Online Encyclopedia


Online Encyclopedia
Originally appearing in Volume V23, Page 193 of the 1911 Encyclopedia Britannica.
Spread the word: it!
CO2 in the blood being only an occasional and relatively unimportant factor in the regulations. The phenomenon of "apnoea" or complete cessation of natural breathing which occurs after forced breathing, was attributed mainly to the already mentioned distension effect through the vagus nerves. To go further back still, it was even supposed that the rate and depth of breathing, and the percentage of oxygen in the inspired air, determine the consumption of oxygen and formation of carbon dioxide in the body, just as the air-supply to a fire determines the rate of its combustion. This old belief is still often met with—for instance, in the reasons given for recommending "breathing exercises " as a part of physical training. It is evident that if the breathing did not increase correspondingly with the greatly increased consumption of oxygen and formation of CO2 which occurs, for instance during muscular work, the percentage of oxygen in the air contained in the lung cells or alveoli (alveolar air) would rapidly fall, and the percentage of carbon dioxide increase. The inevitable result would be a very imperfect aeration of the blood. Investigation of the alevolar air has furnished the key to the actual regulation of breathing. Samples of this air can be obtained by making a sudden and deep expiration through a piece of long tube, and at once collecting some of the air contained in the part of this tube nearest the mouth. By this means it has been found that during normal breathing at ordinary atmospheric pressure the percentage of carbon dioxide (about 5.6% on an average for men) is constant for each individual, though different persons vary slightly as regards their normal percentage. The breathing is thus so regulated as to keep the percentage of carbon dioxide constant; and under normal conditions this regulation is surprisingly exact. The ordinary expired air is a mixture of alveolar air and air from the " dead space " in the air passages. The deeper the breathing happens to be, the more alveolar air there will be in the expired air, and the higher, therefore, the percentage of carbon dioxide in it, so that the expired air is not constant in composition, though the alveolar air is. If air containing 2 or 3% of carbon dioxide is breathed, the breathing at once becomes deeper, in such a way as to prevent anything but a very slight rise in the alveolar carbon dioxide percentage. The difference is scarcely appreciable subjectively, except during muscular exertion. The effect of 1% of carbon dioxide in the inspired air is so slight as to be negligible, and there is no foundation for the popular belief that even very small percentages of carbon dioxide are injurious. With 4 or 5% or more of carbon dioxide, however, much panting is produced, and the alveolar carbon dioxide percentage begins to rise appreciably, since compensation is no longer possible. As a consequence, headache and other symptoms are produced. If, on the other hand, the percentage of carbon dioxide in the alveolar air is abnormally reduced by forced breathing, the condition of apnoea is produced and lasts until the percentage again rises to normal, but no longer. Forced breathing with air containing more than about 4% of carbon dioxide causes no apnoea, as the alveolar carbon dioxide does not fall. If oxygen is breathed instead of air there is no appreciable change in the percentage of carbon dioxide in the alveolar air, and no tendency towards apnoea. Want of oxygen is thus not a factor in the regulation of normal breathing. During muscular work the depth and frequency of breathing increase in such a way as to prevent the alveolar carbon dioxide from rising more than very slightly. It is still the carbon dioxide stimulus that regulates the breathing, although with excessive muscular work other accessory factors may come in to some extent. Under increased barometric pressure the percentage of carbon dioxide in the alveolar air no longer remains constant; it diminishes in proportion to the increase of pressure. For instance, at a pressure of 2 atmospheres it is reduced to half, and at 6 atmospheres to a sixth; while at less than normal atmospheric pressure it rises correspondingly unless symptoms of want of oxygen begin to interfere with this rise. These results show that it is not the mere percentage, but the pressure (or " partial pressure ") of carbon dioxide in the alveolar air that regulates breathing. The pressure exercised by the carbon dioxide in the alveolar air is of course proportional to its percentage, multiplied by the total atmospheric pressure. It follows from this law that at a pressure of 6 atmospheres 1% of carbon dioxide in the inspired air would have the same violent effect as 6% at the normal pressure of r atmosphere. To take a concrete practical application, if a diver whose head was just below water were supplied with sufficient air to keep the carbon dioxide percentage in the air of his helmet down to 3% at most, he would be quite comfortable. But if, with the same air supply as measured at surface, he went down to a depth of 170 ft., where the pressure is 6 atmospheres, he would at once experience great distress culminating in loss of consciousness, owing, not to the pressure of the water, which has trifling effects, but to the pressure of carbon dioxide in the air he was breathing. The air supply must be increased in proportion to the increase of pressure if these effects are to be avoided, and ignorance of this has led to the common failure of diving work at considerable depths. The foregoing .facts enable us to understand the regulation of breathing under normal conditions. The pressure of carbon dioxide in the alveolar air evidently determines that of the carbon dioxide in the arterial blood, and the latter in its turn determines the carbon dioxide pressure in the respiratory centre, which is very richly supplied with blood. The centre itself is extremely sensitive to the slightest increase or diminution in carbon dioxide pressure; and thus it is that the alveolar carbon dioxide pressure is so important. That the stimulus of carbon dioxide is from the blood and not through nerves is proved by many experiments. The function of the vagus nerves in regulating the breathing is apparently to, as it were, guide the centre in the expenditure of each separate inspiratory or expiratory effort; for as soon as inspiration or expiration is completed the inspiratory or expiratory effort is cut short by impulse proceeding up the vagus nerve, and much waste of muscular work and risk of injury to the lungs is thereby prevented. Under ordinary conditions the regulation of carbon dioxide pressure in the alveolar air ensures at the same time a normal pressure of oxygen, since absorption of oxygen and giving off of carbon dioxide normally run parallel to one another. If, however, air containing abnormally little oxygen is breathed, the normal relation between oxygen and carbon dioxide in the alveolar air is disturbed. A similar state of affairs is brought about by any considerable diminution of atmospheric pressure. Not only does the partial pressure of oxygen in the inspired air fall, but this fall is proportionally much greater in the alveolar air; and the effects of want of oxygen depend on its partial pressure in the alveolar air. It has been known for long that any great deficiency in the proportion of oxygen in the air breathed increases the depth and frequency of the breathing; but this effect is not apparent until the percentage of oxygen or the barometric pressure is reduced by more than a third, which corresponds to a reduction of more than half in the alveolar oxygen pressure. In contrast with this an increase of a fiftieth in the alveolar carbon dioxide pressure has a marked effect on the breathing. Along with the increased breathing caused by deficiency of oxygen there is more or less blueness of the skin and abnormal effects of various kinds, such as partial loss of sensibility, memory and power of thinking. Long exposure often causes headache, nausea, sleeplessness, &c.—a train of symptoms known to mountaineers as " mountain sickness." That the primary cause of " mountain sickness " is lack of oxygen owing to the low atmospheric pressure there is not the slightest doubt. Lack of oxygen is thus not only an important, but also an abnormal form of stimulus to the respiratory centre, since it is accompanied by quite abnormal symptoms. A further analysis of the special effect of lack of oxygen on the respiratory centre has shown that this effect still depends on the partial pressure of carbon dioxide in the alveolar air. The lack of oxygen appears, in fact, to have simply increased the sensitiveness of the centre to carbon dioxide, 59 that a lower partial pressure of carbon dioxideexcites the centre, and the breathing is correspondingly increased. By prolonged forced breathing so much carbon dioxide is washed out of the body that the subsequent apnoea lasts until the oxygen in the alveolar air is nearly exhausted. The subject of the experiment becomes very blue in the face and is partially stupefied .by want of oxygen before he has any desire to breathe. The probable explanation of these facts is that want of oxygen does not itself excite the centre, but that some substance—very probably lactic acid, which is known to be formed abundantly —is produced abnormally in the body during exposure to want of oxygen and aids the carbon dioxide in exciting the centre. It is known that the blood becomes less alkaline at high altitudes, and that acids in general excite the centre. A person on a high mountain thus gets out of breath much more easily than at sea-level. The extra stimulus to the centre during work still comes from the extra carbon dioxide formed, but has a greater effect than usual on the breathing. If the extra stimulus came directly from want of oxygen the person on the mountain would probably turn blue and lose consciousness on the slightest exertion. By analysing the alveolar air it can be shown that after a time even a height of 5000 to 6000 ft., or a diminution of only a sixth in the barometric pressure, distinctly increases the sensitiveness of the respiratory centre to carbon dioxide, so that there seems to be a. slow accumulation of acid in the blood. The effect also passes off very slowly on returning to normal pressure, although the lack of oxygen is at once removed. The blueness of the skin (" cyanosis ") produced by lack of oxygen is due to the fact that the haemoglobin of the red corpuscles is imperfectly saturated with oxygen. Haemoglobin which is fully saturated with oxygen has a bright red colour, contrasting with the blue colour which it assumes when deprived of oxygen. According to the existing evidence the saturation of the haemoglobin is practically complete under normal conditions in the lungs, or when thoroughly shaken at the body temperature and normal atmospheric pressure with air of the same composition as normal alveolar air. As the partial pressure of the oxygen in this air falls, however, the saturation of the haemoglobin becomes less and less complete, and the arterial blood assumes a more and more blue tinge, which imparts a blue or leaden colour to the skin, accompanied by the symptoms, already referred to, of lack of oxygen. Normal arterial blood in man yields about rq volumes of physiologically available oxygen for each roo volumes of blood. Of these 19 volumes about 181 are loosely combined with the haemoglobin of the red corpuscles, the small remainder being in simple solution in the blood. Venous blood, on the other hand, yields only about 12 volumes. The combination of haemoglobin with oxygen is only stable in the presence of free oxygen at a pressure of about. that in normal alveolar air. As this pressure falls the compound is progressively dissociated. From this it can be readily understood why the blood loses its oxygen in passing through the tissues, which are constantly absorbing free oxygen, and regains it in the lungs. The marked effects produced by abnormal deficiency in the pressure of oxygen in the alveolar air are also readily intelligible; for even although the arterial blood still contains sufficient oxygen to cover the normal difference between the oxygen content of arterial and that of venous blood, yet this oxygen is given off to the tissues less readily—i.e. at a lower pressure, and thus fails to supply their demands completely. It is evident also that in pure air at normal pressure increased ventilation of the lungs does not appreciably increase the supply of oxygen to the blood, whereas in air largely deprived of its oxygen, or at low pressure, the increased alveolar oxygen pressure produced by deep breathing helps greatly in saturating the blood with oxygen, and may thus relieve the symptoms of want of oxygen. Hence it is that the increased sensitiveness of the respiratory centre to carbon dioxide, and consequent increased depth of breathing, at high altitudes compensates to a large extent for deficiency in the oxygen pressure. Addition of carbon dioxide to the inspired air produces exactly the same result. Indeed Professor Angelo Mosso was led by observation of the beneficial effects of carbon dioxide at low atmospheric pressure to attribute mountain sickness to lack of carbon dioxide, a condition which he designated by the word " acapnia. " When impure air is vitiated, not only by deficiency of oxygen, but also by carbon dioxide, the carbon dioxide causes panting, which not only gives warning of any danger, but prevents the alveolar oxygen percentage from falling in the way it would do if the carbon dioxide were absent. In this way the carbon dioxide greatly lessens the danger. To give instances, air progressively and very highly vitiated by respiration is much less likely to cause danger if the carbon dioxide is not artificially absorbed, and not nearly so dangerous as the great diminution of atmospheric pressure (and consequently of oxygen pressure) which occurs in a very high balloon ascent. Indeed the dangers of a very high balloon ascent are notorious, and a number of deaths or very narrow escapes are on record. Just as oxygen forms a dissociable compound with the haemoglobin of the blood, so does carbon dioxide form dissociable compounds. One of these compounds appears to be with haemoglobin itself, and another is sodium bicarbonate, which is far more easily dissociated in the blood than in a simple watery solution, owing to the presence of proteid and possibly other substances which act as weak acids and thus help the dissociation process. The whole of the carbon di-oxide can therefore be removed from the blood by a vacuum pump, just as the whole of the oxygen can. Venous blood contains roughly speaking about 40 volumes of carbon di-oxide per 'co of blood, and arterial blood about 34 volumes. Of this carbon dioxide only about 3 volumes can be in free solution, the rest being loosely combined. The conveyance of carbon dioxide from the blood to the lungs is thus readily intelligible, as well as the fact that any increase or diminution of the pressure of carbon dioxide in the alveolar air will naturally lead to a damming back or increased liberation of carbon dioxide from the blood, and that by forced breathing carbon dioxide can be washed out of the blood to such an extent that a pro-longed cessation of natural breathing (apnoea) follows, since even in the venous blood the partial pressure of carbon di-oxide has become too low to excite the respiratory centre. It will be evident from the foregoing that in order to supply efficiently the respiratory requirements of the tissues not only must the breathing, but also the circulation, be suitably regulated. In hard muscular work the consumption of oxygen and output of carbon dioxide may be increased eight or ten times beyond those of rest. Unless, therefore, the blood supply to the active tissues were correspondingly increased, deficiency of oxygen would at once arise, since the amount of oxygen carried by a given volume of the arterial blood is very limited, as already explained. It is known that the supply of blood to each organ is always increased during its activity. This. increase can, for instance, readily be seen and measured in the case of contracting muscles or secreting glands; and the volume and frequency of the pulse are greatly increased during muscular work. But while it is evident enough that the flow of blood through the body is determined in accordance with the metabolic activities of each tissue, our knowledge is as yet very scanty as to the means by which this determination is brought about. Probably, however, carbon dioxide may be nearly as important a factor in the regulation of the circulation as in that of breathing. Just as the rate of breathing was formerly supposed to determine, and not to be determined by, the fundamental metabolic processes of the body, so the circulation was supposed to be another independent determining factor; and under the influence of these mechanistic conceptions the direction of investigation into the phenomena of respiration and. circulation has been largely diverted to side issues. Since the circulation, no less than the breathing, is concerned in the supply of oxygen to and removal of carbon dioxide from the tissues, it can readily be understood that defective circulation, such as occurs, for instance, in uncom-pensated valvular affections of the heart, may affect the breathing and hinder the normal respiratory exchange. Conversely, also, defects in the aeration or oxygen-carrying power of the blood may be compensated for by increase in the circulation. For instance, in the very common condition known as anaemia, where the percentage of haemoglobin, and consequently the oxygen-carrying power of the blood, is often reduced to a third or less, the respiratory disturbances may be so slight that the patient is going about his or her ordinary work. A miner suffering from the now well-known " worm-disease," or ankylostomiasis (q.v.), may be working under-ground, or a housemaid suffering from chlorosis may be doing her work, with only a third of the normal oxygen-carrying power of the blood. There seems to be no doubt that in such cases an increased rate of blood circulation compensates for the diminished oxygen-carrying power of the blood. It is well known that at high altitudes a gradual process of adaptation to the low pressure occurs, and the shortness of breath and other symptoms experienced for the first few days gradually become less and less. This adaptation is .partly, at least, due to a marked increase in the percentage of haemoglobin in the blood, though probably circulatory and perhaps other compensatory changes are also involved. In connexion with respiration the action of certain poisons is of great interest. One of these, carbon monoxide, is of very common occurrence, and causes numerous cases of poisoning. Like oxygen, it has the property of combining with the haemoglobin of the blood, but its affinity for haemoglobin is far more strong than that of oxygen. In presence of air containing as little as *05% of carbon monoxide, the haemoglobin will become about equally shared between oxygen and carbon monoxide, so that, since air contains 2o.9% of oxygen, the affinity of carbon monoxide for haemoglobin may be regarded as about 400 times greater than that of oxygen. The blood of a person breathing even a small percentage of carbon monoxide may thus become gradually saturated to a dangerous extent, since the haemoglobin engaged by the carbon monoxide is for the time useless as an oxygen-carrier. Air containing more than about o.r % of carbon monoxide is thus more or less dangerous if breathed for long; but the blood completely recovers in the course of a few hours if pure air is again breathed. The poisonous action of carbon monoxide can be abolished by placing the animal exposed to it in oxygen at an excess pressure of about an atmosphere. The reason for this is that, in consequence of the increased partial pressure of the oxygen, the amount of this gas in free solution in the blood is greatly increased in accordance with Dalton's law, and becomes sufficient to supply the tissues with oxygen quite independently of the haemoglobin. Even at ordinary atmospheric pressure the extra oxygen dissolved in the blood when pure oxygen is breathed is of considerable importance. Carbon-monoxide poisoning is the chief cause of death in colliery explosions and fires, and the sole cause in poisoning by lighting gas and fuel gas of various kinds. Its presence in dangerous proportions may be readily detected with the help of a small bird, mouse or other small warm-blooded animal. In such animals the respiratory exchange is so rapid that symptoms of carbon-monoxide poisoning are shown far more quickly than in man. The small animal can thus be employed in mines, &c., to indicate danger from carbon monoxide. A lamp is useless for this purpose. There are various other poisons, such as nitrites, chlorates, dinitrobenzol, &c., which act by disabling the haemoglobin, and so cutting off the oxygen supply to the tissues. Between the air in the air-cells of the lungs and the blood of the lung capillaries there intervenes nothing but a layer of very thin, flattened cells, and until recently it was very generally believed that it was by diffusion alone that oxygen passes inwards and carbonic acid outwards through this layer. Similar simple physical explanations of processes of secretion and absorption through living cells have, however, turned out to be incorrect in the case of other organs. It is known, moreover, that in the case of the swimming-bladder of fishes oxygen is secreted intq the interior against enormous pressure. Thus, in the case of a fish caught at a depth of 4500 ft., the partial pressure of the oxygen present in the swimming bladder at this depth was 127 atmospheres, whereas the partial pressure of oxygen in sea-water is only about 0.2 atmosphere. Diffusion can therefore have nothing to do with the passage of gas inwards, which is known to be under the control of the nervous system. The cells lining the interior of the swimming bladder are developed from the same part of the alimentary tract as those lining the air-cells of the lungs, so that it seems not unlikely that the lungs should possess the power of actively secreting or excreting gases. The question whether such a power exists, and is normally exercised, has been investigated by more than one method; and although it is not possible to go into the details of the experiments, there can be no doubt that the balance of the evidence at present available is in favour of the view that diffusion alone is incapable of explaining either the absorption of oxygen or the excretion of carbon dioxide through the lining cells of the lungs. The partial pressure of oxygen appears to be always higher, and of carbon dioxide often lower, in the blood leaving the lungs than in the air of the air-cells; and this result is inconsistent with the diffusion theory. As to the causes of the passage of oxygen and carbonic acid through the walls of the capillaries of the general circulation, we are at present in the dark. Possibly diffusion may explain this process. II. Although we cannot trace the exact changes which occur when oxygen passes into living cells, yet it is possible to obtain a clear general view of the origin and destiny of the material concerned in the process, and of the physiological conditions which determine it. The oxidizable material within the body consists, practically speaking, of proteids (albumen-like substances, with which the collagen of connective tissue may be included), fats and carbohydrates (sugars and glycogen). All of these substances contain carbon, hydrogen and oxygen in known, though different, proportions, and the former also contains a known amount of nitrogen and a little sulphur. Nitrogen is constantly leaving the body as urea and other substances in the urine and faeces; and a small but easily measurable proportion of carbon passes off in the same manner. The rest of the carbon passes out as carbon dioxide in respiration. Now carbohydrates and fats are oxidized completely in the body to carbon dioxide and water. This follows from the fact that, practically speaking, no other products into which they might have been converted leave the body except carbon dioxide and water. Moreover, a given weight of carbohydrate requires for its oxidation a definite weight of oxygen, and produces a definite weight of carbon dioxide. There is thus a definite relation between the weight of oxygen used up and the weight of carbon dioxide formed in this oxidation. The same is true for the oxidation of fat and of proteid, allowing in the latter case for the fact that the nitrogen, together with part of the carbon and hydrogen, passes out as urea, &c., in an incompletely oxidized form. From all this it follows that if we measure over a given period (r) the discharge of nitrogen from the body, (2) the intake of oxygen and (3) the output of carbonic acid, we can easily calculate exactly what the ultimate destiny of the oxygen has been, and at the ultimate expense of what material the carbonic acid has been formed. What the intermediate stages may have been we cannot say, but this in no way affects the validity of the calculation. If, during the period of measurement, food is taken, the basis of the calculation is still substantially the same, as the oxidizable material in food consists of practically nothing else except proteids, carbohydrates and fats. Liberation of Energy.—From experiments made outside the body, we know that in the oxidation of a given weight of proteid, carbohydrate or fat, a definite amount of energy is liberated. In the article on DIETETICS it is shown that precisely the same liberation of energy occurs in the living body, due allowance being made for the fact that the oxidation of proteid is not quite complete. The following table shows the respiratory quotients (the respiratory quotient being the ratio betweenthe volume of carbon dioxide formed and that of oxygen used up) and energy expressed in units of heat (calories) liberated per gramme of carbon dioxide produced and oxygen consumed in the living body during the oxidation of proteid, fat and a typical carbohydrate: Substance oxidized. Respiratory Calories per Calories per quotient. gramme of gramme of CO2 pro- oxygen duced. consumed. P •78 o 2.78 3.00 Fat roteid r•o 2.59 3.56 Cane-sugar . In the oxidation of non-living substances the rate varies, within wide limits, according to that at which oxygen is supplied. Thus a fire burns the faster the more air is supplied, and the higher the percentage of oxygen in the air. It was for long believed that in the living body also the rate of oxidation must vary according to the oxygen supply. It has been found, however, that this is not the case. Provided that a certain minimum of oxygen is present in the air breathed, or in the blood supplied to the tissues, it is, practically speaking, indifferent whether the oxygen supply be increased or diminished: only a certain amount is consumed. It might be supposed that the reason for this is that the available oxidizable material in the body is limited, and that if the food supply were increased there would be a corresponding increase in the rate of oxidation. This hypothesis is apparently supported by the fact that, when an increased supply of proteid is given as food, the amount of nitrogen discharged in the urine is almost exactly correspondingly increased, so that evidently the oxidation of proteid increases correspondingly with the supply. Similarly, when carbohydrate food is given, the alteration in the respiratory quotient shows that more carbohydrate than before is being oxidized. Closer investigation in recent times has, however, brought out the very striking fact that, if oxidation be measured in terms of energy liberated by it in the body, it makes but little difference, other things being equal, whether the animal is fasting or not. If more proteid or carbohydrate is oxidized at one time, correspondingly less fat is oxidized, but the total energy liberated as heat, &c., in the body is about the same, unless the diet is very excessive, when there is a slight increase of oxidation. Even after many days of starvation, the rate of oxidation per unit of body weight has been found to remain sensibly the same in man. When more food is taken than is required, the excess is stored up,'chiefly in the form of fat, into which carbohydrate and possibly also proteid are readily converted in the body. When less food is taken than is needed, the stock of fat is drawn upon, and supplies by far the greater proportion of the energy requirements of the body. During the performance of muscular work oxidation is greatly increased, and may amount to ten times the normal or more. Even the slight exertion of easy walking increases oxidation to three times. When the energy represented by the external work done in muscular exertion is compared with the extra energy liberated by oxidation in the body, it is found, as would be expected, that the latter value largely exceeds the former. In other words, much of the energy liberated is wasted as heat. Nevertheless the muscles are capable of working with less waste than any steam or gas engine. In the work of climbing, for instance, it has been found in the case of man that 35 % of the energy liberated is represented in the work done in raising the body. Muscular work, if at all excessive, leads to fatigue, and consequent rest. On the other hand, unnatural abstinence from muscular activity leads to restlessness and consequen* muscular work. Hence on an average of the twenty-four hours the expenditure of energy by different individuals, with different modes of life, does not as a rule differ greatly. The rate of oxidation per unit of body weight varies consider-ably according to size and age. If we compare different warm-blooded animals, we find that the rate of oxidation is relatively to their weight far higher in the smaller ones. In a mouse or small bird, for instance, the rate is about twenty times as great as in a man. The difference is in part due to the fact that the smaller an animal is the greater is its surface relatively to its mass, and consequently the more heat does it require to keep up its temperature. The smaller animal must therefore produce more heat. Even in cold-blooded animals, however, oxidation appears to be more rapid the smaller the animal. In the case of man, oxidation is relatively more than twice as rapid in children than in adults, and the difference is greater than would be accounted for by the difference in the ratio of surface to mass. Allowing for differences in size, oxidation is about equally rapid in men and women. It was for long believed that the special function of respiratory oxidation was (I) the production of heat, and (2) the destruction of the supposed " waste products." Further investigation has, however, tended to show more and more clearly that in reality respiratory oxidation is an essential and intimate accompaniment of all vital activity. To take one example, secretion and absorption, which were formerly explained as simple processes of filtration and diffusion, are now known to be! accompanied, and necessarily so, by respiratory oxidation in the tissues concerned. The respiratory oxidation of an animal is thus a very direct index of the activity of its vital processes as a whole. Looking at what is known with regard to respiratory oxidation; we see that what is most striking and most characteristic in it is its tendency to persist—to remain on the whole at about a normal level for each animal, or each stage of development of an animal. The significance of this cannot be over-estimated. It indicates clearly that just as an organism differentiates itself from any non-living material system by the manner in which it actually asserts and maintains its specific anatomical structure, so does it differentiate itself from any mere mechanism by the manner in which it asserts and maintains its specific physiological activities. (3) MOVEMENTS OF RESPIRATION Normal Respiration.—If the naked body of a person asleep or in perfect inactivity be carefully watched, it will be found that the anterior and lateral walls of the chest move rhythmically up and down, while air passes into and out of the nostrils (and mouth also if this be open) in correspondence with the movement. If we look more closely we shall find that with every uprising of the chest walls the membranous intercostal portions sink slightly as if sucked in, while at the same time the flexible walls of the abdomen bulge as if protruded by some internal force. If respiration be in the slightest degree hurried, these motions become so marked as to escape the attention of no one. The elevation of the chest walls is called inspiration, their depression expiration. Inspiration is slightly shorter than expiration, and usually there is a slight pause or momentary inaction of the chest between expiration and the following inspiration. Apparatuses for measuring the excursion of a given point of the chest wall during respiration are called thoracometers or stethometers. Apparatuses for recording the movements of the chest are called stethographs or pneumographs. Frequency of Respiration.—The frequency of respirationduring perfect rest of the body is '16 to 24 per minute, the pulse rate being usually four times the rate of respiration; but the respiratory rhythm varies in various conditions of life. The following are the means of many observations made by Lambert Adolphe Quetelet (1796-1874): at the age of one year the number of respirations is 44 per minute; at 5 years, 26; from 15 to 20 years, 2o; from 25 to 30, 16; from 30 to 50, 18.1. Muscular exertion always increases the frequency of respiration. The higher the temperature of the environment the more frequent is the respiration. Paul Bert (1833-1886) has shown that with higher atmospheric pressures than the normal the frequency of respiration is diminished while the depth of each inspiration is increased. The frequency of respiration diminishes until dinner-time, reaches its maximum within an hour of feeding, and thereafter falls again; if dinner is omitted, no rise of frequency occurs. The respiratory act can be interrupted at any stage, reversed, quickened, slowed and variously modified at will, so long as respiration is not stopped entirely for more than a short space of time; beyond this limit the will is incapable of suppressing respiration. Depth of Respiration.—The depth of respiration is measured by the quantity of air inspired or expired in the act; but the deepest expiration possible does not suffice to expel all the air the lungs contain. The following measurements have been ascertained, and are here classified according to the convenient terminology proposed by John Hutchinson (181r-1861). (I) Residual air, the volume of air remaining in the chest after the most complete expiratory effort, ranges from 100 to 130 CO. in. (2) Reserve or supplemental air, the volume of air which can be expelled from the chest after an ordinary quiet expiration, measures about loo cub. in. (3) Tidal air, the volume of air taken in and given out at each ordinary respiration may be stated at about 20 cub. in. (4) Complemental air, the volume of air that can be forcibly inspired over and above what is taken in at a normal inspiration, ranges from about Too to 130 cub. in. By vital capacity, which once had an exaggerated importance attached to it, is meant the quantity of air which can be expelled from the lungs by the deepest possible expiration after the deepest possible inspiration; it obviously includes the complemental, tidal and reserve "airs, and measures about 230 cub, in. in the Englishman of average height, i.e. 5 ft. 8 in. (Hutchinson). It varies according to the height, body weight, age,. sex, position of the body and condition as to health of the subject of observation. Vital capacity is estimated by means of a spirometer, a graduated gasometer into which air may be blown from the lungs. The residual air, which for obvious reasons cannot be actually measured, may be estimated in the following way (Emil Harless, 182o-1862; Louis Grehant, b. 1838). At the end of ordinary expiration, apply the mouth to a mouthpiece communicating with a vessel filled with pure hydrogen, and breathe into and out of this vessel half a dozen times—until, in fact, there is reason to suppose that the air in the lungs at the time of the experiment has become evenly mixed with hydrogen. Then ascertain by analysis the proportion of hydrogen to expired air in the vessel and estimate the amount of the air which the lungs contained by the following formula: : V+v= p : roo ; v(roo-p) V= p where V=volume of air in the lungs at the time of experiment, v= volume of the vessel containing hydrogen, p = proportion of air to hydrogen in the vessel at the end of the experiment. V, then, is the volume of air in the lungs after an ordinary expiration; that is, it includes the residual and the reserve air; if we subtract from this the amount of reserve air ascertained by direct measurement, we obtain the 100-130 cub. in. which Hutchinson arrived at by a study of the dead body. Volume of Respiration.—It is clear that the ventilation of the lungs in ordinary breathing does not merely depend on the quantity of air inspired at each breath, but also on the number of inspirations in a given time. If these two values be multiplied together we get what might be called the volume of respiration (Athmungsgrosse, Isidore Rosenthal, b. 1836), in contradistinction to depth of respiration and frequency of respiration. Various instruments have been devised to measure the volume of respiration, all more or less faulty for the reason that they compel respiration wider somewhat abnormal conditions (Rosenthal, Gad, Peter Ludwig, Panum (182o-1885), Ewald Hering (b. 1834). From the data obtained we may conclude that the respiratory volume per minute in man is about 366 cub. in. (6000 cub. centim.). In connexion with this subject it may be stated that, after a single ordinary inspiration of hydrogen gas, 6–10 respirations of ordinary air must occur before the expired air ceases to contain some trace of hydrogen. Types of Respiration.—The visible characters of respiration in man vary considerably according to age and sex. In men, while there is a moderate degree of upheaval of the chest, there is a considerable although not preponderating degree of excursion of the abdominal walls. In women the chest movements are decidedly most marked, the excursion of the abdominal walls being comparatively small. Hence we may distinguish two types of respiration, the costal and the abdominal, according to the preponderance of movement of one or the other part of the body wall. In forced respiration the type is costal in both sexes, and so it is also in sleep. The cause of this difference between men and women has been variously ascribed (a) to constriction of the chest by corsets in women, (b) to a natural adaptation to the needs of child-bearing in women, and (c) to the greater relative flexibility of the ribs in women permitting a wider displacement under the action of the inspiratory muscles. Certain Concomitants of Normal Respiration.—If the ear be placed against the chest wall during ordinary respiration we can hear with every inspiration a sighing or rustling sound, called " vesicular," which is probably caused by the expansion of the air vesicles; and with every expiration a sound of a much softer sighing character. In children the inspiratory rustle is sharper and more pronounced than in adults. If a stethoscope be placed over the trachea, bronchi or larynx, so that the sounds generated there may be separately communicated to the ear, there is heard a harsh to-and-fro sound during inspiration and expiration which has received the name of " bronchial." In healthy breathing the mouth should be closed and the ingoing current should all pass through the nose. When this happens the nostrils become slightly expanded with each inspiration, probably by the action of the M. dilatatores naris. In some people this movement is hardly perceptible unless breathing be heavy or laboured. As the air passes at the back of the throat behind the soft palate it causes the velum to wave very gently in the current; this is a purely passive movement. If we look at the glottis or opening into the larynx during respiration, as we may readily do with the help of a small mirror held at the back of the throat, we may notice that the glottis is wide open during inspiration and that it becomes narrower by the approximation of the vocal chords during expiration. This alteration is produced by the action of the laryngeal muscles. Like the movements of the nostril, those of the larynx are almost imperceptible in some people during ordinary breathing, but are very well marked in all during forced respiration. The Mechanics of Respiration.—The thorax is practically a closed box entirely filled by the lungs, heart and other structures contained within it. If we were to freeze a dead body until all its tissues were rigid, and then were to remove a portion of the chest wall, we should observe that every corner of the thorax is accurately filled by some portion or other of its contents. If we were to perform the same operation of removing a part of the chest wall in a body not first frozen we should find, on the other hand, that the contents of the
End of Article: CO2

Additional information and Comments

There are no comments yet for this article.
» Add information or comments to this article.
Please link directly to this article:
Highlight the code below, right click and select "copy." Paste it into a website, email, or other HTML document.