j^« |JK gata; I. to VIII., first to eighth pair of cranial nerves. pons very large white nerve-cords are seen passing upward and forward to the cerebrum from the organs lying below ("cerebral peduncles" or crura cerebri'). Around each of these cords winds a flat band, the optic tract; these tracts STKUCTUKE OF THE SPINAL COBD AND BEAIN. 57 come together in front to form tlie optic commissure, from which the two optic nerves arise. On each side of the deep longitudinal fissure stretches the olfactory tract, end- ing in its bulb. The intercranial part of this nerve is really a projecting portion of the brain. (For the other bodies on this surface, see Fig. 20.) Upper Surface of the Cerebrum. — On top, the cerebral hemispheres present the appearance of gray nervous matter arranged in folds, called " convolutions " or gyri. These are separated by " fissures " or sulci, of varying depth. A considerable difference exists in the develop- ment of the different convolutions, and in the strength with which the different fissures are marked. The details of this aspect of the brain vary in each individual, and even in the two hemispheres of the same brain. Some sulci and their corresponding gyri appear with a marked regularity in the more fundamental stages of the foetal brain. They have been divided therefore into primary, secondary, and even tertiary, classes. Lobes of the Cerebrum. — By means of the primary sulci the hemispheres of the brain have been " mapped out " into five territories, called Lobes. These are the (1) Frontal, (2) Parietal, (3) Temporal or Sphenoidal, or Tempero-sphe- noidal, (4) Occipital, and (5) Central or Insula, or Island of Ren. The frontal lobe is divided from the parietal, on its upper and lateral surface, by the Fissure of Rolando (sulcus centralis') ; and on its lower surface from the tem- poral lobe by the horizontal branch of the Fissure of Sylvius. The parietal lobe is divided from the temporal, for the greater part, by the Fissure of Sylvius, and from the occipital lobe, on its median sui-face completely, and on its upper surface only very incompletely, by the parieto- occipital fissure. The temporal lobe is separated from the frontal and parietal as already described; but the boun- dary between it and the occipital lobe is very ni-defined. 58 PHYSIOLOGICAL PSYCHOLOGY. Fia. 22. sphenoidal convolutions- 10 sunerim- ii I'w'^Ti ■ ^ ,o ■■ f"? ^' '■ ^' "nfenor temporo- ., P, V, a, four arectent convolutions.' ' ' ^^' '°^'™'' °°°'P"*' convolutions; STRTJCTUKE OF THE SPINAL COKD AND BEAIN. 59 The Island of Meil lies concealed beneath the frontal, pari- etal and temporal lobes ; it consists of a few short convolu- tions which may be disclosed by drawing aside the margin of the Fissure of Sylvius. In describing the other lobes the boundaries of the occipital lobe have been sufficiently defined. Snlei and Gyri of the Upper Surface. — The frontal, tem- poral, and occipital lobes all have three principal con- volutions arranged in nearly parallel tiers (superior, middle, and inferior). On each side of the Fissure of Rolando are the two central convolutions, sometimes called "ascending frontal" and "ascending parietal." Among the sulci, the Fissure of Sylvius is much the most important. It exists in the foetal brain at the third month, and is made by folding the entire hemisphere into an arch. The Fissure of Rolando is also always present in the human brain, and is rarely or never bridged over by a secondary gyrus. It appears in the foetus as eariy as the end of the fifth month. (For further details, see Fig. 22.) Median Aspect of the Cerebrum. — On this aspect of each hemisphere appears an important convolution which arches around the corpus callosum, and is separated from the first Fig. 23. — Convolutions of the Inner and Tentorial Surfaces of the Left Hemisphere, i, t, 1, calloso-marginal fissure; Z, Z, calcarine fissure; m, m, hippocampal fissure; n,n, collateral fissure; PO, parieto-oooipital fissure; 17, 17, marginal convolution; 18, 18, gyrus fomicatus; 18', quadrilateral lobule; 19, hippocampal gyrus; 19', its recurved end ; 25, occipital lobule ; 9, 9, inferior temporo-spbenoidal convolution. 60 PHYSIOLOGICAL PSYCHOLOGY. frontal convolution by a deep and constant fissure (the sulcus calloso-marginalis') ; it is called from its shape, gyrus fornicatus. Its back end curves downward and forward under the name of gyrus hippocampi, to the inner tip of the temporal lobe. The passage without break of one of these convolutions into the other is considered by Ecker to be a most important difference between the hemispheres of the brain of man and those of the ape. (For further details, see Fig. 23.) Spaces and Bodies within the Cerebral Mass. — By cutting off successive slices of the cerebral hemispheres their general internal structure may be exposed. Beneath the corpus callosum, and roofed over by it, is a space in the interior of each hemisphere. These cavities are called lateral ventricles. They are separated by a thin trans- parent wall (the septum lucidum'), and are moistened by a serous fluid. The roof of another cavity, the third ventri- cle, is formed by an expanded fold of the pia mater (velum interpositum'). Each lateral ventricle has a central space and three curved prolongations, or cornua, (the anterior, the posterioi;, and the descending,) which extend into the cerebral mass. On the floor of each lateral ventricle the exposed por- tions of the great "basal ganglia" of the cerebrum are visible. Two large pear-shaped bodies of gray color are here seen, with their broad extremities directed forward into the anterior cornua of the ventricle, and their narrow ends outward and backward. They are called, from their striped appearance when cut open, "striate bodies" or corpora striata. Between the diverging portions of these bodies are certain ovoid masses called "optic thalami." Each thalamus rests upon one of the crura cerebri ; on its outer and back part are two small elevations (corpora genioulata, internum and externum). Along the floor of the descending cornu of the ventricle, the inner surface of the STBUCTUEB OP THE SPINAL COED AND BKAIN. 61 gyrus fornicatus, doubled upon itself like a horn, appears as a white eminence (^hippocampus major or "horn of Fio. 24. — Baaal Oanglia of the Cerebrum seen from above. (Henle.) Ccl, genu of the corpus caJloBum; Ce, corpus striatum ; Vsl, ventricle of the septum lucidum; Cf, column of the fornix; St, stria terminalis; Tbo, optic thalamus; and Ts, its anterior tubercle ; Com, middle commissure between the thalami and over the third ventricle; Fv, pulvinar; Cn, conarium or pineal gland ; Cop, corpus quadrigeminum. Ammon "). An arch-shaped band of nerve-fibres, called the " fornix," is situated beneath the corpus callosum. It consists of two lateral halves which, in front, form two pil- 62 PHYSIOLOGICAL PSYCHOLOGY. lars that descend to the base of the cerebrum and become the corpora albieantia. Behind and between the optic thalami, and resting on the back sm-face of the crura Fig. 25. — A Deeper Dissection of the Lateral Ventricle, and of the Velum Interposi- tum. a, under surface of corpus callosum, turned back; 6, &, posterior pillars of the fornix, turned back; c, c, anterior pillars of the fornix; d, velum interpositum and veins of G-alen; e, fifth ventricle; /,/, corpus striatum; ^, ^, taenia semicircularis; A, A, optic thalamus; ^, choroid plexus; £, taenia hippocampi ; m, hippocampus major in descending coruu; 7t, bippocampuB minor; o, eminentia collateralis. cerebri, are four eminences in two pairs, called corpora quadrigemina ; the front pair are the nates; the back pair, testes. The structure of some of the bodies already referred to requires a yet more detailed description, in order to even an elementary knowledge of the superior cerebro-spinal mechanism. STEUCTUEB OF THE SPINAL COED AND BEAIN. 63 The Cmra Cerebri. — These strong peduncles of the brain ascend from the pons to the optic thalami and the striate bodies. The fibres which constitute them are arranged in two groups, separated by the gray matter of the substantia nigra. The front j portion is called crusta. Of its /'^^'""^ fibres an important part is contin- /^ f^' \ uous -with the longitudinal fibres *x^ * i \. from the pyramids of the medulla ; f \v^ j >^ A it receives other fibres from the \^'' J/\ J substantia nigra. Many of the fig. 26. -tecUon through fibres of the crusta run to the 'Z,^£roi^t^Zftl\ nuclei of the striate bodies and ^?''te°'c™B'"LUri^- "tl^ terminate there; but some radiate "^ntum of the cru^ cerebri. upward through the internal capsule directly to the cere- bral cortex. The back portion of the crus is called tegmentum. Some of its fibres come from the anterior column of the cord and radiate upward to the optic thalami. These fibres are diffused. Others are collected into more well-defined tracts, — especially a tract coming from the superior pe- duncle of the cerebellum, and passing forward over the anterior end of the fourth ventricle (see Fig. 19). The formatio reticularis is continued into the tegmen- tum, and some fibres appear to arise in its cells. The Striate Bodies. — Each corf us striatum consists of two masses, the upper one of which (nucleus caudatus') projects into the lateral ventricle ; the lower one (nucleus lenticularis') is embedded in the white substance of the hemisphere, and constitutes the principal part of the body. These two are separated by an important layer of white matter called the " internal capsule." The details of the structure of the striate bodies are insufficiently made out. The nucleus caudatus receives from the capsule, on the side turned toward it, several 64 PHYSIOLOGICAL PSYCHOLOGY. bundles of fibres. All parts of the nucleus lenticularis are pervaded with bundles of white fibres. Some bundles pass into it from the adjacent parts of the capsule ; some connect it with the caudate nucleus; some radiate from it toward the cerebral cortex. The gray matter of this organ has free nerve-nuclei distributed through it. The striate bodies have apparently a special connection with the frontal and parietal lobes, but also with some con- volutions of the temporal lobe and the Island of Reil. The Optic Thalami. — The gray matter of this organ is subdivided into several parts, Cap- »"''• t int. Clatiatrtini Thalam. opt- eorp: eallasi Hinlerer Thail del jfucteu» eaiidattcfi Comu posteriua Fig. 27. Fig. 28. These and the following two Figures show the arrangement of the white and gray suhBtance in the interior of the cerebrum. (All four are from Q-egenbaur.) Fig. 27. — Horizontal Section through the Right Hemisphere. Fig. 28. — Frontal Section through the Cerebrum in front of the Fornix. Posterior surface of the section displayed. SO that two or more nuclei are distinguished by different authorities. This subdivision is only partial, however; the organ is therefore, perhaps, best described as a mass of gray nervous substance, with multipolar and fusiform cells, and everywhere traversed by nerve-fibres. Its external and STEUCTaKE OP THE SPINAL CORD AND BRAIN. 66 under surfaces are not free, but are united with other parts of the brain. On the outer surface of the optic thalami is the white matter of the internal capsule, composed of fibres diverg- ing from the crusta into the hemispheres. With these fibres mingle those which radiate from the interior of the organ itself. According to a recent authority, the thala- mus is the primary centre of the optic nerve. Fio. 29. Pis. 30. Vie. 29. — Frontal Section througb the Bight Hemisphere of the Cerebrum in front of the Commissura Mollis. Posterior surface of section displayed. Fig. 30. — Frontal Section through the Cerebrum back of the Commissura KoUis. Front surface of section displayed. The Corpora Qnadrigemina. — In the interior of the front pair the most characteristic portion of this organ is found; it is a layer of fine nerve-fibres running longitu- dinally, between which are small, scattered nerve-cells. In the external strata of these bodies multipolar cells are abundant ; in the interior, at the sides of the Sylvian aqueduct, is a collection of gray matter which is continu- ous with the lining of the third ventricle. The third and fourth nerves originate in the nervous substance which lies along this " Aqueduct " (see Fig. 20). Layers of the Cerebral Cortex. — The general arrangement of gray nervous substance upon the surface, and of white 66 PHYSIOLOGICAL PSYCHOLOGY. matter within, is adhered to in all parts of the cerebral cortex. But the form and distribution of the nerve-cells Fio, 31. — Section through the Cerebval Cortex of Man, prepared with Osmic Acid. *Vi- (Sohwalbe) . /, principal external, and II, internal, layer ; x, layer lying as a limit between the two; m, medullary Buhstance sending out bundles of nerve-fibres into //; I, layer poor in cells, but with an external plexus of nerve-fibres (la) ; 2, layer of small, and 3, of large, pyramidal cells; 4, inner layer of small nerve-cells. are different in different regions and in different layers of the same region. As a rule the cortex has five layers or lamince. The entire thickness is, in the adult, from -^ to ^ inch. The first layer shows delicate nerve-fibrils run- STRUCTURE OF THE SPINAL CORD AND BRAIN. 67 ning parallel to the surface and interlacing with a few small branching nerve-cells. The second and third layers contain a large number of pyramidal or spindle-shaped cells. In the second layer the cells are about ^ruv i'lch in diameter, and are closely pressed together. In the third are fewer cells, but they increase in size to perhaps xwir or ^-^ inch, and have their long axes perpendicular to the surface. The fourth layer contains large numbers of small, globular, and branching cells ; the fifth, spindle- shaped bodies with long tapering processes, and smaller irregular cells, very compactly accumulated. The number of nerve-cells in the cortex is enormous. In a bit of its substance, 1 millimeter square and ^ milli- meter thick, 100 to 120 have, on an average, been counted. Modifications of the arrangement just described are found in certain regions of the cerebral cortex. In the occipital lobe, for example, the number of layers is increased, by intercalating granule layers, to seven or eight. In cer- tain layers, called "motor," large cells (named by Betz " giant-cells " ) resembling those in the anterior horns of the spinal cord, are found. Conjecture and research are at work with the question, whether certain of these layers, and the cells they contain, are not more distinctively sen- sory, and certain others more distinctively motor. This subject is very important in its relations to our entire conception of the nature and fimctions of the cerebral mechanism. But as yet histology, even when aided by physiological experiment, has determined nothing definite. Systems of Cerebral Nerve-fibres. — The nerve-fibres of the white substance of the brain are of three classes, according to the destination of the fascicles into which the fibres are gathered. There are the (1) downgoing or peduncular, (2) the commissural, and (3) the arcuate (^fibrae propriae). The peduncular system of nerve-fibres connects the cere- brum with the lower parts of the encephalon. This system 68 PHYSIOLOGICAL PSYCHOLOGy. is called the corona radiata; it is the "blossoming out" of the nerve-fibres on their way between the hemispheres and the lower ganglia. Looked at from above, this system represents the contracting of the downgoing nerve-tracts as they are narrowed into the internal capsule and then taken on to the crura cerebri. (See Fig. 27.) A con- siderable portion of this system, however, terminates in the optic thalami and the striate bodies. The principal tract of the commissural system of cerebral fibres is formed in the corpus callosum. This system con- nects the two hemispheres of the brain. That the fibres of the corpus callosum are not wholly commissural, follows from the fact that, since this commissure lies above the plane of the corona radiata, the peduncular system, on its way to the hemispheres, here intersects with the commissu- ral. A smaller commissure (the anterior^ passes below the lenticular nuclei of the striate bodies and connects the con- volutions around the Sylvian fissure. The system of arcuate fibres of the cerebrum connects the gray matter of more or less distant convolutions of the same hemisphere. These fibres may often be described as a " garland-like interweaving " of two convolutions around the sulcus between them. In certain localities, where the fascicles into which the fibres are gathered are strongly marked, they have received special names ; such are the fasciculus uncinatus, which crosses the bottom of the Sylvian fissure, the fillet of the gyrus fomicatus, extending longitudinally in that convolution, etc. The function of the arcuate fibres is plainly that of joining into a diversified unity the different portions of each cerebral hemisphere. STKUCTUEE OF THE SPINAL COBD AND BKAIN. PATHS OF NERVOUS IMPULSES IN THE SPINAL COED AND BRAIN. The foregoing brief description of the cerebro-spinal nervous system shows that it is a mechanism constructed so as to afford " Tracts " or " Paths," to a greater or less degree distinct, for the transmission of nervous impulses. It is, however, only to a very limited degree that histology alone, or even when helped by embryology and pathology, can make out precisely where these paths lie. (Several of those belonging _to the spinal cord, that are more distinctly traceable by the histological method — for example, the pyramidal tract, both crossed and uncrossed, the direct lateral cerebellar tract, the paths of the anterior and of the posterior nerve-roots, etc. — have already been described.) In the brain also it is thought by eminent authorities that certain chains of nervous organs, in which the gray masses are successively connected by nerve-cords between, can be pointed out. With this in view, three collections of nervous matter (the locus niger, and the two nuclei of the striate bodies), with the bundles of nerve-fibres which bind them together, have been called "ganglia of the crusta" by Meynert. Another chain, consisting of the tegmentum, the red nucleus (a collection of large pigmented cells near the Sylvian Aqueduct), the corpora geniculata, and the optic thalami, has been proposed by the same authority. It is only, however, when the aid of physiology (patho- logical and experimental) is summoned, that much prog- ress toward certainty can be made in determining paths of nervous impulse within the cerebro-spinal axis. Even with this aid, there is still room for conjecture and uncer- tainty. Anticipating additional evidence which will sub- sequently be more fully described, we now indicate the probabilities concerning certain of these paths. Paths in the Roots of the Spinal Cord. — The honor of the 70 PHYSIOLOGICAL PSYCHOLOGY. truly " epoch-making " discovery, that the anterior roots of the spinal cord are motor and the posterior, sensory, must be divided between Sir Charles Bell and Magendie. This discovery may be said to have opened the door to modern experimental physiology. The demonstration of the fact is performed by dividing these roots, respectively, and then observing the physiological results. When a posterior root is divided, all the structures supplied by the divided nerve lose their sensibility; while the muscles supplied by the corresponding anterior root continue to be thrown into action by the will and by reflex stimulation. In this case also, stimulation of the central end of the divided root produces sensory effects ; but stimulation of the peripheral end produces no motion. When an anterior root is divided, on the contrary, the muscles supplied by the nerves of this root cannot be made to act by will ; but no sensory paraly- sis is produced. Moreover, stimulation of the peripheral end of the nerve will now throw the muscles into contrac- tion ; but stimulation of the central end will produce no effects. Thus far, then, the paths in the spinal cord may be said to be distinctly traceable. Paths in the Anterior Columns of the Cord. — The general arrangement of the motor paths in the anterior part of the spinal cord is maintained throughout. Histology has shown us, however (p. 40), that the two halves of the cord are bound together by the commissures ; this fact sug- gests a crossing, at least partial, from one side to the other, of the nervous impulses. Experiment upon the lower animals seems to show that a partial crossing of the motor paths takes place in the cord. In man's case, most if not all of this crossing from side to side, so far as the paths of voluntary motion are concerned, occurs very high up, if at all, in the spinal cord. But the structure of this organ is such as plainly to provide for an intermingling of the paths of the sensory and the motor roots at about STRUCTURE OP THE SPINAL CORD AND BRAIN. 71 the same level. Thus its character as a pile of centres is maintained. Paths in the Posterior Columns of the Cord. — In the posterior parts of the spinal cord are the paths by which the sensory impulses chiefly run from the posterior roots up to the brain. These paths also seem to undergo a par- tial crossing from one side to the other in the cord. In the lower animals, according to the evidence of experiment, the sense of feeling is retained after the cord has been cut entirely through from the front to the posterior columns. Stimulation of these columns produces signs of pain and other sensory effects. Some investigators would confine the paths, by which sensory impulses of touch pass along the cord, to the posterior columns ; they would assign to the gray matter of the cord, the paths for impulses giving rise to sensations of pain. Others consider that these columns conduct sensory impulses only so far as the nerves from the sensory roots pass through them ; it is then the gray substance which carries these impulses upward. The conclusions by which some experimenters locate motor, and even voluntary motor, paths in the posterior columns, are extremely doubtful. Paths in the Lateral Columns of the Cord. — In these por- tions of the spinal cord both the paths of motor and those of sensory impulses are found. As to the former there is little or no dispute. As to the existence of sensory paths in the lateral portions of the cord, the evidence of experi- ment is somewhat conflicting: but, on the whole, it seems to favor an affirmative conclusion. This conclusion accords well with histology. It must be remembered that, when we speak of " paths in the spinal cord," we are not to think of a perfectly fixed and rigid course like that of the iron rails upon which a locomotive runs. No nerve-commotion, when started in any portion of the cord, is necessarily and under all circum- 72 PHYSIOLOGICAL PSYCHOLOGY. stances compelled to take one, and only one, path to its destination. Secondary paths, besides the primary and more ordinary paths, exist in abundance. A considerable work of substitution, especially as regards the tracts along which the sensory impulses move, may then take place. Even in the case of the voluntary motor tracts in man, although such a work of " substitution " apparently does not take place, a certain latitude of movement from a straightforward course undoubtedly exists. Paths in the Brain. — The evidence already presented from histology indicates that certain tracts, probably motor, pass from the crusta through the internal capsule, without entering the basal ganglia, into the frontal and parietal convolutions. Other tracts, which are probably sensory, run through the tegmentum, enter the thalamus and sub- thalamic region ; then, after being redistributed, emerge to find their way to the temporal and occipital lobes. How well physiological experiment agrees with this general conclusion, we shall see subsequently. The paths by which the sensory impulses travel in the brain must be exceedingly intricate ; for the phenomena connected with all sensory disturbances are very compli- cated and often conflicting. For example, if a sensory cranial nerve is severed, the different functions of feeling pain, of pressure, and of temperature, and the power of localization, in the region supplied by that nerve, are all lost. But disease of the cerebro-spinal axis may impair one or more of these functions, and leave the other intact. Again, loss of the sense of temperature and of the mus- cular sense rarely occur separately; but muscular sense frequently disappears and the sensitiveness of the skin to pressure is retained. The paths both of motor and of sensory impulses, cross in the region of the pons Varolii and medulla obloiagata. All the paths of both kinds lie very close to each other in STRUCTURE OF THE SPINAL CORD AND BRAIN. 73 the white nervous substance surrounding the basal gan- glia. There is considerable recent evidence ^ to show that the tracts followed by impulses of muscular sensation pass through the posterior columns or cornua of the spinal cord, and are gathered into more or less distinct bundles, at the back part of the internal capsule, before they diverge to enter the hemispheres. The confidence with which M. Luys, in his work on the " Brain and its Functions," has localized the paths of motor impulses wholly in the striate bodies, and those of the different sensory impulses, olfactory, visual, tactual, auditory, in those four centres of the optic thalami which he distinguishes, cannot be maintained. The tendency of modern investigation is to place more emphasis upon the fibrous nerve-matter surrounding these organs as furnish- ing paths for the conduction of both kinds of impulses. Substantially, in its more obvious outlines, as we have described it, but with an infinite and indescribable com- plexity of details, is constructed this marvellous mechan- ism of the nervous system. Some of the more particular functions of its parts will be investigated in other con- nections. But even this description shows its fitness to serve as a physical basis for the equally indefinite and indescribable complexity of the mental life. How many variations of fundamental types do the organs of the phys- ical mechanism display ! And of how many kinds, shades, degrees of intensity, and modes of local coloring, are our sensations and their representative images susceptible ! The immense variety and essential unity of this physical basis suggest a corresponding variety in unity of the psychical life. 1 Bastian, in Brain, April, 1887, p. 69 f. CHAPTER III. STRUCTURE OF THE ORGANS OF SENSE AND MOTION. In the general division of labor among the organs of the nervous system, certain groups of cells at the surface of the body become especially sensitive to external stimuli. These cells accordingly acquire the special function of receiving and modifying the action of such stimuli, and thus of setting up in the conducting nerves a neural pro- cess which is propagated to the central organs. Every " end-organ," therefore, looks both inward and outward. Significance and Kinds of End-organs. — The en'd-organs are divided into two classes: first, end-organs of sense, and second, end-organs of motion. The former are in general made up of cells, which, posteriorly, pass into nerve-threads that are gathered into the nerve of special sense ; and which, anteriorly, develop conical or fusiform processes. The simplest type is, then, a hair-like process extending outward and connected by a sensitive cell with a nervous filament extending inward. Only a small part, however, of what are called "the organs of the special senses " belongs to the nervous system. The greater part (as, for example, of the ear, the eye, the skin) consists of mechanical contrivances designed to modify the exter- nal stimulus, while conducting it to the truly nervous mechanism. End-organs of Smell. — That portion of the mucous mem- brane of the nose which clothes the upper region of the nasal cavity, and is called regio olfactoria, contains the 74 OE6AKS OF SENSE AND MOTION. 75 end-organs of smell. Two different kinds of cells are here discovered. By Max Schultze and most other investigators, one of these is consid- ered non-nervous and epithelial, the other nervous and olfactory. The epithelial cells are the larger, have an oval nucleus of consid- erable size, and extend through the whole epi- thelial layer. The olfactory cells are spindle- shaped, with a round nucleus, and very fine long processes. Other investigators believe that this difference is not a fixed distinction of kinds, but is rather indicative of different stages of development. The exact relation of the fibrils of the olfactory nerve (which is the specific nerve of smell, and really, in part, a lobe of the brain) to the epithelium of this region is not made out. It seems probable, however, 'that they do not pass directly into factory ceiu and ^ J r J EpUheliiil Cells the processes of the end-organ cells, but are from the Mucous ■*■ o Membrane or the lost in a network whose interstices are filled f°^-J",('-- i^^- ter Schultze.) up with nerve-granules. The contrivance for bringing the stimulus to the end- organs of smell is comparatively simple. It is necessary only that a current of air, in which the stimulating parti- cles float, should be drawn over the mucous membrane of the region. Since in expiration the current is carried past th^ sensory parts without striking them, smelling is almost entirely confined to inspiration. When "snuffing," we increase the amount and force of the air drawn into this region, by first creating a partial vacuum in its cavity. End-organs of Taste. — On the upper surface of the root, and on the borders and apex of the tongue, and in some cases, on the anterior portion of the soft palate, are found certain papillae. Of those papillae which contain the end- organs of taste two kinds are distinguished ; the circumval- 76 PHYSIOLOGICAL PSYCHOLOGY. Fig. 33. — Transverse Section through a Pap- illa Oircumvallata of a Calf. Showing the ar- rangement and distribution of the gustatory bulb. ">/,. (Engelmann.) latce and the fungiformes. The circumvallate papillae are composed of connective tissue invested by epithelium arranged in plates (lamince). At the sides where the epithelial layer is thinner, the end-organs of taste form a zone which extends up- ward to the level where the papillae are no longer protected by the lateral wall. In the fungiform papillae these organs ap- pear in the epithelium which covers their upper surface, and in the sur- faces of the sides. The Gustatory Flasks. — These structures, sometimes called " gustatory knobs " or " bulbs " occupy flask-shaped cavities in the papillae, which they completely fill. Their lower part rests on connective tissue; their upper part or neck has an opening or pore, of from -^^^ to twi^ inch in diameter. Each flask consists of from fifteen to thirty long, thin cells, ar- ranged like the leaves of a bud around its axis. The margin of the pore is formed by bringing several cells together. The gustatory flasks also are composed of two kinds of cells, one epithelial, the other ^ms- tatory. The epithelial or " investing " cells are long, nar- row, bent, spindle-shaped, with a nucleus well marked. The outward end is pointed, the central end branching. The gustatory cells are thin, long, and highly refractive of Fig. 34. — Gustatory Bulbs from the Lateral Qua tatory Organ of the Babbit, 'v/-^, (Engelmann.) ORGANS OF SENSE AND MOTION. 77 Fig. 35. — a, Isolsited Gustatory Cells, from the Lat- eral Organ of tbe Kabbic; &, an Investing and Two Gus- tatory Cells, isolated but still in connection, '^'^/i. (En- gelmann.) light, with nearly the whole body occupied by an elliptical nucleus. The body of the cell is elongated into two pro- cesses ; the upper one of which is broad, but bears a short, fine point, like a stiff hair. This point lies ,» in a canal and rarely projects from the pore of the flask. The glosso-pha- ryngeal nerve is the principal nerve of taste ; but the lingual branch of the trige- minus is thought by some to take part in sensations of this sense. The nerve is distributed to the back of the tongue, then enters the papillae where it forms a minute plexus interspersed with nerve-granules. Its connection with the nerve-cells of sense is probably indirect, through this plexus and its granules. End-organs of Touch. — The sensory nerves distributed to the skin, the organ of touch in the larger sense of the word, terminate either in free end- fibrils or in special structures called "tactile corpuscles" or " end-bulbs." These special structures have been named after different investigators. The three or four different kinds may, however, be considered as modifications of one type. The first of these structures Fig. 36. — Corpuscle of Pacini (or Vater) from the Mesentery of the Cat. (After Frey.) a, nerve with its sheaths; h, system of tunics constituting tbe capsule of the corpuscle ; c, axial canal, in which the nerve-fibre ends. 78 PHYSIOLOGICAL PSYCHOLOGY. to be discovered (more than a htindred and fifty years since, by Vater) was the so-called " Corpuscle of Pacini " (or Vater). These bodies consist of layers of connective tissue, arranged ^ 4ffik l%= concentrically, and most closely packed near the centre. The layers i- J) ""^^ MK surround a cavity containing a soft, nucleated material into which the nerve penetrates. The axis-cylinder of the nerve-fibre which enters the The bulb In FiQ. 37.— Bnd-bulbs from Eje'"'iA^r^mk^Tf, ^^^^ ^® finely fibrillated. form'r^coirwHh'n'fhetnd' cousists of granular substance. The 'nfr've^fibre of"5 ends ^an these bodies abound in the palms within in the form of a knot. ^f ^j^g j^^j^^ ^^^ ^^^ ^^^^^ ^f ^^ie fcct ; but especially in the palmar surfaces of the fingers and toes. In some places they are visible to the naked eye as a minute grain ^ to ■!■ inch in diameter. The "End-bulbs of Krause" are similar to the foregoing structures. They are small capsules of connective tissue in which nuclei can be de- tected, and in which the nerve- fibrils seem to terminate in a coiled mass or bulbous ex- tremity. The " Corpuscles of Wag- ner" (or Meissner) are oval- shaped bodies bearing some resemblance to a miniature fir- cone. The nerve-fibres appear like " creeping roots," to wind beneath the papillse of the skin and, interpenetrating them here and there, to terminate in the corpuscles. Within Fig. 38. — Corpusclesof Touch. (Af- ter Frey.) a, from the soft skin of the duck's hill ; b and c, from the papillse of the tongue of the same animal. ORGANS OF SENSE AND MOTION. 79 the corpuscles, the fibrils are described as forming two or three coils and joining together in loops. Of 400 papillae counted in ^ inch square, on the third phalanx of the index finger, these corpuscles were discovered in 108. They are from -g^y; to Yhs inch long. Since the surface of the skin is everywhere sensitive to pressure and to temperature, but these special structures are not found everywhere, it follows that they cannot be the sole organs of touch. As has already been said, the free nerve-fibrils must act as the end-organs of those sen- sations which belong to the whole surface of the body. Research and conjecture have not yet succeeded in assign- ing special functions to any of the varieties of the end- organs of touch. STRUCTURE OF THE EYE. With the exception of the ear, the end-organ for the sensations of light and color is by far the most elaborate and complicated. It is obviously adapted to be the instru- ment of an intellectual and "geometrical" sense. Con- sidered as a whole, its plan may be stated in one sentence as follows: The Eye is an optical instrument of the nature of a water camera obscura, with a self-adjusting lens, and a concave sensitive membrane of nervous matter, on which an image is formed. Its structure affords a practical solution of several problems. Among these the first is of a purely mechanical sort, and may be called " the problem of protection." Coats or Tunics of the Eyeball. — Three coverings sur- round the eye, one of which, in part, acts also as a refract- ing medium. (1) The external coat consists of two parts, (a) the Sclerotic (or " white of the eye ") a firm, fibrous membrane of connective tissue intermingled with elastic fibres ; and (6) the Oornea, or translucent front one-sixth part, which rises and bulges in the middle like a watch- 80 PHYSIOLOGICAL PSYCHOLOGY. glass, and which is covered with conjunctival epithelium. (2) The second coat also consists of two parts, (a) the Choroid, which is of a dark brown color, due to the pres- Fig. 39. — Horizontal Section through the Left Eye. Vi- (Schematic, from Gegenbaur.) ence of pigment cells ; and (6) the Iris, a circular, disk- shaped diaphragm, in the form of a lens, which is bathed with aqueous humor and has in its centre a circular aper- ture (the "pupil"). The anterior border around the iris consists of the ciliary muscle and the ciliary processes. ORGANS OF SENSE AND MOTION. 81 (3) The Retina is the third, or inner coat of the eye. It is a delicate membrane, consisting of nine or ten layers, of exquisite transparency and almost perfect optical homo- geneity. Its inner surface is moulded on the vitreous body, and it extends from the entrance of the optic nerve nearly as far forward as the ciliary processes. Befracting Media of the Eye. — The eye has four translu- cent refracting media. These are (1) the Cornea, already spoken of as the anterior one-sixth of the outer tunic of the eye. (2) The Aqueous Humor fills the space back of the cornea and is divided by the iris into two chambers, of which the front one is the larger. It is limpid and watery, but holds certain salts in solution. (3) The Crystalline Lens is situated between the iris and the vitreous body. It is transparent, biconvex, with its antero-posterior diameter about one-third less than the transverse diameter. It con- sists of a capsule and an enclosed body, is of "buttery consistency " and made up, like an onion, ef concentric layers. (4) The Vitreous Humor consists of a number of firm sheets, between which fluid is contained, built into a body that is, optically considered, transparent and nearly homogeneous. It is a gelatinous form of connective tis- sue. Though it occupies most of the bulk of the eyeball, it has comparatively little physiological significance. Appendages of the Eyeball. — Of the accessory parts of the eye, — eyebrows, eyelids, lachrymal glands, etc., — only the muscles have any interest to physiological psychology. Of these there are six which are attached to the eyeball, somewhat like a bridle to a horse's head. Four of these muscles spring from the bony wall near the point where the optic nerve enters, extend through the length of the socket, and pass directly to the eyeball, where they are attached to it, — one above, one below, one on the outer, and one on the inner side (the recti ; internal and external, superior and inferior'). The other two muscles of the eye 82 PHYSIOLOGICAL PSYCHOLOGY. are called oblique. The superior internal oblique, instead of running directly to connect with the eyeball, passes through a ring, then turns round, and is attached obliquely Fig. 40. — MuBclesof the Left Human Fig. 41. — Muscles of the Left Human Bye, Eye, seen from above, rs, rectus supe- seen from the outside, ir, Zevoior of the upper rior; re, rectus ezternus ; and H^, rectus eyelid, which covers the rectus superior, rs; internus; 08^ superior oblique, with its re, os, as in the preceding figure; ri/", rectus tendon i, which runs through the mem- inferior; oi, inferior oblique, branous pulley u, at the inner wall of the cavity of the eyeball. to the upper surface. The other oblique muscle begins at the inner wall in the socket, passes under the eyeball, and is attached to it opposite to the superior oblique muscle. Formation of the Retinal Image. — The problem which is to be solved by the end-organ of vision, in its most general form, may be stated as follows : A mosaic of localized sen- sations of light and color must he so constructed that changes in the quantity, quality, local coloring, and sequence of these sensations shall be interpreted as the size, shape, locality, and motion of external visible objects. The most important part of the solution of this general problem falls upon the retina. And the most important problem, subordinate to the general problem, is the " formation of an image " upon the retina. But this problem is an optical one. It is solved by the translucent refracting media of the eye. The four media of the eye constitute a system of refract- ORGANS OF SENSE AND MOTION. 83 ing surfaces, each of which is separated from the one adjoin- ing by a circular cut, as it were, in the whole refracting substance. Thus the "image" of the first refracting sur- face of this system of surfaces becomes an " object," as it were, for the second refracting surface ; the second " image " an " object " for the third surface, and so on. In tracing the coui'se of the rays through these media two things must chiefly be taken into the account. They are (1) the indices of the refraction of the refracting media, and (2) the geometrical form and position of all the limiting surfaces. Our knowledge of the indices of the refracting media of the eye is derived by taking the average result of an examination of a number of eyes supposed to be normal. In this calculation the lens is, of course, much the most important of all the media. But the structure of the lens is such (see Fig. 42) that the index of refraction is not Fio. 42. — Median Section through the Axis of the Lens of the Eye. (Schematic, after Babuchin.) homogeneous throughout. Each layer has its own index, and the amount of the index of each layer increases toward the kernel of the lens. By this contrivance the entire work of refraction done by the lens is made greater than the work which could be done by a homogeneous lens with an index of refraction equal to that of its most highly 84 PHYSIOLOGICAL PSYCHOLOGY. refracting part. The mean index for tlie lens of the normal eye may be given = 1.4545. The mean index of refraction for the cornea is given at about 1.3507 ; of the aqueous humor at 1.3365-1.3420; of the vitreous body at 1.3382-1.3485. The position and form of the surfaces of the refracting media can be only approximately determined. Three of these surfaces are most important, — namely, the anterior of the cornea and the two surfaces of the lens. The convexity of the anterior surface of the cornea is greater toward its edge than at its vertex, where it resembles a section of an ellipsoid. The images formed when the pupil is expanded are thus made sharper. No observable refraction takes place at the back surface of the cornea. Accommodation of the Eye. — The ability to alter the re- fracting conditions of the eye for varjdng distances of the object is called its "power of accommodation." This adjustment obviously cannot take place like that of the photographer's camera obscura, where a considerable change can be made in the distance of the lens from the screen on which the image is formed. It is, in fact, effected by changing the convexity of the lens, — principally, if not wholly, at its anterior surface. This may be demonstrated by several methods of experiment. Indeed, when accommo- dation is taking place, the pupil may be seen to contract and to draw its edge forward. Helmholtz calculated the amount of this movement at j-^ to -^ of an inch. The mechanism for accommodation of the eye to varying distances of the objects must be in control of the brain, for the accommodation is voluntary ; it must also consist of muscles that lie within the eyeball. The most generally accepted hypothesis of its action has hitherto been that proposed by Helmholtz. It is assumed that the lens, when at rest, is in a state of tension by its own elastic power. It is kept flattened by the radial tension of the suspensory ORGANS OF SENSE AND MOTION. 85 ligament (sometimes called the zonula, — a structureless membranous body, interposed between the ciliary part of the retina and the vitreous humor, and radiating outward). The mechanism for withdrawing the tension consists of the Conjunctiva cgrneac Proc. ciliarix roAiarer circulixrcr Cilianwslcel Fie. 43. Sectional View of tbe ConnectionB of the Cornea, Ciliary Muscle, Ciliary FroceBsea, etc. '%. (Gegenbaur.) ciliary muscle, the fibres of which are fixed at one end, at the edge of the cornea. When this muscle contracts, the other ends of the fibres are drawn toward its fixed ends ; they thus relax the tension of the suspensory ligament by p ullin g in the opposite direction to this tension, and the lens is allowed to bulge out by its own elastic forces. More recent researches, however, seem to emphasize the elasticity of the vitreous body as an important factor in accommodation. For near distances, the contraction of the circular fibres of the ciliary body increases the pressure in the vitreous body, driving it into the spaces at the side of the lens, and bringing the suspensory ligament into a stronger convexity. It is the oculo-motor nerve which furnishes in the poste- rior strands of its roots, the fibres that serve the ciliary muscle. Their place of origin is in the back part of the 86 PHYSIOLOGICAL PSYCHOLOGY. floor of tlie third ventricle. This place lies very close to that where stimulation produces contraction of the inter- nal rectus muscle, the use of which is connected with adjustment for near distances. Thus all the mechanism of accommodation is made to work together for the production of the image on the retina. [It is assumed that the laws of optics under which the formation of the image takes place are known, or are to be acquired, from the proper sources. J Outer surface. ft> — Layer of pigment cells. ■ Layer of rods and cones. .Membrana limitans externa. .Outer nuclear layer. — Outer molecular layer. y — Inner nuclear -layer. . luner molecular layer. J ... Layer of nerve-cells. ' — Layer of nerve-fibres. . .Membrana limitans interna. Inner surface. Fio. 41. — Diagrammatic Section of the Human Eetina. (Schultze.) ORGANS OF SENSE AND jMOTION. 87 i H Given the formation of the image upon the retina, it is further re- quired, in order to vision, that the proper physiological processes should be set up within this organ. For it is the Retina which contains the nervous elements by whose ac- tion the system of refracted rays is changed into a mosaic of nerve- commotions. We require then a detailed description of the struc- ture of this wonderful organ. Layers of the Retina. — The ner- vous and other elements of the / retina are arranged in ten layers, — counting from within outward and backward, — the names and order of which are given in the accompanying schematic represen- tation (see Fig. 44). By no means L- all the retinal substance is nervous. Numerous radial fibres, which pene- tra'te its entire thickness, are of connective tissue, in the gaps of which the true nervous elements lie embedded. These gaps are ,, particularly large in the second, U~- third, fifth, and seventh layers. Figure 45 shows (diagrammati- cally) the connections of the true nervous elements as they (^ extend through the retinal layers. ^ We notice here (a) the retinal fibres of the optic nerve, lying parallel to the surface. They are Fio 45 — Diagrammatic representation of the Connections of the Nerve-flbres in the Retina.' (Scbnltze.) The niimbers have the same reference as in rig. 44, 88 PHYSIOLOGICAL PSYCHOLOGY. non-meduUated, extremely fine, arranged in ray-like bun- dles, radiating on all sides from the place of entrance of the nerve. Next (6) come the ganglion-cells, resembling the multipolar nerve-corpus- cles of the rest of the cerebro- spinal system, which form the principal part of layer No. 3. These cells at ■^ j| v _|{;/^ Vw^^-rv the " yellow spot" are eight or ten deep, but diminish in number toward Fio. 46. — Diagrammatic Section of t^^ Ora ServatU. the Posterior Part of the Ketina of a Pig. ^ n 'P'Kq vic^t. «»«/,. (Schultze.) 7, part of outer nu- Kyj -l-ne uei- clear layer; 8, membrana limitans ex- ._r__,„ cAa-rv^a-ni-o terna; 9, rods and cones. Each of the VOUb eiCmenuS cones, which are in very close apposition, -.i: i „ "vr a contains in its inner segment a highly re- 01 layer 1>I 0. tt fractile body, the function of which is i i -i unknown. probably con- sist of extremely fine filaments connected with the processes of the ganglion-cells. (d) Each nucleus-like body in No. 5 is thought to be connected by fibres, both in- ward and outward, (e) In the outer molec- ular layer (No. 6) are numerous filaments of nervous character, while its star-shaped cells are probably not nervous. (/) In layer No. 7 are many nucleus-like bodies connected by radial fibres with the nervous elements of the rod-and-cone layer. Accord- ing to their connection they are called rod- |Ss' of^thl surfacl granules and cowe-granules, respectively. ?e°^gths"lff th^tate"- Rods and Cones of the Retina. — The struc- (Scruu™';) T^e ture of layer No. 9 is particularly interesi> °„"r fsTroken 'into ing. It consists of a multitude of elongated ^^b/ren?.'"" "' '"" Fig. 47. — Rod and Cone from the Hu- man Ketina, pre- served in perosmic OEGANS OP SENSE AND MOTION. bodies arranged side by side, like rows of palisades. These bodies are of two kinds, — one cylindrical, extend- ing the entire thickness of the layer (^^ inch long), and called "rods"; the other, flask-shaped, shorter, and called " cones." The inner ends of both bodies are supposed to be continuous with the fibres of the outer nuclear layer. Each rod, or cone, is composed of an inner and an outer segment or limb. The inner limb appears under the microscope protoplasmic, and feebly refractile. The outer limb is highly refractile. It has been thought that an extremely minute nerve-filament is drawn through the axis of these bodies. It is assumed that they are con- nected with the fibrils of the optic nerve by means of the retinal nerve-cells and radial fibres. In close connection with the rods and cones stand the flat six-fiided cells of the pigment-epithelium. In the cen- tre of the eye only cones appear, which are here of more slender form and increased length ; so that not less than 1,000,000 are set in ^ inch square. Not far from the centre each cone is surrounded by a crown-shaped border of rods. Toward the ora serrata the cones become rarer. The Yellow-spot and th( Blind-spot. — The place oi clearest vision, and the physiological centre of tht eye, is the "yellow-spot' (macula luted) ; it is of oval shape, about ^ inch long, with a depression in the centre (Jovea centralis). About \ of an inch interior from the middle of this spot is the place where the ,S Ch Fig. 48. — Bguatorial Section of the Right Eye, BhowiDg the Papilla of the O^tic Nerve, the Blood-vessels radiatiog from it, and the Macula Lutea. y,. (Henle.) B, eclerotlc; Ob, choroid ; and B, retina. 90 PHYSIOLOGICAL PSYCHOLOGY. optic nerve breaks into the retina. This place is called the " blind-spot," because it can be shown to be inoper- ative in vision. Its size varies considerably for different eyes (-j-V to -^ inch long). It is wanting in all the ner- vous elements. Photo-chemical Processes in Vision. — The physiological or nervous process concerned in vision can be shown to begin in the rods and cones at the back part of the retina. Indeed, by throwing a strong light through the cornea, and causing it to be reflected within the eyeball before it reaches the nervous elements of the retina (an experiment devised by Purkinje), it is possible to perceive the arbores- cent figure formed by the shadow of the blood-vessels expanded on the front part of our own retina. Yet the rods and cones are not directly irritable by light, so as to produce visual sensation, — at least, not unless the inten- sity of the stimulus be so great as to be injurious to the nervous substance itself. How, then, can we see the feeblest rays of the moon as reflected from white paper? A photo-chemical process has accordingly been assumed to result from the direct action of the light ; and this process it is which acts as the immediate stimulus of the nervous elements. Yet after many careful experiments, especially by Kiihne and his pupils, it is difficult to establish the nature of the photo-chemical process concerned in vision, or of the par- ticular pigments upon effecting changes in which the pro- cess is dependent. Any theory involves, chiefly, these two things : fiist, the decomposition by the light of some sub- stance found in the epithelial elements of the retina ; and, second, the action of the decomposition-products thus gained upon the protoplasm of the nervous end-organs. There seem to be decisive objections against making the pigmentum nigrum, or the "visual purple," or any other known pigment, the only substance whose decomposition ORGANS OF SENSE AND MOTION. 91 by the light is necessary to vision. And as to the nature of the changes produced in the rods and cones by the decomposition-products of any pigment, we are really igno- rant. A photo-chemical theory of vision seems, therefore, to be a desideratum rather than a scientifically established and definite fact. The most patent thing revealed by the structure of the eye is its adaptation to serve as the organ of a highly differ- entiated system of intellectual and " geometrical " sensory impressions. The fuller significance of this truth cannot be understood until we have made a detailed study of those series of sensations — infinitely varied, delicately shaded, quickly successive, and speedily and firmly fusing together, as it were — which result from the activity of this organ of sense. STRUCTURE OF THE EAR. The End-organ of Hearing, like that of vision, so far as the principal part of its bulk is concerned, consists of mechanical contrivances for applying the stimulus to the genuine nervous elements of the special sense. The true end-organ is the mechanism of epithelial and nervous cells which is connected with the terminal fibrils of the special nerve of this sense. A brief description of the non-nervous structure is, however, desirable for our purpose. The entire Ear consists of three parts, or ears; these are the external ear, the middle ear, or tympanum, and the inner ear, or " labyrinth," as its complicated structiire causes it to be named. I. The External Ear. — Exclusive of the plate of carti- lage which projects from the side of the head, the external ear consists of two parts. These are the concha and the external meatus. (1) The concha is a rather deep hollow of a shell-like shape. It is probably of little or no use in sharpening our acoustic perceptions ; although it appears 92 PHYSIOLOGICAL PSYCHOLOGY. to be of some service in discerning the direction of sound. (2) The external meatus is a curved passage leading from the bottom of the hollow of the concha to the drum of the ear. Its most obvious office is the protection of the ear- drum ; though it may also modify certain tones by its own resonant action, — strengthening the high ones and dead- ening the low, in some degree. If we place a resounding body iu contact with the teeth, the intensity of the sensation of sound is much increased. This appears to us to be due to direct conduction through the cranial bones ; but it is more probable that the principal path of conduction is indirect, through the ear- drum and small bones of the middle ear to the fenestra ovahs. II. The Middle Ear, or Tympanum. — This part of the or- gan of hearing is an irregular cuboidal chamber, situated in the temporal bone, between the bottom of the meatus and the inner ear. Its outer wall is the membrana tympani or " drum " of the ear. In the inner wall, which separates it from the labyrinth, are two 1 openings called "windows" — the fenestra ovalis and the fenestra rotunda. Near its anterior part it opens into the Eustachian tube. And an irregular chain of bones — called auditory hones — Fie.49.-Drumof the Right Ear with S*^^**'^^^ ^''^OSS the Cavity fH^Se7™?;Sda*ty4aniftBu:6: ^^^^^ ^^s outcr to its inner chian tube; *, tendon of the tensor tym- Tvall pani muscle out off close to its insertion ; ma, anterior ligament of the malleus ; Mcp, rri,- TVTo-mlirQTiQ TTrm-nani ai- its head; and Ml, its long process. Stp, ■'■^^ memorana lympaiU, OP Spir^ tympanica posterior. jj^,^ ^^ ^^^ jjg^j. _ rpj^-g ^^^_ brane consists of three layers, — an external, and internal mucous, and the intermediate membrana propria. The ORGANS OF SENSE AND MOTION. 93 last is the true vibrating membrane, and is composed of unyielding fibres arranged both radially and circularly. A flat membrane, evenly stretched, whose mass is small in proportion to the size of its surface, is easily thrown into vibration by acoustic waves striking against one of its surfaces. It responds readily to tones which approach its own fundamental tone, but is scarcely at all affected by divergent tones. Such a membrane, therefore, cannot repeat a motion which consists of a series of harmonious partial tones. In order to perform such a service, a mem- brane must be so arranged and connected as to have no preponderating tone of its own. For this the ear-drum is prepared by two devices : (1) It is drawn inward into a funnel-shaped form by being attached to one of the auditory bones (the handle of the malleus) ; and (2) it is loaded with a chain of bones so as to have no trace of a tone of its own (see Fig. 60). Thus is secured for it the property of taking up the vibrations of a large scale of tones. Moreover, since the apex of its funnel bulges inward, the ear-drum serves to concentrate the force of the vibra- tions from all sides. Loading it with the auditory bones serves also to dampen its vibrations and prevent them from continuing too long. This gives speed to the rate at which definite auditory sensations can be repeated. The Eustachian Tube. — This opening from the middle ear into the mouth is of indirect but important physiolog- ical service to auditory sensations. In its normal position the tube is neither closely shut nor wide open. When we swallow, it opens and thus effects a renewal of air in the middle ear, maintains an equilibrium of pressure on both sides the drum, and conveys away the fluids that collect in its cavity. But if it remained constantly wide open, we should be likely to hear our own voices as a roaring sound, and the passage of the air in and out during respiration would effect the tension of the drum. 94 PHYSIOLOGICAL PSYCHOLOGY. The Auditory Bones. — The chain of bones which stretches across the tympanic cavity consists of three members, — the Malleus ("hammer"), the Incus ("anyil"), and the Stapes (" stirrup "). The malleus has a head separated by a con- stricted neck from an elongated handle. The latter is attached to the centre of the membrana tympani. The incus has a body and two processes. On the front surface of its body is a saddle-shaped hollow. Its short process is bound by a ligament to the posterior wall of the tym- panum. Its long process ends in a rounded projection (^os orbicular e). The stapes has a head and neck, a base and two crura. These are put together so as to give it the stirrup-shape, which its name implies. The manner in which these bones articulate may be fairly well seen by the accompanying - figure (No. 50). The auditory bones are moved on each other at their joints by two or three mus- cles, — especially by the ten- sor tympani. This muscle is inserted into the malleus, Y ' near the root, and serves to FIG. 60. -Bol. of the Ear, as seen m ^^g^^^^ the tympanic mcm- i'STanv;?r/o7wSir?b-iyVhfslt;! brane by drawing the malleus and II the long, process ; c, Us body, and irrnrnrrl TVip npmietif" TriVirn. pi. the process for articulation with the mwaru. ±ne aCOUStlC VlDra- Bi&pes (processus orbicularis). M, Mai- fi„„(, imnnrtprl >nr +Tio Tnom leus (hammer), of which Mc is the neck; llOUS, impariea Dy tUC mCm- Mep, the head; Ml, the long process; and 1,^5,^,0 tn tVip«p hnnps nrp Tint Mm, the manubrium; S, stapes (stirrup), UiaUBLU uneht! UUntJh, die UOl with its capituium,cp. longitudinal but transverse. They do not, however, resemble the vibrations of a stretched cord or a fixed pin. The bones vibrate as a system of light, small levers, with a simultaneous motion around a common axis. The vibrations are sympathetic, and vary greatly for tones of different pitch and similar intensity. OEGANS OP SENSE AND MOTION. 95 General Office of the Middle Ear. — By this part of the organ of hearing the acoustic waves are transmitted to the inner ear, while their character is greatly modified. Mod- ification is necessary to prepare the stimulus for the organ- ism of the inner ear. The waves in the air, when they reach, the ear-drum, have a large amplitude but a compara- tively small intensity. Their motion must be changed into one of diminished amplitude and increased intensity. But the transmitting, vibrating media must also have the power of answering to the different tones of any pitch perceptible by the ear. m. The Internal Ear, or Labyrinth. — It is in this mar- vellously complex organ that the terminal fibrils of the auditory nerves are distributed and the end-organs of hear- ing are placed. It consists of three parts — the Vestibule, the Semicircular Canals, and the Cochlea. Each of these parts is, as it were, made twice over — once in the form of channels cut in the petrous bone (the osseous vestibule, etc.), and again in the form of a membrane (the membra- nous vestibule, etc.) suspended in the bony cavity, but only partly filling it. No. 3. ■vpj Fio. 51. — No. 1, OsaeouB Labyrinth of the Left Ear, from below; No. 2, of the Bight Ear, from the ioside ; No. 3, of the Left Ear, from above. (Henle.) Av, aque- duct of vestibule ; Fc, fossa of the cochlea ; Fee, its fenestra (rotunda) ; Fv, fenestra of the vestibule (ovalis); ha, external ampulla; h, external semicircular canal; Tsf, tractus spiralis forammosus ; vaa, ampulla of the superior semicircular canal; vc, pos- terior semicircular canal ; and vpa, its ampulla. (A) The Vestibule of the Ear. — The central cavity of the inner ear, called -the "vestibule," is the earliest 96 PHYSIOLOGICAL PSYCHOLOGY. and most constant part of the labyrinth. In its outer wall is the fenestra ovalis ; its anterior wall communicates with the scala vestihuli of the cochlea. The membranous vesti- bule is composed of two sac-like dilatations — the upper and larger called utriculus, the lower saoculus. (B) The Semicircular Canals. — These curved channels open into the utriculus. They are three in number, about one inch in length and ^V iiich in diameter. They have a regular relative position — their planes being at right angles to each other — as indicated by their names — superior, posterior or vertical, and external or horizon- tal (see Fig. 51). Near the vestibule they dilate into the so-called ampullce. Both the osseous vestibule and the osseous canals contain a fluid (^perilymph') in which the corresponding membra- nous parts — themselves distended with a fluid (ew- dolympK) — are suspended. The oifice of this fluid is very important in the trans- mission and further modi- fication of the acoustic waves. ((7) The Cochlea. — This wonderful organ is shaped like the shell of a common snail, about \ inch long. It, too, consists of a mem- branous sac in an osseous cavity. The whole passage is imperfectly divided into two canals by a partition- wall of bone (the lamina spiralis'), which winds 2^ times around an axis (the modiolus'), like a spiral staircase* Fee Fis. 52. — OsBeouo Cochlea of the Right Ear, exposed from in front. Vi- (Henle.) t, section of the division-wall of the cochlea; tt» upper end of the same; Fee, fenestra; H, hamulus; Md, modiolus; Ls, lamina spira- lis. ORGANS OP SENSE AND MOTION. 97 Uj/amenttiiii spiratt Stria vascularis Lamina iaailarii Of these canals, the one which faces the base is called scala tympani; the other, which opens into the vestibule is the scala vestihuli. At the apex they com- municate through a small hole (the helicotremdy. From the free edge of the spiral lamina to the outer wall the interval is bridged over by the basilar mem- brane. Still another membrane (memr Fio. 53. — section through one of the Colls of the brane of Reissner-) *""'"'''■ "^^ (s^'-^'-' "- ^egenbaur.) arises from a crest attached to the same lamina and ex- tends to the outer wall, so as to make a minute canal between itself and the basilar membrane. In this canal — called scala intermedia, or ductus cochlearis, or " canal of the cochlea " — the nervous end-organs of the cochlea are found. The Organ of Corti. — On that surface of the basilar mem- brane which is directed toward the small canal of the cochlea is placed a structure which consists of a wonder- ful arrangement of cells. Some of these cells are curved, elongated, and placed in two groups — an inner and an outer (rods, ot fibres of Corti). The two are arranged (as shown by Fig. 54) so as to make a bow or arch over an exceedingly minute canal (canal of Corti) between them and the basilar membrane. These rods of Corti increase in length from the base to the apex of the cochlea. Each rod rests upon one or two of the transverse fibres of the basilar membrane. The Organ of Corti is separated from the endolymph of the, ductus cochlearis by the so-called 98 PHYSIOLOGICAX, PSYCHOLOGY. membrana tectoria. (For the shape and position of the "hair-cells," inner and outer, the supporting cells, etc., see the accompanying diagrammatic representation.) Fio. 54. — Organ of Corti in the Dog. ^^/x. (Waldeyer.) 6 — c, homogeneous layer of the basilar membrane; u, its vestibular layer; v, its tympanal layer; d, blood-vessel; /, nerves in spiral lamina ; g^ epithelium of spiral groove ; A, nerve-hbres passing toward inner hair-cells z, X;; Z, auditory hairlets on inner hair-cells; I — Zj lamina reticularis; m, heads of the rods of Corti jointed together; the iuner rod seen in its whole length; the outer one broken off; n, cell at base of inner rod; p, q, r, outer hair-cells; s, a cuticular process probably belonging to a cell of Belters ; i, lower ends of hair-cells, two being attached by cuticular processes to the basilar membrane; w, a nerve-fibril passing into an outer hair-cell; z, a sustentacular cell of Belters. Distribution of the Auditory Nerve. — The auditory nerve, on approaching the labyrinth, divides into several portions. In the vestibule, branches are distributed to the utriculus, the sacculus, and each of the three ampullae. In a ridge in the wall of each of these dilatations (the crista acoustica) columnar and fusiform cells are found, with processes from the latter of which the fibrils of the auditory nerve are brought into connection, — probably by means of that minute network of fibrils with which we have already become familiar in the other end-organs of sense. The so-called " auditory hairs " are found by recent observers to be connected with the columnar cells ; they form, on the inner surface of the epithelium of the ridge a " thick-set ORGANS OF SENSE AND MOTION. wood." Calcareous particles exist (the otoliths or "ear- stones ") in both saccule and utricle, lying embedded in contact with the nerve-epithelium. The cochlean branch of the audi- tory nerve pierces the modiolus and gives off lateral branches which pass into channels in the osseousspirallamina. Here the fibres radi- ate to the membra- nous lamina and are connected with a ganglion of cells. Beyond this gangli- on they lose their me- dullary sheath, and become extremely fine axis-cylinders, the delicate fibrils of which are proba- bly connected with the nerve-plexus of the fibrils from the cone-cells of the organ. Special Office of the Labyrinth. — The greater bulk of the inner ear, like the whole of the other principal parts, is used to transmit and modify the acoustic waves. We have seen that the membranous labyrinth is filled with, and sus- pended in, a fluid medium. Molecular oscillations of this fluid can scarcely, however, serve as the direct stimulus of the nerve-elements. Its dimensions are so very small in comparison with the length of the acoustic waves as trans- mitted by the shocks of the stirrup at the fenestra ovalis, pulla Fio. 55. — Scheme of the N'erve-endiQgs in the Am- illse. (After Budinger.) 1, membranous wall of the ampullae, with a Btructureless border 2, through which the nerve-fibre 3, sends its axis-cylinder 4; 5, plexiform connection of the nerve-fibres; 6, auditory cells; 7, sup- porting cells; 8, auditory hairs. 100 PHYSIOLOGICAL PSYCHOLOGY. or the pulsations of air at the fenestra rotunda, that the movement of the entire fluid mass would be practically instantaneous. Several places may be pointed out, how- ever, into which the waves of the fluid could retreat, by the membranes yielding, or by itself running into pores and through the passages connecting the various parts. The friction of these movements back and forth, especially when increased by the action of the otoliths, might thus irritate the nerve-elements. The basilar membrane would undoubt- edly be thrown into vibration by the unequal pressure of the fluid, and thus the nervous structures situated upon it might be irritated. In all this, however, much is still a matter of doubtful conjecture. A still more difficult question to answer is this : How does the ear manage to analyze the acoustic influences? This organ plainly is not contrived so as to reproduce changes in the form of acoustic oscillations in such manner that these changes can be made apparent to the eye or to touch. But our experience with " clangs," or the musical notes of ordinary kind, seems to require that we should find in the ear some sympathetic vibratory apparatus. Such apparatus must suffice for all kinds of noises, and for all musical tones, and for simultaneous hearing of several tones as harmony, and for so rapid a succession of different sensations as occurs when we hear a melody, or even the crackling of an electric spark at intervals of 0.002 sec. It has been for some time commonly assumed that the vestibule and semicircular canals are the organs for hear- ing noises, and the cochlea for hearing musical sounds. This differentiation of function is certainly suggested by the marked differences in the structure of their parts. The otoliths and hairs of the former do not seem adapted for regular sympathetic vibrations. The rods of Corti and radii of the basilar membrane suggest that here is the apparatus needed for acoustic analysis. ORGANS OF SENSE AND MOTION. 101 Recent investigations, however, tend to show that the physiological distinction between noises and sounds will not hold with sufficient rigor. The two seem to pass into each other by insensible gradations. It has been found possible to make a series of sharp noises, like a watchman's rattle, as often as 600 times per second, without producing a musical tone, if all extra accompanying sounds are com- pletely dampened. On such grounds it has been concluded that we hear noises and tones with the same organ. It was first argued by Helmholtz that the fibres of Corti — some 3000 in number and arranged in rows on the basilar membrane like the keys of a piano-forte — are just suitable for the required sympathetic vibrations. But these rods are stiff and not easily vibratory, and their office seems to be that of supporting the hair-cells. Birds, moreover, can appreciate musical notes but have no rods of Corti. It has therefore been proposed by Hensen and others to regard the radii of the basilar membrane as themselves graded to pitch. These radii, by moving up and down, might excite the conical hair-cells, whose number is sup- posed to be about sufficient to satisfy the demands of musical analysis. More recently still, it has been conjec- tured that, since the arches of Corti at the base of the cochlea are small and little spread, and those at the upper end are larger ajid much spread, the size and the shape of these structures may approximately compensate each other ; this would make it possible for all of the arches to vibrate to each of the fibres of the basilar membrane (like the sounding-board of a piano). We are obliged to confess that our knowledge of the minutest structure of the ear, and especially of the manner in which it performs its functions, is exceedingly frag- mentary. As one principal investigator remarks: "It is possible that the working of this apparatus may be altogether different from any of our present conjectures." 102 PHYSIOLOGICAL PSYCHOLOGY. END-ORGANS OF MOTION. The motor nerves of animals have their peripheral con- nection with either electrical organs, or secretory glands, or muscular fibre. A very brief consideration of the last case, only, will suf&ce for our present purpose. After a motor nerve has entered the substance of the so-called voluntary or striated muscle it breaks up into nerve-twigs between the muscular fibres. The axis-cyl- inders of the nerve-twigs lose their medullary sheath, and subdivide into fibrils, which form a flat branching mass within certain disk-shaped bodies inside the sheath of the muscle-fibre (the sarcolemma). These bodies are the so-called " motor end-plates." In the non-striated (or non-voluntary) muscles, the nerves subdivide into very minute plexuses of nerve-fibres, which are distributed in the connective tissue that separates the muscular fibres. The shape and structure of the motor end-plates are different for different animals, and even for different mus- cles of the same animal. The mode of the termination of the nerve in the muscle is thought to be somewhat dis- tinctive of the different parts of the muscular structure. Sometimes the axis-cylinders are enlarged, with granular corpuscles attached or adjacent. Sometimes a granular mass, with its nuclei, forms a kind of floor for the terminal nerve-fibres ; and this eminence may be either elongated, elliptical, or circular. CHAPTER IV. DEVELOPMENT OF THE NERVOUS SYSTEM. In that living germ, in which the life of the indi\ddual human being originates, there is no apparent distinction of bodily organs, or of physical and psychical activities. To scientific observation this germ seems " undifferentiated." But it undergoes a development, and before it can be sub- jected to ordinary observation it has unfolded itself into an elaborate organism. The course of this development . can be traced, in man's case, only very imperfectly by even the most patient embryological investigation. Fortunately, however, the very first things in the life of the other mammals, and even of the chick (the most con- venient subject of study for embryology) are in most important respects similar to those of man's earliest devel- opment. This knowledge, indirectly derived, is constantly being more and more supplemented by direct microscopic inspection of the human embryo, at various stages in its life. Thus a sketch of the principal outlines of man's pre-natal evolution is made possible. A few points selected from such a sketch, and having reference especially to the nervous system, furnish helpful suggestions with regard to certain questions in physiological psychology. EARLIEST DEVELOPMENT OF THE BODILY LIFE. The following brief description of the earliest develop- ment of the animal body is chiefly taken from the detailed embryology of the common fowl.^ 1 For further study of this subject, Foster and Balfour's Elements of Embryology, London, will probably be found most accessible and ser- viceable. 103 104 PHYSIOLOGICAL PSYCHOLOGY. The Ovarian Ovum. — The immature egg (ovarian ovum) of any animal presents the characters of a simple cell. It appears as a naked protoplasmic body containing in its interior a nucleus (^germinal vesicle^, and within this a nucleolus (the germinal spof). It is enclosed in a capsule of epithelium called the " follicular membrane." The principal difference between the ovum of a mammal and that of a bird consists in the amount and distribution of the food-yolk. The human ovarian ovum is only -j-sr to j-5-5^ inch in diameter, because it contains so little food- yolk ; but this small supply is uniformly distributed. As the ovum matures, its body grows in size, and gran- ules appear in the interior. As these -earliest granules enlarge, others appear at the periphery. The germinal vesicle travels toward the surface, and accessory germinal spots make their appearance. The cells of the follicular membrane — at first a single row — now become two or three deep. The superficial layer of the ovum is converted into a striated membrane. Between this and the cells of the follicular membrane another membrane (the vitelline) afterward appears. The striated membrane disappearing, the vitelline remains alone. But the essential constituent of the body of the ovum is an active, living protoplasm. Impregnation and Segmentation. — The spermatozoon, or male fecundating element, may itself be considered as a cell, the nucleus of which is its head. Impregnation takes place by the entrance of a spermatozoon into the ovum, followed by the fusion of the two. On entering, the substance of the tail of the spermatozoon mingles with the protoplasm of the ovum, while the head enlarges and also fuses with a portion of the ovum, thus constituting the nucleus of the impregnated egg. In this actual fusion of substance derived from both parents, provision is made that the offspring shall partake of the physical and psy- chical characteristics of the two. DEVELOPMENT OF THE NERVOUS SYSTEM. 105 Segmentation, or " yolk-cleavage," follows fecundation of the ovum. This process consists in dividing the ovum into a number of cells from which all the cells of the full- grown animal are the lineal descendants. The germinal disk of the ovum is thus broken up into a large number of rounded segments of protoplasm, called the blastoderm. The upper segments being smaller than those beneath, the beginning of two layers is thus made. This distinction is then made more obvious by the segments of the upper layer arranging themselves side by side into a membrane of columnar, nucleated cells; but the segments of the lower layer continue granular, and form a close, irregular network of cells. As the process of segmentation goes on, the differences among the ova of different species of animals become more clearly marked. The mechanical explanation of this is, in part at least, the difference in the amount and distribution of the food-yolk. The Blastodermic Vesicle. — A narrow cavity now appears between the two layers of the ovum, which soon extends so as to separate them completely, except in the region near a small circular area. In this area the inner mass of the ovum has remained longer than elsewhere exposed, before the outer cells closed over it. The enlargement of the ovum, and of the cavity between the layers, gives the whole structure the appearance of a vesicle vdth a thin wall. It is therefore now called the blastodermic vesicle. Its walls are for the most part composed of a single row of outer flattened cells ; while an inner lens-shaped mass of cells appears attached to a portion of the inner side of the outer layer. As the vesicle grows rapidly, this inner mass becomes, on the whole, flattened so as to spread out on the inner side of the outer layer. But its central part remains thick, and forms an opaque spot, which is the beginning of the area where the embryo is to form (the embrt/onic area"). 106 PHYSIOLOGICAL PSYCHOLOGY. Fig. 56. — Vascular Area and Embryonic Area of the Embryo of a Babbit, seven days old. ^/i. (KSlliker.) 0, o, the vascular or opaque area; ag, embryonic area; pr, primitive Streak and groove; rf, medullary groove. The Three Layers of the Embryo. — In tlie area just described there are first formed two distinct strata. Of these, the upper one consists of rounded cells, which lie close to, and become fused with, the flattened outer layer ; it is then called epiblast. The lower one consists of flattened cells, and is called hypoblast. We have thus a double-walled sac (the gastruld). Between these two strata, or layers, a third soon makes its appearance ; from its position it is called mesoblast. These three layers are found in the embryo of all ver- tebrate, and of most invertebrate, animals ; and from them all the different parts of the animal organism are developed. The history of the life of every animal, thus constituted, is in its earlier stages a history of the evolution of these layers. The hypoblast is the secretory layer ; and from it almost all the epithelial lining of the alimentary tract, and DEVELOPMENT OF THE NERVOUS SYSTEM. 107 of its glands, is derived. From the mesoblast come the skeletal, muscular, and vascular systems, and the connec- tive tissue of all parts of the body. But it is the develop- ment of the epiblast which most concerns us. For from it is evolved the central and peripheral nervous system, the epidermis, and the most important parts of the organs of sense. From the beginning, then, the skin in which the end-organs of touch and of the other sensations are situated, as well as all the organs of special sense, constitute, with the brain and spinal cord, one interconnected mechanism. DEVELOPMENT OF THE NERVOUS SYSTEM. The process of differentiating the layers of the blasto- derm is intimately connected with another, in which the foundations, as it were, of the nervous structure are laid. This is — The Formation of the Medullary Groove. — A short sickle- like thickening, due to a forward propagation ("linear Pf ,_ Pt s s Fie. 97. — Primitive Streak of tbe Embryo of a Babbit, eight days and nine hours old. 2™/i. (Kolliker.) No medullary groove has yet been formed, ax, primitive streak ; pr, primitive groove; pf, primitive fold; ect, ectoderm (or epiblast); mes, mesoderm (or mesoblast); en<, entoderm (Aj^oMosO. proliferation ") of epiblastic cells in a straight line, occurs near the junction between the pellucid and the opaque areas, and stretches inward upon the embryonic area. It is called the primitive streak. Its middle line then shows 108 PHYSIOLOGICAL PSYCHOLOGY. a shallow furrow called the primitive groove (see Fig. 57). In the embryonic area, to the front of the primitive streak, the axial part of the epiblast thickens; two folds arise along the boundaries of a shallow groove ; the folds meet in front but diverge behind, enclosing between them the front part of the streak. These are the medullary folds, — the first definite features of the embryo. The part enclosed between the medullary folds is called the " medullary plate " ; it is the portion of the epiblast which gives rise to the central nervous system. The Notochord and the Neural Canal. — Meanwhile an important change has taken place in the hypoblast, in front of the primitive streak. An opaque line has ap- peared, running forward from the front end of the streak, and stopping short at a semicircular fold near the front part of the pellucid area. This opaque line is a concentration of cells in the form of a cord, — the so-called Notochord. Changes in it are to give rise to the distinctively vertebral structure of the animal. The fold is the future head-fold. And now a portion of the blastoderm in the pellucid area, heretofore nearly flat, is " tucked in," with the form of a crescent. Looked at from above, this tuck appears as a curved line along the margin of the medullary groove. Thus the blastoderm becomes at this spot folded in the form of the reversed letter 2 (the "head-fold" already referred to). The upper limb of the fold grows forward, the lower backward. As the fold enlarges, the crescentic groove deepens and its overhanging margin rises above the level of the blastoderm. . Meanwhile the medullary folds increase in height and lean over toward the middle line. They soon come into contact in the brain-region, although they do not at once coalesce. They thus form a tubular canal, called the Neural (or medullary) canal. By the closing of the folds in the head-region, the open medullary groove has now DEVELOPMENT OP THE NERVOUS SYSTEM. 109 The front end of the ■A bl become converted into a tube, which is closed in front but remains open behind. Formation of the Cerebral Vesicles, neural canal has a more rapid growth than the rest. It swells into a small bulb or vesicle, whose cavity is con- tinuous with the neural canal, while its walls are formed of the epiblast. This bulb is the first "brain-bud," or cerebral vesicle. From its sides the processes of the optic vesicles soon grow out. Behind the first vesicle a second, and behind the second a third, is soon formed. Thus there come to be three brain-buds, or germinal brains. From them are to develop the fore- brain, the mid-brain, and the hind-brain. Flexure of the Neural Canal. — The fore-brain vesicle, or front part of the neural canal, becomes bent downward through inequalities of growth. By increase of this flexure the front portion is folded down so that the second vesicle, or mid-brain, pro- jects in front of it. AU the subsequent development of the nervous system is connected with the growth of the three cerebral vesicles and the flexure of the neural or medullary canal. Development of the Cerebral Vesicles. — From the front Fio. 58. — Foi-e-part of an Embryo-chick at the end of the second day, viewed from the Dorsal Side. y,,,. (Kolliker.) Vh, fore- brain; Ablj ocular vesicles; Mk, mid- brain ; HJi, hind-brain ; JT, part of the heart seen bulging to the right side; Fom, vitel- line veins; Mr, medullary canal, spinal part; Mr', medullary wall of the mid-brain ; Uw, proto-vertebral somites. 110 PHYSIOLOGICAL PSYCHOLOGY. part of the fore-brain the vesicles of the cerebral hemi- spheres swell out. Each of these lateral brain-buds has a cavity which is continuous with the cavity of the fore- brain. The cavities on either side become the lateral Mid-brain Foramen ^onroi N. opt. Infundi- N. trig. Fig. 59. — A, Brain of an Embryo of the Rabbit. B, Brain of an Embryo of the Ox. In both cases the side-wall of the left beinisphere is removed. (After Mlhalkovics.) ventricles of the brain. The original vesicle of the fore- brain, having ceased to occupy its front position, develops into the parts around the third ventricle. The front portion of the third cerebral vesicle is now marked off by a constriction ; thus the hind-brain is separated into two parts, — the rudimentary cerebellum with the pons in front, and the rudimentary medulla oblongata. The following table — taken from the ninth edition of Quain's Anatomy, exhibits the relation in which the de- veloped parts of the brain stand to its fundamental rudi- ments : — DEVELOPMENT OP THE NEEVOUS SYSTEM. Ill I. Anterior Pri- mary Vesi- cle. II. Middle Pri- mary Vesi- cle. ni. Posterior Pri- mary Vesi- cle. 1. Prosencephalon, Fore-brain, Thalamen-cephalon, 2. Inter-brain, Mesencephalon, 3. Mid-brain, '4. Epencephalon, Hind-brain. Metencephalon, 5. After-brain. Cerebral Hemispheres, Corpora Striata, Corpus Callosuiu, Fornix, Lateral Ventricles, Olfactory Bulbe. Thalami Optici, Pineal Gland, Pituitary Body, Third Ven- tricle, Optic Nerve (prima- rily). (Corpora Quadrigemina, Crura Cerebri, Aqueduct of Sylvius, Optic Nerve (secondarily). (Cerebellum, Pons Varolii, an- terior part of the Fourth Ventricle. fMedulla Oblongata, Fourth I Ventricle, Auditory Nerve. The Cranial and Spinal Nerves. — Along the cerebro-spi- nal cavity — formed as described above — various changes in the lining of the epiblast take place. This lining is thickened at the side so that the cavity comes to resemble a narrow vertical slit. The sides and floor of the canal of the cerebral hemispheres are also much thickened. But in the region of the third and fourth ventricles the roof of the canal becomes very thin. The cranial nerves spring out of a band, composed of two plates, which connects the dorsal edges of the neural canal with the external epiblast. The fusing of the two plates makes the band into a crest on the roof of the brain. As the roots of the cranial nerves grow centrifugally and become established, the connecting crest is partially obliter- ated. In its earliest stages a cross-section of the brain-tube is essentially like the spinal cord in its internal structure. The region of the medulla leads the others in the develop- ment of the nerve-paths. 112 PHYSIOLOGICAL PSYCHOLOGY. In the spinal cord, the posterior roots of the nerves appear as out-growths of a series of naedian cell-processes i on the back side of the cord. Recent discoveries indicate that the motor- nerves of the cord, as well as of the brain, arise from the fundamental plate. Development of the An- terior Fortionsof theBrain. — Further details con- cerning the later changes in thehind-brainandmid- brain may be omitted. But the fate of the an- terior portion of the cere- bro-spinal substance re- quires a brief further description. Of its two divisions (see the Table, p. Ill), the posterior (thalamen-cephalon), is at first a simple vesicle, formed of spin- dle-shaped cells with walls of nearly uniform thickness. Its floor gives rise to the optic chiasm and the optic nerves ; its sides become thickened into the optic thala- mi ; its roof gives rise to the pineal gland and surrounding structures. Fig. 60. — Head of the Embryo of a Sbeep, cut through the middle. Vi* (Kolliker.) «, under jaw; z, tongue; s, septum narium; occipitale oasilare; th, thalamus opticus; vt, roof of the third ventricle; cp, posterior commissure; mft, mid-brain divided by a fold into two parts ; /, falx cerebri; ff, terminal plate of the fore-brain. At the prolongation of the line of fm is the foramen of Monro, t, tentorium cerebelli ; cl, cerebellum ; p2, plexus of the fourth ventricle. occipitalis Fio. 61. — Brain of a Six-months Human Embryo. Natu- ral size. (Kolliker.) ol, olfactory bulb ; fs, fissure of Syl- vius; c, cerebellum; p, pons Varolii; /, flocculus; o, olive. DEVELOPMENT OF THE NERVOUS SYSTEM. 113 The larger portion of the development from the anterior primary vesicle constitutes the rudiments of the cerebral hemispheres. Here a floor and a roof must be distin- guished. The former develops into the striate bodies by the thickening of its walls. The latter forms the hemi- spheres proper. One principal characteristic of the mammals is the early enlargement of the cerebral hemispheres. In the human embryo they are even by the tenth week much larger than all the rest of the brain. They grow from before back- ward and thus cover up, one after the other, the optic thalami, corpora quadrigemina, and cerebellum. This physical evolution is indicative of the future intellectual superiority and intellectual growth of the human being. We have already seen how early and significant is the formation of the most important convolutions and fissures. By the end of the seventh, month of embryonic life, the principal features of the cerebral hemispheres are already definitely fixed. Development of the Eye. — Lateral growths of the brain- buds, called the "optic vesicles" (see p. 109), ^ j ^ j give rise to the ner- vous parts of the eye. ' These vesicles are c originally connected with the sides of the ^^^ 62. — Longitudinal Sections of the Eye of an a „. 1 1 „„„4„i„ Embryo, in three stages. (Erom Remak.) 1, com- nrSt cerebral vesicle mencement of the formation of the lens I, by depres- i_ T_ _j. J -J Bion of a part of ft, the corneous layer; «, r, the prim- by snort ana Wiae jUve ocular vesicle is doubled back on itself by the T, , •, .1 depression of the commencing lens. 2, the depression StalKS ; tney tnen for the lensls now enclosed, with the lens beginning ,- T . 1 . to be formed on the inner side; the optic vesicle is stand at nearly right more folded tack. S, a tUrd stage, in which the sec- , ondary optic vesicle g I begins to be formed. angles to the embryo. The stalks become narrow and form the rudiments of the optic nerves. At the same time the rudiments of the retina are formed from the vesicles themselves. 114 PHYSIOLOGICAL PSYCHOLOGY. The bulb of the optic vesicle is then made into a cup by- doubling it upon itself. The lens of the eye is formed by thickening some of the superficial epiblast and involuting it inward over the front of the optic cup (see Fig. 62). This involution has at first the form of a pit ; then of a closed sac with thick walls ; then of a solid mass. The subsequent development of the eye depends upon the fact that the walls of the optic cup grow more rapidly than does the lens, and that their growth does not take place equally in all portions of the cup. The different layers of the retina are formed by differen- tiations of the anterior wall of the" hind portions of the optic cup. Here the cells multiply rapidly, and undergo important changes while the wall is thickening. In its early development this wall resembles the brain in its structure. It may, indeed, be considered as a part of that organ. Development of the Ear. — The organ of hearing originally appears on either side of the hind-brain as an involution of the external epiblast, sunk in a mass of mesoblast. It is then shaped as a shallow pit with a wide-open mouth. As the mouth closes up, the pit becomes a vesicle, the otic vesicle. The walls thicken, and the cavity enlarges. Its shape becomes triangular, with the apex of the triangle directed inward and downward. This apex is elongated into the cochlear canal. Part of the vesicle is stretched out into a long narrow process (recessus vestibuW), and from the wall of the main body protuberances grow which become the vertical semicircular canals. Other protu- berances are stretched out and curved into other parts of the organ. The ^ cochlear canal is further elongated and curved. When it has reached two and a half coils, the thickened epithelium of its lower surface forms a double ridge, from which the Organ of Corti is developed. Histogenetio Development of the Nervous System. — All DEVELOPMENT OF THE NERVOUS SYSTEM. 115 the coarser differentiations of structure which have been described are but the expression, as it were, of secret and exceedingly minute changes, called " histogenetic." Deli- cate threads of nervous tissue have been laid down, and nerve-cells have been propagated (^proliferated') along defi- nite lines. The white matter of the spinal cord first appears in four patches at the front and back of either half. The individual fibres appear like small dots in these patches. The gray matter is formed by differentiation of the principal mass of- the walls of the medullary canal. The outer cells lose their epithelial character, and become converted into true nerve-cells, with nerve-fibres as prolongations. The nerve-fibres remain for a considerable time without the medullary sheath. The early histogenetic development of the brain back of the cerebral hemispheres is very like that of the spinal cord. In the floor a superficial layer of delicate nerve- threads is early formed. In the fore-brain the walls of the hemispheres become divided into two layers, the inner of which unites with the fibres of the crura cerebri to give rise to most of the white matter of the hemispheres. The outer layer of rounded cells becomes further differentiated, the deeper part, with its multiplication of numerous cells, forming the principal mass of the gray matter of the cortex. Thus does the impregnated ovum, by a process of evolu- tion, become developed into that wonderful complexity of organs which constitutes the body of the human child. Of the correlated psychical development of the embryo little or nothing of a scientific character can be known. But the physical process by which the nervous mechanism comes into being is, nevertheless, suggestive for the student of physiological psychology. To some of the conjectures and speculations which legitimately follow from our still meagre knowledge of human embryology, we shall return at another time. CHAPTER V. GENERAL PHYSIOLOGY OF THE NERVES. The question, What can Nervous Substance do? natu- rally follows the consideration of its chemical and formal constitution. An answer to this question, however, can- not he gained by inference from our knowledge of the anatomy and histology of the nervous system. It is only by indirect processes of observation and experiment, combined with no little conjecture, that even the beginnings of a clear scientific conception of the functions of this system can be found. The attainments of science on this subject may conveniently be stated under two heads, preceded by a brief introduction. We consider then, first, certain modes of activity belonging to all nervous matter as such ; and, second, — with more of detail, — the "Nerves as Conductors," and the "Automatic and Reflex Functions of the Central Organs." Functions of the Nervous Elements, — Nerves and nerve- cells have certain properties in common; within certain limits both can do the same things. These properties may perhaps be summed up in the two words Excitability (or "irritability") and Conductivity. Both are capable of becoming, under the action of stimuli, the subjects of a specific kind of molecular motion called " neural." When stimulated or irritated, they originate, as a unique func- tion, the process which may be called " nerve-commotion." But both nerve-fibres and nerve-cells can also propagate this peculiar kind of molecular agitation from point to point ; they can conduct nerve-commotion. 116 GENERAL PHYSIOLOGY OF THE NERVES. 117 Nerve-commotion is never, of course, an uncaused event. The causes that excite or irritate the nervous elements to exercise their peculiar function are called '■^stimuli." Stimuli are of two kinds, external and internal. The former comprise all such modes of energy as excite nerve- commotion by acting on the peripheral parts of the ner- vous system, — the terminal nerve-fibrils or end-organs of sense. Internal stimuli act directly upon the substance (nerve-cells) of the central organ ; they consist in general of changes in the blood-supply, — increased or decreased oxygen, presence of drugs, etc. General Office of the Nervous System. — In all forms of animal life, except the lowest, the action of the nervous system constitutes a chief characteristic of their difference from all forms of plant life. Plants as well as animals have contractile tissue ; but the former never have nervous tissue, not to say, a nervous system. The unique functions of this system, as possessed by all the higher animals, can perhaps best be summed up in the one word, "concate- nation." The linking, or chaining together — as it were — of distant and different physical organs and systems, and of the action of the other parts of the body with the phenomena of psychical life, is the unique function of the nervous mechanism. In the plant, for example, every part acts directly and slowly upon contiguous parts only, for the effecting of those changes upon which its life and growth depend. But in the case of the animal, by the mediation of the nervous system, an effect produced in one part of the body may make itself quickly felt in every other part. A draught of cold air, for example, strikes some portion of the surface of the body. Immediately, the heart and lungs modify their action ; the muscles contract ; the secre- tions are distm-bed; a shudder runs through the body; and perhaps the mind is seized with a vague feeling of 118 PHYSIOLOGICAL PSYCHOLOGY. fear. Thus changes which involve the tissues and func- tions of almost all the organs of the body are accomplished by the mediation of the nervous system. MORE PARTICULAR FUNCTIONS OF ALL NERVES. We have already seen (p. 32 f.) that the plan on vrhich the nervous system develops leads to a threefold economy of organs. In this threefold economy, the office of con- ducting the nerve-commotion between the end-organs and the central organs has been assigned especially to the nerves. The function of conducting neural molecular agitations belongs, indeed, as an essential function, to all nervous substance. But the nerve-fibres, as bound together into nerves, possess, in all normal conditions of the nervous system, this office pre-eminently. Moreover, it is only as exercised by the nerves that the laws of neural conduction can be at all satisfactorily examined by direct experiment. Physiological Distinctioiis in the Nerves. — If we considered only the different effects produced by conducting nervous processes along the different nerves, we should be com- pelled to divide the nerves into a variety of classes. In this way it has been proposed to distinguish " nerves of motion," " nerves of inhibition " (or check upon the action of other nerves), " nerves of secretion," " nerves of nutri- tion" (trophic nerves), "centripetal nerves that have no sensory function," and " sensory nerves," whose irritation may result in conscious sensation. The question arises at once, however, whether the differ- ent obvious results which follow irritating all these nerves are due to real differences in the functions of the nerves themselves, or to differences in the structures and connec- tions where the nerves terminate. We do not consider the electrical current which passes along different wires essen- tially different, because it may be used to write a message, light a jet, ring a bell, or cause the legs of a frog to twitch. GENERAL PHYSIOLOGY OF THE NERVES. 119 For reasons which need not be mentioned, it has been customary to reduce all possible classes of nerves to two, according to the direction in which they perform the ser- vice of conduction. Those nerves which conduct nerve- commotion outward from the nervous centres are called "efferent," or "centrifugal," or "motor." Those nerves which conduct nerve-commotion inward toward the ner- vous centres are called " afferent," or " centripetal," or "sensory." Afferent and Efferent Nerves. — The attempt has further been made to reduce the two foregoing kinds of nerves to one class. It is properly claimed that a difference in direc- tion does not necessarily prove an essential difference in function. In favor of such a difference, however, it has been urged that heat is a stimulus of afferent but not of efferent nerves ; and that a constant current passing along an efferent nerve, so long as there are no sudden changes in its strength, does not make the attached muscle con- tract, while such a current does seem to excite impulses in a sensory nerve. On the other hand, the rate of conduction in both kinds of nerves seems to be about the same ; and, indeed, most of the laws, to which we are about to call attention, apply, essentially unchanged, to both. It may further be urged that even more marked differences than those referred to above can be accounted for by the differences in the sources of the stimulation. A molecular disturbance, which would be quite powerless to stir the sluggish muscle-fibres when transmitted to them by a motor-nerve, might occasion pro- found changes in the sensitive ganglion-cells of the central organ when transmitted to this organ by a sensory nerve. Various attempts have been made, more or less success- fully, to demonstrate by experiment that motor and sen- sory nerves can be made to discharge each other's functions. Some experimenters have succeeded in uniting the central 120 PHYSIOLOGICAL PSYCHOLOGY. end of the sensory nerve of the dog's tongue with the peripheral end of the motor nerve, on the same side. Others have reversed the course of the nerve-fibres in the tail of a rat, by bending this appendage over, and planting its end in the back. If these experiments are not quite satisfactory, those of Kiihne may fairly be said to be con- clusive. He showed that if we divide the broad end of the sartorius muscle of the frog into a forked shape, the same stimulation will ascend the fibrils of one tine of the divided muscle, and descend the fibrils of the other tine. No good reason appears, then, why we should not con- sider all nerves as essentially alike in their powers of conduction. It is upon this assumption that the science of so-called " General Nerve-Physiology " is built up exper- imentally. In building up this science, the efferent nerves of frogs have been chiefly used for purposes of experiment. A preparation of such a nerve with a muscle attached is the subject whose behavior is investigated. The Nerve-Muscle Preparation. — The most convenient form of machine for experiment in "general nerve-phys- iology " requires a freshly dissected (gastrocnemius or other) muscle of a frog with the attached (sciatic or other) nerve. Such a preparation can be kept alive for some time in a cool moist chamber. By the simple contrivance of connect- ing the end of the muscle with a lever, arming the lever with some means of making a mark (pen, bristle, or needle), and bringing its point to bear on a travelling surface (plain or smoked paper, or glass), the time and amount of the contractions of the muscle maybe recorded. Stimulation may be accomplished with any kind of irritant, but for obvious reasons the electrical current is preferable as a rule. It may be applied under the greatest variety of conditions, and to any point in the nerve, and with any degree of intensity. The line traced by the armed end of the lever, as it rises GENERAL PHYSIOLOGY OE THE NERVES. 121 and falls with the contractions of the muscle, is called the " muscle-curve." This curve is a measure of the observ- able effect produced by irritating the nerve. If the elec- trical current flows with the course of the nerve toward the muscle, it is called " descending," or direct ; if it flows in the opposite direction, it is "ascending," or inverse. The movement of the muscle which follows closing of the current is called the "making contraction," or "closing contraction " ; that which follows its opening is called the " breaking contraction," or " opening contraction." When the single stimulations are repeated with sufBcient rapidity, the spasms fuse into one prolonged effort of the muscle, known as "tetanus," or "tetanic contraction." The nerve may then be said, with the muscle, to be " tetanized." CONDITIONS OF THE FUNCTIONS OF NERVES. Of the conditions under which alone the nerves are capable of exercising their functions the most important are the following three : — (1) Vitality of the Nerves. — ■ A nerve cannot act as the conductor of a nerve-commotion unless it is alive. The process of conduction is not therefore merely mechanical, like that of electricity along a wire, but is physiological and vital. The death of the nerve is not, however, simul- taneous with that of the body from which it is taken, or of the muscle to which it is attached. On the contrary, by careful treatment, it may be preserved alive for some time after excision. Since the nerve, unlike the muscle, has no death-rigor, it is difficult to say precisely when it is dead. The existence of electrical phenomena in the nerve for some time after it has ceased to excite the muscle is thought by some authorities to show its continued vitality. Nerves, when dying, exhibit two marked changes of excitability. Immediately after being cut, the excitability of the nerve increases, and afterward sinks to zero by sue- 122 PHYSIOLOGICAL PSYCHOLOGY. cessive stages of diminution. The course of these changes is different for different portions of its length. Again, the lower portion of the cut nerve seems to preserve a given degree of vitality for the longest time. Hence "Valli's principle: Nerves die from the centre to the periphery. Closely allied to the foregoing changes are those which take place when the cut nerve remains in its place in the living animal organization (in situ'). For such a nerve the law of increased irritability immediately after section seems, in most cases, to hold good ; the application can be tested, however, only in the case of motor nerves. Nerves, cut in situ, lose their vitality after a time — in warm- blooded animals, of three or four days, but in cold-blooded, of a week or more. A fatty or granular degeneration (discovered by Waller in 1850) takes place in nerve-fibres that are severed from the central organ ; and this degener- ation proceeds from the place of section to the extreme peripheral portion of the fibre. A cut nerve remaining in situ may be regenerated by the axis-cylinders growing out of the central portion and running into, and between, the sheaths of Schwann of the peripheral portion. According to some authorities the conductivity of the nerve is then regained earlier than its power of local irritability. (2) JJse of Oxygen by the Nerves. — As compared with the end-organs and the central organs, or even with the muscles, the nerves are relatively independent of the pres- ence of oxygen for the exercise of their physiological func- tion. The irritability of the nerves continues almost as long in a moist vacuum, or in indifferent gases, as in the air. It may be argued, however, from the marked depend- ence of nervous tissue, in general, upon a supply of arte- rial blood, as well as from the general mechanical theory of the nervous system, that some oxygen is an essential condition of the activity of the nerves. (3) Recovery from Exhaustion. — The nerves, when " ex- 6ENEBAL PHYSIOLOGY OF THE NEEVES. 123 hausted " — as it is said — cannot perform their physiologi- cal functions. But exhaustion of the nerves is difficult to distinguish from exhaustion of the central organs or of the end-organs. In the case of the nerve-muscle machine Bernstein thinks he has shown that by far the greater part of the effects of prolonged and severe stimulation is due to the mwscZe-element in the machine; and that exhaus- tion in the nerve comes on much more slowly than in the muscle. Indeed, some have gone so far as to hold that the nerve is not exhausted at all, but resembles in this regard a metallic wire. But more recent researches seem to show — as indeed we should expect on grounds of general theory — that a prolonged tetanizing current may fatigue the nerve, even when the end-apparatus continues able to perform its functions. Ordinarily, however, when we are tired nervously, it is the central organs, or end-organs, rather than the conducting nerves, that are tired. PHYSICAL PROPERTIES OF THE NERVES. The phenomena called forth by irritating a nerve depend upon the character, amount, and method and place of application, of the stimuli employed. This dependence suggests certain truths as to the physical properties of the nerves as conductors. Mechanical Properties of the Nerves. — The elasticity, ductility, cohesion, etc., of nerves are of little interest to the student of physiological psychology. More pertinent is the fact that all kinds of mechanical attacks upon the nerves excite them, and are followed by pain in case of the sensory nerves, and by contraction in the case of motor nerves. Tetanus may be produced with a toothed wheel or hammer. A certain suddenness of the shock seems necessary to excite the nerve. Pressure on a nerve may be gradually increased until its power of conductivity is lost, without exciting it. Slight pressure or traction 124 PHYSIOLOGICAL PSYCHOLOGY. seems to increase the irritability and speed in conduction of the nerve. Yet nerves may be cut so suddenly as not to excite them. Thermic Properties of the Nerves. — Little is known as to the specific heat of nerves, or as to their power to conduct heat. But the effect of heat on the function of these organs is very marked. The results of experiment differ as to the degree of heat which will act as a stimulus upon the nerves. Considerable changes in the medium temperatures appear to have no effect. One investigator found that suddenly warming a nerve to about 35°- 40° C. occasioned a spasm in the attached muscle ; and warming to a higher degree produced tetanic convulsions. Long ago that noted authority, E. H. Weber, showed that heat and cold do not produce sensations when applied directly to the sensory nerve-trunks in man. For this peculiar sensory effect the intervention of end-organs seems necessary. Four periods have recently ^ been distinguished in the effects produced by heat upon the irritability of the motor nerve. These are thus described: (^) Gradual increase to a maximum of irritability, — viz. 32.75°-39.25° C. ; (^) then gradual diminution of irritability to its total loss ; (C) condition of no irritability, a period coming at any point within 5° above the temperature of maximum con- traction ; and finally (i)) development of heat-rigor. Warmth increases the immediate expenditure of energy in an excised nerve, and so hastens its death ; cold delays this expenditure and so conserves the nerve. Chemical Properties of the Nerves. — The effect of most chemical agents on the nerve is to destroy without exciting it. Changes of the amount of water in the substance of the nerve affect its functional activity. A slight drying raises its irritability; and drying also produces contrac- 1 By Charles L. Edwards, in Johns Hopkins Studies from the Bio- logical Laboratory, June, 1887. GENERAL PHYSIOLOGY OP THE NEKVES. 125 tions ending in tetanus. Swelling the nerve decreases its irritability to the point of entire loss. Certain acid and alkaline solutions also affect the nerve very much like drying it. Some organic substances, like \irea, sugar, and glycerine, irritate the nerve. The principle seems to be, that all chemical stimulation of the nerves is closely con- nected with the destruction of the nervous tissue. Electrical Properties of the Nerves. — The resistance of living nerves to the electrical current is probably about the same as that of the muscles ; it has been given at 50,000,000 times that of copper wire. The conductivity of the nerve has also been given as, on the average, 14.86 times that of distilled water. The excitatory effect of the constant current upon the nerves follows the principle stated by that great explorer, du Bois-Reymond, in 1845. This effect, as measured by the contraction-curve of the muscle, does not correspond to the absolute value of the intensity of the current at each moment, but to the change in this value from moment to moment ; and the effect is greater the less the time in which changes of the same magnitude in the current occur, or the greater their magnitude in the same length of time. The essential fact is that constant currents, while they remain constant, do not irritate the nerve; variations in theSe currents do irritate it. Even upon the sensory nerves it is not certain that the constant current itself, apart from changes in its strength, can have much excitatory effect. The sensory experiences which follow such a current are chiefly due to changes induced in the end-organs and central organs. Excitatory Effect of the Constant Current. — If we experi- ment with the electrical current upon a nerve we find that its excitatory effect is dependent upon the direction in which the current flows. The following table, by Pfliiger, gives the results reached by a large number of observers : — 126 PHYSIOLOGICAL PSYCHOLOGY. Strength or Ascending Cukkent, Descending Cubkent. COBRBNT. Making. Breaking. Making. Breaking. Weak .... Medium. . . Strong. . . . Contraction Contraction Rest Rest Contraction Contraction Contraction Contraction Contraction Rest. Contraction. Rest or weak Contraction. The most important point in this table is more clearly brought out by the following recent summary ^ of results : " Up to a certain strength of current a stimulus will give contraction when the cathode lies next to the muscle (i.e. the current is descending), which will give no contraction when the anode is in that position (current ascending). Above this strength the reverse holds, and a stimulus which is followed by contraction when the excitation has to pass the anode, evokes no response when it has to pass the cathode." The excitatory effect of the electrical ctirrent upon the nerve is also dependent upon the strength of the current. It increases with the strength, from the lowest observable point, until it soon reaches a maximum ; after this, further increase of the effect of the current is to be recognized only by the expanding of this condition of higher irrita- bility — called " electrotonus " — over the extra-polar parts of the nerve. The effect of the current upon the nerve also depends upon the length of nerve excited, and upon the angle at which the stimulus is applied. Up to a certain limit — fixed by different investigators at from t^ to I- inch — the excitatory effect increases with the length of the nerve through which the current flows. The electrical current 1 An Article by 6. N. Stewart, in Journal of Physiology, Oct., 1889. GENERAL PHYSIOLOGY OF THE NERVES. 127 apparently does not excite the nerve at all when it flows through it precisely at right angles to the nerve's axis. The duration of the current also influences its effect as a stimulus. It would appear that the current must act upon the nerve for at least about 0.001|- of a second in order to excite it. By cooling the nerve this " sluggish " period may be increased to nearly 0.02 of a second. In ordinary circumstances, however, it is thought that the action of the stimulus for 0.017-0.018 second will cause the muscles to contract as much as the same strength of current when constantly applied. Electrotonus of the Nerves. — When the nerve of a nerve- muscle machine is under the influence of a constant cur- rent of electricity, very important changes in its condition are observed, as respects both its excitability and its con- ductivity. The general fact that such changes are pro- duced as the effect of applying the constant current is undoubted. But the precise nature of some of these changes is disputed ; while no theory has as yet been de- vised which will satisfactorily account for them all. We shall confine our account at present to the briefest possible statement of the more important alleged facts ; any refer- ence to theory which seems desirable at all, will be made in another connection. The changed condition of a nerve, as respects its phys- iological function, which is produced in it by a constant electrical current, is called "Electrotonus." "Pfliiger's law," so-called, states the case, in general, as follows : The excitability of a nerve under the action of the constant cur- rent is increased in the catelectrotonized region (that is, on both sides of the cathode, or negative electrode, — the point where the current leaves the nerve), and diminished in the anelectrotonized region (that is, on both sides of the anode, or positive electrode, — the point where the current enters the nerve). This law is said, by a chief modern 128 PHYSIOLOGICAL PSYCHOLOGY. authority, to hold good of all kinds of stimulus, and in all cases. The electrotonic effect of the constant current upon the nerve, like its direct excitatory effect, is influenced by the strength of the current, by its direction, its making and breaking, and by the length of the nerve through which the current flows. The changes called " electrotonic " occur in the region of the negative pole (cathode) imme- diately upon making the current; they then quickly but slightly increase and afterwards fall off more slowly again. In the region of the positive pole (anode) the changed condition develops more slowly until it reaches a maxi- mum, and then gradually diminishes. Recent researches have led some investigators to hold that the conductivity of the nerve is changed by the con- stant current in a different manner from its excitability. In its electrotonic condition — that is to say — the conduc- tivity of the nerve is found to be less around the cath- ode than around the anode. From this the conclusion is drawn that the origin of the process of excitation of the nerve is not like its propagation. Into the refinements of this change, when the stimulus of the electrical current is applied to different parts of the nerve outside, or within, the poles (inter-polar and extra-polar) we cannot enter. PROCESSES EVOKED IN CONNEOTIOHr WITH THE FUNCTION OF NERVES. When the nerve is excited certain processes connected with its physiological action are indicated, in a more or less obvious way. Mechanical Processes in Excited Nerves. — No appreciable mechanical changes, like the contraction of the muscle- fibre, can be detected in excited nerves. Whatever changes occur are invisible and impalpable. In the nerve-cells, however, mechanical changes can be detected as the result GENERAL PHYSIOLOGY OF THE NERVES. 129 of excitation. Repeated and prolonged excitation of the ganglion-cells of the posterior root results in shrinkage of their nucleus and of the cell-protoplasm 5 and in chang- ing the nucleus from a smooth and regular to a jagged and irregular outline. The large cells show most of this effect of being obliged to do work; the small cells little or none at all. The average shrinkage of the large cells is found to be measurable as — in some cases — from 24 to 36 per cent.^ Thermic Processes in Excited Nerves. — If any rise of tem- perature is produced in a nerve by irritating it, the amount is exceedingly small. Helmholtz concluded that his means of detecting heat to within a few thousandths of a degree showed no such change in excited nerves. On the other hand, nervous excitation appears to produce a perceptible change of temperature in the centres of the brain; and this change can scarcely be due wholly to increased flow of arterial blood. But to this subject we shall return in another connection. Chemical Processes in Excited Nerves. — The most obvious indications that chemical processes are concerned in the physiological functions of excited nerves are certain changes in "reaction," or in taking stains, which some observers claim to have found. It has been asserted that, after extreme exertion caused by cramping in cases of strychnine-poisoning, the nerves have an acid reaction. Very recently two observers, on comparing two frogs, one of which was killed after resting and the other after having the eighth nerve stimulated for an, hour, found several per cent, more nuclei staining red in the stimulated than in the rested pair of nerves. Evidences of marked chemical changes in the nervous centres, due to work done there, are of course not wanting. But the direct experimental evidence for the same thing in the nerves is still incomplete. 1 See the American Journal of Psychology, May, 1888, pp. 479 ft. 130 PHYSIOLOGICAL PSYCHOLOGY. Electrical Processes in Excited Nerves. — In 1843 du Bois- Reymond found what he considered direct experimental proof of the existence of electrical currents in the nerves. It had, of course, been previously conjectured that nerve- commotion is a phase of electricity. But this experi- menter discovered that, if we cut a nerve and then apply an electrometer to it, the cross-section is negative toward the longitudinal surface of the nerve. The current, which is thus shown to be flowing in the nerve, from the cut end to the equator, is called " natural nerve-current," or " current of rest." Its electro-motive force is greater, the larger and thicker the nerve. In the sciatic nerve of a frog it is given at from 0.022 to 0.046 of a Daniell's cell. It continues for some time after the irritability of the nerve is lost. The same investigator found that the " current of rest " is diminished in energy by tetanizing the nerve. This swing of the needle which measures the " current of rest," back- ward toward zero when the nerve is repeatedly irritated by passing through it an interrupted current, is called " nega- tive variation." It shows that the electro-motive force of the nerve is diminished by the nerve being excited. The bearing of these phenomena also upon a general mechanical theory of the nervous system will be referred to in other connections. LAWS OF CONDUCTION IN THE NERVES. Only a very few statements can be made, with respect to the physiological function of the nerves as conductors, which are properly entitled to the dignity of being called " laws. " And these laws are determined almost wholly by experiments with the motor nerves of frogs. There is evi- dence, however, in respect to certain of the more simple forms of activity, that all nerves conduct nerve-commotions in essentially the same way. Relations of Magnitude between Stimulus and Besult. — On GENERAL PHYSIOLOGY OF THE NERVES. 131 attempting accurately to compare the amount of the stim- ulus applied to the nerves with the amount of resulting nervous impulse, great difficulties are encountered. There is indeed no absolute measure for either of the values which it is desired to compare. Electricity is the only stimulus of the nerves that admits of a fairly approximate measure- ment by objective standards. The effect produced by stimulation is almost wholly manifested in organs with which the nerve is connected, rather than in the nerve itself. It does not, therefore, admit of easy direct measurement. Measuring the result of the stimulus in the nerve by the amoimt of contraction produced in the connected muscle, we find it to be (as has already been indicated, see p. 126) within certain limits, directly proportional to the amount of the stimulus. Two remarkable apparent exceptions to this law are noted : (1) On increasing the amount of the stimulus beyond the point necessary to produce the first maximum contraction, another stage is reached in which the effect further increases, in proportion to the stimulus, until a second maximum is gained. (2) In some circum- stances, after reaching the first maximum, the effect dimin- ishes with the increase of the stimulus, then rises on further increase until the second maximum is reached. The excitability of the different nerves is different, and of different localities of the same nerve under different circumstances. It is usually greater in winter than in summer. In the cut nerve it is greater near the cross- section. The reflex effects of stimulating a sensory nerve are said to be greater the nearer the central organ the stimulus is applied. The lower part of a nerve is found more excitable under the ascending, the upper under the descending, induction-current. Stmunation of Stimulations in the Nerves. — In order to keep the successive waves of nerve-commotion apart, an 132 PHYSIOLOGICAL PSYCHOLOGY. interval of about j^ second must elapse between the re- peated stimulations. Otherwise they fuse, and tetanus results. If tliis interval is observed, the combined effects of the different stimulations may be piled, or " summed " up, in the nerve. They may then be seen in superimposed contractions of the muscle. This law is also important for a mechanical theory of the nervous system. Speed of Nervous Impulses. — In 1844 the great physiolo- gist Miiller declared it forever, impossible to know the speed of the nervous impulses. In 1850 Helmholtz an- nounced the speed as from 26.4 meters (86.6 feet) to 27.25 meters in motor nerves. Subsequent researches have, in the main, confirmed the conclusion of Helmholtz. The rate of nervous impulses varies greatly, however, under different circumstances. By changes in temperature results can be obtained in the motor nerves of man, varying from 98 feet to 295 feet per second. The general conclu- sions for the sensory nerves favor numbers lying between 98 and 131 feet per second. As soon as the strength of the stimulus rises above a certain limit, the speed of the resulting impulses appears to increase with the strength of the stimulus. In the spinal cord and in the brain the speed of the ner- vous impulses is, in general, much slower than in the peripheral nerves. This is due to the far greater com- plexity of these organs, and to the accompanying possibility of the impulses spreading into side tracts, as it were : in brief, the greater variety and amount of the work that must be done. Exner calculated the speed of motor impulses in the cord at 11 to 15 meters (about 36 to 49 feet). The sensory impulses of the cord, he thought, travel at the average rate of about 8 meters (about 26^ feet). Sensations of touch arise, as we all know by expe- rience, later than sensations of pain, when we are struck with a missile, or burned with a brand. Some have main- GENERAL PHYSIOLOGY OF THE NERVES. 133 tained, therefore, that the speed of the former is to that of the latter as from 27 to 50 meters compared with 8 to 14. The argument is not conclusive, however, since we do not know the length of the paths by which the impulses that produce the two kinds of feeling travel, or the kind and amount of cerebral action which they respectively involve. Integrity of the Nerves Necessary. — The slightest separa- tion of the parts of a nerve, even if its cut ends are left in the closest mechanical contact, destroys its power to conduct nervous impulses. Nerve-commotion, unlike the electrical current, cannot jump the smallest gap in the ner- vous structure. The ancients knew that tying a nerve prevents its action. They explained the fact by saying that the flow of nervous fluid was thus hindered. So also does the fineness of the localization which belongs to the organs of motion, and especially to the organs of special senses, like the skin and eye, indicate the physiological isolation of the nerve-fibre during its course between the end-organs and the central organs. It would further seem that the law of the " specific energy " of each nervous ele- ment is connected with the assumption necessary to explain the phenomena attendant upon the starting and propagat- ing of nervous impulses in the conducting nerves. But to this subject of the " specific energy " of the nervous elements we shall recur at another time. When speaking of conduction in the nervous substance of the spinal cord or of the brain, we are not to think of the nerve-commotion as moving along one fixed path, after the exact analogy of the far simpler case of the nerve-muscle machine. The spinal cord does not act as a "perfectly isomorphic medium " for the transmission of nervous impulses. Its extremely complex structure has shown us that it is not adapted to act as such a medium. The case of the brain in this regard is even more complicated. After all the thousands of experiments which have been 134 PHYSIOLOGICAL PSYCHOLOGY. performed in what is called "nerve-physiology," we are not yet in a position even to indicate with scientific exacts ness a complete mechanical theory of those molecular wave-like disturbances which the application of stimuli produces in a single nerve attached to a muscle. How much less then do we know of the molecular science of the nerve-commotions in the cord and in the brain? Yet that the nerve-commotions are molecular wave-like changes there can be no doubt. And these changes are connected with, but are not identical with, those mechanical, ther- mic, chemical, and above all, electrical processes, which have just been described. CHAPTER VI. REFLEX AND AUTOMATIC NERVOUS FUNCTIONS. When a physiological function is occasioned in some peripheral nerve, independently of a so-called act of will, by the stimulation of some other peripheral nerve, this function is said to be "reflex." In other words, reflex action is the result of the secondary stimulation of one nerve, through a central organ, by the primary stimulation of some other nerve. On the other hand, all excitations of the nervous system which originate in the nervous centres themselves are called "automatic." Doubtless this term must often be used to conceal our ignorance of the real origin of a neural process ; doubtless also, many processes, which a't first sight appear to be automatic, are afterwards discovered to originate reflexly. Notwith- standing this, nervous impulses which result in the move- ments of the muscles, or in the inhibition of such move- ments, apparently originate in the central organs under the action of internal stimuli. But such automatic activi- ties belong distinctively to the central ganglia of the brain. Kinds of Reflex Action. — Theoretically, various kinds of reflex nervous action are supposable in the nervous system. Thus two motor nerves might be combined through a central organ (" co-motor reflexes ") ; or an excitation arising in a motor nerve might, without an act of will, be transferred to one or more sensory paths ("reflex-sen- sory"). As to the existence of "co-sensory reflexes" — or cases where the excitation of one peripheral sensory nerve, 135 136 PHYSIOLOGICAL PSYCHOLOGY. through the central organ, occasions the excitation of another locally different, peripheral, and sensory nerve — there can be little doubt. The nose, for example, may be made to tickle by looking at the sun ; and strong rubbing or squeezing of one muscle may sometimes occasion pain in muscles located far away on the surface of the body. But only one kind of reflex functions of the nervous system seems hitherto to have been made available for purposes of scientific experiment. These are the so-called "reflex- motor," or "sensory-motor." Such terms are given to reflex action when a motor nerve is stimulated in a secondary way, through a central organ, by applying stimulus to sensory nerve-endings. We must carefully guard ourselves, however, from the misconception that lurks in the word "reflex." The effect of the central organ is never that of simply turning back, or reflecting, a nerve-commotion from one path to another. On the contrary, the passage of a nerve-commotion through a central organ profoundly modifies both the condition of the organ and the character and distribution of the nerve- commotion itself. THE SPINAL CORD AS A CENTRE OF REFLEX MOTOR ACTIVITIES. Our previous survey of the structure of the spinal cord suggests the truth that it is specially adapted to act as a central organ in the exercise of a variety of reflex-motor functions. The older investigators assumed a great variety of special mechanisms, consisting of distinct systems of sensory and motor nerve-fibres, with connecting cells, appro- priated for the sole purpose of executing the various kinds of reflexes. Modern investigation tends rather toward the view that there are no special elements of the cord appro- priated merely to reflex-motor functions. The whole structure of the organ is such as to adapt it in all its parts, for this as well as for other nervous activities. KEFLEX AND AUTOMATIC NEKVOUS FUNCTIONS. 137 The Eeflex-Hotor Machine. — The most convenient piece of mechanism for experimenting to discover the laws of spinal reflexes is the so-called " brainless frog." This preparation consists of the spinal cord, still alive, but sep- arated from the brain by section below the medulla oblon- gata. If the flank of such a frog be touched, a slight twitching of the muscles, which lie immediately below the spot of the skin thus stimulated, will result as a reflex motion. If a hind leg be stretched out and pinched it will respond in a purposeful way to withdraw itself from the irritation. Increasing the strength of the stimulus will elicit reflex motions involving the fore leg of the same side ; then both legs of the other side ; and perhaps, finally, all the muscles of the body. If when one leg of a brain- less frog is irritated it is at the same time prevented from movement, the cord of the preparation will sometimes use the other leg to accomplish the purpose of removing the irritation. Phenomena, similar to those obtained from the brainless frog, may be obtained from other brainless animals. A decapitated salamander, when the skin of one of its sides is pinched, will bend that side into a concave shape. It was for some time supposed impossible to obtain simi- lar phenomena from the spinal cord of mammals. And, indeed, the spinal reflexes in the case of a mammal whose cord has been disconnected from his brain, are, immedi- ately after section, comparatively weak and purposeless. But if such an animal be kept alive for some time, then strong, varied, and purposeful movements will follow sensory stimulation of the skin of the parts below the place of section. After some weeks or months, reactions resembling those described in the case of the frog begin to appear. Moreover — a very significant fact ! — it is found that the breed, sex, age, and training of the animal deter- mine the character of these reflex brainless movements. 138 PHYSIOLOGICAL PSYCHOLOGY. Strength and Suddenness of Spinal Eeflexes. — Continuous irritation of the skin of a brainless animal, if slowly ivr creased, does not give rise to reflex movements. But a much smaller degree of stimulus, if suddenly applied, will call forth such movements. Repetition of the shocks with the electrical current is much more effective than is the constant current, in starting spinal reflexes. And here we are reminded again of the law already announced, as holding good in " general nerve-physiology " (see p. 125). A decapitated frog may be placed in water, and the water slowly heated until its life is destroyed, without its show- ing any reflex activity. Even normal frogs — though with much more difficulty — can be heated, either locally, vnth one leg in the water, or all over, while sitting on a cork in a cylinder of water, without causing motion, if the increase of temperature be gradual enough. Speed of Spinal Reflexes. — It has already been said (p. 132) that the process of conduction suffers a delay while passing along the spinal cord. The time of " cross-con- duction" in the cord also seems to depend upon the strength of the stimulus. By increasing its strength — it has been calculated — the time consumed by the spe- cifically central processes may be diminished from 0.055 to 0.047 of a second. With very strong stimulus the time occupied by the central processes becomes almost too brief to detect. Condition of the Spinal Cord. — The character of the resulting reflex movement is very largely determined by the condition of the cord when the sensory impulses enter it. Lesion increases the excitability of the part below the lesion. Some drugs heighten and some depress its excit- ability. When the cord is poisoned with strychnine, for example, the slightest stimulation will cause such an explosion in it, and such a diffusion of energy along unaccustomed paths, that convulsive cramping is occa- KEFLEX AND AUTOMATIC NERVOUS FUNCTIONS. 139 sioned over the entire body. Changes of temperature also seem to affect the reflex-motor activities called forth by stimulating the spinal cord. Effect of Locality. — The extent and character of the spinal reflexes depend, in a remarkable way, upon the locality to which the stimulus is applied. The most remarkable difference is perhaps that evoked by irritating some spot on the skin of the brainless animal, and then comparing the result with that obtained by applying the stimulus directly to the trunk of the sensory nerve. A slight irritation of the skin may result in the extended movement of many muscles, combined in a purposeful way. The direct stimulation of the trunk calls forth irregular movements in a few muscles only. What par- ticular reflex actions follow stimulation, depends upon the particular locality of the skin to which the stimulus is applied. Laws of Spinal Reflexes. — The most general rule of the reflex-motor activities of the spinal cord may be stated in terms like the following : The irritation of a sensory nerve with a small degree of stimulus gives rise to reflex move- ments which originate in the cord on the same side, at about the same altitude as that at which the sensory impulses enter the cord; increased stimulus gives rise, also, to movements that arise on the other side of the cord, at about the same altitude ; a still greater increase, to those that originate on both sides of the cord, with the prefer- ence, however, to the same side. It would seem, then, that the nerve-commotion which enters the cord is dispersed, first, along the network of cells and fibres near the point of entrance on the same side ; then, across, at the same altitude, to the other side; then, up and down on both sides of the cord. Excitation, started anywhere in the cord, tends to radiate in all directions, but with the preference for certain paths 140 PHYSIOLOGICAL PSYCHOLOGY. marked out by the structure and habits of the cord. Hence the spinal cord has been called " the organ for the dispersion of irritation." Alleged Automatic Functions of the Spinal Cord. — This organ is not capable of " irregular automatism," — that is, of such spontaneous excitation as takes place in the higher centres of the brain, on volition. It does, however, dis- charge certain functions that are less certainly reflex than those which have already been considered. What is called the "tonic action" of the cord upon the muscles is a marked instance of such functions. The fact that this action does not simultaneously contract all the muscles connected with the cord, or any one set of them with the same energy as any other, throws some suspicion on its automatic character. Moreover, we can often ascribe the " tone " of the muscles of the brainless animal to the action of subtle influences, such as movements of the air, etc., upon the surface of the skin. If a brainless frog be hung up, after having the sciatic plexus cut on one side, the muscles of the leg on the other side have the better " tone." But the same flaccid condition of the muscles on the cut side exists when only the sensory roots of this plexus are cut. The circulation of the blood in a brainless animal seems to be in a measure dependent upon the condition of the spinal cord. Hence it is claimed that so-called " vaso- motor centres " exist in the cord. Circulation may continue with regularity in a decapitated frog ; but the removal of any considerable part of the cord aff