption of a cer- tain amount of each kind of nutrient daily. (See p. 70.) If the amount of food taken by a person in a day is only a little in excess of what is required, this excess is readily taken care of by the action of the bowels. If, however, too much is taken, the intestines become clogged and over- worked, the food is not properly digested, poisons are de- veloped and not removed, and all the evils of biliousness and indigestion follow. For this reason a balanced, moderate diet, taken at regular intervals, becomes abso- lutely necessary to a satisfactory operation of the digestive and absorbent system. Americans are especially prone to over-eat, and many of the ills of our nation have their source in this intemperance in eating. EDDY. PHYS. — 10.X. DIGESTION IN THE LOWER ANIMALS. The process of digestion (as a preliminary step in the changing of food into living matter or protoplasm) is to be met with wherever we find living cells. In man the system has reached its most complex development, but if we examine the various groups of animals, we find that in every group the process of changing food into soluble sub- stances by the aid of enzymes is quite as important and necessary a step in metabolism as it is in man, though the organs which perform this action may be much simpler of structure. It is interesting, then, to study the methods of digestion in the lower forms, in a comparative way, from the light they throw upon the development of the complex human system. In the pages that follow a few types have been selected to bring out the manner in which the human sys- tem has been developed from lower types. Digestion in one-celled animals.— In animals like the amoeba and paramcecium (one-celled animals) the food is taken into the cell in the form of solid particles. In this respect the protozoans differ from all higher animals. Digestion proper, therefore, in these animals occurs in the cytoplasm of the cells themselves, and to distinguish it from that of other forms is called intracellular digestion. 1 Chapters X, XIV, XVII, XX, XXII, XXIV, XXVII, and XXXI, are intended for study in connection with a course in zoology. In con- nection with laboratory work in zoology they furnish necessary data for comparison. When not preceded by such laboratory work they should be omitted. 146DIGESTION OF THE AMCEBA 147 While the methods of food taking of one-celled animals differs greatly, some engulfing the food like the amoeba, others having a distinct opening or mouth for the purpose, and rows of cilia to direct the water currents toward the mouth, the digestion proper is practically similar in method in all. The amoeba will serve for a type. Digestion of the amoeba. —With the ingested food is always taken in some water which collects about the food and forms a food bubble (food vacuole) in the cytoplasm. Soon the cytoplasm around the food vacuole begins to pour into it a secretion which contains hydrochloric acid and probably an enzyme. This secretion is therefore a sort of gastric juice. At the same time the vacuole is kept in motion by the streaming of the cytoplasm. Under the combined actions of this secretion and movement, the solid particle is broken up and dissolved and is then ready to be assimilated directly by the cytoplasm. In this way new protoplasm is built up, and the animal increases in size and strength. In certain one-celled formsthe food vacuoles move in a per- fectly definite path (see Fig. 49) through the cy- toplasm, and this move- ment probably serves to distribute the digested food uniformly, as well as to break it up. In such cases the resemblance to a primitive circulatory system is evident. Another way in which these one-celled forms differ is in the different nutrients which they are able to dissolve. F.V. F.v. Fig. 49 —Path of food particles in a para- mcecium ; JST, nucleus ; M, mouth ; F. V., food vacuoles. Note the decrease in size of the food vacuoles as the food is digested.148 DIGESTION IN THE LOWER ANIMALS The amoeba, for example, cannot digest cither starch or fats, while other forms have no difficulty in doing so. This is accounted for on the supposition that the amoeba can produce no starch or fat dissolving enzymes, while the other forms do produce them. Digestion in multicellular animals. — As an animal in- creases in number of cells it becomes manifestly more difficult for the food to reach the cells. In such animals we should expect to find various adaptations to remedy this difficulty. Hydra and sponges. In the little animal known as the hydra, and in sponges, the cells which form the animal are arranged in layers about a central cavity. (See Fig. 50 ) Fig. 50— Longitudinal section of a Hydra; 6, bud which will form a younghydra; ba, base by which it is attached when not creeping ; ia, mouth ; ov, ovary with an egg > sp, sperm ary. Idie cells lining this cavity are provided with hairlike projections, or cilia, which are in constant motion, and by their motion create a current which draws the waterDIGESTION IN MULTICELLULAR ANIMALS 149 into the cavity and causes it to circle about, thus bring- ing the food which it may contain in contact with all the cells which line the cavity. Many of these cells give off digestive secretions which dissolve the food matter con- tained in the cavity, and thus prepare it for absorption. Such cells represent simple glands, and since the solution takes place outside of the cells it is called extra-cellular digestion, and the cavity becomes a sort of digestive cavity or tract. All animals except the protozoans have extra-cellular digestion. In the forms mentioned (hydra and sponge) the waste is thrown out of the lining cells into the same cavity from which they absorb the food and is removed by the outgoing currents of water. Worms. In these forms we have for the first time a definite alimentary tract or canal. In some, this canal Fig. 51 — Diagram of a longitudinal section of an earthworm, showing the central digestive tube. is open only at one end, while in others it forms a con- tinuous tube running the length of the body with a true mouth and anus. Such a continuous tube is found in the earthworm. In this animal the slitlike mouth at the front of the body opens into an expanded, muscular walled cavity just back of it called the pharynx. By the action of these muscular walls the food is sucked into the cavity through the mouth, after it has been150 DIGESTION IN THE LOWER ANIMALS broken up by the lip. From this pharynx, the food is forced backward by true peristaltic movements of layers of muscles in the walls of the alimentary canal until it reaches the anal opening. In traversing this canal it passes through several modified regions where it is acted Fig, 52 — Cross section of earthworm, showing central intestine (1) and the dorsal fold or typhlosole (T); D, dorsal bloodvessel; K, ventral bloodvessel; B. C.t body cavity. upon by special apparatus or secretions, and in this way the digestible material dissolved out and absorbed through the walls of the canal into the body cavity of the animal, where a fluid similar to the blood in our bodies receives it and distributes it to all parts of the body. In the earthworm the regional division of this tract is of par-DIGESTION IN MULTICELLULAR ANIMALS 151 ticular interest in showing how the needs of the animal have brought about adaptations of a straight tube to digestive purposes. For example, the widening at the pharynx is a modification to suit the earth burrowing habits of the animal and permit the taking in of food of a solid character. The saclike enlargement back of the pharynx, called the crop, permits the food to be stored for a time in the tract, and enables the animal to eat more at one time than he can digest, and thus give the parts a chance for rest. The thick walled portion back of the crop, called the gizzard, is merely a portion whose muscle layers are thickened to form a special grinding organ for macerating the food, since the animal possesses no teeth. Furthermore, all these modifications are made in a tube of essentially the same general structure as our own tracts, since it has an inner layer of secreting cells, a middle muscle layer, and just outside it is a blood fluid to receive the absorbed food, as in our own body. In fact we might almost say that the earthworm’s digestive tract is a living model of what a simple human tract might have been before it was modified into stomach glands, etc. In the earthworm, therefore, we have a distinct advance over the hydra and sponge in that certain cells have been set aside to make a tube in which to digest the food for the whole body. Further, the manner in which this tube is constructed and modified indicates the way in which the complex system of glands, cavities, and organs of the higher animals could have been developed from such a simple foundation to fit the needs of the animal. As we pass from the earthworm to the higher animals the in- creased amounts of food necessary to furnish a much152 DIGESTION IN THE LOWER ANIMALS larger number of body cells, the character of the food required by these cells, the opportunities for getting food, and many other factors combine to produce these greater modifications of structure. The nature of these adapta- Fig. 53 — Section of a fish ; st, stomach ; i, intestine ; <7, gills. Davison, Zoology. tions to increased needs, however, is not different in kind from that indicated in the earthworm. A few types will illustrate the direction which these modifica- tions have taken in different animals. Modifications in the digestive tube of the fish.— In this animal, the skin that lines the mouth has produced out- growths of a horny character called teeth, which prevent the food from slipping backward when seized. These are not like our teeth in that they are not set in sockets, and are found all over the surface of the mouth. In the mouth also, is found a fleshy motionless projection called the tongue, which has in it nerves of taste. It has no movement and does not aid in swallowing like ours. Back of the mouth is a widened space or pharynx, in the sides of which four slits are developed for letting in water to the breathing organs or gills. From the pharynx theMODIFICATIONS IN THE FROG 158 food passes backward into a U-shaped portion or stomach. From this stomach the remainder of the tube or intestine passes at first forward and then turns and extends straight backward to the anus. Opening into this forward tube are a number of blind tubes called pyloric creca. There is also a liver with a large gall bladder connected to this part of the intestine by a duct. The cieca and liver secrete digestive fluids which they pour into the alimentary tube. The principal advance of this form of tract over that of the worm is seen in the two devices employed to increase the digestive surface, namely, by the folding and by the extension of the walls at points to form external glands. Other adaptations to the new needs are seen in the devel- opment of a tongue and teeth in the mouth, and in the increase in size of the stomach portion. Modifications in the frog.— The most striking adapta- tions in the digestive tract of the frog are the mouth structures, the increased coiling of the intestine, and the division of the tube into the stomach, small and large intestine. In addition to the liver the frog also possesses a pancreas. The mouth of the frog contains no teeth with sockets, but there is a single series of projections around the upper jaw which serve, as those of the fish do, to prevent the escape of the food when swallowing. In the frog, the tongue, unlike that of the fish, is movable, and plays a distinct part in swallowing. Unlike ours it is fastened at the front with the tip pointing down the throat. The end is sticky and can be extended to catch insects, which it snaps back into the throat. The large intestine ends in an enlarged space called a cloaca into which open also the egg or sperm ducts and the154 DIGESTION IN THE LOWER ANIMALS urinary tubes. The anus by which this cloaca opens in this animal, therefore, is the place of discharge of other matter besides indigestible food. The advance of this animal over the fish is seen in the increase of digestive surface Fig. 54 — Section of frog; t, tongue ; l. mouth; st, stomach ; p, pancreas : d. duo- denum ; si, small intestine ; r, rectum, and b, bladder, opening into a common cavity, the cloaca; ur, ureter, and ovd, oviduct, also opening into the cloaca; k, kidney ; or, ovary ; lie, liver. Davison, Zoology. by coiling, the placing of the teeth in relation to the jaws, and the greater tendency to regional arrangement. Adaptations in warm-blooded animals.—- In these animals (with the exception of birds) are found for the first time, true teeth in sockets. All such teeth may be divided into four classes (incisors, canines, premolars, and molars) as in man, and are placed uniformly upon the jaws. The kind which predominates in any animal is determined by the kind of food which it eats. Thus, squirrels and rats have the incisors greatly developed for gnawing, while the flesh caters are marked by the predomi- nance of canines, and the grass eaters by the molars.ADAPTATIONS IN WAEM-BLOODED ANIMALS 155 In birds the teeth are replaced by horny sheaths, the mastication of the food being provided for by a thick walled portion of the digestive tract called the gizzard. Fig. 55 — Skulls, showing teeth ; A, monkey ; B, horse, a grass eater (note number of molars); <7, dog, a flesh eater (note the prominent canines) ; Z), rodent, «* gnawer (note the prominent incisors). While the other parts of the digestive tract are, in general, similar to that in man, the stomachs of the various forms show interesting modifications. The stomach of the rabbit and other rodents, and of the carnivora, is almost identical with that of man. The birds show a curious adaptation to compensate for the lack of teeth and the uncertainty of food supply. In these animals the stom- ach is in three parts. First, at the base of the gullet is a sac called the crop in which the food may be stored156 DIGESTION IN THE LOWEK ANIMALS until needed by the body. By means of this organ the bird is enabled to eat enough at one meal to last for some time. Succeeding this is a compart- ment called the giz- zard. When food is needed it is passed from the crop into this gizzard which has an outer wall of thick muscles and an inner horny lining. Inside it, too, are stones and other hard objects which the bird swal- lows for the purpose of grinding the food thoroughly to pre- pare it for the ac- tion of the digestive fluids. When thor- oughly ground up, the food passes into the third compart- ment, or true diges- tive stomach, where the gastric juice is secreted. The ruminants or cud-chewing animals show still another curious modification. In these animals, such as the cow, the stomach is divided into four compart- Fig. 56 — Digestive tract of a bird; a, esophagus ; b, crop ; c, proventriculus ; cl, gizzard ; e, liver; /, gall bladder ; y, pancreas ; h, duodenum ; i, small intestine; k, caeca; m, ureter; w, oviduct ; o, cloaca.ADAPTATIONS IN WARM-BLOODED ANIMALS 157 ments. The first and largest is called the rumen, and into this the grass mixed with saliva passes without mastication as it is eaten. From this rumen the food is passed into a second compartment with a honeycombed wall and called the reticulum. After the grazing is over the food is returned to the mouth from this reticulum in the form of boluses or. round balls called the cud. This cud is then thor- oughly masticated by the molars and finally swallowed again. This time it passes directly into a third compartment called the psalterium, which is ridged, and strains the food into the fourth compartment, the abomasum or rennet stomach. In this abomasum the actual digestion takes place, and here is secreted the gas- tric juice. Such an arrangement as this is very evidently an adaptation to compensate for the greater difficulty of digestion of such food as grass and hay; but it is to be noted that in none of these cases is the purpose and general method of digestion essentially different. In all, the various structures are working to one common end, namely, the transforming of food into soluble matter. And the ultimate method employed is the action of fer- ments or enzjunes secreted by cells located in the walls or connected with the digestive tract. Fig. 57 — Stomach of a sheep (ruminant); e, esophagus ; Ru, rumen ; R, reticulum ; Ps, psalterium ; A, abomasum ; P, pylorus; P, duodenum.158 DIGESTION IN THE LOWEK ANIMALS In all the warm-blooded animals the intestines are much longer and more coiled than in the cold-blooded forms, and the time required for the digestion of vegetable matter is responsible for the greater length of the intes- tines in the herbivorous animals as compared with the carnivora. In conclusion, we may note that all modifications of digestive methods are merely adaptations to meet special conditions. These adaptations may take the direction of specialized structures for producing digestive fluids or preparation of food for the action of fluids, or they may be directed toward increase in surface and the insurance of more complete digestion. In every case, however, the transformation of food into soluble matter as a prelimi- nary to absorption is the end sought, and the ultimate chemical means is the action of special compounds called enzymes secreted by specialized cells.XI. BLOOD ANJ) LYMPH. After digestion and absorption the food is ready for the use of the cells. Absorption has placed this food in the blood for delivery to the cells, and it is in its character as a carrier of digested nutrients and other substances to the cells that we are to investigate the functions and structure of the blood. Blood vessels. —The blood in our bodies is contained in a set of tubes called blood vessels, and these vessels are so constructed and connected as to form a completely closed system. In other words, while the blood absorbs matter into itself and gives out matter from itself to the cells of the body, this exchange takes place through the walls of the blood vessels, and never, except in case of accident (bleeding), does the blood (as blood) escape from the system of closed tubes. In this respect it is like a train on a circular track. We may load goods into this train and .take goods from it, but the train itself keeps on continually and, except in case of accident, never leaves the track. In our body the blood is kept in con- stant movement within the tubes by the beating of the heart and for this reason the blood vessels and their con- tents are spoken of collectively as the circulatory system. If it were not for this constant motion it would be mani- festly impossible for food taken in from the alimentary tract to reach the other parts of the body. Before con- sidering the way in which this liquid is kept in circulation we shall first consider its composition. 1591G0 BLOOD AND LYMPH Blood structure. — We may readily obtain human blood for study at any time by pricking the end of the finger with a clean needle. (See Ex.'XLII.) If we examine a little of the blood so collected on a flat surface we note that the drop is not of the same color throughout. At the edge of the drop it shows a yellowish color while the center is dark red. With the compound microscope we find that this difference in color is due to the fact that blood con- sists of two distinct parts, a yellowish liquid called the plasma, and a vast number of tiny round disks called corpuscles. The red color of the blood is due to the pres- ence of these corpuscles (red corpuscles). If we use the higher powers of the microscope we find that these red corpuscles are actually biconcave disks (see Fig. 58) Fig. 58 — Blood corpuscles ; A, red corpuscles in rouleaux ; a, a, colorless corpus- cles (X400); B, red corpuscles, highly magnified; C, view of edge; Z), three quarters view; E, red corpuscle swollen with water ; Ft O, Ht distorted red cor- puscles. massed together in bands or rouleaux. At the same time, we may be able to make out one or more colorless bodies of irregular shape floating among these bands of disks. These latter are called white corpuscles to distinguish them from the other more numerous form. They differ from the other corpuscles in having a nucleus. Both kinds of corpuscles are cells, but the red corpuscles haveSTUDY OF BLOOD 161 lost their nucleus. The presence of these cells allows us to consider the blood as a true tissue. In general, then, blood is a tissue consisting of a yellowish liquid (plasma) in which float a large number of non- nucleated cells (red corpuscles) and a few nucleated cells (white corpuscles). Properties of blood.—If we mount a drop of blood on a glass slide and allow it to stand for a few moments, at the end of that time we note two distinct changes in its appearance. First, it is darker in color, and second, its surface is covered with a film which, when pricked with a needle, shows an elastic consistency. This formation is called a clot. If we examine the needle puncture on the finger we shall find that it has ceased to bleed, and is covered with a similar clot. If we wash this clot off the finger, bleeding begins anew, and this gives us a hint of an important use for this clot formation. An examination with the microscope of the clotted blood on the slide shows that it is actually a complicated network of slender threads in which are entangled the corpuscles. Evidently these threads must have their origin in the plasma, and to understand their nature it is necessary to make a careful examination of the plasma. For this purpose a larger quantity of blood is necessary. Study of blood. — Fortunately for our purpose, the blood of all warm-blooded animals is practically identical in structure with human blood. This fact permits us to use fresh-drawn blood of a pig or cow. (See Ex. XLIII.) First, place some blood in a bottle and let it stand for a time exposed to the air. Put the rest in a basin and whip it vigorously with twigs. Pour the whipped blood into another bottle. While these two bottles are standing we EDDY. PHY3. —11162 BLOOD AND LYMPH may examine the twigs with which we did the beating. These will be covered with a stringy elastic substance which, when washed, is seen to consist of glistening white strands or fibers. These fibers are composed of a sub- stance called fibrin, and are identical in composition with those which were noted in the clot of human blood. Furthermore, if we test this fibrin with our nutrient tests, we shall find that it responds strongly to one test only, namely, that for proteid. In other words, fibrin is a coag- ulated proteid which was present in the plasma of fresh- drawn blood in a liquid state. If we examine the bottled blood after it has stood for awhile we shall find the con- tents of the bottle filled with fresh-drawn blood in a differ- ent condition from the whipped blood. The former will be found to have solidified into a clot of the same shape as the bottle, and to be floating in a small quantity of a straw-colored liquid (serum). The whipped blood will show no clot, and is in the same condition as when we put it in the bottle. If we pour off the serum from the first bottle, and then break the bottle so as to remove the clot entire, we may examine this clot and study its structure. Such examina- tion shows it to be made up of a solid mass of fibrin fibers which inclose masses of corpuscles. The corpuscles give it its color. The whipped blood fails to clot because we have removed its fibrin by whipping. From these observations we must conclude that the serum is plasma whose fibrin has been removed, and that the presence of this fibrin is necessary to the formation of a clot. Our experiments thus far have succeeded in separating only one of the constituents of blood plasma, namely, fibrin.STUDY OF BLOOD 163 If our nutrient tests are applied to the serum which was poured from the bottle containing the clot they fur- nish the following interesting results: First. They show it to be rich in proteid. Second. They indicate small amounts of fats and grape sugar. Third. They show the presence of salts. Fourth. They show that starch is absent. Fifth. The serum gives a weak alkaline reaction. These results justify the following conclusions in regard to the composition of blood plasma: First. The proteid tests show that the plasma contains other proteid matter beside the fibrin, since that was removed by the clotting, and that this proteid matter does not coagulate like fibrin when the blood is removed from the living body. Second. The small amount of grape sugar in circulation at any time is what we should expect from our study of the formation of glycogen in the liver (see p. 142), while the small amount of fat present shows that most of the fat poured into the blood by the lacteals does not enter into the general circulation but is used up by the living tissues. Third. The absence of starch is accounted for by the fact that all the starch was changed by digestion into grape sugar and enters the blood in that form. Fourth. The presence of salts explains the osmotic pres- sure of the blood, accounts for its alkaline reaction, and serves to keep certain proteids in solution. Finally, the presence of these particular nutrients shows in an indisputable way that it is the plasma of the blood which is the carrier of digested nutrients.164 BLOOD AND LYMPH While our experiments show in a general way the com- position of the blood, we must go to the chemist for a care- ful analysis of its structure. Such chemical analyses show the plasma to be composed of the following substances: COMPOSITION OF PLASMA. Plasma ' Water . fFibrinogen Proteids . \ Paraglobulin (serum globulin) C Serum albumin Extractives - (Fats ! Sugars 1 Urea [ Acids Salts f Chlorides I Carbonates ! f I Sulphates I Phosphates j Sodium Potassium Calcium | Magnesium L Iron (_ Enzymes and unknown substances Blood proteids. — Fibrinogen is fibrin in the liquid state. It is, of course, absent in the serum. It varies in amount from .22% to .40% in human blood, and to it is due the clotting power of the blood. (The causes of clotting will be discussed under a separate heading, see p. 169.) Serum albumin is the form of proteid which is supposed to have been formed from the digested proteids of the food. It forms 4.5% of the blood of man, and is supposed to be the form of proteid used in supplying the tissue cells. Paraglobulin forms 3.1% of the blood of man. Its origin and function are at present unknown. The two proteids, then, whose functions are definitely known are serum, albumin, and fibrin. Blood extractives.—The fats and sugars are evidently absorbed from the digestive fluids, and as nutrients are carried to the tissues for use as energy producers. Urea and acids are merely forms of waste matter given off byTHE SOLID MATTERS IN THE BLOOD 165 the cells to the blood, and their presence indicates another function of blood, viz., the removal of wastes. (See p. 352, Excretion.) Blood Salts. The mineral salts are probably absorbed in the same way as the other nutrients, and their presence accounts for the osmotic pressure of the blood as well as for the alkaline reaction. This last is mainly due to the presence of the salt, sodium carbon- ate. Certain proteids (notably the globulins) are kept in solution by the presence of salts. Water of blood. — This liquid is taken in by osmosis from the digestive tract and other parts of the body, and the liquid character of blood is mainly due to it. It forms 78% of the entire blood. The solid matters in the blood- —Having now a general idea as to the nature of the liquid plasma, let us turn to the solid parts. We have already noted that blood con- tains two kinds of corpuscles, the red and the white. In addition to these solids, blood contains a third set of bodies called blood plates. These three solids are only slightly heavier than the plasma, and the movement of the blood keeps them pretty thoroughly mixed with the plasma. A knowledge of their origin and function is necessary to a clear understanding of the blood functions. Red corpuscles. These bodies to which the red color of the blood is due, are shown by the microscope to be biconcave, circular disks which are usually banded to- gether in rolls like a roll of coins. (See Fig. 58.) They are very small, having an average diameter of about .0077 millimeter, and while their number varies greatly with the condition of health the average is five million to a cubic millimeter of blood in males, and four million and a half per cubic millimeters in females. When viewed singly they are seen to be of a vellowish-red color, but in masses they appear deep scarlet. This red color is due to a pigment which they contain called haemoglobin. Haemo-166 BLOOD AND LYMPH globin gives them an important function as carriers of oxygen from the lungs to the tissues, and their physiology is largely concerned with the action of this pigment. In structure they are found to consist of a colorless substance called the stroma, which* gives shape to the cor- puscles, and in this stroma is deposited the hemoglobin together with some water and salts. They contain no nucleus, and are to be considered as cells which have been modified to meet the needs of the blood. Haemoglobin. This pigment consists of two chemical compounds, haematin and a proteid body. The haematin has the power when in contact with oxygen to combine with it and to form an unstable compound which breaks down when it reaches the tissues, and gives up its oxygen to the cells. This power of haematin to unite with oxygen is due to the iron which forms a large part of its chemical composition. An interesting demonstration of the power of the red corpuscles to take up oxygen is furnished when- ever we mix blood with air. For example, shake violently a tube half filled with blood. The increased redness of the shaken mixture is due to the taking up of oxygen. In the body the red corpuscles are brought in contact with the air in the lungs, and the oxygen which they pick up is carried by them to the tissues and there given up to the cells as needed. It is evident, then, that one function of the red corpuscles is to carry oxygen to the tissues. This, however, is only one of their functions. Haemoglobin forms compounds not only with oxygen but also with carbon dioxide. This compound is called car- bohemoglobin to distinguish it from oxyhemoglobin. This compound is formed when the carbon dioxide is absorbed from the tissue cells. In this way the red corpuscles areORIGIN OF RED CORPUSCLES 167 not only enabled to carry oxygen to the tissues but also to remove part of the waste carbon dioxide from them. In summary, then, we may say that the two uses to the body of the red corpuscles are the carriage of oxygen to the tissues and the removal of carbon dioxide from them. The red corpuscles of man and of all mammals, except the camel family, are biconcave, circular disks without nuclei. In the camel family, they are elliptical in shape. They vary, however, in size, and this variation often enables experts to determine the character of blood stains in murder trials. The red corpuscles of fishes, amphibia, reptiles, and birds are oval in shape and usually nucleated. FIG. 59—Various types of red corpuscles. Origin of red corpuscles. — The fact that the human corpuscles are without nuclei shows that they are cells which are in a degen- erating state and hence cannot last long in circulation. Since168 BLOOD AND LYMPH they are so necessary to the life of man it is evident that there must be a constant renewal of the supply from some source. This source is found in the red marrow of bones. Here are produced nucleated, colorless cells, called erythroblasts, which multiply and form daughter cells. These daughter cells finally produce hsemoglobin in their cytoplasm, and thus form nucleated red corpuscles. These lose their nucleus and finally enter the blood stream in the form we have recognized as red corpuscles. The old corpuscles are continually breaking down, and after undergoing dissolution in the blood stream, the liberated haemoglobin is discharged by the liver as bile pigment. It was once thought that their destruction took place in the spleen, but there is no evidence to prove this and it is now believed that this destruction may take place anywhere in the blood stream. In a normal human being the supply and loss is so balanced as to 'maintain a fairly constant number of corpuscles in the blood. The number of red corpuscles in the blood varies not only with the sexes, as already noted (see p. 165), but also with other con- ditions. They increase greatly with the altitude, supposedly to compensate for the reduction in air supply. They often decrease in number in disease causing a condition known as anemia. At present the remedies for lack of blood corpuscles usually take the form of tonics, that is, the giving of medicines, usually rich in iron, which stimulate the formation of both erythroblasts and haemoglobin. Exercise in the open air is a far better remedy than any drug for this condition. White Corpuscles.—These bodies are not nearly so num- erous as the red corpuscles being only five thousand to seven thousand per cubic millimeter. They are colorless amceboid cells having a nucleus and no definite cell out- line, moving about in the plasma in much the same man- ner as an amoeba, which they strongly resemble. They are of two kinds: those with granular cytoplasm are more numerous and are called leucocytes ; those without granules in the cytoplasm are called lymphocytes. The leucocytesTHEORIES OE CLOTTING 169 develop from bone marrow; the origin of lymphocytes is uncertain. The leucocytes have various functions. Some, called phagocytes, absorb disease-causing bacteria as then- food, and thus protect the body from disease. Others aid in the absorption of fats and proteids from the digestive tract. They also aid in coagulation or clot forming and finally help to maintain the normal supply of proteid in the blood plasma. Blood plates.— These are cir- cular or elliptical bodies smaller than red corpuscles. They are more numerous than the white corpuscles, and have been shown to be capable of amoeboid move- ments. They disintegrate rapidly when removed from circulation, and hence little is known of their structure. Their principal function appears to be their action in clotting, which we will discuss under that head. Theories of clotting.— In the beginning of the chapter we noted that blood removed from circulation tended to solidify into a clot. We noted also that the foundation of this clot was the coagulated fibrinogen or fibrin, of the blood plasma, and that the failure of whipped blood to clot was due to the removal of this fibrinogen by whipping. These experiments, however, merely give us an explanation of what the clot is made of, namely, of fibrinogen. The fibrinogen is made to coagulate and forms fibers of solid proteid matter called fibrin, between whose strands are caught the corpuscles, and to whose shrinking is due the squeezing Fig. GO—A phagocyte (white cor- puscle), A, engulfing a bacte- rium, B; 1, 2, 3, steps in the process.170 BLOOD AND LYMPH out of the serum or defibrinated plasma. What the cause is that suddenly produces this coagulation of fibrinogen, and the conse- quent formation of fibrin fibers, the experiment failed to explain. While this question has not yet been fully answered the theory which is generally accepted at the present time may be stated as follows: The experimental facts upon which the theory is based are three. First, the coagulation of fibrin is not due to exposure to air, as it takes place with equal readiness when air is not present. Second, it occurs only when there are present calcium salts, fibrin- ogen, and an enzyme called thrombin. Third, the enzyme throm- bin is not a part of the normal composition of plasma, but is furnished under special conditions by the combined action of the blood plates, the leucocytes, and the calcium salts. The formation of the clot involves the following actions: First, the breaking down of the blood plates and leucocytes in the presence of calcium salts results in the formation of a chemical compound or enzyme called thrombin. The enzyme reaction of this thrombin causes the coagulation of fibrinogen into fibrin. The reason that this action occurs only when the blood is removed from the blood vessels is probably due to the fact that the blood plates and leucocytes do not disintegrate in sufficient numbers while the blood is in the body to produce much thrombin. It is also possible that, in blood which is inclosed in blood vessels there may be compounds among the unknown constituents of plasma which neutralize the thrombin as fast as it is formed. Conditions affecting the blood. — Since the blood fur- nishes the oxygen and food supply of the tissues, it is evident that it must be maintained in such a condition as to perform these functions. Anything, therefore, which interferes with the number of corpuscles, the presence of haemoglobin, the oxygen content of the corpuscles, or the food content of the plasma, is bound to interfere with its normal functions. The conditions which have been found to be essential to a healthy condition of bloodBLOOD TEMPERATURE 171 are an abundance of fresh air to secure sufficient oxygen supply, plenty of sleep to prevent too great a drain by the tissues upon the food and oxygen it carries, exercise to allow the proper distribution of food and oxygen to all parts of the body and the removal of waste, and nutri- tious food for diet to maintain the proper food content of the plasma. This latter condition also affects the red corpuscles as well as the plasma, as without iron it is impossible to form hremoglobin. Amount of blood in the body.—- We might expect that the constant giving up of food and oxygen by the blood would cause its quantity to vary. So far as known, however, the loss of these substances and the acquiring of wastes is so balanced that the quantity is fairly constant, and in a healthy person amounts to about one thirteenth of the total weight of the body. The amounts in different parts of the body, however, vary. Roughly, it is distributed in the following proportions:1 Heart and Lungs....................................25% Liver................ ... .........25% The Skeletal Muscles 25% The Other Organs...................................25% Blood Temperature.— Heat is constantly being pro- duced in all of the animal tissues of the body, and at the same time heat is being given off to the outside air from the surface of the body. The amounts produced and lost vary in different parts of the body. Now, the blood is passing constantly from one part of the body to the other, and hence necessarily carries warmth from the parts where greater amounts of heat are produced to places where there is less. In this way it tends to equalize the tempera- ture in all parts of the body, and the temperature of the 1 Foster, Physiology.172 BLOOD AND LYMPH blood is therefore the average temperature of the whole body. In man the production of heat and the loss to the outside of the body is so nicely balanced that the average or blood temperature remains almost constant. This being the case, any rise in this temperature indicates a more or less serious disturbance in the body. In some animals, such as the fishes, frogs, and reptiles, the loss of heat is so great compared with the production that the temperature of the blood is only slightly above that of the surrounding air or water. Such animals feel cold to the touch, and are spoken of as cold-blooded animals. They are also called variable temperature animals, since their average temperature varies with that of their surround- ings. In birds and mammals, including man, the amount of heat produced is greater and the loss so nicely balanced that their average temperature is not only kept constant, but at the same time is higher than that of their surround- ings and does not vary with the outside temperature. Such animals feel warm to the touch and are therefore called warm-blooded. In man the temperature of a healthy body remains at about 9S° Fahrenheit, and though it may increase to 103° or 105° in violent exercise, this is only temporary and it sinks in a few hours to normal. A continued temperature of over 100° is usually an indication of fever. This fact enables the physician to discover at once the presence of fever by taking a person’s temperature. In birds the temperature is much higher than in man, these animals having an average temperature of 105° Fahrenheit. Summary of blood action.— The blood is usually spoken of as the nutritive fluid of the body, since its main function is to supply food or nourishment to the tissues. It has, however, other func-SUMMARY 173 tions as we have seen, and we may summarize its functions as follows: (а) It carries to the tissues the food which has been prepared by digestion, and introduced into the plasma by absorption. (б) It carries oxygen, which it takes from the air in the lungs, to the tissues and there exchanges it for carbon dioxide which it carries away. The red corpuscles are the carriers in this case. (c) It carries from the tissues to the lungs, kidneys, and skin, the wastes of the body. Such wastes include carbon dioxide, urea, acids, etc., and are removed from the body by the organs mentioned. (d) It is the medium for the transportation of the internal secretions of certain glands. (e) It distributes the heat produced in various parts of the body and thus tends to equalize the body temperature. Blood in other animals.— Since the function of the blood is the distribution of food and oxygen, and the collection of wastes, it is evident that animals composed of a single or few7 cells wall have no need of such a fluid. In animals like the amoeba, and even in animals of more complicated cellular structure, such as the hydra, sponges, and jellyfishes, the digested food and the air are brought in direct contact with the cells, and hence in these animals there is no blood and no need of such fluid. As animals increase in complexity, some of their cells are necessarily so placed that the food is no longer brought in direct contact with them. In such animals some system of distribution becomes necessary, and hence most animals higher than those mentioned have some kind of fluid that answers the purpose of the blood in our bodies. In many this blood fluid is not inclosed in tubes and is without corpuscles. It is to be compared rather with the plasma than with the red blood of man. Crayfishes, crabs, and insects have such a fluid, and it is colorless. In the earthworm the blood fluid is red, but has no red corpuscles, the ha3moglobin being dissolved in the fluid itself. It is only when we reach the vertebrate or backboned animals that we find blood containing corpuscles specialized for the carrying of oxygen and carbon dioxide. In the lower forms of vertebrates these corpuscles are nucleated. Only in birds and mammals do we find the non-nucleated corpuscle.174 BLOOD AND LYMPH The Lymph. In the pages preceding we have attempted to make it clear that the materials required by the tissues are brought to them by the blood, and that the wastes (carbon dioxide, urea, etc.) given off by the tissues are finally conveyed by the blood to organs which remove them from the body. In the lower animals where the blood fluid is not inclosed in tubes, and bathes all the tissues, this exchange between tissue cells and blood fluid is a comparatively simple matter. In man and the higher animals the fact that the blood is contained always in closed tubes makes it less easy to see how this exchange takes place. The explanation of this exchange involves a study of another important liquid of the body known as the lymph. All the tissue elements of the body are bathed in this liquid. Furthermore, the lymph spaces are connected by a set of tubes known as the lymphatics, which permit the flow of the lymph from one part of the body to another. Origin and structure of the lymph. —Examination of the composition of the lymph contained in the lymph tubes and spaces shows that it is practically identical in struc- ture with the plasma of the blood, and in fact it is sup- posed to be plasma which has in some way passed out of the blood vessels and become a tissue bathing fluid. It was once thought that the lymph in the tubes and the lymph in the spaces were in direct communication. Recent investigation has shown that the lymph tubes are closed as completely as the blood vessels, but the lymph contained in them is identical in structure and origin with that in the spaces. How has the plasma that is contained in the blood vessels been able to escape through the wallsORIGIK AND STRUCTURE OF THE LYMPH 175 into the lymph spaces, and how does the lymph of these spaces pass into the lymph tubes? The answer to this question involves the whole mechanism by means of which the tissue elements receive their food and get rid of their wastes. In Figure 61 the relation of a lymph tube, blood tube, lymph space, and tissue element (cell) to one an- other is indicated in a diagrammatic way. Let us ex- amine this diagram for a moment, to see what explana- tion it suggests for this problem. The first point to be noted is that between the lymph in the space and the plasma in the blood tube there is only the thin membrane of the tube wall. Secondly, between the lymph in the space and the liquid contents of the cell is a second membrane, the cell wall. Finally, between the lymph in the space and that in the lymph tube is a third membrane, the lymph tube wall. If we recall what we learned in regard to dialysis and osmosis (see p. 136) it is evident that the above relation satisfies the conditions necessary to those processes. By osmosis the dissolved gases and some of the nutrients of the plasma together with the water, pass through the walls of the blood vessel into the lymph in the space, while in similar manner the wastes in this fluid pass into both the blood vessel and lymph tube. Furthermore, through the cell wall, the176 BLOOD AND LYMPH nutrients of the lymph in the space are exchanged for the wastes of the cell. In this manner then the plasma of the blood tube becomes lymph and is in turn passed on into the tube, which drains it away to other parts of the body. Another feature of this exchange is seen in the action of the corpuscles. It will be noted that the red corpuscles do not leave the blood tube, but the white corpuscles or leu- cocytes are able to pass through tiny spaces in the walls. The reason for this is that the leucocytes are able to change their shape and to push themselves through holes too small to let the rigid red corpus- cles pass. Certain proteids which are not dialyzable also probably pass through similar spaces in the blood tube walls by filtration. Another feature to be considered in this diagram is the disposal of the wastes. If the liquids were station- ary the exchange would go on until all reached the same Fig. 62 — The lymphatic system; a, union of left jugular and subclavian veins, and point of union with the thoracic duct, b; c, right jugular and subclavian veins, and point of union of right lymphatic system ; d, receptacle for food absorbed from the intestine; the oval bodies are glands.LYMPH FLOW 177 composition. The fact that the lymph tubes are continu- ally draining off the liquid they receive, and that the plasma is being constantly renewed, together with the vary- ing activity of the cell, makes the composition of the three liquids subject to constant change. It is this change that gives a definite direction to the flow. While it is true that some substances pass from the lymph tubes into the space, and from the space into the blood tube, the prevailing direction of flow is from blood tube to space to lymph tube, and in this way most of the cell wastes are carried into the lymph tubes. What then becomes of these wastes which enter the lymph tubes? The lymphatics. —The lymph tubes just referred to are only the ultimate tiny endings of a branch- ing network of tubes called the lymphatics. From all parts of the body these tiny tubes (lymph capillaries) are collected together, and finally empty their contents into two large trunks on each side of the body. These trunks are of different size, the one on the left side with its branches being much the larger. It is this left branch that includes the system of branches called the lacteals (see p. 135). Both the right and left branches open finally into two veins on the right and left sides of the neck. In short, then, the waste laden lymph is finally drained from all parts of the body and poured into the blood again at such points that the wastes must pass through the lungs before entering the general circula- tion and are finally eliminated by the kidneys. Lymph flow. — Unlike the blood system, there is no EDDY. PHYS. —12 Fig. 63 —Lymph tubes, showing valves, d.178 BLOOD AND LYMPH apparatus for pumping lymph. The lymph vessels are, however, provided with, valves which allow it to flow only toward the veins. While the flow is irregular and slow the pressure in the capillaries is sufficient to keep it in motion, and the movements of muscles that press upon the tubes also aid in driving it onward to the veins. Since the lymph is laden with waste, it will be recognized that anything which interferes with this flow is bound to disturb the health of the body by the over-accumulation of waste in the spaces. Muscular exercise is necessary, since it maintains the lymph flow. At irregular intervals the tubes open into spongy bodies, known as lymph glands or nodes. These nodes are spongy masses of fibers filled with cor- puscles similar to the white cor- puscles of the blood. They are supposed to have two functions, first, the removal of certain wastes from the lymph, such as bacteria, which the corpuscles devour, and second, the produc- tion of the corpuscles them- selves. In other words, these glands filter the lymph as it passes through them, and tend to remove harmful bac- teria in this way. Occasionally these nodes become over- charged with poisonous material, and break down, forming abscesses or boils. Fig. 64 — A lymph node.SUMMARY OP THE LYMPH RELATIONS 179 Summary of the lymph relations. — From the preceding it is evident that the most important action of lymph is to act as a middleman for the exchange of nutrients and wastes between the blood on the one hand and the cells on the other. Furthermore, the waste matter is in turn taken up by the lymphatics, which drain it off into the veins. In cases of dropsy the failure of the drainage system to remove these wastes causes an accumulation of lymph in the spaces, and often causes the body to swell to an enormous extent. This relation of the lymph spaces and lym- phatics also explains why medicines injected under the skin are so quickly distributed to all parts of the body, the lymphatics taking it up and distributing it to the tissues much more quickly than if it had entered the system through the digestive tract. Certain parts of the body pour into the lymph more of their wastes and secretions than others, and this accounts for the varia- tion in composition of lymph in different parts of the body, but the foundation of the fluid is the blood plasma.XII. CIRCULATION. Anatomy of the Heart and Blood Vessels. Enough has been said in our study of the blood to make clear that its function is the distribution of food and oxygen to the tissue cells, and the removal of waste from these cells. In other words, its purpose in the body is distribution. Furthermore, it is clear that in order that distribution be made to all parts of the body, the blood must penetrate to all these parts, and its flow to and from these points must be continuous. Such conditions are satisfied by inclosing the blood in tubes of varying size and structure, so arranged as to enter all parts of the body, and connected in such a way that a pump placed in one part of the system will force the blood in a constant stream from the pump to these places and back to the pump again. In our body this closed system of blood vessels is spoken of collectively as the circulatory system, while the pump which keeps the blood in motion is called the heart. The various kinds of tubes are called veins, arteries, and capillaries, according to the direction of blood flow. Let us first examine the structure and workings of the heart, or pump. Anatomy of the heart.—(See Ex. XLVII.) This organ is a hollow, muscular sac about the size of a large pear, lying in the chest cavity just above the diaphragm, and immediately behind the lower two thirds of the breast- bone (see Fig. 65). It is shaped somewhat like a cone 180ANATOMY OF THE HEART 181 with the smaller end below and slightly to the left of the breastbone. It is this position of the point that makes the Fig. 65 —External view of the heart; L, L, lungs; RA, right auricle; LA, left auricle; RV, right ventricle; LV, left ventricle; IVC, inferior vena cava; SVC, superior vena cava; Ao, aorta; PA, pulmonary artery ; Tr, trachea. beating more distinctly felt on the left side of the body, but the bulk of the organ is in the middle of the cavity. The heart is surrounded by a sac called the pericardium.182 CIRCULATION This is conical like the heart, but the broad end is down- ward and attached to the diaphragm. Both the inside of the pericardium and the outside of the heart are covered with smooth slippery membranes, while between these two layers is a fluid which moistens the surfaces and lessens the friction between them as the heart beats. The heart is also lined internally with fibrous membrane while the walls between these outer and inner membranes are composed of a peculiar kind of striped muscle tissue whose cells differ markedly from the striped cells of ordinary voluntary muscle. These muscular walls contain nerves, blood vessels, and connective tissue. Their thick- ness varies markedly in different parts of the organ. Internal structure of the heart. — (See Fig. 66.) On opening the heart, it is found to be completely separated into two distinct halves by a longitudinal partition or septum. Each of these halves, which are right and left in posi- tion, is divided by transverse partitions into an upper and lower cavity. The cavities which are at the broad end of the heart are thin walled and flaplike in appear- ance. From a fancied resemblance to the ears of a dog they are called auricles. The other two cavities and the thick walls of muscle which surround them, make up the bulk of the heart. These latter cavities are called Fig. 66 — Relation of cavities in one side of the heart.EXTERNAL. ANATOMY OF THE HEART 183 ventricles. In the partition between the right auricle and right ventricle is an opening guarded by a three-cornered valve known as the tricuspid, valve. The left auricle and left ventricle communicate by a similar opening, and the valve which guards this opening is called the mitral valve from its resemb- lance to a miter. The auricles re- ceive the blood from veins, and this blood passes from the auricles into the ventri- cles through the, valves. The ven- tricles in turn, by contraction of their muscular walls, force this blood out of the heart into arter- ies and thus over the body. It is the pumping ae- Fig. 67 — External view of the heart; 1, right ventricle; 2, left ventricle ; 3, riglit auricle ; 4, left auricle ; 5, aorta; 6, pulmonary artery ; 7, innominate artery; 8 carotid arteries; 9, subclavian artery; 10, supe- rior vena cava 11, pulmonary veins. stitutes what is known as the heart beat. The two sides of the heart beat in unison. External anatomy of the heart. — (See Fig. 67.) When the heart is examined from the outside, the position of these four internal cavities is seen to be outlined by grooves tion of these ven- tricles which con-184 CIRCULATION on the surface. The two flaplike auricles at the top of the heart are separated from the ventricles by a deep groove, while a shallow furrow marks the division of the right and left ventricles, and the position of the septum. On the outside may also be seen a number of large blood vessels which extend from the top of the heart. In addition to these there are smaller blood vessels (the coro- nary system) which branch over the surface of the heart and supply it with blood. The outside is also more or less coated with fat. Relation of blood vessels and heart cavities. — If we push bristle seekers through the external blood vessels, and then carefully dissect away the two sides of the heart we can make out the course of the blood through these vessels and the heart. Let us first examine the right side. (See Fig. 68.) Following the course of the seekers we find that two large veins, called the superior vena cava and inferior vena cava, open into the right auricle, while at the back of the auricle is the opening of a small vein (the coronary vein). Blood which enters the heart from these three veins passes Fig. 68—Fight 6ide of the heart (dis- sected) ; 1, superior vena cava ; 2, infe- rior vena cava; 3, right auricle; 4, tricuspid valves ; 5, papillary muscle fastening the threadlike chorda? ten- din®, which are, in turn, fastened to the edges of the valve; 6, pulmonary artery; 8, aorta; 9, branches of the aorta; 10, left auricle; 11, ventricle; 12, semilunar valves.BLOOD VESSELS AND HEART CAVITIES 185 through the opening guarded by the tricuspid valve into the right ventricle. This tricuspid valve is so constructed as to open to admit the blood from the auricle when the ventricle walls expand, and to prevent its flow backward into the auricle when the ventricle contracts. It is in the form of a curtainlike flap which is fastened by chords {chordae tendince) to raised prominences on the ven- tricle (;■papillary muscles). If we follow the seekers from the right ventricle they extend outward through a single artery known as the pulmonary artery, since it conveys blood from the ventricle to the lungs. This artery has at the base series of valves or pockets of a half-moon shape {semi- lunar valves). These valves are so arranged as to per- mit the flow of blood from the ventricle into the artery when the ven- tricle contracts, and pre- vent its backward flow when the ventricle relaxes. The left side of the heart shows the following connec- tions (see Fig. 69): Into the left auricle open four veins (the right and left pulmonary veins) so-called because they bring blood to the auricle from the right and left lungs. Fig. 69 — Left side of the heart (dis- sected) ; 1, pulmonary veins ; 2, papil- lary muscles ; 3, cut surface of the left ventricle walls ; 4, semilunar valves at the entrance to the aorta ; 5, pulmo- nary artery ; 6, mitral valve ; 7, aorta ; 8, vena cava; 9, branches of the aorta.186 CIRCULATION From this auricle the blood passes into the left ventricle through an aperture similar to that between the right auricle and ventricle. This opening is guarded by the mitral valve, which is fastened and operates in a manner similar to the tricuspid valve. From the left ventricle a single large blood vessel opens, called the aorta or great artery. This, like the pulmonary artery, is guarded from back flow at its base, by semilunar valves. Near its base, also, separate two smaller arteries (the coronary arteries) which convey blood to the substance of the heart itself. Arteries, veins, and capillaries. — (See fix. XLIV.) In the preceding paragraphs we have used the terms artery and veins to indicate blood vessels, but without distin- guishing clearly between them. What is the meaning of the terms artery, vein, and capillary as applied to blood vessels? An examination of the back of the hand will show us a number of dark, bluish colored lines, which give no sensa- tion of beating when we place the finger upon them. These lines are actually blood tubes lying underneath the skin, and their color and lack of beat or pulse indicates that they are veins. If now, we place our finger on the inside of the wrist we shall find that there is a certain spot where a regular beating or pulse ic felt. That spot marks the presence of an artery near the surface, and if we look closely we can make out a thin line less blue than the surrounding veins. While almost all parts of the body surface show blue veins the pulse spots are very few, the most prominent being located on the inside of the wrists, the temples, and the under side of the knee. This indicates that the arter- ies are less near the surface than the veins and are thusarteries, veins, and capillaries 187 better protected. The necessity for this is found in the fact that the bleeding of a cut artery with its pulsing blood is much more difficult to stop than a bleeding vein. If we prick any spot on the skin with a needle we get bleeding, even though no arteries or veins are visible. This is due to the fact that a network of tiny tubes, too small to see with the eye, and called hair tubes or capilla- Fig. 70 — Capillary circulation in the web of a frog’s foot (X 100); a, Z>, small veins, d, capillaries in which the corpuscles are seen to follow one another in single series ; c, pigment cells in the skin. ries, because of their tiny dimensions, connect veins and arteries. The relation of arteries, veins, and capillaries may be observed to good advantage by stretching the living web of a frog’s foot or tadpole’s tail under the objective of a compound microscope. (See Ex. XLY. and Fig. 70.) The blood vessels, which are transparent, show188 CIRCULATION clearly the streams of blood plasma and corpuscles in motion. With closer attention we can make out which tubes are arteries, veins, and capillaries. The blood in the arteries flows in spurts or pulses. The capillaries show a steady streaming motion and are so small that the corpuscles move in almost single file through them. The veins are as large as the arteries but the blood in them flows in a steady stream. If, now, we observe the direc- tion of flow in the several vessels, it will be seen that the blood in the arteries divides and passes into the small capillaries, while from them in turn it passes on into the veins and through these toward the bod}' again. In other words, the capillaries are the connecting links be- tween the arteries and veins, and in passing through them the blood loses its spurting method of flow and is converted into a steady continuous flow. The difference in color between the arteries and veins is explained by the color of the blood in the two tubes. The blood in the arteries is bright red, being full of oxygen, while that in the veins has a darker color owing to the larger proportion of carbon dioxide present. This latter blood is not blue, and that color is due to certain light effects. Summary. 1. Arteries convey blood from the heart, and veins return the blood to it 2. Arteries contain much oxygen and little carbon dioxide, veins contain much carbon dioxide and little oxygen. 3. Blood in arteries moves in spurts (that is, it pulses), while that in capillaries and veins moves in a steady stream. 4. Arteries are better protected than veins by being in general placed farther below the surface. (This does not apply to the main vein trunks.)STRUCTURE OF BLOOD VESSELS 189 Structure of blood vessels.—(See Ex. XLYI.) Arteries. A cross section of an artery (see Fig. 71) shows it to be made up of three layers of tissue. The outer coat is composed of fibrous con- nective tissue, which is tough, and a strong pro- tective layer. This coat contains also some elastic fibers that enable it to expand under the pulsing of the blood. The middle layer consists of a mixture of unstriped muscle cells and elastic fibers, and is thicker than the other two layers. The inner layer is made of flattened, epithelial cells, and between it and the middle layer are a varying number of bands of membranous and elastic tissue. Veins. The structure of the veins is practically identical with that of the arteries (see Fig. 72) ex- cept that the outer coat is thickest and the two inner layers are much less de- veloped. Owing to this difference they are much less elastic, and readily collapse when empty. The Fig. 72 —Cross section of a vein ; c, outer fibrous sheath ; m, middle muscle and elastic tissue layer; e, inner epithelial cell layer. Fig. 71 — Cross section of an artery; c, fibrous connective tissue, outer sheath; m, muscle cells and elastic fibers; e, epithelial cells, with bands of tissue separating them from the middle layer.190 CIRCULATION toughness and thickness of the outer coat gives them great strength. Many veins also possess semilunar valves formed by pockets located at intervals on the inner surface of the tubes. These are so arranged (see Fig. 73) that the edge toward the heart is free and prevents a backward flow in the tube by pres- sing out and bridging the tube under backward pressure. These valves are found chiefly in the superficial and muscle supply veins. The ar- teries have no such valves except at the entrance into the heart. The location of these valves may be recognized by the knotted ap- pearance of the veins at these points. Capillaries. As the arteries decrease in size the outer and middle coats gradually disappear until, in the capillaries, only the inner layer semi] unar is left. This is composed of flattened cells valves. cemented together at the edges. Such a structure is evidently well adapted to the process of osmosis, dialysis, and filtration.XIII. CIRCULATION (continued). The Flow of Blood, and Blood Systems. In the preceding chapter we have seen the way in which the various parts of the circulatory system are put together. In the present chapter we shall investigate the cause of the blood flow and its course in the blood vessels. Heart action. — (See Fig. 66.) Enough of the structure of the heart has been outlined to show that the actual driving of the blood into the arteries is due to the con- tractions of the ventricles. Before these ventricles contract it is necessary that they be filled with blood, and this requires a preliminary contraction on the part of the auri- cles. The order of these contractions is as follows: The action begins at the mouth of the large veins (the venae cavae). The blood is flowing steadily from these veins into the auricles and dilating them. Next the tricuspid and the mitral valves are forced open by the blood pressure in the auricles and the ventricles fill. Now the auricles contract and send a wave into the ventricles dilating them still further and checking the flow from the veins by back pressure, while the mitral and tricuspid flaps float upward. As the ventricle becomes fully filled, it, in turn, begins to contract, and these flaps are suddenly pressed upward until they cover the opening tightly and prevent further flow from the auricle into the ventricle and also back flow from the ventricle into the auricle. The chordae tendinae prevent the flaps from being forced back through the opening. The blood in the ventricles is now under 191192 CIRCULATION pressure and forcing down the semilunar valves at the mouths of the arteries; the blood from the ventricles enters these arteries with a spurt. While this ventricle contrac- tion is going on, the auricles begin to expand and the blood flows from the veins into them. Before they finish ex- panding the ventricles also begin to expand, while the semi- lunar valves of the arteries close to prevent back flow from them. During this period of expansion the mitral and tricuspid valves begin to open, and both auricles and ven- tricles to fill with blood. This goes on until the auricles begin to contract again, and this cycle is repeated indefi- nitely. The position of the valves may be summarized as follows: (a) As the auricles contract, the mitral and tricuspid valves are open; the semilunar valves are closed. (b) As the ventricles contract, the mitral and tricuspid valves are shut tightly. The semilunar valves are open and blood is entering the arteries. (c) During the pause before the next contraction the mitral and tricuspid valves are open to permit the filling of ventricles, while the semilimars are closed to prevent back flow from the arteries. The name of diastole has been given to the period when the auricles and ventricles are expanding, while the period of contraction of these organs is called the systole. Between these two periods there comes a time when neither is con- tracting or expanding. Using these terms the various stages of a “ cardiac cycle ” or heart beat may be ex- pressed as follows: First. An auricular systole. Second. A ventricular systole. Third. Heart pause (both organs in diastole).PULSE 193 Sounds of the heart.— If one listens carefully to the heart beat, by placing the ear against the chest, two distinct sounds may be distinguished. The first is lower in pitch than the second and of longer duration. It is supposed to be produced by two factors, the vibration of the tricuspid and mitral valves in closing, and the vibration of the contracting muscle mass. The second sound is sharper and is due to the closing of the semilunar valves. The first sound is essentially a systolic sound, while the second is dia- stolic. These two sounds are of great aid to the physician in deter- mining cases of heart trouble. For detecting variations in this sound an instrument called the stethoscope is used, and this method of examination is called auscultation. Pulse.— When the ventricles contract the blood is forced into the arteries with a spurt. If the arteries, capillaries, and veins were rigid tubes this sudden pressure would be transmitted instantaneously throughout the entire system, and a quantity of blood would enter the auricles exactly equal to that which left the ventricles. Owing to the elasticity of the walls of the blood vessels this is not the case. Part of the pressure is used up in expanding the walls. Since the arteries are subject to the most direct effect of this systole the reason for their thicker walls is apparent. This elastic character of the arteries also results in the pressure being transmitted in a wave instead of directly. The results of this may be pictured as follows: As the ventricle contracts the ex- pressed blood stretches that portion of the artery nearest the heart. As the ventricle expands and this pressure is removed this enlarged part of the artery begins to contract slowly, and the pressure is transferred to another point further along in the artery. As the next systole occurs this process is repeated, and thus the effect of the pressure or wave is gradually transmitted throughout the whole EDDY. PHYS. — 13.194 CIRCULATION arterial length. In this tvay each portion of the artery wall expands to a maximum and then contracts, at inter- vals corresponding to the systolic and diastolic periods of the heart. The disappearance of this wave in the capillaries and veins is explained when we consider how the arteries branch into smaller and smaller tubes, which are farther and farther removed from the seat of pressure until in the capillaries the wave has become so split up as to be barely perceptible, and in the veins is converted into a steady flow. In this respect it is like a water wave, which becomes less apparent the farther we go from the center of the disturbance. It is this wavelike character of the flow of the blood in the arteries that enables us to feel a pulse at points where the arteries are near the sur- face. What we actually count in taking the pulse is the time between successive periods of pressure and relaxa- tion, and since these correspond to expansion and contraction of the heart they enable us to detect the rate of beat of the heart. ' The physician uses the pulse point in the wrists and temples to furnish him Avith valuable information about the heart and circula- tion. Pulse rate.— The rate of the pulse or heart beat is subject to great variation under different conditions of health and disease. It is this fact that makes it a valuable indicator of the body con- dition to the physician. Age, sex, size, atmospheric pressure, all have a distinct effect upon the pulse rate. The average rate for the adult man is seventy beats per minute, and for the adult woman seventy-eight to eighty per minute, showing that sex affects the rate. Tall individuals have a slower rate than short persons, rvhile large bodies have a slower rate than small bodies. The human rate is most rapid in infancy, sinks rapidly at first, and then more slowlyPULMONARY CIRCULATION 195 to a minimum in adult life, rising slowly as we enter extreme old age. The following table gives the average rates: At birth........................140 beats per minute. Infancy.........................120 Childhood.......................100 Youth ...........................90 Adults....................... 75-70 “ Old age..........................70 “ Extreme age...................75-SO “ Systems op Circulation. From what has been said it follows that the blood which passes through any given chamber of the heart must ulti- mately return to that same chamber. Owing, however, to the division of the heart into two parts, which beat in unison, and the branching of the blood vessels, the actual course of the blood requires some further explanation. For convenience, the blood vessels that are concerned in the transfer of the blood from the right to the left side of the heart, and that have their branches in the lungs, are spoken of collectively as the pulmonary system. Those concerned with the transfer of blood from the left side of the heart to the right side, and whose branches take it all over the body, are spoken of as the systemic system. Pulmonary circulation.— When the blood enters the right auricle it is partly exhausted of its oxygen and is dark and purplish in color, owing to the abundance of car- bon dioxide which it contains. In this condition it enters the right ventricle, and when that organ contracts it is forced into the 'pulmonary artery which in turn breaks up into two branches leading to the right and left lung, and these into smaller branches within the lungs. These pul-196 CIRCULATION monary arteries convey the blood from the right ventricle directly to the lungs. In the many capillaries, into which the branches of the pulmonary artery divide in the lungs, the blood takes on a new supply of oxygen and loses its carbon dioxide. These capillaries are collected into largerTHE SYSTEMIC CIRCULATION 197 and larger veins and finally bring back the oxygen laden blood to the left auricle by four veins (two from each lung). These veins are called the right and left pulmonary veins respectively. From the left auricle the blood passes into the left ventricle, and in this manner the transfer of blood from the right to the left side of the heart is brought about. In our study of the lungs we shall discuss more in detail the manner in which it loses its carbon dioxide and takes on oxygen. In this circulation it will be noticed that it is the pul- monary artery which contains the carbon dioxide laden blood, while the pulmonary veins contain the bright red, oxygen laden blood. Owing to this fact the only universal definition that will apply to all arteries and veins is one which is based on the direction of blood flow. Arteries convey blood away from the heart, veins toward the heart. The systemic circulation. — From the left ventricle this oxygen laden blood is pumped into a large artery called the aorta. The aorta subdivides into many branches which carry the blood to all parts of the body. In the tissues these arteries subdivide into capillaries, and these, in turn, collect into veins through which the blood finally enters two large veins (the superior and inferior venae cavce) and is returned to the right auricle by them. In this system the blood is transferred over the body and from the left to the right side of the heart. Since it is this system that carries food and oxygen to the tissues and transfers their wastes and carbon dioxide to the excretory organs, the relations of the branches is important to a clear understanding of the blood flow of this system. As it leaves the heart, the great aorta bends first to the left, then downward through the rear part of198 CIRCULATION Kxq. 75 —The arterial systemTHE SYSTEMIC CIRCULATION 199 the chest cavity, finally piercing the diaphragm and en- tering the abdominal cavity. The change in direction as it leaves the heart results in an arch called the arch of the aorta. From the top of this arch on the right side is given off a large branch which, in turn, soon divides into two parts. One of these two parts (the right carotid artery) supplies blood to the right side of the head. The other part passes under the collar bone, down the arm, and at the elbow divides into the radial and ulnar arteries thus supplying the right arm with blood. It is the radial artery whose pulsings are felt in the wrist. From the top of the arch on the left side are given off two branches, one of which (the left carotid) supplies blood to the left side of the head, while the other branches as on the right side and supplies blood to the left arm. From the aorta, near the heart, arise also the coronary arteries which, branching over the surface of the heart, supply the tissues of that organ with blood. As the aorta curves downward through the chest cavity it gives off branches which supply the lung tissues, the esophagus, and the muscles of the ribs and spinal column. From the aorta in the abdomen are given off five main branches. The first (cceliac axis) divides into three parts which supply the stomach, spleen, and liver respectively. Two other branches (the mesenteries) carry blood to the walls of the intestines. The remaining two branches carry blood to the kidneys. The main trunk of the aorta con- tinues to the rear of the abdominal cavity and finally divides into two branches which each enter a leg and supply these extremities with blood. All these branches of the aorta subdivide first into smal- ler arteries and then into capillaries, which are in turn col-200 CIRCULATION lected into veins by which the blood is returned to the right auricle of the heart. The blood supplied to the head by the carotids is re- turned to the heart by two large veins (the jugulars) pass- ing down each side of the neck; while from the arms come two other veins. These four veins finally unite to form the superior vena cava which opens into the right auricle of the heart. In a similar manner various veins return the blood from the lower part of the body and unite in one common vein (the inferior vena cava) which also opens into the right auricle of the heart It is this lower part of the venous system which collects the food from the digestive system. The portal circulation. — The veins which are found in the walls of the stomach and intestine do not convey the blood directly to the in- ferior vena cava. After collecting the food from these organs these veins are collected into a single large vein (the portal vein) which passes di- rectly to the liver. Here this vein splits up into capillaries again, which in turn collect again into a second vein (the hepatic vein). This latter pours its blood into the inferior- vena cava. It is to be noted in this case that the capil- laries of the liver arise from and unite again into veins, and that the food is thus forced to pass through two sepa- rate sets of capillaries before entering the heart. This branching of the portal vein into capillaries is the only Fig. 76 — Diagram of the portal circulation.THE LOADING OF THE BLOOD WITH NOURISHMENT 201 example in the body of a vein’s connection with another vein through capillaries. In all other cases the capillaries connect veins and arteries. It is this arrangement of portal-vein capillaries which permits the liver to act as a storehouse of carbohydrate material, and to regulate the amount of that material which shall enter the general circulation at any given time. The renal circulation. — The blood which is returned to the left side of the heart by the pulmonary veins has been freed of its carbon dioxide only. Hence the blood which passes into the aorta is still rich in urea and other body wastes brought to the heart by the venae cavae. Much of this waste is diverted into the artery branches which carry blood to the kidneys (the renal arteries). In these organs, by means which we shall discuss more fully under the excretory system, this waste is removed, and the renal veins, therefore, return to the inferior vena cava blood which is nearly free of body wastes. Changes in blood composition. — In passing through the various systems of circulation, portal, renal, systemic, and pulmonary, the blood undergoes distinct changes in composition. Having in mind the parts of the body which these systems supply, we may now summarize these changes as follows: The loading of the blood with nourishment. — 1. Food enters the blood at two points in its circulation. (a) All kinds of food, except the fats, enter the capil- laries of the stomach and intestines and are collected and carried to the liver by the portal vein. In the liver, the capillaries into which this vein divides permit the liver cells to separate from the rest of the202 CIRCULATION blood the carbohydrate, together with certain poisons (see p. 142). The carbohydrate thus separated is either stored in the liver as glycogen, or given out as needed in the form of grape sugar to the hepatic vein. This hepatic vein passes this grape sugar and the other unchanged nutrients into the general cir- culation by opening into the inferior vena cava. The poisons are passed into the liver secretion or bile and returned to the intestine with that fluid for removal with the feces. (b) Fats are collected by the lacteals, and enter the veins by way of the thoracic duct at a point just under the left collar bone where the left jugular and left arm vein join. Thence they pass into the heart by the superior vena cava. 2. Oxygen enters the blood in the lungs, and is con- veyed to the heart by means of the pulmonary veins. We shall consider this phase of the circulation more in detail in our study of respiration. The loading of blood with waste. — All veins carry wastes (COs, urea, etc.), which they collect from the tis- sues. This waste either enters the blood directly by way of the capillaries, or else enters the lymphatics and thus the veins by the thoracic duct or right lymphatic duct. The right lymphatic duct enters at the joining of the right jugular vein and the right arm vein. Unloading of nourishment by the blood.— All arteries1 carry food and oxygen to the tissues which they supply, and deliver these substances to the cells of the tissues 1 The pulmonary veins and arteries are exceptional in that the pul- monary veins carry oxygen laden blood, and the pulmonary arteries, carbon dioxide laden blood.ARTERIAL AND VENOUS BLOOD 203 through the thin walls of the capillaries in the manner already described (see p. 175). The plasma composition indicates that it is the part of the blood which carries the food to the tissue cells. Unloading of wastes by the blood.— The carbon dioxide given off by the tissues is removed from the blood almost entirely in the lungs. It is noteworthy that the lungs do not free the blood of wastes in general, and on that account should not be said to 'purify the blood. In other words, their great function is to exchange oxygen for carbon dioxide or aerate the blood. Most of the urea and true wastes of the body are removed by the kidneys and the skin, this waste being collected from all parts of the body by the veins and conveyed to these organs by special arteries, such as the renal artery to the kidneys. Arterial and venous blood.— From the description given above it is readily seen that the blood in the arteries and veins varies in composition in different parts of the body. In general, however, we may say that the veins contain less oxygen and more carbon dioxide than the arteries. The redder color, due to the larger proportion of oxygen present, has led to the custom of representing arteries by red, and veins by blue lines in a diagram of circulatory systems. The notable exceptions to this rule are the pulmonary veins and arteries. The pulmonary vein con- tains very little carbon dioxide, while the pulmonary artery contains very little oxygen. In general, then, the terms arterial and venous are used to indicate relative amounts of oxygen and carbon dioxide in the blood. In that sense the pulmonary vein contains arterial blood, and the pulmonary artery, venous blood.204 circulation Regulation of Blood Flow. Heart control. — The amount of blood which passes through a given part of the circulatory system at any one time is, of course, regulated primarily by the heart beat. This beat, which is produced by rhythmic contractions of the heart muscles, is ordinarily very regular. Its rate, however, is subject to certain external influences. For example, a decrease in blood pressure in the arteries is associated with an increase in heart rate and vice versa. Muscular exercise also increases the heart rate. Even so little an effort as a change in position is sufficient to change the rate, that of a standing position being higher than that of a sitting or reclining one. The purpose of these changes in rate is readily under- stood when we consider that tissues which are active are using up more food and creating more waste than still tissues, and consequently require a greater flow to supply this demand. Mental states and emotions, blood content, and tem- perature, all affect the rate of heart beat. Some lower this rate, others increase it. Some, like temperature, cause an increase up to a certain limit, and then above that limit produce a decrease. Physicians take advantage of these factors and introduce into the blood certain drugs which may accelerate or lower the rate of beat, and thus can control it to a certain extent in cases of disease. The manner in which all these special conditions are able to affect the heart is through what we call nerve action. The muscle walls of the heart are connected by nerves to other parts of the body, and when one set of nerves is stimulated by any of the above means, the nerve-ARTERY CONTROL 205 ends In the heart muscle receive this impulse and increase the rate of contraction and expansion of the cells. When another set of nerves is stimulated, they, in turn, cause the contraction to be reduced in rate. These are like telegraph wires connecting stations. One station is the heart where messages are received. The others are found in the brain and in the body tissues, and the messages sent from these latter points control the action of the heart. Of the two kinds of nerves those that decrease the rate are called heart inhibitors; and those that in- crease the rate, heart accelerators. By their aid the rate of flow is adjusted to the needs and conditions of the tissue cells. Artery control.— The walls of the arteries, as well as of the heart, are supplied with the ends of what are called the vaso-rnotor nerves. Under the normal action of these nerves the walls of the arteries have a certain elasticity which responds to the blood pressure; stretching when the pressure is increased and contracting as the pressure is removed. When these vaso-motor nerves are stimu- lated further the effect produced may be a still greater contraction of the arter}r walls and a consequent decrease in the blood content of the contracted artery, or a paralysis of the walls which causes them to lose their elastic tension and stretch. In this way it is possible for conditions, external or internal, to affect these nerves and produce stimuli, which, in turn, control the blood supply of any given part of the body. The nerves which, when stimu- lated, cause the contraction of the w’alls are called the vaso-constrictors. Those which, when stimulated, pro- duce paralysis of the walls and thus allow them to stretch are called the vaso-dilators. This stretching and contract-206 CIRCULATION ing of the artery walls with the consequent increase and decrease in blood must necessarily cause the capillaries, whose walls are also elastic, to become stretched. Hence variation in the tension of the artery walls is always accompanied by similar variations in the blood supply to those capillaries which the arteries feed. The conditions which may thus affect these vaso-motor nerves are many. Heat to a certain limit, exercise, the presence of food in the digestive tract, etc., all affect the vaso-dilator nerves and cause the artery walls to relax. Cold, on the contrary, affects the vaso-constrictors and causes the contraction of the walls. Mental emotions in some cases cause relaxation of the walls, and in other cases, contraction. Thus, blushing is due to overcharg- ing of the facial capillaries from relaxation of those artery walls which supply these capillaries; while paleness is due in the same manner to contraction of the supplying artery walls. In both cases the mental emotions are responsible for the nerve stimulus. Injury to tissue cells invariably results in a relaxation of the walls of the arteries supplying the affected part, and thus permits an increase in the blood supply to the injured area. Such an increase is spoken of as congestion. Congestion and inflammation. — A simple case of con- gestion is seen when we scratch the skin with a pin, the scratched surface becoming reddened from the action of the injured cells upon the artery supply. If the injury is severe and the cells are killed, swelling often follows the redness, and painful sensations may result. Such congestion, with its attendant swelling, is merely an indication of the first steps in the repair of the injury. The flow of blood to the injured part is soon followed, inCONGESTION AND IMELAMMATION 207 this case, by the outpouring of plasma and white blood corpuscles (leucocytes) through the capillary walls into the lymph spaces. This extraordinary flow is due to changes in the cement which holds together the capillary cells com- posing the walls, and may in some instances be accompanied by the outpouring of red corpuscles. In simple injuries the result of this outpouring of plasma is to stick together the old cells and cause them to grow and unite, while the leucocyt.es and injured cells are absorbed anti the injury thus repaired. If, however, the injury is extensive or bacteria are in- troduced into the injured area, a more extensive action is involved. The growth of the bacteria or the nature of the injury may cause the death of other cells in the neighbour- hood, and the formation of a mass of dead tissue results. In this event the white corpuscles proceed to kill or absorb the bacteria and dead tissue cells with the formation of a mass of matter usually designated as pus. At the same time new cells are formed and a new tissue grows in place of the old one. This pus may be removed by the lympha- tics or absorption into the blood vessels, or it may soften the tissue about it and discharge, as in an abscess. In the latter event, the white corpuscles either devour the bacteria and cells, or give off ferments which dissolve them. Meanwhile the living cells about the injured part arc ex- cited to growth and reproduction, and thus new cells pro- duced which unite and replace the old tissues. The processes here are evidently two, the dissolving and removal of old matter through the agency of the white corpuscles, and the formation of new tissue cells to replace the injured ones. The manner in which the old matter is removed varies with the location of the injury. These208 CIRCULATION two processes of repair are called inflammatory processes, and inflammation is usually defined as “ the local attempt at repair of an injury,” Examples of simple inflammatory processes.—When the mucous membrane is injured by exposure to cold, there results a conges- tion in the membrane. This congestion may be complicated by the presence of bacteria. In such congestion the formation of pus is accompanied by an increased activity and secretion of the mucous cells. The combined secretion and pus collects in the passages inclosed by these membranes and forms the discharge common to colds. This congestion of the mucous membrane may be brought about by the action of the cold upon the vaso- motor nerves which control the artery supply to the skin. In this event their action is to cut off the blood supply to the skin, and the excess of blood thus cut off is forced to accumulate in the mucous membranes and produce congestion. If this congestion and inflammatory process is confined to the nasal passages the result is called a cold in the head, or catarrh. If in the throat, it may result in swelled tonsils and a sore throat. If in the bron- chia passages, it is called bronchitis, while in the lungs it may produce pneumonia. When certain bacteria attack the throat membranes the result may be severe. Such an instance is seen in diphtheria. Another example of inflammatory processes is found in the pimples and boils. In these cases, bacteria are introduced in some way into the underlying layers of the skin where they cause the death of the surrounding cells. Congestion results, and the leuco- cytes dissolve the dead cells and devour the bacteria forming a greater or less quantity of pus. The pus thus formed softens the surface layers of skin and forms a head or weak spot on the surface through which the pus may ultimately be discharged. “Taking cold in a boil” is actually a case of accidental intro- duction of bacteria into a partly discharged boil, and the resultant formation of new pus, and has no connection whatever with cold. The introduction, by the blood, of bacteria into the internalTREATMENT OP CUTS AND BRUISES 209 tissues, internal disturbances of function, and many other causes, may bring about so extensive a congestion and accumulation of poisonous matter that the body is unable to remove it. In that event the presence of this matter in the body may produce various forms of disease, such as fevers, growths, etc. Treatment of cuts and bruises. — When the capillaries are cut the bleeding is very slow and is usually easily checked by the natural clotting of the expressed blood. The cutting of an artery or vein, however, is a much more serious matter, and owing to the spurting of the blood the cut artery is more difficult to treat than the vein. If the artery or vein is small the mere pressing together of the walls with a bandage may check the flow sufficiently, to permit the formation of a clot. When the artery or vein is large the blood pressure is too great to allow the clot to form and special measures are necessary. The object of all such measures is to secure sufficient pressure upon the blood vessel to force the cut edges together and diminish the blood flow sufficiently to permit the formation of a clot. Owing to the direction of flow in an artery, this pressure must be applied on the side of the cut nearer the heart. In case of a vein the pressure should be applied on the side farther from the heart. In some cases simple bandaging fails to stop the flow and the clot is washed away as fast as it forms. In such cases an application called a tourniquet is resorted to. This consists of a band- age applied on the proper side of the cut and twisted tight with a stick so that the walls of the blood vessels are pressed together at a point above the cut. The blood being shut off from the cut ends, a dangerous loss of blood is thus prevented. In emergencies persons may thus be saved from bleeding to death since all the tools required are a EDDY. PHYS. —14.210 CIRCULATION strip of cloth and a stick. The tourniquet is of course only a temporal'}’ measure and is to be employed only as an emergency remedy until the surgeon can treat the wound. Surgeons usually close large wounds by tying the ends of the cut blood vessels with clean catgut. Such bindings are called ligatures. When the ends of a vein are cut they may grow together again and thus circulation be restored through the original channel. Artery ends, on the contrary, do not grow to- gether but the ends close and the branches which are in communication above and below the cut enlarge and replace the original channel. All cuts should be kept clean and free from bacteria. Weak solutions of certain germicides can be obtained from druggists and used to wash and protect the cut from these dangerous visitors. Blood poisoning is one of the serious results of neglect of these precautions. The healing of a cut or bruise will take place without pain or inflammation if kept clean. Effect of alcohol upon the circulation. — The most marked effect of small amounts of alcohol is the dilation of the surface capillaries and their filling with blood. The result is a sensation of warmth or glow due to the stimu- lation of the surface heat nerves. Alcohol produces this effect by its action on the nerve centers, which, in turn, stimulate the vaso-dilator nerves of the arteries that supply these capillaries, and not by any direct action upon the blood vessels. In this case the sensation of warmth is deceitful, since the bringing of the blood from deeper parts of the body to the surface permits the escape of heat and a slight lower- ing of the body temperature.EFFECT OF ALCOHOL UPON THE CIRCULATION 211 In larger doses alcohol produces effects in one of two ways: (a) It may cause the deatli or degeneration of the cells with which it comes in contact. For example, it may cause the cells of the heart to become charged with fat and thus weakened, or it may make the cells of the artery walls brittle and unable to withstand pressure, or finally, it may actually poison and destroy the cells of certain tissues. (b) When brought in contact with the cells of the tissues it may so injure them that congestion and inflam- mation results. This inflammation may cause a disturb- ance of the natural action of the congested part due to the swelled condition, or it may stimulate unnatural growth on the part of the surrounding tissue cells. Another bad result of this congestion may be the stretching of the arte- ries to such an extent that they are unable to recover their natural size, and thus- produce continued congestion in the part affected. The extent of the injury which alcohol may produce in the body by either one of these means, that is, directly or indirectly, is a subject for a medical text, and we need only outline here the method of the action. In brief, then, it is evident that alcohol in either large or small doses acts upon the nervous centers and through them upon the vaso-dilator nerves, producing abnormal distribution of the blood, and that in large doses it is actually poisonous to the cells with which it comes in contact. While the extent of the injury produced by small doses may be only temporary, continued use or excess causes such changes in the distribution of blood as to be extremely dangerous to the system.XIV. CIRCULATION IN THE LOWER ANIMALS.1 In the one-celled animals, such as the amoeba, the food is digested in the cell protoplasm itself, and oxygen is absorbed directly from the surrounding air or water. In like manner the wastes and carbon dioxide are removed from the cell and cast into the surrounding medium. In such animals a circulatory system is evidently unneces- sary. In fact, the only suggestion of it is found in the movement of the protoplasm in the cell by means of 'which the absorbed food is distributed throughout the cell. This, however, is entirely an internal cell circulation, and not a system for bringing food to the cells as in man. In the sponges and coelenterates each cell is able to obtain its air supply and food supply directly, and no special circulatory system is necessary. When we come to animals possessing many layers of cells it is evident that some cells will be so placed as to be unable to secure food and air directly or to rid themselves of waste. Some special apparatus, therefore, becomes necessary to supply these cells with food and to remove their wastes. In this way there arises the circulatory system as we have considered it in our definition. Some of the systems in the lower animals are interesting from the light they throw upon the development of our own complex network of tubes. Insect circulation. — In the grasshopper, we see one of the simplest forms of a circulatory system. In this animal 1 See Footnote on p. 146, Chapter X, 212INSECT CIRCULATION 213 the food passes from the digestive tract directly through the walls of the stomach and intestine into the body cavity. Here it would remain were it not for a tube with several swellings along its surface extended along the dorsal surface of the body cavity. This tube, with its swellings, is a sort of muscular pump or heart, closed at the rear end and open in front. The swel- lings expand and contract rhythmi- cally. As they expand the liquid of the body cavity is sucked in through openings along the sides. These open- ings are protected by valves which open inward. As the swellings contract these valves close, and the inclosed fluid is thus forced forward and out the front end of the tube toward the head. In this manner the body fluid is kept in motion, and by means of the cavities between the tissues it flows around and bathes all the tissues, and finally returns to its starting point, thus sup- plying the body cells with food and receiving their wastes. This waste is absorbed by kidney tubes which extend into the body cavity from the intes- tine, and the absorbed fluid is excreted through the intestine with the fa?ces. The liquid circulated contains no red corpuscles, as the air supply is furnished by an entirely independent set of tubes. There are no arteries or veins, the blood flow- ing through the spaces of the body from the mouth of Fig. 77 —Portion ot an insect’s blood tube (dissected); a, b, muscular walls ; d, valves between the compartments; c, valve covering the general opening into the body cavity; through this valve the liquid enters.214 CIRCULATION IN THE LOWER ANIMALS the tube to its starting point. The heart is simply a contractile structure whose office is to keep the liquid in motion. The earthworm. — In this animal we have a slight advance over the insect. It possesses a true closed sys- tem of tubes conveying a red blood. It consists of two tubes running lengthwise of the body, one dorsal and one ventral. The dorsal blood vessel can be easily made out through the transparent skin of the living earthworm, and observation shows that the red liquid which it con- tains moves in pulses toward the head. At the anterior Fig. 78 — Diagram of a longitudinal section of an earthworm ; h, one of the fine pulsating arches which pump blood from the dorsal blood vessel into tlid ventral blood vessel. end of the animal are five pairs of arches or blood vessels, which branch from each side of the dorsal blood vessel and, encircling the esophagus, open below it into the ven- tral blood vessel. These pairs are arranged, one behind the other, like rings from the sixth to the eleventh segment. These five pairs of tubes are called the aortic arches, and they pulsate in unison, forcing the blood to flow from the dorsal blood vessel into the ventral. The course of the blood, then, is from the rear of the body along the dorsal blood vessel to the head; downward, through the arches to the ventral vessel, and through* this toward the rear of the body again. Both dorsal and ventral bloodTHE EISH 215 vessels give off branches, and while the exact course of the blood flow from the ventral vessel is at present un- known, it is supposed that in some way it is ultimately brought back into the dorsal vessel and thus completes its circuit. The branches of the two main vessels sub- divide into capillaries, and by means of these the blood is brought into contact with the various tissues of the body, and thus food is distributed and wastes collected. The red color of the blood is due to the hremoglobin dissolved in the fluid, the only corpuscles present being a few white ones. Besides this inclosed fluid, we find the body cavity filled with a similar liquid to that found in the grasshopper, and it is known that part of the absorbed food enters the inclosed system and part is distributed by the cavity fluid, so that we actually have two methods of circulation in this one animal. The fish. — It is only in the vertebrate animals that we find a true blood with red corpuscles. The simplest vertebrate system is found in the fish. In this animal the heart is a simple structure consisting of a single ventricle and auricle, and is located in a pericardial cavity separated from the rest of the body organs by a thin mem- brane. The blood is driven from the ventricle through an artery (the aorta) to the gills. Here the artery branches into capillaries and through the thin walls of these is effected the inter- change of oxygen and carbon dioxide between the water d Fig. 79—Plan of circulation in a fish ; a, auricle ; b, ventricle; c, branchial arte- ry; e, branchial veins ; (I, gills ; /, aorta; g, vena cava.216 CIRCULATION IN THE LOWER ANIMALS and the blood. These capillaries ultimately return the blood to a dorsal artery which branches and carries the blood to all parts of the body, where the division of the branches into capillaries permits the giving up of food and the removal of the waste of the tissues. These capillaries are now collected into veins, and the blood is returned by these tubes to the auricle of the heart to repeat this cycle indefinitely. As in man the food of the digestive tract is absorbed by capillaries which unite to form the portal vein, and this absorbed food is carried to the liver by this vein. Here the portal vein produces capillaries, and these, after branching throughout the liver, unite again into the hepatic vein and are carried directly to the auricle by this tube. In this circulation the single auricle and ventricle cor- respond to the right side of the human heart, and the heart receives and expels only the venous blood. The friction in the gill capillaries retards the flow so greatly that the circulation in the arteries of the fish is very sluggish. The red corpuscles of the fish are nucleated, and it also possesses a lymphatic system. The frog.— In the frog the blood is aerated in the lungs, and the heart lias two auricles and one ventricle. The right auricle, as in the fish, receives blood from the veins which bring it from all parts of the body to this division of the heart. The left auricle receives aerated blood from the lungs. It is to be noted that the frog has advanced over the fish in having the aerated blood returned to the heart before pumping it over the body, and as we would expect, this arrangement allows a little faster flow through the arteries. Since, however, there is only one ventricle, the two kinds of blood found in the auricles must become mixed in this chamber, and the arteries in consequence areTHE BIRDS 217 filled with mixed blood. By an ingenious arrangement of arteries and valves the blood is not evenly mixed, but the part which is pumped to the head is richest in oxygen, that to the body organs next richest, and that to the lungs and skin is poorest in oxygen and richest in carbon dioxide. As in the fish, the red corpuscles are nucleated. In its early life the frog breathes by gills. The reptiles.— These animals have, in general, the same sort of circulatory system as the amphibians. There is, however, one modification in the struc- ture of the heart that shows an ad- vance. In most reptiles the ventricle is partly divided by a partition, and this partition tends to keep separate the aerated and non-acrated blood so that the aorta which arises from the left side of the ventricle receives blood freer of carbon dioxide than the pulmo- nary arteries which arise from the right side. This partition is interesting as a step toward complete separation of the two kinds of blood. The red corpus- cles are nucleated. The birds.— In these vertebrates the ventricle is completely divided by the partition, and the bird heart, like that of man, is four chambered. This' fact alone would enable us to classify birds as higher forms than reptiles. The rate of flow in birds is much more rapid than in man, and the temperature of the blood is much higher. This is due Fig. 80.— Plan of circu- lation in amphibians and reptiles ; a, right auricle receiving ve- nous blood from the system ; 6, left auri- cle receiving arte- rial blood from the lungs; c, common ventricle; d,f, c, sys- temic arteries, capil- laries, and veins ; the arrows indicate the streaming of tween 1 See Footnote, p. 146, Chapter X. 294CAUSES OF LOCOMOTIOH 295 in our present discussion we are more directly concerned with locomotion. Causes of locomotion in animals.— The locomotion of all animal forms can usually be explained as due to one of four causes: (a) A change of position due to the action of some force outside themselves, such as currents of water or air. This sort of movement is, of course, passive movement as far as the animals are themselves concerned, since they take no part in its production. (b) A change of position due to change in the cell com- position, as a result of which there is a gain or loss in specific gravity. Such change allows the buoyant force of the water to move the body up or down. This is not a passive movement, since the changes in composition determine the action of the buoyant force. (c) A change in position due to active secretion on the part of the cells composing part or all of the animal, and an accumulation of this secretion in such amounts and in such a manner as to force the body forward. (d) A change in position due to expansions and con- tractions of part or all of the cell bodies composing the animal, these movements being so arranged as to produce a movement of the whole animal. The first cause needs little discussion, as it is due to no physiological action <3f the animals affected. The other three need some explanation to make their action clear. Locomotion due to changes in specific gravity. — Changes of position of this character are usually confined to water- dwelling forms. Protoplasm is slightly heavier than water, and an animal, therefore, which rests upon the bottom and has no apparatus for raising itself must re-296 MOVEMENT IN LOWER ANIMALS main there unless it can produce such changes in its com- position as to make it lighter than the water it displaces. Certain one-celled animals have the power to accumulate carbon dioxide within the cell bodies to such an extent as to decrease their specific gravity. Thus, the body be- comes lighter than the liquid it displaces, and the buoyant force of the water floats it upward. Further, they can discharge this gas, increase their specific gravity, and sink, as they have become heavier than the liquid displaced. In this way many one-celled, water-dwelling animals are able to change their position without the aid or use of special organs of locomotion. In the higher animals, Fig. 127 — Swim bladder of the fish. the swim bladder of the fish and the connection of the lungs of the bird with cavities of the body, by means of which air can be introduced into the body, are apparently parallel cases, but it must be noted that in these cases the decrease in body specific gravity is due to the action of special muscles and not to changes in the composition of the protoplasm of the cells. The result, however, is the same. Locomotion due to secretion. —There are certain forms among the one-celled animals and plants that are able to creep over surfaces without contractile movements, or any apparent organs of locomotion. Examination shows thatLOCOMOTION DUE TO CONTRACTILITY 297 in certain eases this creeping movement is due to active secretion. In one case which has been especially well studied (the one-celled gregarines), it has been shown that, out of the food supplied to it, the protoplasm of the cell secretes a slimy substance, just as the cells of the salivary glands produce saliva. This slimy substance is forced out of the cell body and accumulates at the rear of the animal. Here it sticks to the surface and hardens, and thus by its accumulation and resistance forces the body along slowly before it. In the higher forms of animals we find no cases of motion due entirely to secretion. In the snails, however, the body secretes a slimy mucus which forms a surface for the action of the muscular foot and which we see as a shiny trail when it hardens. The common house fly also secretes a sticky fluid on the bases of the foot, by means of which it is enabled to walk over smooth surfaces. The web of the spider over which it travels, and by means of which it raises and lowers itself, is a secretion of certain glands of the body. These cases are evidently not cases of motion due to secretion alone, but simply instances of where secretion aids other methods. Motion due to secretion alone is confined to a few one-celled forms. Locomotion due to contractility.— Practically all of the free-moving animal forms owe their ability to change position to the power of contractility exercised by the protoplasm of part or all of the cells of the body. We can distinguish three kinds of locomotion due to the contractility of protoplasm: (a) Amoeboid movements. (b) Ciliary movements. (c) Muscular movements.298 MOVEMENT IN LOWER ANIMALS Amoeboid movements. —In the study of the amoeba it was noted that this animal moves by means of pseudo- podia. These pseudopodia are merely extensions of the body protoplasm into which the rest of the protoplasm flowed. In other words, the amoeba extends its proto- Fig. 128 — Various shapes of an amceba in motion. plasm at a definite point on the surface, and then, by con- traction, forces the rest of the protoplasm to flow in that direction. The point where the pseudopods are pro- duced varies with conditions and, since the animal has no cell wall, theoretically these pseudopodia may be pro- duced anywhere. In general, however, they are so formed as to carry the animal toward food or away from irrita- ting causes, and this fact shows that the expansions and contractions are in some way a response to a stimulus. The result of this peculiar kind of contractility of proto- plasm is to produce a slow creeping motion, and all animals which move in this slow creeping fashion with the aid of pseudopodia are said to show amoeboid movement. In some of the forms of protozoans which have walls about the body, amceboid movement is still possible by extending the pseudopodia through openings in the walls. The little shelled form known as arcella and common in all fresh water is an example of such a type. (See Fig 130.) In the higher forms we find no cases of amceboid move- ment on the part of the animals as a whole; but often the cells of a part of the body may exhibit this method ofCILIARY MOVEMENTS 299 motion. In our own body, for example, the movement of the white corpuscles is a case in point. To this amce- Fig. 129 — White corpuscles escaping from blood tube by amoeboid motions. boid movement these corpuscles owe their power to pass through spaces in the walls of the blood vessels through which the smaller but rigid, red corpuscles cannot pass. Ciliary movements.— Amoeboid movement is at best a very slow method of changing position, and is interesting mainly in showing us that free protoplasm has the power of contractility. We should expect, then, to find in the higher protozoans a more rapid method of progression. Such a modification in motile organs we do find in many of the pro- tozoans, and these organs are called cilia. In the arcella, movement is produced by pseudopodia which are projected through an opening in the cell wall, and these can be extended and withdrawn as necessity requires. -Shell ’seudopod Fig. 130—An arcella creeping by pseudopod.300 MOVEMENT IN LOWER ANIMALS In the slipper animalcule or paramoecium (see Fig. 131), which is a higher type of protozoans than the arcella, the body wall is covered with permanent projections of protoplasm called cilia from their resemblance to hairs. These hairlike structures are composed of protoplasm like the pseud- opodia, but, unlike them, they cannot be withdrawn, and in contracting and expanding they move at a different rate and in only two directions. As it contracts the cilium is bent quickly and sharply backward; relaxing, it re- turns much more slowly to an upright position. Many cilia, acting in concert all over the body, push the body through the water just as oars do a boat. A peculiarity of ciliary action, however, is that they do not all contract at once, but the movement of one follows that of the one preced- ing in definite order, so that in action they resemble the waving of a field of grain in a wind, each cilium corresponding to a single stalk of the grain. Very rapid movement is made possible by these Fig. 131 — Paramce- cium ; c} cilia; cvr contractile vac- uole ; n, micro-nu- cleus ; m, macro- nucleus; g, gullet; /, food vacuole; z, modified cilia (tri- chocysts). Fig. 132 — Ciliary movement. structures, and their permanent form permits the body to be completely covered with a wall, and at the same time this rigid wall furnishes a resistant structure which the cilia can push through the water much more readily than would be possible if the body were plastic.CILIAKY MOVEMENTS 301 Sometimes the cilia are replaced by a single long pro- jection which performs movements of a whip-lash char- acter. Such modifications are called flagella, and are found Fig. 134—Colony of flag- ellated protozoans with Fig. 133 — A flagel- two flagella, lated protozoan ; F, flagellum; 2V, nu- cleus ; By body of cell. in many one-celled plant and animal forms. (See Figs. 133 and 134.) In many of the higher animals, including man, are found cells provided with similar, ciliary projections, but here the action of the cilia is not to produce movement of the cells themselves. On the contrary, their function here is to produce currents in the liquid that bathes them or to remove obstacles from the surface of the cells. Their action may be compared to the motion of the oars when302 MOVEMENT IN LOWER ANIMALS the boat is tied. In such case, the result is that the water, and not the boat, is moved. In many of the lower animals the digestive cavities are lined with such ciliated cells, and so the currents of water which they create cause the food, floating in the water, to be brought within reach by the currents set up. In our bodies, the walls that line the bronchial tubes are lined with such ciliated cells, and here they serve to free the surface of the membrane from particles of dust which might produce harm if they were allowed to remain. In fact, it is only in the case of the protozoans and a few of the young stages of the lower animals that cilia are used to move the body of the animal itself. In all other forms their function is the setting up of currents or the removal of particles. Cilia and flagella are to be con- sidered, then, as merely a modified portion of protoplasm in which the power of rhythmic contraction and expan- sion has been specially developed. They are chiefly interesting, therefore, as indicating the manner in which protoplasm can become developed into a motile organ. Muscular movement.— Muscle cells differ from ordinary cells in that they contain certain modifications of pro- toplasm called fibrillse. These fibrilla: are thin threadlike fibers embedded in the fluid protoplasm of the cell, and may be variously arranged as to direction, but are always par- allel to one another. In certain instances these fibrillse are divided into segments by cross lines or striae, and these Fig. 135 — Ciliated epi- thelial cells lining the bronchial tubes; a, basal membrane ; b, spherical epithelial cells; c, cili- ated cells ; , dorsal blood vessel; /, intestine; V, ventral blood vessel; N. V., ventral nerve cord. cavity or are absorbed into the blood circulation. Those wastes of a nitrogenous character which remain in the cav- ity of the segment pass directly into the ciliated end of one or the other of the nephridia, and, after action by the cells, pass out of the pore as a secretion. Similarly, the wastes which entered the blood are eventually brought to the branches which surround this tube, and, being absorbed from the blood by the secreting cells, are made into a secre- tion, and in turn pass down the duct and out the pore. InVERTEBRATE KIDNEYS 881 this way each of these coiled tubes absorbs wastes either from the body cavity or blood, transforms it into a secre- tion, and pours it into a duct and out the pore. The action is the same in every way as that of a single tubule and cap- sule of the kidney. (See Fig. 171.) The kidney of the clam is a coiled tube almost identical in its structural features with this nephridium of the earthworm, while the green glands of the crustacean are supposed to act in a similar manner. Kidneys of insects.— These animals show a modification of the nephridium plan in that the tubules are arranged in a ring about the intestine. The ends of the tubes float in the body cavity of this animal, and absorb the waste from that, while their mouths open, not to the outside, but into the in- testine, so that the nitrogen wastes pass out with the faeces through the anus. Vertebrate Kidneys.— In all the vertebrates we find some form of kidney. These kidneys diff er from a nephrid- ium mainly in that they are a combination of many secreting tubules under one covering, and they differ from that of man only in the manner in which the ducts termi- nate. For example, the ducts of the bird and reptile kidney open into an enlargement of the rectum (lower end of in- testine) called the cloaca, and in these forms the anus is Fig. 177 — Kidney tubes of an insect; k, kidney tubes; a, esophagus; b, crop; c, d, stomach; e, intestine; /, large intestine; g, anus.382 SKIN STRUCTURE AND EXCRETION the common external opening for the kidney and the intes- tine. In mammals the openings are for the most part dis- tinct, the kidney opening being called the urinary opening. The development of the bladder is simply a device for pre- venting continuous secretion and permitting the animal to control this operation of removal from the body. In general, then, whether we are dealing with the nephridium of the worm or the sweat gland of man, the means of disposal of waste are practically identical in action, and the kidneys are to be considered as simply a collection of separate excretory glands enormously increased in number to provide for the increase in waste, and having a common duct for removal.XXV. THE NERVOUS SYSTEM. Structure. The study of the preceding pages has taught us the part played by the various organs and structures in our body in the operation of tearing down and building up the human machine, and the manner in which these operations keep it in running shape. We have also seen that all these systems are interrelated, the work of one being directly dependent upon the activity of another, while in our life we are conscious of the fact that these many parts are all acting harmoniously and with comparatively little attention on our part. For example, in so simple an operation as walking, every system in our body is con- cerned. The muscles contract and expand, and in so doing produce motion, consume oxygen and nutriment, and give off wastes and carbon dioxide. The necessity for food must be met by a quickened flow of blood, while the in- crease in appetite warns us to supply the digestive system with more nourishment. The lungs mus.t expand and con- tract to furnish the blood its oxygen, and the excretory system must increase its activity a little to dispose of the wastes. Even in our sleep these systems are all more or less active, and the needs of one part of the bodjr can be met only by special effort on the part of all the others. Manifestly such a harmonious action would be utterly impossible unless the organs had some means of com- munication by means of which the needs of one might be 383384 THE NERVOUS SYSTEM Plate IY.— Nervous system.GENEKAIj COMPOSITION OP THE NEKVOUS SYSTEM 385 communicated to the others. Some central system of con- trol must also exist, in order that the various activities of the body may be directed by us. The system of organs in our body by means of which this direction and control are exercised is spoken of collectively as the nervous system. The relation of this system to voluntary and involuntary actions and the structure of the parts will be considered in the following paragraphs. General composition of the nervous system.— The ner- vous system may be said to consist of three parts: nerve centers, nerves, and nerve-end organs. The nerve centers in our bodies are the brain, the spinal cord, and little patches of nervous matter scattered in different parts of the body, called ganglia. The nerves arise from these centers, and with their branches penetrate all organs and tissues of the body. Finally, each nerve ends in some form of organ called an end organ. Some of these organs are very simple in structure; others, such as those at the ends of the sight and hearing nerves, are exceedingly complex. For convenience, the cord and brain, together with the nerves which arise in them, are spoken of as the central nervous system, while the sets of ganglia on each side of the spinal column, with their nerve connections, are called the sympathetic system. All nervous centers are, how- ever, in communication, and these two systems are sepa- rated in name for convenience of study, and not because of any true separation in the systems. In general, then, the brain and cord may be likened to the “ central ” of a telephone system, the nerves to the wires, and the end organs to the individual telephones. Such an arrangement makes it necessary that all communi- EDDY. PHYS.—25386 THE NERVOUS SYSTEM cations from one organ to another, or from one part of an organ to another, be made through the brain and cord. If we enlarge our idea of this arrangement and conceive of the brain and cord as not only switching messages from one part of the body to another, but also of being able to modify the character of these messages, we shall have a true picture of the manner in which the brain and cord control and harmonize the actions of all parts of our bodies. The nervous system of the frog. — A dissection of the ner- vous system of the frog is easily made, and since it is essentially similar in kind to ours, a study of this system forms a good in- troduction to the study of the human system. As in our body, the brain is inclosed in the skull, and is con- tinued downward as the spinal cord through the spinal cavities of the vertebra. In this posi- tion the system is well pro- tected from external pressure. If this bony covering be carefully removed so as not to injure the contents or its connections, a structure, such as shown in Figure 178 is seen. Fig. 178 — Nervous system of frog; a, cerebral hemispheres (fore- brain) b, olfactory lobes; c, eyes; d, midbrain; e, optic lobes; /, hindbrain (cerebellum); <7, medulla; h, i, spinal cord; I to X, first to tenth pair of cranial nerves; 1 to 10, spinal nerves.THE FROG’S BRAIN 387 The frog’s brain.— The brain may be divided into three parts, called respectively, the forebrain, midbrain, and hind- brain. From these parts of the brain are given off ten pairs of nerves called cranial nerves. These nerves are made up of fibers called nerve fibers. Those cranial nerves whose fibers bring impulses or messages to the brain are called afferent or sensory nerves. Those whose fibers con- Fig. 179 — Frog brain; /, top view; A, olfactory nerves; B, olfactory lobes; C, D. forebrain; E, midbrain; F, optic lobes; G, hindbrain; H, medulla; /, cord; J, K, L, M, N, nerves; II, ventral view; III, lateral view. vey impulses from the brain are called efferent or motor nerves. Several of these nerves contain both efferent and afferent fibers, and such are called mixed nerves.388 THE NEKVOUS SYSTEM The forebrain.— This consists of two elongated bodies lying side by side and called the cerebral hemispheres. Together, they constitute the cerebrum. At their ex- treme front are two swellings, from which a pair of cranial nerves extend to the nose. The fibers of these nerves are afferent, and carry sensoiy impulses from the nose to these lobes of the brain. They give to the animal the sense of smell. On this account the lobes of the brain from which these nerves arise are called the olfactory lobes, and the nerves are called the olfactory nerves. The olfactory are the only nerves connected directly with the cerebral hemispheres. The midbrain.— Just back of the hemispheres is the midbrain. This is covered on the top and sides by two swellings called the optic lobes, so named because the sensory nerves from the eye enter them. They are the receiving points of sensations of light. The optic, or sight nerves (the second pair of cranial nerves) enter these lobes on the under side, and are crossed so that the nerve from the right eye enters the left lobe, and vice versa. These optic lobes also communicate with the forebrain by means of which we will speak later, but the optic nerves have no direct connection with any part of the brain except these lobes. The third pair of cranial nerves also arises from the midbrain, and, unlike the first two pairs, it carries impulses from the brain instead of to it (that is, it is a motor or efferent nerve). The nerve fibers of this third pair control four of the six muscles which move the eyeballs and also some of the internal eye muscles. They are called the oculo-motor nerves. The hindbrain.—The rest of the brain, which extendsTHE SPINAL CORD OF THE FROG 389 backward to the beginning of the spinal cord, is called the hindbrain. It shows two regions. The transverse ridge just back of the optic lobes is called the cerebellum, while the wedge-shaped portion back of this, from which the spinal cord arises, is called the medulla oblongata. Seven pairs of cranial nerves arise from this medulla. The fourth pair, known as the trochlears, are very small, and carry mo- tor impulses to one of the muscles which move the eyeballs. The fifth pair, or trigeminals, are very large. They belong to the class of mixed nerves. Some of their fibers bring sensory impulses from the teeth and face to the brain. Other fibers carry motor impulses to the muscles of the jaws and eyelids. The sixth pair, or abducents, carry mo- tor impulses to a muscle of the eyeball. The seventh pair, like the fifth, are facial nerves of a mixed character, and carry impulses to and from certain parts of the face. In the higher animals and in man this nerve is purely a motor nerve, and controls the muscles which give expression to the face. The eighth pair, or auditory nerves, are sensory, and carry impulses of hearing from the ears. The ninth pair, or glossopharyngeal, supply the tongue and pharynx, and are mixed nerves. The tenth pair, or the vagus nerves, have the widest distribution of any. They are also mixed nerves, and their fibers supply the lungs and air passages, the heart and blood vessels, and the organs of digestion. Finally, the centers of all these cranial nerves are in communication with parts of the forebrain, and the con- sciousness of sensations has its seat in that organ, though the nerves are not directly attached to it. The spinal cord of the frog.— The cord of the frog, which is merely an extension of the medulla, is much shorter than that of man. From it are given off some ten pairs390 THE NERVOUS SYSTEM of spinal nerves whose fibers are distributed to various parts of the body. . Figure 178 illustrates the method of branching of these nerves. Comparison of the frog’s spinal cord and brain with that of higher forms.— The spinal cord and brain are to be con- sidered as one structure, the brain lobes and swellings being simply modifications of the more uniform spinal cord. These modifications become much more marked in the higher forms, and instead of lying directly behind one an- other the increase in size and the lack of space compels these brain lobes to become more or less twisted and doubled upon one another. If we keep the parts of the frog’s brain in mind, however, we shall have no difficulty in recognizing the homologous parts of the human central system, as, aside from the folding and variation in size, they are almost iden- tical, part for part. Let us now examine the human brain with this com- parison in mind, and see if we can identify the structures found there.1 The human brain.— The brain of man is inclosed in that part of the skull called the cranium, and it fills this part completely. At the base of this cranium is an opening called the foramen magnum through which it is continued downward as the spinal cord. Various apertures in this bony box permit the entrance and escape of the cranial nerves. Brain membranes.— When the bony plates of the cra- nium are removed, the brain is found to be covered with three membranes. The outermost (the dura mater) has 1 The brain of the sheep is easily obtainable, and is so nearly identical with that of man in structure that it serves well for laboratory work. (See Ex. LXIII.)BRAIN MEMBRANES 391 a roughened outer surface which fits closely to the inner surface of the bones, and forms the inner periosteum of these bones. The brain side of this membrane is smooth, Fig. 180 — Human brain in place; C, C, wrinkled surface of the cerebrum; Cb, cerebellum; M.Ob, medulla; B, bodies of vertebrae, and 6', their spinous processes inclosing the spinal cord. and the whole membrane is thick and fibrous. It lines the whole cranial cavity, and ingrowths from this membrane support and invest all parts of the brain. At the foramen magnum it is continued as the outer coat of the cord. The blood vessels which supply this membrane are, for the most392 THE NERVOUS SYSTEM part, distributed on the side next to the bones. In fact, their principal function Is to supply nourishment to the bones, not to the brain. A second membrane, called the arachnoid, fits against the smooth inner surface of the dura mater. This is a delicate, transparent, serous membrane. Beneath it, enveloping the brain itself, is a third, thin membrane, called the pia mater. This pia mater is composed of a fine network of small blood vessels held together by tissue, and these blood vessels supply the nourishment and remove the wastes of the brain itself. It is practically impossible to remove this membrane, as it covers all the folds of the brain. These three membranes and the cranial bones furnish a very efficient protection for the delicate matter of the brain proper. The brain proper.— Like the frog brain, it consists of three portions, forebrain, midbrain, and hindbrain, but these parts are so bent and doubled upon one another that they form a figure similar to an interrogation point (?). The two hemispheres of the forebrain occupy the bulk of the cranial cavity and correspond in position to the top, front, and part of the back of the curved portion of the interrogation point. The olfactory lobes are tucked away at the very front of the curve. In man, the forebrain con- stitutes nine tenths of the bulk of the entire brain. The midbrain, with its optic lobes, is projected forward in a position corresponding to the curve at the top of the stem of the interrogation point. When viewed from the under side, this projection brings the optic lobes just back of the olfactory lobes. The hindbrain has a much more developed cerebellumEXTERNAL FEATURES OF THE HUMAN BRAIN 393 than the frog’s has, and this organ is here divided into two lobes by a longitudinal partition. Underneath the cere- bellum, and extending downward, is the wedge-shaped medulla. In comparison with our figure the hindbrain occupies the position of the stem of the interrogation point, and if this were continued downward the extension would represent the spinal cord. External features of the human brain.— Viewed from the top, the two hemispheres of the forebrain (the cere- brum) are seen to be much convoluted and folded, thus increasing then surface without increasing the space occu- pied. The two hemispheres are pressed closely together, the line of separation being marked by a fissure (the longitudinal fissure). Viewed from the bottom we can make out the con- tinuation of this fissure back to the midbrain, and on each side of it appear the olfactory lobes. (See Fig. 182.) Just back of these lobes are the crossed optic nerves (the optic chiasma) which enter the optic lobes. Back of this chiasma is a band of white matter (the pons varolii) which connects the two lobes of the cerebellum and covers the top of the medulla. Underneath this pons, the medulla divides into two stalks (the crura cerebri) which spread out to form the cerebral hemispheres,394 THE NERVOUS SYSTEM The cerebellum shows a wrinkled surface different in appearance from the folded cerebrum. Between its lobes Olfactory Bulb---- {to which is attached the Olfactory Nerve) Pituitary Body____ Optic Nerve---- Optic Chiasma----- Oculomotor Nerve-—^ Trochlear Nerve--- Trigeminal Nerve--- Pons Varolii--- Abducens Nerve- - Facial Nerve— Auditory Nerve- __ Glossopharyngeal Nerve__ Vagus Nerve— Spinal accessory Nerve— Hypoglossal Nerve- Medulla Oblongata--- First Spinal Nerve— Cerebellum— Spinal Cord— Second Spinal Nerve— Fig. 182 — Human brain viewed from below. is seen the lower part of the medulla whose lower end nar- rows and is continued to form the spinal cord. Along the under side can also be made out twelve pairs of cranial nerves, two more than in the frog’s, brain. The first ten pairs are connected to the same regions as those of the frog and have the same functions. The eleventh and twelfth arise from the medulla and are known respec- tively as the spinal accessory and hypoglossal. The spinal accessory is a motor nerve and supplies the muscles of the shoulders. The twelfth, or hypoglossal, is also motor andINTERNAL FEATURES OF THE HUMAN BRAIN 395 supplies the muscles of the tongue. The first spinal nerve of the frog corresponds to this nerve in action. Internal features of the human brain.— If we split the brain in two by a vertical cut, parallel to and a little to the left of the longi- tudinal fissure, we can make out the following parts on examining the right side. (See Fig. 184.) The fissure is seen to con- tinue downward to a white fibrous body which binds the two hemispheres to- gether. This is called the corpus callosum. This band curves in such a way as to inclose a space actu- ally outside the brain and often called the " fifth ” ventricle. This fifth ven- tricle is further inclosed by two thin membranes (the septa lacida). The forward curve of this connecting body below the fifth ventricle forms the roof of the midbrain, and this part is called the fornix. Under the fornix is the cavity of the midbrain or the third ventricle. This third ventricle communicates with cavities in the right and left hemispheres (the second or lateral ventricles) by two openings in the fore part (the foramen Monroe). In the figure, the opening into the right ventricle only appears. The midbrain cavity is also extended forward in a funnel- shaped tube called the infundibulum. Fig. 183 — Origin of cranial nerves; H, hemispheres; C. S. corpora striatum; P, pineal body; Pt, pituitary body; C. Q. cor- pora quadrigemina; Cb, cerebellum; Thy optic thalamus; M, medulla; / to XII, the pairs of cranial nerves; Sp, 1, 2, spinal nerves.396 THE NERVOUS SYSTEM The lateral ventricles in man are frequently named the first, and second ventricles. Extending from one side to the other of the third ven- tricle is a solid cord called the median commissure. The sides of the cavity are formed by the optic lobes, or thalami, Foramen of Monro „ „ \ Third Ventricle Corpus Callosum \ , Pineal Bod,J \ Fornix \ \ \ Cerebrum -Corpora Quadrigemina Cerebellum Optic Chiasma Pituitary Body / Oculomotor Serve / Tons Varolii Aqueduct / / Medulla Oblongata / Fourth Ventricle f-----Spinal Cord Fig. 184 — Vertical section of brain. and its floor is formed by the crura cerebri, or stalks, of the cerebral hemispheres. We have already noted that these stalks are the forward extensions of the medulla. At the rear of the third ventricle is a passage called the iter, or aqueduct, leading to the fourth ventricle, a small cavity in the hindbrain. The top and sides of this iter are formedINTERNAL EEATDRES OE THE HUMAN BRAIN 397 by four little bodies called the corpora quadrigemina. Its floor is the medulla and the pons Varolii. The fourth ventricle projects upward toward the cerebellum, and the internal structure of this organ shows a branching, treelike core of white matter (the arbor vitce) surrounded on the outside by gray matter. The lobes of the cerebellum are only partly separated, the fissure dividing them being very shallow. The floor and sides of the fourth ventricle are formed by the medulla, and this cavity is continued backward as the neural cavity of the spinal cord. From the above description it will be seen that the brain is actually a hollow organ with a continuous cavity surrounded by various outgrowths or walls and398 THE NERVOUS SYSTEM connecting all regions of the brain with the neural cavity of the spinal cord. In life this cavity (the ven- tricles) is filled with a liquid known as the cerebrospinal fluid and the entire cavity is lined with a layer of ciliated epithelium. The third, fourth, and lateral ventricles are also supplied with blood from a net- work of blood vessels called the choroid 'plexus. Rupture and hemorrhage of the blood vessels of the lateral ventricles pro- duces pressure upon the inside of the hemispheres and is one cause of apoplexy. If we cut slices at right angles to the longitudinal fissure in either hemisphere, we find that, like the cerebellum, the solid part of the cerebral hemispheres is composed of two kinds of matter, white and gray, surrounding a central cavity (the lateral ventricle). The out- side of the hemispheres and folds is called the cortex and is composed of gray matter; the white matter forming the core of this part of the brain. The human spinal cord. — This cord, which is an extension of the medulla, extends from the foramen magnum through the spinal cavities of the ver- tebrae to the articulation of the first and second lumbar vertebrae. From here it narrows off into a slender filament which runs back to the end of the neural canal behind the sacrum. It is nearly cylindrical in form, being slightly flattened Fig. 186 — The human spinal cord and its branches.THE HUMAN SPINAL COED 399 dorso-ventrally, and has an average diameter of three quarters of an inch, with a length of about seventeen inches. It is largest in the neck region from the third to the first dorsal vertebrae, and there is a second expansion Fig. 187—Cross section of spinal cord; 1, anterior fissure; 2, posterior fissure, 3, central canal; 4, 5, commissures; 9, posterior root; 10, anterior root; a, e, gray matter. The outer layer is the pia mater, covering the white matter and send- ing into it blood vessels. in the last dorsal vertebra. Externally, it is covered with three membranes like the brain: the dura mater, arachnoid, and pia mater. The dura mater is not in close contact with the bones,400 THE NERVOUS SYSTEM as in the brain, but is separated from their periosteum by a layer of fat and a network of blood vessels. It does not send partitions into the cord as it does into the brain. The arachnoid is also not in close contact with the pia mater, but forms a loose bag around it. The pia mater closely invests the surface of the cord and, as in the brain, supplies it with blood. The cord itself shows two fissures, a wide shallow anterior one and a narrow deep posterior one, thus dividing it into two right and left halves. The posterior fissure is not a true groove but rather a partition of connective tissue. Each half of the cord gives off at intervals, from the top and bottom, nerve roots which unite at a short distance from the cord into common nerve trunks. (See Fig. 188.) These trunks or spinal nerves are arranged in pairs in the cervical and dorsal regions and emerge through openings between the vertebrae. At the lower end of the cord they are crowded together in parallel bundles. There are, in all, thirty one pairs of these nerves. Each spinal nerve trunk splits up into man)^ smaller nerves and these into still smaller nerves which are distributed to various partsINTERNAL STRUCTURE OF THE CORD 401 of the limbs and trunk, thus bringing all parts of the body in communication with the cord. They are also connected on each side of the cord with the system of nerve centers called the sympathetic ganglia. (See Fig. 189.) Internal structure of the cord. — If we sec- tion a portion of the cord in such a manner as to split a pair of spinal nerves (see Fig. 188), we can make out the following de- tails: Like the brain, the cord is seen to consist of white and gray matter with this difference, that, in the cord, the white matter is on the outside. The inner gray matter is in the form of a letter H. Two horns of this II project forward (the anterior cornua) and two backward (the pos- terior cornua). The cen- tral part, correspond- ing to the cross bar of the H, shows a small cavity in its center which is the continuation of the ventricle cavities of the brain. From the posterior and anterior cornua emerge nerve fibers which unite to form the roots of the Fig. 189 — Roots of a dorsal spinal nerve and its union with the sympathetic system; c, c, anterior fissure of cord; «, anterior root; /», posterior root with spinal ganglion; s, sympathetic, and e, its double connections with spinal nerve a' by a white and a gray filament. EDDY. PHYS. — 26402 THE NEEVOTJS SYSTEM spinal nerves. The places where these roots emerge appear as four grooves on the surface of the cord. The posterior roots show a swelling just back of the point where they unite with the anterior fibers. This swelling is called the spinal ganglion. Experiment has shown that the anterior roots are composed mainly of efferent or motor fibers and convey motor impulses to the muscles, while the fibers of the posterior root are afferent or sensory and bring sensory impulses to the cord. The spinal nerves which contain both kinds of fibers are, therefore, mixed nerves. Structure of a spinal nerve trunk and fiber. —If wre cut a cross section of a spinal nerve we find it made up of many bundles of nerve fibers. The entire trunk is sur- rounded with a sheath of connective tissue. Inside this sheath, each bundle of fibers is wrapped in a sheath of con- nective tissue called the 'perineurium. We may remove this sheath and separate the bundles into the nerve fibers themselves. In these ultimate nerve fibers, thus separated, we have the actual conductors of impulses, and our dissection shows that the spinal nerve may aptly be compared to the cables of a telephone station where each cable consists of bundles of separate wires, both bundles and wires being care- fully insulated from one another. Examination of a single nerve fiber shows it also to be composed of layers. In the middle is a core, or axis, com- posed of modified protoplasm. This is called the axis cy- linder, and it is this cylinder which transmits the impulse. Surrounding this is a sheath called the medullary or myelin sheath. This sheath varies greatly in thickness in different fibers and is often segmented, forming the so-called “ nodesSPINAL NERVE TRUNK AND FIBER 403 of Ranvier.” Outside this myelin sheath is a thin mem- brane called the neurilemma, and comparable to the sar- colemma of the muscle fiber. Lying under the neurilemma are found nuclei, one for each segment of the medullary layer. These two outer layers of the fiber do not conduct impulses, and while their function has not been absolutely determined, they may be compared to the insulation which surrounds an electric wire. The axis, evidently, is the wire. In some nerve fibers the medul- lary layer is lacking, the axis being sur- rounded by the neurilemma only. All nerve fibers originate either in the nervous system or in the sympathetic ganglia. Those from the former are all medullatcd; some of those of the latter are non-inedullated. All fibers admit also of classification as afferent or efferent, since, in a given fiber, the direction in which the impulse travels is always the same. Mixed nerves, therefore, are composite bundles of afferent and efferent fibers. The words afferent and efferent simply indicate the di- rection in which the impulse travels. Owing to the fact that efferent fibers all carry impulses which produce motion, the words efferent and motor are equivalent in describing a given fiber, while for the same reason the words afferent and sensory may be interchangeably applied to fibers which convey sensory impulses. Finally, the two classes of fibers may be subdivided again on the basis of the effect of their impulses. For _ Node of Ranvier \\-Neurilemma Neuraxon ~~or Axis Cylinder Medullary ~~Sheath Fig. 190—Section of a nerve.404 THE NERVOUS SYSTEM ■Solar Plexics example, a motor fiber transmits an impulse which may retard or increase a motion, wliile a sensory fiber transmits impulses which may increase or decrease sensation. Those fibers, either motor or sensor}', which excite to greater motion or sensation, are called excitatory. Those which decrease motion or sensation are called inhibitory. This inhibition or excitation is supposed to be determined, not by the nature of the im- pulse, but by the tissue in which the fiber ends. The plexus.— The fact just noted, that the spinal nerves are composed of bundles of fibers, explains how it is pos- sible for them to split up into smaller nerves and ultimately into separate fibers, just as we may unravel the strands of a rope. In many regions of the body some of these unravelled nerve fibei’S from several spinal or ganglionic nerves interlace to form a meshwork, and such networks are called plexuses. In this fusion, new combinations of fibers are often made, and nerves extending from a plexus may con- tain fibers from several different spinal or ganglionic nerves. Evidently the stimulation of such a nerve will result in messages to several different parts of the central system, and vice versa. For example, the nerves supply- ing the legs arise from large plexuses, and are thus supplied —'Abdominal Aorta —L. Sympathetic Chain 1—It. Sympathetic Chain Hypogastric Plexus r^JJiao A. Fig. 191 — Tlie solar and hypogastric sympathetic plexuses.CELL THEORY OF NERVE STRUCTURE; NEURA 405 with fibers from many different parts of the central sys- tem. The advantage of such a combination is seen in the manner in which a single nerve is able to coordinate very complex movements, the messages from the different cen- ters of control being concentrated, as it were, by this arrangement. One of the largest plexuses is located just above the pit of the stomach, and is called the solar plexus because smaller plexuses radiate from it like the rays of the sun. A blow on this area paralyses many regions, and may even produce death. Cell theory of nerve structure; neura. — The presence of nuclei under the neuri- lemma at once suggests the idea that nerve fibers bear some relation to cells. Examination of the gray matter of the cord or brain in which these fibers orig- inate gives us further information on this point, and to the relations thus established is due the present theory of nerve structure and action. Preparations of gray matter show it to be rich in nucleated cells of irregular outline. (See Fig. 192.) From the angles of this outline project fine branches of protoplasm, which may subdivide again and again to form treelike growths, or which may continue some distance with- out branching, in the form of a cylinder. The branching outgrowths are called dendrites, while the cylinders are called axis cylinders, or Fig. 192 — Nerve cells from the cortex of the cerebral hemispheres; Ax axis cylinder. axones.406 THE NERVOUS SYSTEM Nerve cells, wherever found, agree in this general struc- ture. In some the production of dendrites is much less than in others. Some also have one axon only, while others may have from two to several. Those cells with one axon arc called unipolar, with two, bipolar, and with more than two, multipolar. If we trace the course of one of these axones we find that it may extend a long distance from the parent cell with or without branching, and, what is most important, such ax- ones are the axis cylinders of the nerve fibers. We see then that a nerve fiber is to be considered as an elongated, axis cylinder, or axon, which has become cov- ered with one or more pro- tective sheaths. When the axis cylinder branches, these branches are called collaterals, while the terminal branches in which they all end are called the terminal brushes. A nerve cell with its dendrites, axones, sheaths, collaterals, and terminal brushes, is spoken of collectively as a neuron. Such a unipolar neuron is shown in Figure. 193. Theory of neuron action. — The action of an amoeba free axis cylinder; b, axis cylinder surrounded by neurilemma alone; c, axis cylinder surrounded by medullary sheath alone; d, axis cylinder surrounded by the sheath and neurilemma and divided into segments (by constrictions called the nodes of Ranvier).THEORY OF NEURON ACTION 407 when touched with a needle point demonstrates that protoplasm is capable of responding to stimulation. Since the result is the same no matter where the protoplasm is touched, it follows that it is also capable of conducting impulses thus set up. In other words, responses to certain forms of stimulation and conduction of impulses is a proD- erty of protoplasm. When we examine the cytoplasm of a nerve cell we find it to be made up of two kinds of proto- plasm. One kind resembles closely that of the amoeba. The other kind seems to be in the form of threadlike fibrils. The axon protoplasm is almost purely of the fibrillar kind. This fibrillar protoplasm is found to be able to transmit impulses much more rapidly than ordinary protoplasm. In other words, it is protoplasm with a highly developed power of conduction. The discovery of this structural peculiarity of nerve cells enables us to think of them as cells whose protoplasm has been specially developed for conducting impulses. Keeping in mind this fact, and also the structure rela- tions in a neuron, the formulation of the theory of neuron action may be expressed in the following laws: First law. An impulse may be started wherever there is protoplasm, but externally stimulated impulses have their origin usually in the terminal brushes of the axones or in the dendrites, while the internally stimulated im- pulses arise in the body of the nerve cells. In general, then, the nerve fiber or axon is not a source of impulses, though in cases of inflammation of the sheaths it may’ become so. Second law. An impulse started in any part of the protoplasm of a neuron is conducted by the fibrillar pro- toplasm to every other part of the neuron protoplasm.408 THE NERVOUS SYSTEM Thus, an impulse started at the end of an axon is trans- mitted directly to the protoplasm of the cell body and through that to the extreme ends of the dendrites and any other axones that may arise from that cell body. Similarly, an impulse arising in the dendrites or cell body is likewise transmitted over the axones to their termina- tions. Bearing in mind that impulses usually originate in one of three places, the cell body, the dendrites, or the ter- minations of an axon, it will follow that the principal func- tion of the nerve fiber is conduction. This characteristic is still further evidenced by the fibrillar character of the axon protoplasm. When two points are connected by two neura the arrange- ment is as follows: The dendrites of one neuron are so placed as to be brought in contact with the terminations of the axon of the second. In this relation the course of the impulse takes one of two directions, and this direction may be formulated in the third law. Third law. An impulse which originates in the termi- nations of the axon of one neuron, and has been conducted to the dendrites of the nerve cell of that neuron, may pass from those dendrites into the terminals of the axon of the second neuron, and thus on to the dendrites in the final cell of the series. Similarly, an impulse originating in the cell body or dendrites of one neuron may pass, by way of the axon terminations, into the dendrites of the connecting neuron, and thus on to the final axon termi- nations of the series. These three laws enable us to explain the method of impulse transmission in all forms of nerve action; while neura may differ in the number of axones, in the length of the axones, and in location, the laws of action given aboveDISTRIBUTION OE NEURA 409 hold true for all classes. In connection with the third law of action it may be noted that whenever there is a passage of impulse from one neuron to another there is at this point a delay in the transmission, and sometimes several impulses must reach a cell before it can overcome this resistance between dendrites and axon contacts and discharge into the next cell. We may compare this delay to the effect of placing a small barrier in the path of a stream. The effect is not to stop the stream, but to check it until the volume of water is great enough to push aside the barrier. The normal rate of nerve discharge is about ten impulses per second. It will be noted finally that the above laws give no explanation of the manner in which nerve impulses origi- nate. The manner in which externally stimulated im- pulses originate will be considered in our study of the sensory end organs. The manner in which impulses originate in the cell body is still unknown. It is believed, however, that just as oxidation of fuel and metabolism account for the contraction of muscle protoplasm, so these nerve impulses set up in the nerve cells result from certain chemical changes in the composition of the cell proto- plasm. The fact that nerve cells require food and become fatigued by overwork justifies this belief, but the exact nature of the metabolic changes involved is unknown. Distribution of neura.— In the central nervous system, the cell bodies of the neura which originate there are found mainly in the gray matter of the cord and in the cortex of the brain. The axones of these cells may extend far beyond the limits of this system, or again they may be confined within the central system. The spinal and cranial nerves represent bundles of axones which410 THE NERVOUS SYSTEM originate from these centra] system cells and connect them with more or less distant parts of the body. Many of the axones found in the white matter of the cord and brain, however, never leave the central system, and the function of these is to connect the brain cells with the cells of the cord or with those of other parts of the brain. In general, then, the gray matter of the central system consists mainly of nerve cells, while the white matter is largely composed of nerve fibers. Each cell gives out from one to several axones, and the entire nervous system may be thought of as very complicated network of inter- connected neura.XXVI. THE NERVOUS SYSTEM (continued). Function. From the nervous system we gain control of the organs of the body, and to its action we owe our power to adjust ourselves to our surroundings in general and to maintain relations with other human beings and objects. In securing to us this adjustment to our envi- ronment, the nervous system manifests many forms of activity. For example, the interconnection of the fibers permits the transmission of messages of control and super- vision. The end organs of various forms give rise to special kinds of impulses, and when these are transmitted to the centers, the structures found there are able to trans- late them into sensations and thus furnish information in regard to our surroundings which is of vital importance in determining the character of the control impulses. Finally, the nerve centers are the origin of those mental operations which we call will, emotion, thought, etc. In short, the verjr individuality of a man is a function of his nervous system. The explanation of the many actions which we group under the common name of mind activities is properly the province of a science called psychology. In the study of the physiology of nerve action we are concerned only with the changes which take place in the nervous ele- ments of structure in giving rise to these operations and the manner in which these changes occur. Owing to the 411412 THE NERVOUS SYSTEM delicacy of the nervous structure and the difficulties in the way Of experiment our physiological knowledge of the nervous system is extremely limited, and, in fact, much has still to be learned of the physiology of nerves and nerve cells. In the following pages we shall simply try to indicate a few of the facts which have been deter- mined up to the present time, the manner in which such knowledge has been acquired, and its bearing upon our common activities. Action, voluntary and reflex.— The action of every organ of our body is dependent upon nerve control. The simplest division that can bo made of the actions of the body separates them into tvro classes. Those which require the exercise of the will and are associated with consciousness are called voluntary actions. Those which take place without any effort of will and may or may not be accompanied by consciousness we call reflex. All such actions as the picking up of objects with the fingers and the avoiding of obstacles in walking, in short, all actions which involve a conscious choice on our part belong to the voluntary type. All such actions as the regulation of the beating of the heart, the withdrawal of the hand from a hot object, etc., in which the action takes place without any consciousness of choice on our part, belong to the class of reflexes. The relation which consciousness may bear to such actions is brought out in the two examples of reflex action cited. In the beating of the heart consciousness is entirely lacking; while in the withdrawal of the burned hand we may be conscious of the pain and the move- ment, but ordinarily the movement has occurred before the pain is felt, and hence, in this case, while conscious-THE KEFLEX AKC 413 ness may accompany the action, it does not determine the action. Many actions which are at first voluntary may become reflex by repetition; and, in fact, one of the funda- mental features of education is concerned in making reflex or habitual, actions which in the child are voluntary and require effort of will on his part. The reflex arc.— The spinal cord is the seat of most of the reflex actions of the organs of the trunk and limbs. The relation of the nerve elements in simple reflexes which have their origin in the cord, is shown in Figure 194, and the relation there shown is typical of all reflex actions. In this figure it will be seen that the posterior root of the spinal nerve has on its surface a swelling which was called the spinal ganglion. This ganglion contains many bipolar nerve cells whose axones extend in different directions. One axon passes out and forms one of the sensory fibers of the spinal nerve. It terminates in some form of an end organ. The other terminates in a brush in the gray matter of the cord. Likewise, in the anterior cornua of the cord is found a unipolar nerve cell whose dendrites are in414 THE NERVOUS SYSTEM contact with the terminal brush of the sensory fiber. The axon of this nerve cell passes out through the anterior root and becomes one of the motor fibers of the spinal nerve. It terminates in a muscle. If, now, the sensory end organ of such a system is stimu- lated, as when the finger is brought in contact with a hot stove, the impulse passes up over the axon or fiber to the ganglion cell and through that is transmitted to the ter- minal brush of the other axon in the cord. From this brush it passes into the dendrites of the motor cell, and this cell in consequence discharges an impulse over its axon to the muscle and causes that to contract. In this simplest form of reflex action only two neura are involved; one neuron conveying the sensory impulse and the other the motor impulse. Most of our reflexes are more complex than this, but their com- plexity consists in the fact that they involve more than two neura and not in any essen- tial difference in ac- tion. The manner in which a single sensory impulse may stimulate several motor fibers is seen in Figure 195. Here the sensory axon splits up in the cord into collaterals, and each collateral dis- charges through a different motor cell. Such an arrange- ment explains how several muscles can be refiexly thrown into action by a single sensory impulse. It must be noted, Fig. 195 — Diagram showing how one sensory nerve may stimulate several motor nerves in a simple reflex.THE VOLUNTARY ARC 415 however, that the production of the impulses is essentially the same as in the simpler case. The reflex arcs of the cranial nerves are similar in action to those of the cord, and involve sensory and motor neura. Inasmuch as in this action the impulse travels in a circuit, the name of reflex arc has been given to such arrangements as above. The voluntary arc.— When we come to examine what takes place in a voluntary action we find that the process is not very different from that of the reflex action, except in the time required for transmission. Figure 196 shows a typical voluntary arc. In such a circuit the path of the416 THE NERVOUS SYSTEM impulse is as follows: End organ to cord, cord to brain centers by connecting neura, brain centers to other brain centers by other connecting neura, and finally back from the last brain center to the motor cell in the cord and over its axon as a motor impulse to the muscles. In such a cir- cuit it often happens that the cord motor cell is so near the sensory fiber brush that a division of the discharge is made, part of the impulse passing over the long circuit noted above, while the other part passes directly over to the motor cell and thus establishes a short-circuited reflex. In traveling the long circuit to the brain no motor discharge can take place until the impulse is returned from that region. Such a circuit is characteristic of all deliberate actions. In the case of the division of the impulse, while part of the im- pulse may continue on to the brain and produce conscious- ness, the short circuited impulse is able to produce the action perhaps even before consciousness is aroused. In the case of the burned finger the importance of this latter arrangement is evident in regard to speed of action. This short circuiting also illustrates how a voluntary action may become reflex, for if gradually the bulk of the discharge follows the shorter circuit the less necessary becomes the long circuits, and it may ultimately be abandoned entirely. In general, then, a voluntary circuit differs from a reflex circuit only in the length and in the number of neura involved. The tracing of all these connecting neura and their distribution in the central nervous system forms a part of the study of the nervous system in which much work still remains to be done. Almost nothing is known as yet about the character of the changes which take place in the brain cells and the manner in which past experiences can be stored and modify the nervous discharges. BySPINAL COED 417 experimenting with lower animals, and by the study of human beings whose nerve tracts have become diseased in part, it has been found possible to locate the tracts where sensory impulses are translated into consciousness, and also the points of origin of control impulses for different organs. Localization of functions in the central nervous system.— Such experimental studies as noted above have demon- strated that a sensory impulse produces no conscious sensation unless it reaches a certain area of the brain, and if this area is destroyed and the transmitting fiber with its end organ preserved intact, no sensation results. Sim- ilarly, it has been found that certain organs and parts of the body are under the control of definite areas of the brain and cord, and these areas must be in a healthy con- dition or the action of these organs is seriously interfered with. The determination of these areas of sensation and control is called the localization of functions, and the method by which results have been obtained is largely through experiment with lower animals. Spinal cord.— If we cut the spinal cord of the frog just back of the medulla, the frog will continue to live and the heart will continue to beat, but he ceases to breathe or swallow. The animal lies flat on his belly and loses all power of voluntary movement or conscious sensation. He is still able, however, to respond to external stimuli with definite movements. Thus, if we irritate the skin, the animal will make definite and skilful movements to re- move the irritating body; if we pinch the toe, the foot is withdrawn. From these and similar experiments it has been conclusively demonstrated that the cord is the seat of those reflex movements which concern responses to EDDY. PHYS.—27418 THE NERVOUS SYSTEM externally applied stimuli, and is also the path of sensory impulses to the brain. In higher animals similarly treated this reflex response is less apparent, and indicates that in such animals the reflex action of the cord is more depen- dent upon control from higher parts of the system. Cerebellum.— Experiments upon the cerebellum of the pigeon develop the following facts concerning this organ. When all the cerebellum is removed the animal can still execute voluntary movements, but these move- ments instead of being definite and coordinated are sprawl- ing, uncertain, and throw the animal into all sorts of grotesque positions, while it is totally unable to fly. From these experiments and others upon higher forms it appears that while the cerebellum does not originate motor im- pulses it does have the power to regulate and coordinate these impulses, and is thus responsible for orderly move- ment. An animal still retains sensation when the cere- bellum is sliced away. Cerebrum.— In the cortex of the cerebral hemispheres is the seat of all the psychical reactions which we include under the name of intelligence, memory, will, and the emotions. It is also the seat of conscious sensation, and in it originate the impulses which produce voluntary move- ments. Involuntary movements or reflexes may, how- ever, take place when these hemispheres are removed. A pigeon whose hemispheres are removed becomes a stupid, drowsy creature, which, when thrown into the air, recovers its equilibrium and flies, and it may even be made to feed by properly applied external stimuli. If starved, how- ever, it becomes restless and pecks aimlessly at the ground, but does not seem to be able to locate or recognize food for itself. In other words, it has lost all those responsesHISTORY OR CEREBRAL LOCALIZATION 419 which depend upon memory of past experiences, that is, intelligent responses. It shows no signs of fear or pleasure, and while it responds to sensory stimuli, these responses are purely reflex and not accompanied by true conscious- ness. History of cerebral localization.— When it was estab- lished that the cerebral cortex was the seat of psychical activities, the question at once arose as to whether all parts of the cortex were equally capable of exercising these faculties, or whether different parts controlled specific faculties. The idea of separate areas for separate func- tions was first presented by Franz Joseph Gall, who divided the cortex into areas of special activity, and put forth the theory that the more developed any mental quality was the larger and more prominent became the cortical area which produced it. Further, since the cortex fits closely to the cranium, the relative prominence would be indicated externally by the shape of the skull. From this position arose the practice of phrenology or the determination of mental qualities by examination of cranial prominences. Opposed to the position of Gall was the view of Flourens, who held that all parts of the cortex were capable of pro- ducing all kinds of mental qualities, and that when one part was removed the remaining parts supplied the quali- ties originally centering in that part. Without going into this controversy we may present the modern views on the subject as follows: First. The general view that functions are localized has been definitely established, and it is possible to map the cortex of the brain and thus indicate areas which, when stimulated, will produce definite actions. Second. The idea of Gall that the more marked the420 THE NERVOUS SYSTEM development of a function the larger and more prominent the area has been proved false. Third. The interconnection of areas has been shown to be so intimate that, although normally specific areas control specific functions, an injury to one part may affect all the others. In other words, the cerebrum is composed of many organs intimately associated with one another and inter- dependent. In general, the left hemisphere controls the Fig. 197 — Diagrams illustrating localization of functions in the cerebrum; /, outer surface of left hemisphere; II, inner surface of same. Motor areas are shaded; Cc, corpus callosum; Fr, frontal lobe; Oc, occipital lobe; Pa, parietal lobe; J?o, fissure of Rolando; Sy, Sylvian fissure; Te, temporal lobe. right side of the body and vice versa. The chief motor and sensory areas of the hemispheres are shown in Figure 197. Functions of the medulla.— This is the center of those reflexes which control the respiratory and circulatory organs. When a cut is made anterior to the medulla the animal continues to breathe and the heart to beat for some time after the operation. If, however, the cut is made at the point where the medulla joins the cord or it is destroyed entirely, heart beat and breathing soon cease and rapid death follows. Other important reflexes i TrFUNCTIONS OF THE SYMPATHETIC SYSTEM 421 are also located in this organ which is the origin of many of the cranial nerves. Functions of the sympathetic system.— This name was originally given to the chain of ganglia extending along each side of the spinal column under the mistaken idea that it was a pathway for all the so-called sympathetic (reflex) actions of distant organs. This system was sup- posed to arise from the brain by branches connected with the fifth and sixth cranial nerves. It is now known that this system consists merely of a collection of ganglia or nerve cells which are connected with one another and with the spinal nerves. While it is customary to limit the term to the ganglia on each side of the spinal column, it actually includes many other ganglia with or without names which lie in different regions of the body. Accord- ing to Professor Langley, the efferent fibers from the sympathetic and related ganglia supply the unstriped muscle tissues, the cardiac muscles and the glands, in short, the muscles of the involuntary organs of the body, such as the walls of the intestines. He has given the name of autonomic to this system to indicate that these fibers are to a certain extent independent of the central nervous system. All the sympathetic ganglia are connected with the central nervous system by medullated fibers from the spinal or cranial nerves, but these fibers do not pass directly to the unstriped muscle tissue. On the con- trary, they transfer their impulses to a ganglion cell and the impulse passes out through a fiber (autonomic fiber) of this ganglion cell which is usually non-medullated. The connection of these fibers with the spinal cord is illustrated in Figure 189. Unlike the fibers which supply the striated muscles, practically the entire set of autonomic422 THE NERVOUS SYSTEM fibers is removed from the control of the will. Under ordinary conditions they are probably excited reflexly, but the course of the afferent fibers is at present little understood. Most of these reflexes are also unconscious actions, and include such movements as the dilation and contraction of the arteries, the movements of the intestines, and the secretory activities of the digestive glands. The entire field of action and method is still very little under- stood. Hygiene of the Nervous System. As has been stated, the manner in which nerve cells generate impulses or give rise to psychical reactions is unknown. We do know, however, that generation of impulses and mental activity is in some way dependent upon the metabolism of the nerve cells. From this, and the fact that nerve cells, like all other cells, are made of protoplasm, it follows that lack of nourishment or of air is bound to affect their activity, overwork to fatigue them, and rest to restore them. Comparison of the nerve cells of a child with those of an adult show that the latter are much more branched and more intimately connected than those of the child, which is another indication of the rela- tion between increased nervous action and metabolism. Nerve food.— It used to be thought that nerve cells required special kind of foods for their development, notably, phosphorus; and fish, which are rich in phosphorus, were supposed to constitute an ideal brain food. It is now known that their protoplasm is built up of identically the same sort of materials as are used by other cells, and hence a food which is nourishing to the laborer is equally so to the brain worker. The important point to be kept in mind is that there must be a sufficient quantity of food, and that theBEST 423 organs of digestion and circulation, etc., must be kept in good order for its preparation and distribution. In short, whatever tends to produce a healthy body also tends, indirectly, to main- tain a good condition in the nervous system. Exercise, therefore, is as valuable to the brain worker as it is to the laborer, and per- haps even more so, in that the latter is apt to neglect muscular activity, while the very work of the laborer tends, to keep his system in order. Air supply.— The chemical changes which take place in the nerve cells are essentially similar to those in the muscle cells, in that the production of energy or action involves the oxidation of cell substance and food. It follows, therefore that not only must the nerve cells receive food but also a plentiful supply of oxygen. The brain responds very quickly to a lack of air by a feeling of dullness and headache, and it is extremely important that our school and living rooms be well ventilated. Exercise that tends to develop the lung power is likewise valuable. Rest.— Like the muscle cells, again, the nerve cells accumu- late wastes and consume food by continued activity. It is imper- ative, therefore, that they be allowed periods of rest when the supply of food may be renewed and the excess of waste removed. Sometimes, as with the muscles, a change of work will permit one set of cells to rest while another is working, and thus a change of occupation is a rest. But no person can work continuously and maintain a healthy nervous system, and for that reason sleep is nature’s greatest remedy for tired nerves. Children whose nerve cells are growing should have plenty of sleep in order that their nerve cells may store up sufficient food in these rest periods to meet the daily needs, and at the same time permit of growth and branching. On this account adults whose cells have attained their normal size need less sleep than children. Nerve cells are like muscle cells in another particular, in that they degenerate by disuse and lose their irritability. The person who is too careful of overtasking his nervous system may carry this care to excess and lose his mental power entirely. It is this need of constant nerve exercise which gives especial value to a general education, since the variety of subjects in such a course insures the normal development of all parts of the system.424 THE NERVOUS SYSTEM Diseases of the nervous system. — Nerves. The sheaths which inclose the axis cylinder may become the seat of inflammation. The result is intense pain and various forms of nervous disease, according to the extent and nature of the inflammation. Among diseases of this nature are included all forms of neuritis, tooth- ache, etc. Spinal cord. — Injuries to the spinal cord or the introduction of certain forms of bacteria may result in the destruction of the whole or part of this organ. The result is paralysis of the parts supplied by the affected portion. Such diseases are usually slow in action and often incurable. Brain. — Anything which affects the nerve cells of the brain, such as pressure, narcotics, worry, lack of work or sleep, etc., is bound to result in disturbed mental action. Such influences may result in sleeplessness, hysteria, lack of self-control, irritability, and even in total loss of control or insanity. Interference with the blood supply and pressure may produce fits or apoplexy. In short, so sensitive is the brain to external influences that its encase- ment in the bony cranium is none too strong a protection for this delicate structure and any interference with its structure is attended with serious results. Alcohol and the Nervous System. Stages in intoxication.— It is a fact known to everyone that alcohol in any quantity affects the nervous system almost immediately. It is noticeable that moderate quan- tities have apparently a stimulating effect upon the various centers of speech, thought, and action. The winebibber becomes exhilarated, his speech flows more rapidly, his wit appears keener, and he is conscious of a sense of freedom from restraint. If the amount of alcohol is increased con- tinuously, this stage of apparent stimulation is soon followed by excessive exhilaration. The subject becomes still more talkative and his freedom from restraint manifests itselfTHE PROBLEM OF THE PHYSIOLOGIST 425 in excesses of various kinds. He laughs immoderately at remarks only slightly humorous and in many ways shows that his powers of judgment are becoming dulled. Grad- ually this dullness spreads. His speech becomes thick and uncertain and a stage of dullness and drowsiness of all his mental faculties follows. He may still be able to walk, but his inclination is to be quiet, and, if permitted to do so, he passes rapidly into a stupid, drunken slumber. Finally, the loss of control over his speech spreads to include the centers of voluntary movement. He loses control of his legs and is no longer able to walk, staggering blindly when he attempts to do so. Such a condition represents what are usually the final stages of acute intoxication or drunk- enness. If, however, the doses of alcohol are still con- tinued they reach a point when the involuntary centers which control the breathing and movements of the heart become affected, and in this case death of course must re- sult. The problem of the physiologist.— The problem of the physiologist is to account for these well-known effects and to tell exactly what is the action of the alcohol at each step of the process. All are agreed that alcohol in large or con- tinued doses ultimately dulls or paralyses the nervous cen- ters, and that this narcotic effect is felt first in the higher centers. After these centers are dulled the paralysis grad- ually spreads until it includes the motor centers and ulti- mately the vital processes, such as breathing and heart beat. Thus, in the description given, it is easy to see that the centers of judgment are the first to become dulled, and that the effect of this is to remove the speech and motor centers from control so that they seem to be stimulated. That this is not a true stimulation, but the first step in426 THE NEBVOUS SYSTEM paralysis, is seen in the speed with which these centers become dulled, while the tendency to attack the higher centers first is shown by the fact that speech is lost before control of movements. In regard to the effect of alcohol in small doses there are at present two different opinions. One school of physi- ologists holds that alcohol in small doses is a true brain stimulant, and that the exhilaration felt as a result of such doses is accompanied by increased keenness of judgment, and that, while temporary in its effect, this effect is not due to loss of control by paralysis of higher centers. The other school claims that alcohol is never a stimulant, but always a narcotic, whether in large or small doses. They explain the evident exhilaration which follows small doses as due to the same causes as the excessive exhilara- tion, namely, that it results from a partial paralysis of the centers of judgment and control. They say that, by re- moving the inhibiting influences from the speech and other centers, such paralysis inevitably causes in these centers a freedom of restraint which manifests itself in apparent stimulation. In short, this school claims that alcohol from its entry into the system is never a stimulant, but always a narcotic in its effect upon the nervous system as a whole. The determination of the true position in this regard must come as a result of experiment, and the results of experiment have so far been extremely conflicting. Not enough has been done as yet to settle the question for either side, and all that can be said at present is that by far the larger number of physiologists take the view that alcohol is always a narcotic, even in small amounts. Alcohol and time reactions.— Many experiments have been made to determine the effect of alcohol on the timeALCOHOL AND DISEASES OE THE NERVOUS SYSTEM 427 required for a nervous impulse to travel over the fibers. All such experiments require the subject to perform a cer- tain muscular action upon receiving a certain sensory stim- ulus, and the measure of the time which elapses between the receipt of the sensory impulse and the performance of the action is taken to indicate the rate of transmission of the impulse —the “reaction time ” as it is called. By com- paring the reaction time of a person under normal condi- tions with the time when under the influence of definite amounts of alcohol the effect of alcohol can be inferred. The results of such experiments are not absolutely con- clusive in regard to the effects of small amounts of alcohol. All, however, tend to show that larger doses of alcohol lengthen the reaction time for all mental processes. The importance of this fact has led all corporations employing men in places where quickness of action, clearness of per- ception, close attention to duty, and the like are required, to forbid absolutely the indulgence in alcoholic drinks while performing such duties. Such actions on the part of corporations are significant warnings to us of the danger of this alcohol habit and its tendency to destroy efficiency. Alcohol and brain workers.— It follows directly from the preceding that alcohol is not of advantage to the brain worker, and leads to confusion of judgment. One unfor- tunate effect of its use is that in dulling certain control centers it often produces in the indulger the delusion that he is actually keener in perception, when actually this apparent keenness may be only an indication of impaired powers. Alcohol and diseases of the nervous system.— Continued use of alcohol lowers the resistance of the body to disease, and it is a well-known fact that hard drinkers often sue-428 THE NERVOUS SYSTEM cumb to diseases from which a normal person would easily recover. Many cases of insanity are reported as due to alcohol, but while it is undoubtedly true that the use of alcohol is in many cases indirectly the cause, the exact relation to this mental disease is not yet definitely known. The only form of mental disturbance which is a direct result of alcoholic indulgence is the violent disturbance known as delirium tremens. It is also a well-known fact that nervous weakness can be transmitted by heredity, and thus children of drinking parents often develop mental disease which is due to this inherited weakness.XXVII. THE NERVOUS SYSTEM IN THE LOWER ANIMALS.1 Nervous tissue, wherever it exists, is in the form of nerve cells and fibers, as in man. Frequently the nerve cells are collected in masses which are called ganglia, and Fig. 198 — A nerve ganglion. The round bodies are nucleated nerve cells; the fibers are nerves. in this chapter the use of the word ganglion may be taken to mean a collection of nerve cells. The fibers are always developed as processes of the nerve cells. Protozoans and coelenterates.—The one-celled animals can evidently have no true nervous system. Their protoplasm, however, is sensitive to stimuli and conducts impulses. Such a condition is interesting mainly as showing the rela- tion of ordinary protoplasm to the specialized protoplasm of the nerve cell and as indicating the origin of the ner- vous system. The first appearance of nerve cells and 1 See Footnote, p. 140, Chapter X. 429430 THE NERVOUS SYSTEM IN THE LOWER ANIMALS fibers is found in the ccelenterates. In the hydra are cer- tain scattered cells which are partly muscular and partly nervous in function. In the jellyfishes we find a network of fibers and cells scattered over the disk. The general purpose of such systems seems to be to give sensitiveness to the animals and thus aid them in securing food. Echinoderms.— In the starfish and its allies we have a true system. In these animals we find a ring of five gan- glia surrounding the mouth and connected by a nerve cord. Each ganglion gives rise to radial nerves which extend along the arm opposite the ganglion. The peculiarity of this system is that it actually consists of three systems, each of which has its own combina- tion of cells and fibers and sup- plies a special part of the body. These three systems are abso- lutely independent of one another in action. We may compare the nervous control of the starfish to a city with three absolutely in- dependent telephone companies, each company with its own “central” and wires. Such an arrangement is absolutely unknown in any other group of animals. Mollusks. — The nervous system of such animals as the clam and its allies consists of several ganglia located in different parts of the body and interconnected by nerve trunks. Each ganglion controls special parts of the body, and this arrangement is spoken of as a scattered system of control. The arrangement in the clam consists of two Fig. 199 — Diagram of nervous system of starfish; r, nervous ring around mouth; 11, radial nerves to each arm, ending in the eye.MOLLUSKS 431 compound ganglia (cerebro-pleural) located on the right and left sides of the esophagus and connected by a nerve cord called a commissure. Each of these ganglia actually consists of two cell masses. The two cerebral masses give off nerves which control the head parts while the pleural masses control the neigh- boring parts of the body. Each com- pound ganglion gives off also two nerve trunks which pass backward along the same side of the body as the ganglion from which they originate. The right and left trunks of one pair unite with a gang- lion (visceral) near the siphon end of the animal. The right and left trunks of the other pair unite with a ganglion in the foot, called the pedal ganglion. The pedal gang- lion sends out fibers which direct the movements of the foot, while the visceral ganglion, through its fibers, controls the internal organs and the parts near it. In such a system it is evident that we have control stations located near their seat of control and, unlike those of the starfish, all are interconnected into one system by the nerve trunks. We may compare such a system with a city telephone system in which each district has its own “central” but all the centrals are connected. In the snail we find an interesting modification of the clam system. Here the main ganglia are collected in a ring about the esophagus, w'hile additional ganglia are found scattered through the body. In the cuttlefish this Fig. 200 — Ner- vous system of clam; c, cere- b r o-p 1 e u r a 1 ganglia; c', cerebral com- mis8ure; p, pedal ganglion; ps, visceral ganglion; p' and psf, nerve trimks.432 THE NEEVOUS SYSTEM IN THE LOWEJR ANIMALS ring of ganglia is actually inclosed in a cartilage box, thus giving it a still closer resemblance to a brain. Worms.— In the worms we find two main ganglia usually located in the head end and united by a com- missure. In the simpler worm forms these ganglia are the only ones in the body, and fibers from them supply all parts of the body. In the earthworm these two ganglia are represented by a single two-lobed mass located just above the esopha- gus and called on this account the supra-esophageal Fig. 201 — Nervous system of earthworm; g, ganglion; v.n. ventral nerve cord. ganglion. From this compound mass or “ brain ” a ring of cord encircling the esophagus unites it with a ganglion mass under the esophagus called the sub-esopha- geal ganglion. Finally, a double cord extends from this latter ganglion along the ventral side of the body. This cord shows a series of small ganglia at intervals corres- ponding to segments. Each of these small ganglia is the source of motor and sensory fibers which supply that particular segment, and these segment ganglia are the seat of the reflex actions. It is not difficult, then, to compare such a system with the one in man. The supra-esophageal ganglion may be compared to the brain, the ventral cord to the spinal cord, and the smaller ganglia to the centers of reflex actions and sources of spinal nerves in the cord.VEKTEBIIATE SYSTEMS 433 This brain ganglion and cord constitute a central system in distinction to the fibers which arise from the small ganglia and which correspond to our peripheral system. Arthropods.— In the lower arthropods, such as the lower forms of crustaceans, and the wormlike larva1 of insects, the nervous system is practically identical with that of the earthworm. In the higher artlnopods such as the higher crustaceans and insects, the plan is the same, but many of the ganglia are fused together in the head and thorax making this end of the system much more brainlike in structure. This head mass also gives nerves which supply the eyes and antennae much as our cranial nerves are given off from the brain. The ventral cord shows fewer ganglia, and each ganglion furnishes motor and sensory fibers to larger areas than in the worms. In short, while this system is similar in composition to that of the worms the ten- dency is to centralize control in the head. Vertebrate systems. — In the vertebrates the most remarkable changes are the develop- ment of the true brain as an organ of mental operations, and the transfer of the whole system to the dorsal side of the body where it is protected in the skeletal structures. The essential features of the systems found in the different forms of vertebrates are identical, the variation being in the direction of complexity. Figures 203 and 204 illustrate the manner in which the complex brain of man has been developed, and the relative stages in EDDY. PHYS.—28 l|k Fig. 202—Ner- vous system of a cater- pillar.434 THE NERVOUS SYSTEM IN THE LOWER ANIMALS development seen in the vertebrates lower than man. The feature which strikes the eye as most vital in this Fig. 203—A comparison of vertebrate brains; O.L. olfactory lobes; C.B., cere- bral hemispheres; O.L. optic lobes; C.L. cerebellum; M, medulla; S.C. spinal cord. comparison is that the changes have arisen mainly in order to accommodate the increasing brain surface to the limits of the inclosing cavities. For example, in the Fia. 204 — Comparative wrinkling of cerebral surface of cat (A), and man (R).VERTEBRATE SYSTEMS 435 lower vertebrates the increase in size has been met by doubling and folding one part upon the other, while in the higher forms the wrinkling or furrowing of the cortex gives increased surface. Along with these physical changes has come increased mental power, until, in man, we have an animal preeminent over all others by virtue of the superior mental powers.XXVIII. THE SPECIAL SENSES. Touch, Taste, and Smell. In the preceding chapters we have discussed, mainly, the mechanism by which sensory and motor impulses are conveyed and controlled; and the areas of the central system where these sensory impulses are received and translated into conscious sensation. The manner in which impulses originate and their character requires a study of the arrangements at the surface ends of the afferent fibers. Such arrangements are grouped together mider the name of end or sense organs. They are all specialized struc- tures adapted for response to certain kinds of stimuli only, and are named according to the effect of the impulses upon the consciousness. Thus, organs which produce sensations of sight are called sight organs, those which produce sensations of touch, tactile organs, etc. It must be distinctly borne in mind, however, that all of these organs merely originate impulses, and it is.only when these impulses have traveled over the afferent fibers and stim- ulated certain tracts of the brain cortex that we become conscious of the sensations which they produce. In short, the sense organ is simply a structure adapted for response to a special kind of stimulus; and the impulse sent out by this stimulus, the brain interprets as sight, pain, heat, etc., according to the quality of the impulses. Laws of sensory impulses.— There are three laws of sensory impulses which arc applicable to all kinds of sen- sory fibers. 436LAWS OF SENSORY IMPULSES 437 First law. A sensory fiber wherever stimulated, car- ries only one kind of impulse. Thus a touch nerve fiber, whether stimulated in its end organ or elsewhere always produces the sensation of touch, never that of sight or hearing. Second law. In consciousness an impulse is located as coming from the natural source of such an impulse. Thus, in an amputated leg, pressure upon the cut end of the nerve is felt by the brain as though it came from the original end organs of that nerve. In this way may be explained many of the instances where pains are felt in the toes or fingers even after these parts have been amputated. Third law. The increase in any given sensation from a definite kind of impulse does not take place uniformly with the increase in stimulation. On the contrary the amount of stimulation has to increase by a definite quan- tity before any increase in sensation is felt. A man named Weber investigated this relation between stimuli and sen- sation, and the law is often called Weber’s Law on that account. An illustration will make clear its application. If we hang a thirty-gram weight to the finger we get a certain feeling of pressure. If we now add a half-gram weight no increase in pressure is felt, and it is only when we add a whole gram that the increase is actually felt. If, now, we start with sixty grams we have to add two grams before the increase is felt; that is, one thirtieth of the original weight. In this case, the stimulus increase necessary to sensation is one thirtieth of the original weight. The amount of increase in a stimulus necessary to in- creased sensation varies greatly with different fibers. It is possible to reduce by practice the increase in stimulus438 THE SPECIAL SENSES required by any given fiber. This relation of amount of stimulus to sensation is one of the factors which determine the sensitiveness of a sense center. The tips of the fin- gers, which require much less increase in stimulus to pro- duce sensation than the skin of the back requires, are said to be more sensitive than the latter. Classification of sense organs.— The end of every affer- ent nerve is a sense organ, but for convenience these end organs are grouped under various heads according to the region where they end and the quality of their impulses. The following scheme presents the commonly accepted classification: CLASSES OF SENSE ORGANS. Name of Group Location of End Organs Quality of Impulse r Pressure Cutaneous . Skin and mucous membrane Warmth of the mouth Cold L Pain Internal . . Internal membranes and muscles ' . Pain Muscle sense Hunger and thirst Gustatory Tongue Taste Olfactory Nasal passages Smell Visual . . . Eves Sight Auditory . . Ears Hearing Cutaneous sensations.— It was formerly customary to group all the sensations which were produced by the impulses from the sensory nerve endings in the skin as touch sensations. Modern experiments have shown that touch sensations show four distinct qualities, namely, pres- sure, warmth, cold, and pain, and that this difference inDISTRIBUTION OF CUTANEOUS SENSE AREAS 439 quality depends not upon the nature of the stimulus but upon the particular fiber which is stimulated. In other words, the fibers and end organs which transmit impulses of warmth are as distinct from those which transmit impulses of pressure as they are from those of sight, hearing, and S7nell. For this reason the old term, touch sensation, has been re- placed by the word cutaneous (skin) sen- sations, and the variety of end organs in- cluded under this head are to be considered as four distinct kinds of sense organs. Cutaneous end organs. — The sensory fibers which end in the skin show several varieties of end organs. Some form networks in the outer dermis where the axis cylinders lose their sheaths, and from this network extend out into the epidermis as tiny naked filaments. Others show special- ized structures which may be classified under three heads, as tactile cells, tactile bulbs, or tactile corpuscles. At pres- ent it is impossible to tell by the look of a cutaneous end organ what kind of impulse it will transmit. In other words, unlike the fibers which end in the eyes and ears, we cannot be sure from their structure whether a given form of end organ produces pressure impulses or heat impulses, pain or cold. Distribution of cutaneous sense areas. — The most inter- esting fact concerning cutaneous senses is that all parts of the skin are not equally sensitive to all kinds of stimuli. For example, in a given area of skin there will be spots which respond only to pressure, and not to cold or heat. Other spots in the same area will respond to heat and not to pres- sure, cold, or pain. In short it is possible to map any given Fig ‘205. — A touch corpuscle in a skin papilla; C, corpuscle; iV, nerve fibers.440 THE SPECIAL SENSES portion of the skin and to locate on such a map by different colored dots the position of the different sources of sen- sation. The relation of these spots to the fibers may be summarized as follows: (a) The surface of the skin is di- vided into spots or areas in the center of which is the end- ing of a sensory fiber, (b) Each of the four cutaneous senses has its own areas and fibers, and stimulation of one of these spots produces only one kind of sensation, heat or cold, warmth or pressure, (c) The pain spots are most numerous and the warmth spots least numerous. The ends of the pain fibers are nearest the surface and those of the warmth fibers farthest below the surface. (d) Dis- crimination between two stimuli of the same kind depends upon the stimulation of two distinct fibers. Thus, we feel the pressure of two points as two only when each point stimulates a different pressure fiber from the other, (e) Fibers of a given kind are much nearer together in certain parts of the skin than in others, and sensitiveness in any given direction depends partly upon the nearness together of the fibers. Pressure. — The pressure spots are, next to the pain spots, the most numerous. In those portions of the body which are covered with hair the pressure fibers terminate in a ring surrounding the hair follicle. These rings, therefore, form the end organs. In other parts of the body the tactile corpuscles seem to function as end organs. These cor- puscles are particularly abundant on the tips of the fingers where they underlie the dermal papilla'. It is estimated that on the entire body exclusive of the head region there are about five million pressure spots. These spots are very close together on the tips of the fingers and less near in other parts of the body.PRESSURE 441 The delicacy of pressure sense can be measured in two ways, by determining the least amount of pressure necessary to arouse a sensation, and by locating the least distance apart at which two points can be felt as two. The method of determining the first consists in varying the pressure upon various portions of the body until the least amount necessary to produce a sensation in any given part is de- termined. The results of such tests show that the skin of the face, forehead, and temples is most sensitive of all the skin areas to the feeling of pressure. The back of the hand is more sensitive in this respect than the tips of the fingers. Two milligrams of pressure is necessary to arouse sensation on the forehead, while from five to fifteen milligrams is necessary when applied to the finger tips. Such tests simply determine the relative sensitiveness to stimuli of the parts tested. To test the power of discriminating between two pres- sure stimuli, the method consists in applying to the surface two points and determining how near together these points can be placed and still felt as two. The relative sensitive- ness of different areas of the body to such a test is given in the following table: 1 Area Distance of Points Apart Tip of tongue . 1.1 millimeters Tip of finger 2.3 Middle of palm 8 to 9 millimeters Forehead 22.6 millimeters Back of hand 31.0 Forearm 40.6 Chest 45. Middle of back 07.7 “ 1 N.B. In both methods of test the subject is blindfolded,442 THE SPECIAL SENSES The above results show that while less stimulus is neces- sary to arouse a sense of pressure in the face than in the fingers, the power to discriminate is particularly acute in the finger tips and tongue. This second method is also of service in locating the distribution of nerve ends, since to feel two points as two means that two nerves are being stimulated. Another interesting feature of this discrimination ense is that it can be increased with practice, notably in the case of the blind. This power is probably due to in- creased branching in the nerve ends thus increasing the number of spots on a given area. Temperature sensations.— These are of two kinds, heat and cold, each with separate spots of stimulation and separate fibers. Their location may be determined by passing gently over the skin of the blindfolded subject, metal points which are colder or warmer than the skin surface. As this is done there will be aroused at definite points distinct sensa- tions of heat or cold, which are at the same time entirely distinct from pressure sensations. If a certain area be explored in this manner and the spots located by ink spots, of different color, say red for cold spots and black for warm, the extent of the nerve ends and the relative number of heat and cold nerves for a given area can readily be determined. Figure 206 shows such a map. The results of such tests applied to all areas of the body may be sum- marized as follows: Fig. 206 — A portion of tlie skin mapped for hot and cold spots; white spots, cold; black spots, hot.PAIN SENSATIONS 443 The cold and hot spots are always distinct, some portions of the skin being sensitive only to cold, some to heat. The cold spots are much more numerous and more sensitive to stimuli than the heat spots. The sensitiveness to stimuli varies for different areas of the body, the order of sensitiveness to temperature being as follows: tip of tongue, eyelids, forehead, cheek, lips, limbs, trunk. Tem- perature sense is always relative to the temperature of the skin, that is, objects colder than the skin stimulate cold spots and feel cold, and vice versa. Hot and cold spots may be stimulated by mechanical and chemical stimuli. Menthol, for example, gives a cold sensation. They may also be stimulated from within, the skin feeling hot in case of fevers being a case in point. No distinctive end organs for these sensations have as yet been discovered. Pain sensations.— It used to be thought that pain was the result simply of excessive stimulation of sensory fibers. Thus, if a pressure nerve was stimulated by too great a weight the result was a sensation of pain added to that of pressure but conveyed by the same impulse. It is now believed that pain sensations of the skin at least are pro- duced by special impulses which pass over special fibers and that these fibers are distributed in spots as are the other cutaneous nerve ends. The results of careful experi- mentation in locating and determining the peculiarities of the skin pain spots may be summarized as follows: Summary.— (a) Pain spots are distinct from and more numerous than other cutaneous sense spots. (6) Pain spots of different regions vary greatly in the amount of stimulus necessary to produce a sensation. Those of the eye being much more sensitive than those of the fingers. All, however, require much greater stimulus than those of temperature or pressure, (c) The nerve fibers444 THE SPECIAL SENSES which supply the pain spots have no distinctive end organs so far as known. (d) They may be stimulated by any of the usual forms of cutaneous stimuli, such as temperature or pressure, pro- vided these st muli are of sufficient intensity, (e) The location of a pain stimulus by the brain is aided by the pressure and tem- perature stimuli, parts which have lost these centers still retain- ing the sense of pain but the subject has difficulty in locating the seat of stimulation. Internal Sensations. The membranes of the internal organs, such as the mucous membranes that line the alimentary tract and the lungs and the membranes that invest the muscles or cavities of the body, have no pressure, warmth, or cold fibers. Increase in internal temperature or pressure can be detected, therefore, only by its effect upon the cutan- eous fibers. These membranes are, however, supplied with pain fibers and other fibers whose function it is to supply the brain with information as to the health and needs of these parts. Very little is known of these latter fibers or their end organs, their method of distribution or stimula- tion. The sensations of hunger, thirst, general health- fulness or weakness are all produced by impulses passing over these fibers, and their function as indicators of the condition of our internal organs is extremely important. At present, we know only of the presence of these fibers and admit our ignorance of their end organs or properties. Gustatory Sensations. Distribution of fibers and end organs.— The sensation of taste is produced by impulses sent over special nerve fibers whose ends are distributed over various parts of the mouth cavity and particularly in certain regions ofDISTRIBUTION OF FIBERS AND END ORGANS 445 the tongue. If the tongue is protruded and its surface examined with the aid of a mirror it is possible to dis- Fig. 207 — The tongue; K, filiform papillse; I, fungiform papilke; Z, circumval- late papillse. tinguish many raised points on the surface of its mucous membrane. These raised points are called papillae, and are of three forms—the circumvallate, fungiform, and filiform. Of these, the circumvallate are the largest and least numer- ous. They appear as circular extensions of the membrane and are surrounded by a little depression or ditch. They are only seven to twelve in number, and lie on the upper surface near the root of the tongue.446 THE SPECIAL SENSES The fungiform papilke are so called from their resem- blance to a mushroom, and consist of small, rounded ele- vations supported on short, slender stalks. They are found all over the middle and fore part of the tongue on the upper surface. The filiform papillm are most numerous and the smallest, and appear as little pointed projections scattered all over the upper surface and edges of the tongue. In the walls of the cir- cumvallate papillae, in some of the fungiform papillae, and in portions of the pharynx, palate, and epiglottis, and even in the vocal cords, are peculiar structures known as taste buds. These taste buds are oval bodies composed of a number of over- lapping epithelial cells (protective) surrounding a core of elongated or taste cells ivhich project through a pore at the top some stiff hairlike processes and are connected at the base with the taste nerve fibers. The hairlike extensions constitute the part of these organs which is directly stimulated by the tasted substance. The impulse thus origi- nated is passed through the body of the taste cells and finally stim- ulates the terminal branches of the taste nerve fibers themselves. These taste buds, therefore, may be con- Taste Cells Fig. 209—A, isolated taste bud, from whose upper free end project the ends of the taste cells; B, sup- porting or protecting cell; C, taste cell. Fig. 208—Section of circumvallate papilla.KINDS OF TASTE SENSATIONS 447 sidered as the specialized end organs of the sensory taste nerve fibers. The perception of taste is produced by the connection of these fibers and the brain through the fifth and ninth pairs of cranial nerves. Stimulation and sensitiveness.— These taste buds can be stimulated only by substances in solution, and in dis- solving solids the saliva of the mouth plays an important part in the production of taste sensations. Dry salt or sugar placed upon the tongue remains tasteless until dissolved by the saliva. The sensitiveness of the taste buds to stimuli is influenced by many external factors as well as by the concentration and character of the dis- solved substance. The result of experiments in this direction may be summarized as follows: Summary.—(a) The taste buds are most sensitive at a tem- perature of from 50 to 86° Fahrenheit. Very high or very low temperatures tend to paralyse the taste buds and thus lessen or destroy the sensation of taste. (6) Rubbing the substance to be tasted against the tongue increases the sensitiveness of the taste buds. This accounts for our habit of rubbing the tasted food against the walls of the mouth with the tongue. (c) The sensitiveness varies with the character of the tasted substance. The order of greater sensitiveness in this respect being bitterness, sourness, sweetness, saltness. Kinds of taste sensations.— While our taste sensations appear to be very varied in kind, it has been demonstrated that they are actually only four in number, namely, bitter, acid, sweet, and salt. The bitter taste is most developed at the back of the tongue, the sweet taste at the tip of the tongue, the acid or sour taste at the sides, and the salt taste is nearly equally distributed. Many of our taste sensations are combinations of these four pri- mary sensations with odor or sight sensations This fact can be148 THE SPECIAL SENSES easily demonstrated by closing the nasal cavities and bandaging the eyes when the sensation of taste produced by certain foods will often disappear entirely. This fact also accounts for loss of taste in cases of colds in the head, the nasal passages being clogged by mucus. Certain substances taste sweet when first taken into the mouth and bitter when swallowed. This is due to the fact that these substances are capable of stimulating both sweet and bitter taste buds and also demonstrates that these primary sensations are carried by different afferent nerve fibers. Olfactory Sensations. Distribution of fibers and end organs.— The nasal cavity is located just above the hard palate. It communicates Fig. 210—Section of nose showing outer wall of right nasal cavity; u, b, c, d, interior of nose; k, olfactory bulb giving off nerves to inside of nose. with the external air through the two nasal openings and with the pharynx by a single posterior opening. TheDISTRIBUTION OF FIBERS AND ORGANS 449 walls of this cavity are formed by the inferior maxillary palate, the nasal and ethmoid bones, and is lined through- out with mucous membrane. In this membrane are dis- tributed the fibers of the first pair of cranial nerves (the olfactories), and through these fibers impulses of smell are carried to the olfactory lobes. These fibers end in the upper part of the membrane of this cavity in special end organs of smell. The lin- ing of the nose is, there- fore, divided into a lower or respirator}7 membrane composed mainly of cili- ated epithelial cells, and an upper or olfactory mem- brane, rich in nerve fibers, and devoid of ciliated cells. The end organs themselves consist of the ends of the fibers developed into long, olfactory cells, and these cells are protected on all sides by surrounding nucleated, epithelial cells. The olfactory cells show a long, nucleated cell body some- what like a taste cell, and, like that, ending in tiny hair- like projections at one end, while the other is continuous with the nerve fiber. The hairlike processes are first stimulated, and transmit this stimulus to the fiber which conveys the impulses to the brain. EDDY. PHYS. —29 Fig. 211 — Olfactory cells from lining of nose; O, O, olfactory cells terminating nerve fibers; Ey E, ordinary epithelial cells of the mucous membrane; Hy sensitive cilia affected by odor.450 THE SPECIAL SENSES Stimulation and sensitiveness.— The explanation of the stimulation of an olfactory nerve fiber is as follows: It is supposed that all odoriferous bodies give off par- ticles in the form of vapor, which latter when dissolved in the liquids of the mucous membrane stimulate the sensi- tive hairs of the end organs. The vapor may be given off by a near or distant body, but the question of whether it be smelled or not is determined by the currents of air which carry this vapor. If the air is inhaled it carries this vapor through the nasal passages, where the dis- solved particles produce the sensation in the manner described. Similarly, swallowed bodies may send their vapors up into the nasal passages through the posterior openings, and stimulate in the same manner. The sensitiveness of the olfactory nerves to stimuli varies greatly in different persons and between persons and animals. While in man the sensitiveness is not so great as in dogs and many of the lower vertebrates, it is still very acute. By taking odoriferous substances and diluting them to known amounts it has been shown that the sense is so keen in the average man that one part of camphor in four hundred thousand parts of air can still be readily detected, while odors like musk and vanilla can be diluted to eight and ten million parts of air and still be detected. These olfactory cells, however, soon become exhausted by continuous stimulation and cease to react. Their sensitiveness is also affected by lack of secretion on the part of the membranes. Kinds of olfactory sensations.— Smell sensations are usually classified as agreeable and disagreeable, according as their effect is pleasant or unpleasant, and these effects undoubtedly serve to guard the person from evil atmospheres. This is hardly a trueKINDS OF OLFACTORY SENSATIONS 451 classification, however, as many odors are agreeable to some persons and intensely disagreeable to others. It has not yet been determined whether there are, as in taste, fundamental or primary odors out of which others are compounded, but it seems reasonable to suppose that such is the case, and that special fibers convey special kinds of odor impulses.XXIX. THE SPECIAL SENSES (continued). Visual Sensations. The eyes arc the external organs by means of which visual impulses are produced. So complex are these organs that we must first study their anatomy and some of the laws of light before we can appreciate the manner in which they produce visual impulses. Form and protection of the eyeball. — The eyes con- sist of two nearly spherical bodies called the eyeballs, which, together with certain protective structures, completely fill the two sockets in the facial bones which we call the orbits or eye sockets. Both eyeballs and their accessory parts are identical in struc- ture, hence a description of one will serve for both.1 In front the eyeball is protected by movable folds of the skin called the eyelids. These lids are lined with a 1 A sheep’s skull with the bony parts and eyeballs uninjured serves well for a demonstration of the structure of the eyeballs. (See Ex. LXX.) 452 Fig. 212—Section of eyeball in orbit; frontal bone; 2} cheek bone; 3, eyebrow; 4, eyelids; 5, folds of the conjuctiva lining eye- lids and covering front of eye; 6, 7,8, muscles moving eyeball; c, cornea; a, aqueous humor; it iris; l, lens; r, vitreous humor; ck, choroid layer; r, retina; a, sclerotic layer; v, optic nerve.FORM AND PKOTECTKH OF THE EYEBALL 453 mucous membrane called the conjunctiva,1 which is also folded back to cover the front of the eyeball. The lids are movable, and when lowered form a protective curtain in front of the eyeball. Their edges arc lined with sebaceous glands, which, by secreting oil, prevent their sticking together when closed. A row of hairs also fringes these lids and serves to keep out dust and furnish a slight shade. These lids are closed by the contractions of a cir- cular muscle (the orbicularis) which acts in part reflexly, as when an object is suddenly brought toward the eye or head. As these lids rest against the front of the eyeball, friction would naturally result between the two surfaces when they rub together. The mucus secreted by the membrane partly prevents this by acting as a lubricant. There is in addi- tion a small gland about the size of an almond (the tear gland) located on the outer and upper side of the orbit, and this gland secretes a salty liquid called the tear fluid or the lachrymal fluid. The gland is also known as the lachrymal gland. This fluid flows over the front of the eyeball assisted in -its progress by the motion of the lids until it reaches the inner angle of the lids next to the nose. Here it is drained off into a small duct (the lachrymal chid) which 1 Inflammation of this lining causes conjunctivitis, or pink eye. Fig. 213 — View of right eye and tear appa- ratus; G, tear gland; C, C, upper and lower tear ducts; B, common nasal tear duct.454 THE SPECIAL SENSES empties it into the nasal passages. Not only does this fluid aid in lubricating the surface of the ball and eyelid, but in passing continuously across the exposed part of the eyeball it removes from the surface dust or other foreign particles which may tend to collect there. Sometimes this fluid is poured forth very copiously, as when the membrane covering the eyeball is irritated, or the person is under strong emotion. Under such circumstances the ducts cannot drain it off fast enough, and it accumulates and rolls over the edges of the lids down the cheeks. Under ordinary conditions of flow, the oily secretion at the edge of the lids is sufficient to prevent its escape in this way. Back of the lids the eyeball itself rests upon a fatty layer which lines the orbit. Connected to it is the optic nerve which enters the eyeball from the back through an opening in the socket. This nerve carries the visual fibers to their centers in the brain. The eyeball is also held in place and moved by six muscles. These muscles are controlled by motor impulses sent from the brain along the fibers of the third, fourth, and sixth cranial nerves. The attachment of sides of the eyeball, and, by contracting and relaxing alter- nately, turn the eye toward or away from the nose. Two (the inferior and superior rectus) are attached to the lower in}. VOlique Fig. 214 — Muscles of the eye- ball. these muscles shows an ingenious mechanical adaptation to the structure of the orbit. (See Fig. 214.) Four (rectus muscles) are straight muscles attached at one end to the eyeball and at the other to the bottom of the eye socket. Two (the internal and external rectus) are attached to theSTRUCTURE OF THE EYEBALL 455 and upper surface of the eyeball respectively, and by their action turn the eyeball upward or downward. The other two muscles (the inferior and superior oblique) serve to rotate the eyeball outward or inward. The inferior oblique has its origin near the front of the orbit on the inner side and passing under the eyeball is inserted on the outer side. Its contraction rotates the eyeball outward. The superior oblique has its origin at the back of the orbit like the rectus muscles, passes forward just above the superior rectus and its tendinous end, after first passing through a loop or pulley formed by a fibro-cartilage on the upper edge of the orbit, turns outward over the top of the eyeball, and is inserted in the eyeball at a point on the inner side. By this arrange- ment the contraction of a straight muscle rolls the eye inward. The combined action of these six muscles is responsible for all the movements of the eyeball. The muscles of the two eyeballs contract and relax in pairs, the internal rec- tus of one eye contracting at the same time as the external rectus of the other, and similarly for all the others. In this ■way, both eyes are turned in the same direction at the same time. As has been already mentioned, the eyelids are closed by the orbicularis muscle. The upper lids, however, are pro- vided with elevator muscles attached to the back of the orbit, and these muscles pull the upper lids open farther than would be possible by the extension of the orbicularis alone. Structure of the eyeball.— The eyeball itself, when de- tached from its supports, is a nearly spherical sac of about an inch in diameter. It is covered, except at the points where the optic nerve enters and in front, with an opaque456 THE SPECIAL SENSES white coat known as the white of the eye or sclerotic coat. In this layer are the blood vessels which supply the coat with Fig. 215 — Section of the eyeball; Sc, sclerotic coat; Can, conjunctiva; C, cornea; /, iris; 0, optic nerve; Ch, choroid layer; C.P, ciliary processes; Cm, ciliary muscles and ligament; R, retina; L, crystalline lens; A, aqueous humor; Vt vitreous humor. nourishment. At the front this layer is replaced by the transparent layer of mucous membrane (the infolded con- junctiva) and a transparent modification of the white of the eye called the cornea. Back of this cornea is a colored, circular, muscular curtain called the iris. If we split the eyeball vertically so as to split the optic nerve it shows the following internal structure. The conjunctiva in front folded back from the eyelids. Over the whole surface, and covering the outside of the optic! nerve, is the sclerotic layer just mentioned, with its trans- parent window or cornea in the front. Inside the scleroticSTRUCTURE OF THE EYEBALL 457 is a second layer, called the choroid, composed of loose con- nective tissue, and colored black by pigment particles similar to those in the skin. At the front where the cornea joins the sclerotic this middle layer is thrown into plaits (iciliary processes) beyond which it is continued as a muscular curtain (the iris). This iris is composed of cir- cular and radial unstriped muscle fibers, is colored by pigment, and has a circular aperture in the center (the pupil). At its outer edge it is firmly united with the sclerotic layer by a ligament called the ciliary ligament, and to the ciliary process of the choroid. The contraction of its circular fibers decreases the size of the pupil, while the contraction of the radial fibers reverses this action. By means of this variation in the pupil, the amount of light which enters the eye is kept under control. The third layer of the eye is an extension of the optic nerve called the retina. It consists of the endings of the optic nerve fibers and a supporting layer of connective tissue. The entire layer is very thin and lines the whole interior of the eyeball with the exception of the front, where it stops at the edge of the ciliary process. This retina is the sensitive layer of the eye and the one in which the visual impulses originate. Just behind the iris is a biconvex, transparent body, called the crystalline lens. The front surface of this lens is slightly flatter than the back and is in contact with the inner surface of the iris. Between it and the cornea is a space which is filled with transparent watery liquid called the aqueous humor. The entire cavity of the eyeball back of this lens is filled with a glassy semi-solid mass called the vitreous humor. The lens itself is surrounded by a thin membranous capsule, and is kept in place by a circular458 THE SPECIAL SENSES band (the suspensory ligament) which attaches the edges of the capsule to the ciliary process of the choroid layer. Between the ligament and the edge of the choroid layer arc a series of muscles (the ciliary muscles). When these muscles contract they draw forward the edge of the choroid layer, thus causing the suspensory ligament to relax. This contraction permits the lens to shorten its diameter and become thicker and more bulging. In this way the shape of the lens may be changed, and this affects its image- forming power. Functions of the various eye structures.—The human eye is constructed essentially like a camera, and we can Fig. 216—An eye model. obtain an idea of the function of its parts by study of the parts of a camera or eye model such as shown in Figure 216. This shows a box painted black on the inside and having at the front a small aperture into which may be fitted perforated cardboard disks of different sized aper- tures. Inside the box is a biconvex lens of glass so mounted as to bring its center opposite the center of the cardboard aperture, and to be capable of movement for- ward and backward. It also contains a ground glass screen similarly mounted, and adjustable to forward andFUNCTIONS OF THE VARIOUS EYE STRUCTURES 459 backward movement. If, now, we place this box with the aperture pointing toward an illuminated object and move the lens backward or forward, wc find that for a certain position of the lens there appears on the ground glass screen an inverted picture of that object. In other words, this combination of lens, screen, and aperture, with its light-excluding walls enables us to project inverted images of external objects upon the screen. If we should replace the screen in this apparatus by a sensitive photographic plate we should obtain a permanent impression of the image or picture. Our box would then fulfil all the func- tions of a camera. Now, the human eye is essentially such a sort of image producer as this box or camera. The only difference is that, instead of a sensitive photographic plate capable of one exposure and producing one picture, we have a ner- vous structure, the retina, which receives these impressions and transmits them to the brain as nerve impulses. This retina is always ready to receive impressions; the human camera is always “ loaded.” But the resemblance to the camera does not stop with the retina. The pupil of the eye with its iris serves to regulate the amount of light which shall enter, just as the cardboard disks or stops of a camera do. The sclerotic layer corresponds to the wooden or metal walls of the box, and like those gives shape and strength to the structure.- The black, inner choroid coat, by means of its pigment, absorbs light, and prevents internal reflection, just as does the black paint on the inside of the camera box. Finally, the cornea and lens cor- respond to the lens of the camera, and, like that, produce inverted images upon the sensitive surface or retina. The points of difference between the eye and a camera460 THE SPECIAL. SENSES may be noted as follows: First, the retina and lens of the eye are fixed in position. The external aperture is pro- tected by the transparent cornea and thus makes the eye dust-proof. The air is replaced by the transparent aqueous and vitreous humors, but this does not affect the comparison, since these humors are transparent. They are made necessary by the fact that layers of the eye are not stiff like the wood of the box and would collapse were it not for the presence of these fluids. Finally, the lens and retina are fixed in position, and the eye, therefore, must resort to special means for producing pictures of near and distant objects and bringing them all to focus upon the retina. To understand this last process it is necessary that we first understand how a lens produces images. rays distributed all over the receiving over the receiving Surface, sul£ace' and the total effect is a jumbling together of all these images so that it is im- possible to detect anything but a mass of light. (See Fig. 217 — Scattered images produced by How pictures are formed by lenses. — Every point of a luminous body trans- mits light in the form of rays. These rays pass from the luminous body in straight lines and in every direction. Wherever a ray of light strikes it produces an image of the point from which it came. In the ordinary course of light rays, therefore, images are formed allHOW PICTUKES AEE FOKMED BY LENSES -161 Fig. 217.) Now, each ray is supposed to be made up of vibrations of a substance which fills all space and is called ether. The rate at which this ether vibrates determines the color of the light ray. These vibrating lines or rays when they pass from one transparent substance into another of different density are bent in definite directions, the shape and density of the transparent body determin- ing the direction of bending. Glass structures called lenses are simply masses of transparent matter so shaped as to control the direction of the bending in rays which pass from air into them. By means of them it is possible Fig. 218—Effect of lenses on rays of light; A, a concave lens spreading the rays; B, a convex lens bending the rays to a point. to concentrate the rays which pass through them (con- vex lenses) or to spread the rays (concave lenses). (See Fig. 218.) The point to v'hich rays are converged by a convex lens, or from which they seem to be spread by a concave lens, is called the focus of those rays and the action of the lens is called focussing. By its shape, a convex lens is able to focus the rays which it receives from a luminous body, so that the rays, which come from a certain point on the object shall, after passing through it, all converge at a certain spot. In this vcay the lens is able so to collect the462 THE SPECIAL SENSES scattered rays of a luminous body that a sharp but in- verted image of the object is thrown upon any screen placed at the point of convergence. (See Fig. 219.) In Fig. 219—Convex lens of eye forming an image by converging the rays on the retina. a camera a glass lens serves this purpose. In the eye, the cornea and crystalline lens accomplish the same result. Accommodation.— In a camera with a lens of a definite shape the images formed by the converging rays are focussed at different distances back of the lens according as the objects which produce the rays are near to or far from the lens. To secure a sharp image for any given object such instruments are provided with mechanical adjustments by means of which either the lens or the screen may be moved until the distance between them is such that the screen is exactly at the point of convergence of the rays. Such an adjustment is called accommodation. This accommodation to distant and near objects may also be secured by varying the thickness of the lens, and in the eye it is this latter method that is employed. As has been seen, the cornea and retina (screen) are fixed. By means of the ciliary muscles and suspensory ligament,EFFECT OF THE IMAGES UPON THE RETINA 463 however, the shape of the crystalline lens may be so changed as to secure this accommodation, and the invol- untary movements of these muscles en- ables the human eye to accommodate itself automatical!}'- to near and distant objects. (See Fig. 220.) The eye a self-supporting camera. — In all physical details, then, the eye acts as a camera, and forms inverted images of objects upon the surface of the retina, exactly as the camera does upon the sensitive plate. Being living tissue, however, it requires nourish- ment for its cells, and hence the choroid layer not only serves by its color to pre- vent internal reflection but through its nourishes the tissue cells of the inner eye. Fig. 220 — Dia- gram to illus- trate changes of the lens in accomodation. blood vessels The sclerotic layer also contains blood vessels which supply it with food. Effect of the images upon the retina.— The actual sen- sation of sight is located not in the eye but in the occipital lobes of the cerebral hemispheres where the visual impulses of the optic nerves are ultimately received. The optic nerve fibers in fact originate in these regions, passing from here to the anterior corpora quadrigemina (see Fig. 184, page 396), and are continued from here to a junction called the optic commissure. At this point many of the fibers cross, thus connecting the left eye with the right hemisphere, and vice versa. Some, however, do not cross, so that if the center of vision in one of the hemispheres is destroyed, there will still be sight in both eyes. This crossing of the main fibers is called the optic chiasma. After passing through it the fibers inclosed in two right and left trunks (the optic464 THE SPECIAL SENSES nerves) pass to the back of the orbits, eyeball, and, spreading out their fibers, where they enter the have their endings in the end organs of the retina. The projec- tion of an image upon the retina stimulates these end organs and thus arouses the im- pulses of vision. How the brain cells interpret the impul- ses produced by an inverted image into a sensation of sight is unknown. Structure of the retina. —The optic nerve fibers enter the retina on its anterior surface. Near this surface the fibers connect with the axis cylin- ders of large gang- Fig. 221 — A, Diagram of structure of the retina lioncells. These Cells seen under tlie compound microscope; 2?, the . essential nervous elements; C, the direction of Oil OtnOF CVlin- light stimuli; 1, internal membrane; 2, nerve which. COMiect fibers; 3, nerve cells; 4, terminal brushes of neura; 5, inner nerre cell layer; 6, 2d set With. Other Cells (see terminal brushes; 7, nerve cells; 8, external mem- -r-v <-) pigment layer; h, vitreous humor; i, cornea. Each ocellus points at a different angle and hence while the whole eye is fixed the range of vision is extensive. In such an eye each ocellus forms its own image independ- ently of its neighbours. These images are all collected by the brain into a single perception much as are the images of our two eyes. The nearest approach to the structure found in man and in all other vertebrates among the invertebrate forms is theVISION 485 eye of the squid. Here we find a lens with a humor-filled cavity, retina, and other layers much as in man. In the squid eye, however, the rods are on the side of the retina nearest the lens and they are supplied with optic fibers from the back of the eye. The eyes of the vertebrates are essentially alike in structure. The principal differences are in the means used to secure accommodation and range of vision.XXXIT. THE VOICE. In the study of the ear we noted that sounds are pro- duced by setting in vibration various objects, which vibrations are propagated as waves of air. When the air waves reach the ears they produce impulses of sound. In our study of the respiratory system we noted that in the human body the expired air was capable of producing sounds and that the various modifications of these sounds were called the voice. Combining these two observa- tions we should expect to find in our bodies some struc- ture in which vibrations of different pitch, loudness, and Fig. 227 — The larynx; A, front view; B, side view; C, back view. quality could be set up by expired air. Such an instru- ment, in fact, is found in the.larynx or cartilage box at the 486 *STRUCTURE OF THE LARYNX 487 top of the trachea or windpipe. This box is prominent externally as a projection which we call the Adam’s apple. Structure of the larynx.— The larynx or voice box is composed of cartilages, membranes, and muscles. The front and sides are formed of a single large cartilage (the thyroid). The curved front of this cartilage has a pro- jection at the top like the lip of a pitcher (the Adam’s apple). It curves around on each side in broad, flat sheets, which end above and below as projecting points. The two upper points are attached by ligaments to the hyoid bone and the two lower projections fit into sockets in a second cartilage called the cricoid. This cricoid cartilage is a complete ring. The broad part of this ring forms the back of the larynx, while the narrow part encircles the front of the box just below the thyroid. On the top of the broad part of the cricoid are two triangular cartilages (the arytenoids) whose bases form true joints with the cricoid and whose apices extend upward. By means of the joints all these cartilages can move freely upon one another. Beside these four hyaline cartilages there are five yellow cartilage structures. One, the epiglottis, is attached to the top and front of the thyroid, and projects upward to form a lid for the box. Two others form small horn-shaped projections on the top of the arytenoids and the remaining two are inclosed in folds of the membranous lining of the box. The box is lined with a smooth layer of mucous mem- brane. This layer is smooth, except at the point where the glottis or pharynx opening is located. Here it is thrown outward in two sets of ridges. The upper ridges are called the false vocal cords. They have nothing to do with voice production, but help to close the glottis. The two lower488 THE VOICE ridges are made up mainly of elastic tissue running in the membrane from the front angle of the thyroid cartilage to their attachments at the base of the arytenoid cartilages. These ridges are the true vocal cords. They have a glistening white color and sharp smooth edges. In ordi- Fig. 228—I, The vocal cords in the larynx as seen with larynxoscope; A, when voice is being produced; B, when no voice is produced; e, epiglottis; co, vocal cords; cvs, false vocal cords; a, s, w, elevations due to cartilages; /, root of tongue. II, A, the cords alone when voice is being produced; B, vocal cords when no voice is produced. nary breathing they are separated, forming a V-shaped opening to the glottis. The arytenoids are united to one another and to the cricoid cartilages by bands of muscle. By means of these muscles they may be pulled forward, backward, or to one side. These muscle movements affect the cords attached to the arytenoids, pulling them apart, or by their back- ward movement varying the cord tension. Vocalizing.— In ordinary breathing the arytenoids areLAWS OF SOUND VIBRATION 489 drawn forward and to one side and the vocal cords are thus separated. In this position they do not vibrate, and hence produce no sound. In whispering they are in the same position and the sounds produced are in this case due entirely to the position of the lips and tongue. In talking, shouting, singing, however, these cords are drawrn inward and backward by the cartilage muscles, and the expired air is thus forced through the narrower slit between their edges. This sets the edges in vibration producing sound. How this sound is varied in pitch, quality, and loudness requires some knowledge of the laws of vibrating bodies. Laws of sound vibration. — The production of sound depends upon two factors. The first is a vibrating body. The second is a layer of air between this body and the sense organs which produce the impulses. Under these conditions the vibrations of the body are transmitted by the air in the form of waves. If the body which vibrates does so regularly and continuously, a musical tone is produced. If irregularly or at irregular intervals, the effect is called noise. Sound sensations differ in three characteristics, pitch or tone, loudness, and quality. This variation in sensation finds its explanation in corresponding variations in the manner of vibration of the sounding body. (a) Loudness. Of two vibrating bodies vibrating at the same rate, the one whose vibrations are widest are most audible. Thus, a body struck hard produces wider vibrations and hence a louder sound than when struck lightly. (b) Pitch. The number of vjbrations a body makes in a given time determines the pitch or tone of the sound. Thus, a tuning fork whose prongs vibrate two hundred and fifty-six times a second produces the pitch called middle C of the piano, while the next note above it in the musical scale requires two hundred and eighty-eight vibrations per second. Vibrating bodies differ in size and elasticity, and this difference determines their rate of vibra- tion. A large-sized string vibrates slower, and thus gives a lower490 THE VOICE tone than a small one, while a tightly stretched string gives a higher note than one of the same size but less taut. (c) Quality. A piano string and a violin string may be of the same size and tension and be struck with the same force. In that event they will give the same pitch and have the same loud- ness. We are still able, however, to detect that one is a violin string and the other a piano string. This difference between sound producers of the same pitch and loudness we call quality. It is due to the fact that few bodies vibrate simply and as a whole. Most bodies have not only a vibration as a whole which gives what is called the fundamental note, but separate parts of the bodies vibrate more or less independently producing distinct waves, which combine with the fundamental and are called overtones. No two bodies produce these overtones in the same way, owing to differences in size, shape, or material, and hence the complex tone produced is characteristic of each vibrating body and enables us to distinguish between various sources of sound. Resonance.— The quality of a sound may be changed and its loudness increased by reenforcing the original vibration with others of a similar pitch. Thus, a violin string stretched over a sounding board sets the air of the sounding board in vibrations of the same pitch, reenforcing and altering the quality and loud- ness of the original string vibrations. Such reenforcement is called resonance and the reenforcing body is called a resonator. Application of sound laws to vocalizing.—The vocal cords furnish the vibrating bodies which are struck or set in vibration by the expired air. The air cavities of the throat, pharynx, nasal cavity, and mouth are the resonators. By movements of the larynx cartilages the tension of these cords can be changed, and thus the pitch is regulated. The force with which the air is expired determines the width of vibration and the loudness, while loudness and the quality of the tones may be still further modified by changing the shape of the resonating cavities.HYGIENE OF THE VOCAL ORGANS 491 Articulate speech is the result of voluntary changes in these resonating cavities, and the organs most concerned in producing the changes are the lips, tongue, and teeth. Vowels are open-mouth sounds, with corresponding changes in the lip and cheek positions. Consonants are closed-mouth sounds, and tongue, lips, teeth, and cheeks aid in their production. Harshness, softness, etc., are due to differences in the quality of the vocal cord vibrations and the character of the resonating cavities In a child the resonating cavities and larynx are smaller than in adults. At about thirteen to fifteen years of age the larynx widens rapidly and the cords lengthen. The result is a deepening of tone and a change of voice. The vibrations of the cords are under voluntary control, and it is possible by practice to control these vibrations and the shape of the resonators so as to improve greatly the clearness and quality of the voice. Hygiene of the Vocal Organs. Since the voice is under voluntary control it is important that children be taught to control it properly. This teach- ing must be mainly by example, and the best training is secured by giving the child good examples to follow. Sing- ing lessons are also valuable in teaching by exercise the proper control of the vocal muscles, even though the child never become a singer. Exercise which develops the lung capacity also increases the volume and resonance of the voice. If the lining of the larynx becomes inflamed through colds or irritation the vocal sounds are at once affected. Similarly, inflammation of the walls of the resonators or air492 THE VOICE passages reacts upon the voice. On that account colds of all sorts manifest their presence almost immediately in the quality of the vocal sounds. The ordinary precautions against disease of these passages, therefore, act as pre- ventives of vocal troubles.XXXIII. BACTERIA AND SANITATION. Whenever moist foods, such as milk, eggs, meats, bread, etc., are exposed to warm air for any length of time changes take place in their composition which render them unfit for food. These changes we designate as souring, fermentation, decay, mold formation, etc., according to the character of the change. Suclr changes are produced not by the action of the air, but by the presence in the air of certain vegetable organisms commonly called germs. Analysis of these germs shows that there arc three forms of plants which may cause changes in food, and that these three forms belong to the classes yeasts, molds, or bacteria. While these three forms of plants are very different in their appearance and action, they show striking similarity in the manner in which they bring about changes in foods. Of these three forms, the bacteria are perhaps the most varied in their effect; and since they illustrate the action of all, as well as being vitally connected with the health anti disease of the human ani- mal, we are justified in devoting part of our study of phy- siology to the action of these plant forms. Experimental study of bacteria. — (Sec Fix. LXXII.) If we prepare three test tubes by placing in one chopped hay and water, in another some milk, and in a third some wheat and water, and allow all three to stand uncovered for some time in a warm room, we may observe the following changes in the contents of the tubes: The hay solution becomes turbid, and a scum forms on its surface, while from the mixture comes a disagreeable odor. 493494 BACTEEIA AND SANITATION The milk gradually curdles and becomes sour to the smell and taste. The meat gives off fetid odors and changes in color and appearance. If, now, we mount some of the liquid from any one of these three tubes on a slide under a compound microscope, we find it swarming with curious-shaped little bodies myriad in number and with sharply defined outlines. These bodies are extremely small beings, only zo hoo of an inch in diameter. Examined with the highest powers of the microscope they are seen to consist of a cell wall sur- rounding a bit of protoplasm, while scattered through the protoplasm are tiny particles which color more strongly than the other parts, when stained, and are supposed to correspond to the nuclear material of other cells. These tiny bodies are bacteria. Definition and forms of bacteria.— Study of preparations such as the above, and of other sources, shows that there are many different species of bacteria, all differing in shape and action, but all showing the characteristic wall of cellulose and the scattered nuclear material. It is this cellulose wall that inclines us to place these forms among the plants. We can, therefore, define bacteria as follows: One-celled vege- table organisms having a definite unchanging cell wall and containing nuclear material in the form of scattered frag- ments which is never combined in the form of a single nucleus. Certain forms show cilia and arc capable of move- ment. Others move only as they are carried about by the air or liquids in which they arc present. In identifying and naming bacteria, use is made of the unvarying character of the outline. Experience has also shown that definite actions are always associated with defi-BACTERIA AS LIVING CELLS 495 nite shaped cells. Thus, bacillus tuberculosis is the name given to the particular bacterium which causes tubercu- losis. In this name the bacillus refers to the shape of the cell (bacillus = rod shaped), while the second name indi- cates the specific action of this form. Similarly, stophylococcus pyogenes means spherical cells producing pus formation. Spirillum volutans means corkscrew-shaped cells which whirl in their motion through liquids, etc. (See Fig. 230). Bacteria as living cells. — Bacteria contain no chlorophyll (or green col- oring matter) and, therefore, cannot make starch. On this account they must find their food ready made. Living organisms which obtain their food at the expense of other organ- isms are called parasites, and the liv- ing organisms upon which they live are called their hosts. If, however, the source of food be no longer living, as was the case of the chopped hay in the tube, the organism which feeds upon it is called a saprophyte. Bacteria are all para- sites or saprophytes. Those which take their food from living cells, such as the diphtheria and consumption forms, belong to the parasitic forms, while those which live upon dead matter, such as dead animal or plant structures, belong to the saprophytic forms. Like all living cells, bacteria take in food and oxygen, and give out wastes. They are rendered inactive by cold and the presence of certain chemicals, while most of them are Fig. 229 —Bacteria, highly magnified; a, the germ of typhoid fever, stained to show the cilia; b, a spiral ciliated form; c, a rod-shaped form, in chains; d, a spherical form.— a, b, from Eng- LEit and Prantl.496 BACTERIA AND SANITATION killed by exposure to high temperature, such as the boiling- point of water. They require water to develop in, and when there is no water present they become inactive, and in a dried condition may be easily borne about by the winds. This drying, however, does not kill them, anti when trans- planted to a moist place they become active again and grow. The most remarkable feature of their action is the ra- pidity with which they reproduce. Like all single cells, they reproduce by division. In bacteria, however, this pro- cess of division is much simpler than in higher forms of cells and is accompanied by no nuclear changes. When a bacterium has reached a certain size the walls begin to fold in at the middle, a partition forms, and the two halves break apart to form two individuals. Each of these halves ab- sorbs food until it becomes of normal size and then splits again. In active forms this whole process may be repeated as frequently as once in thirty minutes, and it has been calculated that one bacterium can produce at this rate a colony of over 200,000,000,000,000 in twenty-four hours. It is this marvellous rate of development that makes the introduction of a single disease causing bacterium into our system so dangerous. A single diphtheria bacillus can alone produce little damage, but if allowed to develop the increase may become sufficient to cause serious damage in a short time. Occurrence of bacteria and infection.—Bacteria are found in earth, air, and water, and in fact may be present prac- tically everywhere on the earth’s surface. Certain condi- tions are, however, more favorable to their development than others, and fortunately the malignant forms arise only under conditions which are subject to control.HOW BACTERIA PRODUCE DISEASES 49T How bacteria produce diseases and other effects. — In the act of obtaining their food bacteria may bring about chemi- cal changes in the material from which they take it, thus transforming that material into new and simpler com- pounds. Again, they may give off as wastes, materials which are poisonous to the host upon which they live. Such poisons are called toxins. Finally, they may by rapid reproduction become so numerous as to clog effectually the part of the host where they are present, and thus interfere with the normal action of that part. All bacterial effects may be traced to one of these three causes. For example, the souring of milk is brought about as follows: The lactic bacteria found in milk break up the sugar in the milk in the attempt to obtain food and trans- form it into lactic acid. This acid causes the proteid part of the milk to coagulate or curdle, and gives the sour taste and odor. Again, the bacteria which collect in meat break up the complex proteid matter into simpler compounds and thus reduce it to simple mineral compounds. Both of these examples illustrate the first mentioned effect of bacteria. The action of the diphtheria bacterium illustrates the second method. In this case, the bacterium gives off a toxin, or poison, which enters the blood and disturbs the action of the organs of the body. In the case of diphtheria, a substance has been found which if injected into the blood destroys the effect of the toxin, and is called on that account an anti-toxin. The formation of tubercles by the massing of the tuber- culosis forms in the lungs, thereby breaking down the EDDY. PHYS. — 32498 BACTERIA AND SANITATION tissues and clogging the air sacs, is an example of the third method of action. Frequently a given bacterium combines these methods of effect. In every case, however, the or- ganism produces its effect through some result of its meta- bolism, and these secondary effects, such as poisoning or clogging, decay, and the like, are the result of the attempt of one organism to take its food from another. Beneficial bacterial action.— It must not be thought that all bacteria are harmful. If it were not for decay of waste materials, for example, the farmer would be un- able to raise any crops. Decay in his case means the trans- formation of dead plant and animal matter into simpler compounds, such as are acceptable as food to growing plants, and without bacteria the supply of such food in the soil would be soon exhausted. Again, in the manufacture of many dairy products, such as butter and cheese, the bac- teria perform a very important part in the manufacture. Certain forms of bacteria have also been found which, if introduced into the soil with certain plants, can transform the nitrogen of the air into soluble nitrates which the plants can use, and such forms are extremely valuable. Harmful bacterial action.— The forms of bacteria which are most harmful to man are those which produce disease, and the following list gives us some idea of the diseases which are now known to be due to bacterial action: Bubonic plague Asiatic cholera Fowl cholera Hog cholera Diphtheria Lockjaw Erysipelas Consumption Glanders Typhoid fever Boils and carbuncles The forms of bacteria which are beneficial in causing de- cay of waste material may be harmful to food supplies. TheMEDICAL METHODS 499 methods employed in the preservation of foods are devices employed to prevent the action of bacteria upon food stuffs. Relation of business methods to bacteria.— In the para- graphs on the action of bacteria as a living cell we noted certain conditions which were favorable to their develop- ment, and vice versa. The methods used in the preserva- tion of food depend upon a proper use of the facts there noted. For example, in cold storage methods we have the application of the principle that bacteria are rendered in- active by cold. Foods, therefore, will not decay so long as they are kept sufficiently cold to secure this inactivity. We also noted that bacteria need water to develop in. Dried meats, therefore, and foods preserved in salt or sugar, either are free from this necessary water or have the water re- placed by solutions of salt or sugar, which are not favor- able to the action of the plants. The packing of foods in air-tight cans which have been submitted to temperatures of boiling water meets two conditions unfavorable to bac- teria. The heating of the food and can kills the germs that might be present, while the sealing prevents the bacteria in the ah from entering after the foods are cooled. Certain chemicals also kill bacteria, and it has been customary in the past to make use of some of these as preservatives of food. This latter method should always be carefully in- spected, since often the chemicals used may be in such quantity and of such a nature as to be themselves dan- gerous to health. The pure food law in some of its clauses covers just this ground. Medical methods.— The human body is extremely well protected from disease-causing bacteria, since the only ave- nue of approach to most of its organs is through the blood;500 BACTERIA AND SANITATION and to get into the blood is a difficult matter for bacteria, since the skin is of such structure as to prevent passage of the germs. In the case of wounds or breaks in the skin, these serve as excellent passages into the blood for bacteria. The entrance of bacteria into the body is spoken of as in- fection, and the part where they enter is said to be infected. It becomes, therefore, extremely important that a wound which is healing be kept free from these dangerous germs. Medical men have discovered certain chemicals which when applied to wounds do not prevent healing but at the same time do prevent the development of bacteria. Such substances are called antiseptics. All wounds should be first washed as clean as possible with warm water and then bandaged with one of these antiseptics. In this way antiseptics such as 0.1 per cent and 0.05 per cent solutions of carbolic acid and corrosive sublimate, may prevent such serious complications as tetanus or lockjaw, and blood- poisoning. It should be borne in mind that small wounds are quite as dangerous as large ones in this respect and should be treated as carefully. In operations requiring the use of instruments everything used must be freed from bacteria. The process usu- ally consists in treating the instruments and clothing worn as well as the hands to the action of various chemicals or antiseptics. Thus, the instruments are frequently soaked in solutions of carbolic acid or corrosive sublimate or are submitted to the action of hot steam. Materials made bacteria free in this manner arc said to be sterilized or dis- infected. Both terms indicate the same result, but the word disinfected is more frequently applied to the result of chemical treatment, while sterilizing more often refers to the action of heat. In removing a splinter with a needleSANITATION 501 a simple method of precaution consists in sterilizing the needle by holding it in a flame for a moment. The ban- dages used in binding up wounds should all be ‘sterilized in some manner. Disinfection.— While this term means to free from bac- teria, it is frequently used to refer to the methods in which objects arc freed from disease germs. After a patient has been sick of diphtheria or other germ disease it becomes necessary as a matter of safety to the community that all clothing, furniture, and surroundings of the sick person be freed of germs, since their presence there might mean their introduction into some other person. There are various methods of performing this disinfection, but one of. the simplest consists in sealing up the room and burning sul- phur in it. The fumes of burning sulphur are sufficient to kill all forms of harmful bacteria. There arc other kinds of disinfectants beside sulphur, and any form which kills bacteria is entitled to the name. In use it is imperative that the disinfectant be brought in direct contact with the germs. The mere presence of a disinfectant in a room is not a sufficient safeguard, and often disinfectants are sold which give out an odor but do not necessarily come in con- tact with the germs for a sufficient length of time to kill them. It is quite as important to know how to use a dis- infectant as to know its name. One of the best disinfec- tants is plenty of bright sunlight. Sanitation. It is important that we know how to treat cases of bacteria-caused disease. It is still better to prevent such disease altogether. To a certain extent every one is respon- sible for the proper care of his person and property. In a502 BACTERIA AND SANITATION community, however, there are always some people who are careless in this regard, and unless the enforcement of such care be placed in the hands of specific authorities and backed by legal penalties for disobedience the action of one individual may result in great harm to the community. The department of a government, either city, town, or state, which has in charge the care of the public health is called the Sanitary department. In a city the work of sanitation is usually divided among several departments, for example, The Board of Health, The Street Cleaning Department, etc. The extent to which a city board of health looks after the health of its citizens is seen in the following summary of the duties of such a board: Summary of duties of a board of health.— The enforce- ment of all laws, ordinances, and regulations relating to the safety of human life and the care and protection of health. Exclusive control of the treatment of infectious and contagious diseases and of the public hospitals used for such cases. May cause unsafe or unsanitary buildings to be vacated. May condemn and destroy buildings that are themselves unfit for habitation or that make adjacent buildings unfit. May destroy tainted or dangerous food. May prohibit the sale of adulterated milk or other drinks or foods. May prescribe and enforce rules concerning the sanitation of tenements, lodging houses, shops, and dwellings. May be called upon to deal with sources of disease and infection and to keep the streets and houses free from mat- ter dangerous to health. It must insure the pinity of the drinking water.CONCLUSION 503 Iu short, it has full power in any case where the action of an individual tends to menace the health of the community. Department of street cleaning.— This department is en- trusted with the care of the streets and public highways of the city, and the removal of garbage, rubbish etc., from such streets. Conclusion.— To attempt to indicate all the ways in which a community cares for the health of its individuals would be impossible in the limits of this book. It should be borne in mind by every citizen, however, that unless he himself takes an interest in maintaining his own health such carelessness may react upon the community, and that it is the duty of every individual to inform himself of the laws which govern the healthy action of his own body.INDEX. A. Abdominal cavity, 109, 223. Abducent nerve, 389. Abomasum, 157. Abscess, 207. Absorption, 133. definition of, 133. hygiene of, 145. of nutrients, 142. summary of, 142. Accommodation of eye, 402. Acids, 23, 24. Adam’s apple. Adaptation of digestive organs, 154. of limbs, 261. of skulls, 262. Adenoids, 335. Adipose tissue, 44. Adulteration, 6S. Afferent nerves, 403. Affinity, chemical, 13, 15. Air, composition of, 20. expired, 324, 325. inspired, 324, 325. Air breathers, 341. Air sacs, 319. Albumin, 241. Albuminoids, 53, 241. Alcohol: as a beverage, 82. as a drug, 82. as a food, 75, 76, 77, 79, 85, 86. as a fuel, 78. as a producer of heat and energy, 78, 80, 81, 293. 505 Alcohol — as a solvent, 84, 141. effect upon brain workers, 427. effect upon circulation, 210, 211. effect upon digestion, 130,131. effect upon gastric juice, 130, 131. effect upon muscular activity, 284, 292. effect upon nerve control, 211. effect upon nervous system, 427. effect upon respiration, 338. effect upon saliva, 131. effect upon the heart, 211. effect upon the intestines, 131. effect upon the kidneys, 373. effect upon time reactions, 426. effect upon veins and arteries, 211. narcotic or stimulant, 426. percentage in beverages, 84. percentage in patent medi- cines, S4. relation of study to physi- ology, 75. Alimentary canal, 52, 90. functions of, 93. in earthworm, 150. regions of, 92, 93. Alkalies in saponification, 126. Amceba, circulation in, 212. digestion of, 146, 147. movement of, 298. study of, 33. Amoeboid movement, 297, 298.506 INDEX Ampulla, 474. Amylopsin, 125, 126. Anabolism, 59, 352. Anaerobia (footnote), 341. Anatom y, 11. Anemia, 16S. Ankle, bones of, 233. Ankylosis, 243. of joints, 251. Antiseptics, 500. Anti-toxin, 337. Anus, 129. Anvil bone, 235. Aorta, 186, 197, 199. Aortic arches, 214. Apoplexy, 39S. Appendages of the body, 30. Appendix, vermiform, 129. Aqueduct or iter, 396. Aqueous humor, 457. Arachnoid membrane, 392. Arbor vita?, 397. Arms, bones of, 232. Arterial blood, 187, 188. Arteries, 186-189. control, 205. pulmonary, 185, 195. pulse in, 186. renal, 201, 367, 368. structure, 1S9. Arthropods: body covering, 375. circulation in, 212, 213. excretion in, 381. nervous system of, 433. respiration in, 346. Articulations, 244. Arytenoid cartilages, 487. Asphyxia, 332. Assimilation, 175. Astigmatism, 123. Atlas vertebra, 225, 227. Auditory sensation, 470-479. Auricles, 182-184. action of, 191, 192. Auscultation, 193. Autonomic, system, 421. Axial skeleton, 224. Axis cylinders, 47, 402. Axis vertebra, 225-227. B. Bacteria, 493-503. action of, 498. as living cells, 495. experimental study of, 493. forms of, 494. metabolism of, 496. occurrence, 496. relation to blood poisoning, 210. relation to business methods, 499. relation to disease, 497. relation to fermentation in large intestine, 130. relation to inflammation, 207, 208. relation to medical methods, 499. relation to nervous diseases, 424. relation to respiratory dis- eases, 337. reproduction of, 496. size of, 494. structure of, 494. Ball and socket joints, 230, 231. Bases, 23, 24, 25. Baths and bathing, 363. Beef extract, nutritive value of, 74. Bellied muscles, 267. Biceps muscle, 206, 209. Bicuspid teeth, 95, 96. Bile, 122, 124, 127. Bile duct, 124. Biliousness, 144. Binocular vision, 466. Biology, 10.INDEX 507 Birds, circulation in, 217. digestive system of, 150. heart of, 217, 218. respiration in, 349. skeleton of, 260. Biuret test for proteid, 53. Bladder, 367, 371. Bleeding, 209. Blind spot in eye, 465. Blood, 159-179. absorption into, 134, 135. aerating of, 203, 327. arterial, 1S7, 1SS, 203. changes in lungs, 326. clotting of, 161. cold, 351. composition of, 201, 202. conditions affecting, 170. experimental study of, 161. extractives, 164. flow, 191, 194-196, 204. in other animals, 173. in veins, arteries, and capil- laries, 187, 188. loading of, with nutrients, 201, 202. loading of, with wastes, 202. plasma, 161. poisoning, 210, 365. properties of, 161. purifying, 203, 327. quantity and distribution, 171. relation to circulation, 159. relation to respiration, 326. structure, 160. systems, 191. temperature, 171. unloading of nutrients from, 202. unloading of wastes from, 203. venous, 203. warm, 351. Blood corpuscles, red, 160, 165, 167, 238. white, 167. Blood plates, 165, 169. Blood proteids, 164. Blood vessels, 159. of the heart, 184. of the skin, 356. relation to alimentary canal, 130, 134. See Arteries, Veins, Capillaries. Blowing, 331. Boards of Health, duties of, 502. Body, parts of, 30. Body cavity in earthworms, 150. Body covering of lower animals, 374-376. Body framework, 221. Boils, 208. Bones, 221-243. composition of, 240. diseases of, 248-249. effects of age upon, 241. forms of, 236, 237. fractures of, 250. functions of, 222. regions of, 223. structure, 42, 43, 237-243. Bone cells, 243. Border cells, 113. Botany, 10. Bow legs, 249. Bowel movements, 145. Brain: development, 434. diseases, 424. external features of, 393. forebrain, 388. frog, 388. hindbrain, 388. human, 390. internal features of, 395. membranes, 390. midbrain, 388. structure, 392-397. Breastbone, 223. Breathing, 328. abdominal, 328. causes of, 322, 323.508 INDEX Breathing— costal, 328. habits, 328. hygiene of, 331. modifications of, 341. mouth, 314. movements, 320. rate of, 324. relation of sex to, 324. Bright’s disease, 373. Broken bones, 250. Bronchi, 315. Bronchial tubes, 316. Bronchitis, 208, 336. Buccal cavit3r, 93. Bunions, 249. Burning (combustion), 19. Burns and blisters, 365. C. Caecum, 12S. Csecapylonic in fish, 153. Calcium, 13, 26. Calices of kidney, 368. Canals, semicircular, 473, 477. Canalicule, 43, 241. Canines (teeth), 95, 96. Capacity of lungs, 329. vital, 329. Capillaries, 186, 1S7, 188. in the lungs, 319. seat of absorption, 134. structure of, 189. Capsules (bony), 42. Carbohaemoglobin, 166. Carbohydrates, 26, 53. as fuel, 60. digestion of, 132. Carbohydrates in muscle, 276. oxidation of, 27. Carbon, 16, 25, 26. Carbon dioxide, 16, 26. excretion bv skin, 362. See Respiration. Cardiac cycle, 192. Cardiac muscles, 266, 275. Cardiac nerves, 205. Cardiac orifice (of stomach), 111. Carnivora, teeth of, 155. Carotid artery, 199. Carpals (bones), 232. Cartilage, a form of connec- tive tissue, 42. articulations, 227, 240. in bones, 222, 240. Casein and caseinogcn, 26, 115. Cataract, 46S. Catarrh, 208. Cavities, body, 29. Cells, air, 319. auditory, 476. defined, 33. Cells, forms of, 31. in tissues, 31. muscle, 282, 283, 2S4. nerve, 47, 405, 406. smell, 449. structural units, 32. structure of, 32. taste, 446. Central nervous system, 385. Centrum (of vertebra;), 225, 226. Cerebellum, 393. Cerebellum, function of, 418. Cerebral hemispheres, 392. distribution of gray matter in, 398. Cerebral localization, 419. Cerebro-spinal fluid, 398. Cerebrum, 392, 393. functions of, 418. Cervical vertebrae, 224, 226, 227. Charcoal, 16. Cheek bones, 234. Chemical compounds defined, 14.INDEX 509 Chemical compounds, dis- tinguished from mixtures, 14. Chemical compounds in the body, 25. Chemical compounds, relation to affinity, 13. Chemical elements, 12,13, 28. Chest cavity, 320. Chiasma optic, 393. Chief cells, 113. Chocolate, 74. Choking, 331. Chordae tendinae, 1S5, 19. Choroid layer of eyes, 457. Choroid flexus, 398. Chyle, 141. Chyme, 116. Cigarettes, 88. Cilia in epithelial tissue, 41. in hydra, 14S. use in digestion, 147. use in lungs, 318. Ciliary ligament, 457. Ciliary movement, 297-299. in protozoans, 300, 301. in higher forms, 302. Ciliary muscles, 458. Ciliary processes, 457. Circulation of blood, 180, 211. Circulation effects of alcohol upon, 210. in lower animals, 211, 219. portal, 200. pulmonary, 195. renal, 201. significance of, 218. systemic, 197. systems, defined, 180. Circumvallate papillae, 445. Clavicles, 229, 230. Cloaca, 153, 381. Clotting of blood, 161, 169. Coal, 16. Coccyx, 225. Cochlea, 473. Cocoa, 74. Coelenterates, 341, 375, 429. Coeliac axis, 199. Coffee, 74. Cold-blooded animals, 35. Colds, 20S, 336. Collar bone, 229, 230. Colloids, 137. Colon, 128, 129. Color production, 466. blindness, 466. Combustion, 18,19. in muscles, 282, 283. Complexion, care of, 364. Condiments, 73. Cones and rods, 465. Congestion, 206, 207. Conjunctiva, 453, 456. Connective tissue, 41, 42, 43. Consciousness, 418. Conservation of energy, 62. Constipation, 145. Contractility, 35. as a cause of locomotion, 297. of muscles, 28. Cooking, 70. Coral, skeleton of, 254. Corium, 355. Cornea, 456. Corns, 366. Coronary arteries, 186. system, 184,199. veins, 184. Corpora quadrigemina, 397. Corpora striata, 395. Corpus callosum, 395. Corpuscles, bone, 42. red, blood, 160, 165, 167, 238. white, blood, 167. Cortex of brain, 405. Cortex of kidney, 368. Corti, fibers of, 474. organs of, 475. Costal cartilages, 229. Coughing, 331.510 INDEX Cranial bones, 233. Cranial nerves, 388, 389, 394. Cranium, 234, 390. Cricoid cartilages, 487. Crop, 151,156. Croup, 337. Crura cerebri, 393. Crustacea, skeleton of, 256. Crying, 331. Crypts, 120. Crystalloids, 137. Cud, 156. Cutaneous sensations, 43S, 439. Cuticle, 354. Cuts, 209, 365. Cystic duct, 122. Cytoplasm, 34. D. Defalcation, 144. Deglutition, 107. Delirium tremens, 42S. Dendrites, 47. Dental formula;, 96. Dentine, 97. Dentition, 96. Dermis, 354-356. Dextrose, 54. Diabetes, 143. Dialysis, 136, 13S. Dialyzer, 136. Diamond, 16. Diaphragm, 109, 180, 316, 320, 321.' Diastase, S3. Diastole, 192. Diet, daily, 70. Digestibility of nutrients, 67. relation to food value, 67. Digestion, 90-132. chemistry of, 99. defined, 90. extra cellular, 149. hygiene of, 145. Digestion — in lower animals, Chapter X, 146. of the mouth, 106. of the large intestine, 128, 129. of the small intestine, 118, 125-128. of the stomach, 114. peptic, 114-116. summary of, 132. Diphtheria, 20S, 337. Disinfection, 500, 501. Dislocation of joints, 250. Distillation, S3. Distilled liquors, 83. Division of labor, 49. Dorsal vertebra;, 224-226. Ducts bile, 124. cystic, 124. thoracic, 135, 141. Duodenum, 119. Dura mater, 390. Dust and dusting, 334. E. Ears, bones of, 235. drum, 471. external, 471. hygiene of, 478. inner, 472. middle, 471. passages of, 474. structure and form, 470. vestibule, 473. wax, 471. Earthworm, blood in, 215. circulation of, 214. digestion of, 149. excretion in, 379. Earthworm respiration in, 343. Ectoderm, 375. Ectoplasm, 34. Elastic fiber, 41,XUDEX 511 Elbow joint, 232. Elements, chemical, 12, 13, 28. Emulsion, 127. Enamel (tooth), 97. Endoplasm, 34. Endoskeleton, 221. Energy, conservation of, 62. definition of, 60. kinetic, 62. muscular, 282, 2S3. potential, 61. sources of, 59, 61. transformation of, 61. Enzymes, defined, 102. in muscle, 276. Epidermis, 353, 354. absence of blood in, 305. cells of, 354. nerves of, 355. Epiglottis, 108, 315, 487. Epithelial tissue, 39, 40. Erythroblasts, 168. Esophagus, action of, 109. structure of, 108. Ethmoid bone, 234. Eustachian tube, 107, 471, 472. Evolution, 3S, 21.9. Excretion, 352-373. in lower animals, 374-382. Exercise, forms of, 286. relation to muscles, 285, 286. relation to breathing, 329. Exhaling, 319, 322, 323, 324. Exoskeleton, 221. Extractives, alcoholic, S4. Eye, 452-469. diseases, 469. functions of, 458. hygiene of, 467. muscles of, 454. position of, 452. structure of, 452-455. Eyeballs, 452-455. Eyesockets, bones of, 235. Eyelids, 453. F. Face bones, 233-235. Facial nerve, 389. Faeces, 129, 144. Fangs (tooth), 97. Far sight, 468. Fatigue muscle, 285. Fatigue, nerve, 423. retinal, 466. smell, 450. Fats, 26, 27, 60. absorption of, 140. digestion of, 126, 127. emulsification of, 127. forms of, 55. in bones, 241. in muscle, 276. saponification of, 126. tests for, 55. Fatty acids, 126. Feathers a form of skin, 377. Fehling’s solution, 54, 55. Femur, 232. Fenestra ovalis, 474. rotunda, 471. Fermentation, 82, S3, 102. Fermented drinks, S3. Ferments, 102. Fibers, muscle, 302, 303. nerve, 402, 404. Fibrus and fibrinogen, 26, 162. Fibula, 232. Filiform papillre, 445. Filtration, 139. Fingers, bones of, 233. Fissures of brain, 393. Fish, bones of, 257. circulation of, 215. digestion of, 152. gills of, 215, 348. heart of, 215. respiration of, 348. Flagella, 301. Flavors, 73. Flat bones, 239.512 INDEX Floating ribs, 229. Fontanelle, 243. Foods, 51, 89. cost of, 68, 69. composition, 56-59. determination of fitness, 67. preservation, 499. values, 63. Food accessories, 73. Food stuffs, 53. Food vacuoles, 147. Foot, bones of, 233. Foramen magnum, 234, 390. Foramen Monroe, 395. Forebrain, 388. Fornix, 395. Fractures, 250. Freckles, 354. Frog, bones of, 257, 258. brain of, 3S7. circulation of, 216. digestion of, 153. heart of, 216. nerves of, 3S6, 387. respiration of, 348. Frontal bone, 234. Fuel, 60, 282. Fundus of stomach, 117. Function defined, 8. Fungiform papillae, 445. G. Gall bladder, 122. Gases in the body, 25, 26. Gastric glands, 113. Gastric juice, 114. effect of alcohol upon, 130, 131. Gelatin, 26, 241. Germicides, 210. Germs, 493. Gills, 343, 144, 346-348. Gizzard, 151, 155. Glands, 99-101. in hydra, 149. Glands — lachrymal, 453. lymphatic, 178. oil (see Sebaceous). salivary, 94, 102, 103, 104. sebaceous, 358. sweat, 357. wax, 471. Glomerulus of kidney, 370. Glossopharyngeal nerve, 389. Glottis, 107. Glucose, 54. Gluten, 52. Glycerine, 126. Glycogen, 27. in liver, 142,143.' in muscles, 276. Grape sugar, 54,143. Graphite, 16. Gustatory sensations, 444. H. Habit, 413. Haemocyanin, 344. Haemoglobin, 165, 166. Hair, 358. care of, 364. follicles, 359. nerves of, 358. papillae, 358. Hammer bone, 225. Hand, bones of, 233. Hard palate, 93, 235. Haversian canals, 43, 241, 242 Head, bones of, 233. Headache cures, S9. Health, Board of, 502. Hearing, 470. in lower animals, 482. Heart beat, 183, 191, 204, 205. muscle, 266, 275. nerves, 205. relation to lungs, 312, 316. sounds, 192, 193. structure, ISO, 183.INDEX 513 Heat generation, 14,19. relation to oxidation, 19, 27. sensation, 442. Heel bone, 233. Hepatic artery, 123. cells, 124. duct, 122. vein, 123, 200. Herbivora, 15S. Hiccoughs, 332. Hindbrain, 3S8. Hinge joint, 245. Hip bones, 232. joint, 230, 231. Histology, 9, 39. Host, 495. Humerus bone, 232. Humor, aqueous, 457. vitreous, 457. Hunger, 117. Hydrochloric acid, 113, 116. Hydrogen, 15, 21, 22. Hygiene, 8, 9, 10. of bones and joints, 248. of circulation, 206. of digestion, 145. of the ears, 47S. of the eyes, 467. of muscles, 285, 286. of nervous system, 422. of respiration, 330. public, 501, 502. Hyoid bone, 375. Hypodermis, 375. Hypoglossal nerve, 394. I. Ileocolic valve, 128. Ileum, 119. Iliac bones, 231. Images, 460, 463. Incisors (teeth), 95, 96. Incus bone, 472. Indigestion, 145. Infection, 496. Inflammation, 206, 208. Infundibulum, 395. Inhaling, 319, 322-324. Inhibitory nerves, 205, 404. Insects, circulation of, 212, 213. excretion of, 381. respiration of, 346. Inspiration, 319, 322, 324. Instep, bones of, 233. Intercellular material, 44. Intercostal muscles, 317. Internal sensations, 444. Intestine, small, 118, 119-121. large, 128-130. Intestinal juice, 121, 127, 128. Intoxication, 424. Invertose, 128. Involuntary muscle, 266. Iris, 456, 457. Irritability, 35, 280. Ischium, 231. Iter, 396. J. Jaw bones, 235. Jejunum, 119. Joints, 244-251. diseases of, 249, 250, 251 dislocations of, 250. forms of, 244-248. structure of, 245. Jugular veins, 141, 200. K. Katabolism, 59, 352. Keratin, 26. Kidneys, 366-372. of insects, 381. of vertebrates, 38. sugar, disposal of, 143 Knee joints, 232.514 INDEX L. Labyrinth of ear, 473. Lachrymal bones, 235. duct, 453. glands, 453. Lacing, 249. Lacteals, 135, 141. Lactose, 54. Lacunas (bone), 42, 43, 241. La grippe, 337. Lamellae (bone), 43. Large intestine, 128, 129. Larynx, 315, 486, 4.87. Lateral ventricles, 395. Laughing, 331. Leather, 355, 378. Legs, bones of, 232. Lens, action of, 460. crystalline, 457. Leucocytes, 16S, 207. Levers, 288, 289. Ligaments, function of, 245. transverse, 227. Light, effect of, 460, 461, 463. nature of, 461. Lime, in bones, 240. Lime water and C02, 16. Limula, 361. Lipase, 126. Litmus, 23. Liver, as an absorbent organ, 142. as an excretory organ, 142, 372. as a storehouse of energy, 143. structure of, 121-123. veins of, 201. Liver cells, 123. Liver starch, 142. Lobes, optic, 38S, 392. Locomotion, causes of, 295. due to contractility, 297. due to secretion, 296. due to specific gravity, 295. Locomotion, organs of, 305- 309. Loudness, 489. Lumbar vertebrae, 225, 226, 227. Lungs, blood changes in, 326. capacity of, 329. capillaries of, 319. development, 348-350. excretory action of, 353. structure of, 312-319. Lymph, 174-179. capillaries, 177. exchange, 175. flow of, 177, 178. formation of, 174. glands, 178. nodes, 178. relation to absorption, 135, 175. relation to assimilation, 175. relation to osmosis, 175. relation to plasma, 174. tubes, 177. valves, 177. Lymphatics, in villi, 133, 134. system of, 176. Lymphocytes, 168. M. Malars (bones), 234. Malleus (bone), 472. Malpighi, capsules of, 369. pyramids of, 369. tubules of, 369. Malt, 83. Manuals, 21S, 261, 350. Man, biological position of, 37. Mandibles (bones), 235. Manganese dioxide, 18. Marrow (bone), 237. Maxillae (bones), 235. Meats, cooking of, 71. Medulla oblongata, 394, 420.INDEX 515 Medullary layer of kidney, 369. sheath of nerves, 402. Membrane, mucous, 94, 314. Membranes of brain and cord, 392, 400. of tympanum, 472. synovial, 246. Mercuric oxide, 17. Mesenteric arteries, 199. Mesentery, 119. Metabolism, 59, 352. Metacarpals (bones), 232. Metatarsals (bones), 232. Metallic elements, 23. Microbes (see Bacteria). Midbrain, 388. Milk, 57, 64, 248. Milk glands, 377. Milk teeth, 96. Millon’s reagent, 53. Minerals, 24, 25. Mineral salts, 55, 56. Mitral valves, 183, 186, 191. Mixtures distinguished from chemical compounds, 14. Molars (teeth), 95, 96. Mollusks, 255, 375, 430. Motor nerves, 403. Mouth digestion, 106. Movement, amoeboid, 297, 298. ciliary, 297, 299. in lower animals, Chapter XX, 294. muscular, 297. Mucous membrane, 94, 314. Mucus, 94. Muscles, 265-293. action of, 27S, 287, 288. attachment to bones, 240, 268. bellied, 267. biceps, 206, 209. comparison with engine, 281- 283. composition of, 276, 277. contraction of, 278, 281. Muscles — digastric, 268. efficiency of, 287. fatigue of, 285. forms of, 267-269. heart, 266. involuntary, 265, 266. nerve endings in, 279. non-striated, 266, 274, 275. papillary, 185. rib, 317. striated, 266, 267, 269. triceps, 269. voluntary, 265, 266, 274. Muscle cells, 282, 283, 284. Muscle fibers, 272, 273, 305. fibrils, 274. Muscular movements, 297-302, 303, 304. Muscular tissue, 44, 265, 266. Musical sounds, 489. Myogen, 277. Myopia, 467. Myosin, 26, 276. N. Nails, 359, 360, 365. Narcotics, 86. Nasal (bones), 235. Near sight, 467. Nephridia, 379. Nerve cells, 47, 405, 406. Nerve control, 291. Nerve diseases, 424. Nerve endings in muscle, 279. Nerve fibers, 402-404. excitating, 404. inhibitory, 404. medullated, 403. Nerve muscle preparation, 279. Nerve processes, 47. Nerves, afferent, 403. cranial, 3S9. dermal, 356.516 INDEX Nerves — efferent, 403. epidermal, 354. motor, 403. sensory, 403. vaso-motor, 205. Nervous system, 382-428. central, 385. composition, 385. diseases of, 424. effects of alcohol upon, 424. functions of, 411. hygiene of, 422. in lower animals, Chapter XXVII, 429. sympathetic, 385. Nervous tissue, 46. Neura, 47, 405, 406. theory of action, 406-409. Neural arch, 226. Neural cavity, 225, 226, 397. Neuralgia, 424. Neurilemma, 403. Neuron (see Neura). Neutralization, 23, 24. Nicotine, 87. Nitrogen, 20. Nitrogenous compounds, 26, 52. Nodes of Ranvier, 402. Non-metallic elements, 23. Non-nitrogenous compounds,26. Non-striated muscle, 44,45,266, 274, 275. Nose, bones, 235. Nose, sense organs of, 448. Nucleus, 32, 34. Nucleolus, 34. Nutrients, 52. absorption of, 139. forms, 52, in blood, 163. in bone, 241. j in foods, 57. sources of, 56. uses of, 59. Nutrition, 51-72. O. Oblique muscles, 110. Occipital bone, 234. Ocelli, 483, 484. Oculo-motor nerve, 388. Odontoid, process, 227. Odors, 450. Olfactory lobes, 392, 393. nerves, 388. sensations, 448. Omentum, 111. Opium, 88, 89. Optic chiasma, 393. lobes, 388,392. nerve, 388. thalami, 396. Orbit of eye. Organic chemistry, 16. compounds, 26. Organs, 30, 31. of Corti, 475. Osmosis, 136, 138, 163. Otocysts, 483. Otoliths, 473. Oxidation, 18, 19, 282 (see Respiration). Oxides, 19, 25. Oxygen, 13, 17, 18, 19, 26 (see Respiration). Oxyhaemoglobin, 166. P. Pain, sensations, 443. Palate, bones, 235. hard, 93, 235. soft, 93, Pancreas, 124, 125. Pancreatic juice, 125, 127. Papillae, dermal, 355, 356. hair, 358. nail, 360. taste, 445. Papillary muscles, 185.INDEX 517 Paramoecium, 146, 147. Parasite, 495. Parietal bones, 234. Parotid glands, 102. Patent medicines, 64. Pectoral girdle, 223, 230. Pelvic girdle, 230, 231. Pelvis, 223, 232. of kidney, 368. Pepsin, 113, 115. Peptic digestion, 117. Peptone, 115,137,140. Pericardium, 181. Perilymph, 473. Perimysium, 270, 271. Perineurium, 402. Periosteum, 237, 240, 241. Peristalsis, 109, 125. Peritoneum, 109. Perspiration, 357,362,363. Phagocytes, 169. Phalanges, 232. Pharynx, 94, 107. Phosphorus, 13. Phrenology, 419. Physiology, 7, 8, 9, 10. Physiological chemistry, 10. Pia mater 392. Pigment cells, 354. Pitch, 489. Pivot joint, 247. Plant physiology, 11. as a source of nutrients, 58. Plasma, 161. Pleura, 319. Pleurisy, 336. Plexus, 404, 405. Pneumonia, 208, 336. Poisons, 144. Pons varolii, 393. Portal circulation, 200, 201. Portal vein, 123, 134, 142. Potassium chlorate, 17. Premolars (teeth), 95, 96. Presbyopia, 469. Pressure sensations, 440, 441. Proteid, 52. absorption of, 138, 139. action of trypsin upon, 126. action of gastric juice upon, 114-116. blood, 163, 164. composition of, 26. digestion of, 126, 114-116. forms of, 26, 52. muscle, 276. sugar formers, 144. tests for, 53. Protoplasm, 32-36. movement of, 294. Protozoans, 252, 341, 374, 429, 480. Psalterium, 157. Pseudopodia, 35, 298. Ptyalin, 105. Pubis (bones), 231. Pulmonary artery, 185, 195. capillaries, 196. sacs, 316-319. veins, 185, 197. Pulp cavity (tooth), 97. Pulse, 193,' 194, 195. Pupil of eye, 457. Pure Food Law, 84. Pus, 207. Pyloric orifice, 111. Pylorus, 112. Pyramids of Malpighi, 369. Q- Quality of sound, 490. R. Radial artery? 199. Radius bone, 232. Rectum, 128. Reflex action, 412. arc, 413.518 INDEX Renal artery, 201, 367, 368. circulation, 201. veins, 201, 367, 368. Rennin, 113. Reproduction of cells, 50. Reptiles, 217, 259, 349. Residual air, 329. Resonance, 490. Respiration (see Breathing), 312-339. artificial, 332, 333. effects of alcohol upon, 338. in lower animals, Chapter XXII, 340. organs of, 313, 314, 320, 324. Respiratory organs, 313, 314. diseases of, 335. Rest, 423. Reticulum, 157. Retina, 457, 463-465. _ Ribs, action in respiration, 320. cartilages of, 229. floating, 229. muscles of, 317. processes, 229. relation to thorax, 223. types of bone, 238, 239. Rickets, 248. Rigor mortis, 276. Rodents, teeth of, 155. Rods and cones, 465. Rods of Corti, 475. Round shoulders, 249. Rumen, 157. Ruminants, 156, 157. Running, 290, 291. S. Saccharose, 54. Sacculus, 473. Sacrum, 225, 231. Saliva, action of, 105. composition, 94, 104. effect of alcohol upon, 131. Salivary glands, 94, 102, 103, 104. Salts, absorption of, 141. blood, 163, 164. characteristics, 23-25. Sanitation, 10, 493, 501, 502. Saponification, 126. Saprophyte, 495. Sarcolemma, 45, 272, 273, 274, 277. Sarcoplasm, 45, 273. Sarcostyles, 274. Scapula, 230, 232. Sclerotic layer of eye, 456. Sebaceous glands, 356, 358, 471. Secretion, 99. a property of epithelium, 40. as a cause of motion, 296. Semicircular canals, 473, 477. Semilunar valves, 185,186, 191, 192. Selection of food, 36. Sensation, 436-485. auditory, 470. classification of, 438. cutaneous, 438. gustatory, 444. in lower animals, 480-485. internal, 444. olfactory, 448. pain, 443. pressure, 440. temperature, 442. visual, 452. Senses, special {see Sensation). Sensory organs, development of, 481. Septa, lucida, 395. Serous membrane, defined, 110. Serum, 162, 163. Sheath, medullary, 402. Shoulder blades, 230. Shoulder joints, 232. Sighing, 332. Sight sensation, 452, 465.INDEX 519 Sigmoid flexure, 128. Silver sulphide, 15. Skeleton, 221-251. appendicular, 230. axial, 224. bones of, 222. external, 29, 221, 252. hygiene of, 248. in lower animals, 252-263. internal, 29. 221, 252. Skin, 353-363'. as an excretory organ, 361. of lower animals, 374-382. functions of, 361. hygiene of, 363. modifications of, 353-356. structure of, 353. Skull, 223, 233. uses of, 235, 236. Smell, sensation of, 448-450. in lower animals, 482. Snails, 345. Sneezing, 331. Soaps, 126, 140. Sobbing, 331. Sore throat, 208, 335. Sound, 474, 489, 490. Special senses (see Sensation). Speech, 486. Sphenoids, 234. Sphincter pylorus, 112, 117. Spinal accessory nerve, 394. Spinal column, 222, 224, 228. Spinal cord, 398. center reflex action, 417. connection with sympathetic system, 401. diseases of, 424. functions of, 417. membranes of, 399. of the frog, 389, 390. structure, 401. Spinal nerves, 400, 401, 402. Spinous process, 226. Spiracles, insect, 346. Spitting, 332. Spleen, 168. Sponges, 148,212,237,253,341, 378. Sprains, 250. Spyrogyra, 32. Squameous epithelium, 40. Standing, 290, 291. Stapes (bone), 235, 472. Starch, absorption, 140. composition, 27, 53, 54. digestion of, 105, 126. in blood, 163. Starfish, 254, 342. Steapsin, 125, 126. Sterilizing, 500. Sternum, 228, 229. Stimulants, 74. Stirrup bones, 235. Stomach, as a storehouse, 116. curvatures of, 112. enzymes of, 113. glands of, 113. method of action, 117. structure, 111. Strangulation, 331. Striated muscle, 266, 267, 269, 270, 272-274. Stroma, 166. Structure, 8, 28. Subcutaneous layer, 355. Sublingual glands, 102. Submaxillary glands, 102. Submucous coat, 111. Succus entericus, 127. Suffocation, 331. Sugar, composition, 27. forms, 53-55. in blood, 163, 164. in muscle, 277. tests for, 54, 55. Sulphur, 15. Suspensory ligament, 458. Sutures, 235, 243, 247. Swallowing, 107. Sweat, 356. Sweat glands, 356, 357.520 INDEX Sympathetic nervous system, 421. Symphyses, 243, 247. Synovial fluid, 245, 246. membrane, 246. Systematic circulation, 198-200. Systems, defined, 30. Systole, 192. T. Tarsals (bones), 232. Taste sensation, 444-447. Taste buds, 446. Taste in lower animals, 482. Tea, 74. Tears, 453. Tear glands, 453. Teeth, 94-96. in lower forms, 152-155. structure, 97. use in digestion, 98. Temperature, body, 19, 171, 172, 363. sensations, 442. Temporals (bones), 234. Tendons, 238, 267, 271. Theobromin, 74. Thoracic duct, 135, 141. Thorax, 223. Thyroid cartilage, 487. Thumb, bones of, 233. Tibia (bone), 232. Tidal air, 330. Time reactions, 426. Tissue, 31, 39, 4S. Tobacco, 87, 88, 284. Toes, 233. Tongue, 94, 99, 444, 445. Tonsils, 94. Tonsilitis, 335. Touch, 440, 481. Tourniquet, 209. Toxins, 497. Trachea, 315, 317, 31S. Trachete, insect, 346. Triceps muscle, 269. Tricuspid valves, 183, 185, 191. Trigeminal nerve, 389. Trochlear nerves, 389. Trunk, 29. Trypsin, 125, 126. Tuberculosis, pulmonary, 337. of the joints, 251. Turbinate bones, 235. Tympanum membrane, 471. Tympanum, 471. Typhlosole, 150. U. Ulna, 232. Ulnar artery, 199. Urea, 26, 352, 357, 362, 371, 372. Ureter, 366. Urethra, 367. Urine, 371. Urinary organs, 366. Utriculus, 473. Uvula, 94. V. Vacuoles, 34. Vagus nerves, 389. Valvulae conniventes, 120. Vaso-motor nerves, 205. Vegetables, cooking of, 71. Veins, 186-190. Vena cava, 184, 191, 197, 200. Venous blood, 203. Ventilation, 333. Ventricles (heart), 183, 148, 191, 192. Ventricles of the brain, 395, 396, 397. Vermiform appendix, 129.INDEX 521 Vertebra, 222-240. cervical, 224, 226, 227. dorsal, 224, 225, 226. lumbar, 225, 226, 227. Vertebrates, 222, 252, 376, 433: Vestibule of ear, 473. Vestibular passage, 474, 476. Villi, 120, 134, 135. Viscera, 29. Vision, 452. in lower animals, 4S3. Visual sensation, 452. Vocal cords, 488. organs, 486. Vocalizing, 488. Voice, 486-492. Voice, hygiene of, 491. Voluntary action, 412. arc, 415. muscle, 266. Vomer, 235. W. Walking, 290, 291. Warm blood, 351. Water, absorption of, 141. as a nutrient, 56. breathers, 341. composition, 22, 25. Wax glands, 471. Wind pipe, 315. Wisdom teeth, 96. Worms, 343, 375, 432. Wrinkles, 356. Wrist, 233. X. Xanthoproteic test, 53. X-ray, 220-221. Y. Yawning, 332. Yellow spot of eye, 465. Z. Zoology, 10.OUTLINES OF BOTANY $1.00 By ROBERT GREENLEAF LEAVITT, A.M., of the Ames Botanical Laboratory. Prepared at the request of the Botanical Department of Harvard University Edition with Gray’s Field, Forest, and Garden Flora..................$1.80 Edition with Gray’s Manual of Botany.................................2.25 THIS book icovers the college entrance requirements in botany, providing a course in which a careful selection and a judicious arrangement of matter is combined with great simplicity and definiteness in presentation. The course offers a series of laboratory exercises in the morphology and physiology of phanerogams ; directions for a practical study of typical cryptogams, representing the chief groups from the lowest to the highest ; and a substantial body of information regarding the forms, activities, and re- lationships of plants and supplementing the laboratory studies. «U The work begins with the study of phanerogams, taking up in the order the seed, bud, root, stem, leaf, flower, and fruit, and closing with a brief but sufficient treatment of cryptogams. Each of the main topics is introduced by a chapter of laboratory work, followed by a descriptive chapter. Morphology is treated from the standpoint of physiology and ecology. A chapter on minute structure includes a discussion of the cell, while another chapter recapitulates and simplifies the physiological points previously brought out. The limitations of the pupil, and the restrictions of high school laboratories, have been kept constantly in mind. The treatment is elementary, yet accurate ; and the indicated laboratory work is simple, but so designed as to bring out fundamental and typical truths. The hand lens is assumed to be the chief working instrument, yet provision is made for the use of the compound microscope where it is available. AMERICAN BOOK COMPANY (■74)A NEW ASTRONOMY $!-3° By DAVID TODD, M.A., Ph.D., Professor of Astron- omy and Navigation, and Director of the Observatory, Amherst College. ASTRONOMY is here presented as preeminently a science of observation. More of thinking than of memorizing is required in its study, and greater emphasis is laid on the physical than on the mathematical aspects of the science. As in physics and chemistry the fundamental principles are connected with tangible, familiar objects, and the student is shown how he can readily make apparatus to illustrate them. In order to secure the fullest educational value astronomy is regarded, not as a mere sequence of isolated and imperfectly connected facts, but as an inter-related series of philosophic principles. The geometrical concept of the celestial sphere is strongly emphasized; also its relation to astronomical instru- ments. But even more important than geometry is the philo- sophical correlation of geometric systems. Ocean voyages being no longer uncommon, the author has given rudimental principles of navigation in which astronomy is concerned. The treatment of the planets is not sub-divided according to the planets themselves, as is usual, but according to special elements and features. The law of universal gravitation is unusually full, clear, and illuminating. The marvelous dis- coveries in recent years and the advance in methods of teach- ing are properly recognized, while such interesting subjects as the astronomy of navigation, the observatory and its instruments, and the stars and the cosmogony receive particu- lar attention. *f[ The illustrations demand special mention; many of them are so ingeniously devised that they explain at a glance what many pages of description could not make clear. AMERICAN BOOK COMPANY (181)TEXT-BOOKS ON GEOLOGY By JAMES D. DANA, LL.D., late Professor of Geology and Mineralogy, Yale University Geological Story Briefly Told . ............. . . $1.15 Revised Text-Book of Geology. Fifth Edition (Rice) . .1.40 Manual of Geology. Fourth Edition . . ............5.00 THE present edition of the GEOLOGICAL STORY was the last considerable work of the eminent author’s long life. As geology is emphatically an outdoor science, the student is urged to study the quarries, bluffs, and ledges of rock in his vicinity, and all places that illustrate geological operations. The prefatory suggestions are full of practical help, besides enumerating the tools and specimens desirable for this study. The REVISED TEXT-BOOK OF GEOLOGY is here brought down to the present time as regards its facts, but it still expresses the views of its distinguished author. While the general plan and the distinctive features have been pre- served as far as possible, a few important changes have been made relating mainly to zoological and botanical classifications, to the bearings of geology and paleontology upon the theory of evolution, and to metamorphism. The order of the grand divisions of the science, physiographic, structural, dynamical and historical, remains the same as in previous editions. «[| The present edition of the MANUAL OF GEOLOGY was wholly rewritten under the author’s direction. Owing to the extensive recent investigations, new principles, new theories, new facts relating to all departments of the science, and widely diverging opinions on various questions have all made their contributions to this book. The later tracing of formations and mountain-making, the increased number of fossils, the study of canons and other results of erosion, and the development of petrology are all prominently treated here. AMERICAN BOOK COMPANYCHEMISTRIES By F. W. CLARKE, Chief Chemist of the United States Geological Survey, and L. M. DENNIS, Professor of Inorganic and Analytical Chemistry, Cornell University Elementary Chemistry . $i.io Laboratory Manual • $°-5° THESE two books are designed to form a course in chemistry which is sufficient for the needs of secondary schools. The TEXT-BOOK is divided into two parts, devoted respectively to inorganic and organic chemistry. Diagrams and figures are scattered at intervals throughout the text in illustration and explanation of some particular experi- ment or principle. The appendix contains tables of metric measures with English equivalents. Theory and practice, thought and application, are logically kept together, and each generalization is made to follow the evidence upon which it rests. The application of the science to human affairs, its utility in modern life, is also given its proper place. A reasonable number of experiments are in- cluded for the use of teachers by whom an organized laboratory is unobtainable. Nearly all of these experiments are of the simplest character, and can be performed with home-made apparatus. The LABORATORY MANUAL contains 127 experi- ments, among which are a few of a quantitative character. Full consideration has been given to the entrance requirements of the various colleges. The left hand pages contain the experi- ments, while the right hand pages are left blank, to include the notes taken by the student in his work. In order to aid and stimulate the development of the pupil’s powers of observa- tion, questions have been introduced under each experiment. The directions for making and handling the apparatus, and for performing the experiments, are simple and clear, and are illustrated by diagrams accurately drawn to scale. AMERICAN BOOK COMPANY (161162")A BRIEF COURSE IN GENERAL PHYSICS $1.20 By GEORGE A. HOADLEY, A.M., C.E., Professor of Physics, Swarthmore College A COURSE, containing a reasonable amount of work for an academic year, and covering the entrance require- ments of all of the colleges. It is made up of a reliable text, class demonstrations of stated laws, practical questions and problems on the application of these laws, and laboratory experiments to be performed by the students. The text, which is accurate and systematically arranged, presents the essential facts and phenomena of physics clearly and concisely. While no division receives undue prominence, stress is laid on the mechanical principles which underlie the whole, the curve, electrical measurements, induced currents, the dynamo, and commercial applications of electricity. The illustrative experiments and laboratory work, intro- duced at intervals throughout the text, are unusually numerous, and can be performed with comparatively simple apparatus. Additional laboratory work is included in the appendix, to- gether with formulas and tables. HOADLEY’S PRACTICAL MEASUREMENTS IN MAGNETISM AND ELECTRICITY. $0.75 THIS book, which treats of the fundamental measurements in elec- tricity as applied to the requirements of modern life, furnishes a satis- factory introduction to a course in electrical engineering for secondary and manual training schools, as well as for colleges. Nearly ioo experiments are provided, accompanied by suggestive directions. Each experiment is followed by a simple discussion of the principles involved, and, in some cases, by a statement of well-known results. AMERICAN BOOK COMPANY ('59)DESCRIPTIVE CATALOGUE OF HIGH SCHOOL AND COLLEGE TEXT-BOOKS Published Complete and in Sections WE issue a Catalogue of High School and College Text- Books, which we have tried to make as valuable and as useful to teachers as possible. In this catalogue are set forth briefly ana clearly the scope and leading charac- teristics of each of our best text-books. In most cases there are also given testimonials from well-known teachers, which have been selected quite as much for their descriptive qualities as for their value as commendations. ^ For the convenience of teachers this Catalogue is also published in separate sections treating of the various branches of study. These pamphlets are entitled : English, Mathematics, History and Political Science, Science, Modern Languages, Ancient Languages, and Philosophy and Education. In addition we have a single pamphlet devoted to Newest Books in every subject. % Teachers seeking the newest and best books for their classes are invited to send for our Complete High School and College Catalogue, or for such sections as may be of greatest interest. «jy Copies of our price lists, or of special circulars, in which these books are described at greater length than the space limitations of the catalogue permit, will be mailed to any address on request. ^y All correspondence should be addressed to the nearest of the following offices of the company: New York, Cincin- nati, Chicago, Boston, Atlanta, Dallas, San Francisco. AMERICAN BOOK COMPANY (3H312)Cornell University Library arV141 A text-book in general physiology and an gy and a 3 1924 031 168 705 oiin.anx 31924031168705