■1 t* .v- *:.->. ■■MOB - t ■ • : it ■ KiT^i '*?-'/ - ■ H J LIGAMENTS THEIR NATURE AND MORPHOLOGY LIGAMENTS THEIR NATURE AND MORPHOLOGY BY JOHN BLANLVSUTTON, F.R.C.S. LECTURER ON COMPARATIVE ANATOMY, SENIOR DEMONSTRATOR OF ANATOMY, AND ASSISTANT SURGEON TO THE MIDDLESEX HOSPITAL; ERASMUS WILSON LECTURER, ROYAL COLLEGE OF SURGEONS, ENGLAND. LONDON H. K. LEWIS, 136 GOWER STREET, W.C. 18S7 PRINTED BT ■ . K. LEWIS. 136 GOWEK STRELT, LONDON, W.C. PREFACE. Five years ago. whilst examining" a partially dissected Ostrich, my attention was arrested by the remarkable tendon-like appearance of the Ligamentum Teres in the hip joint. Closer examination induced me to believe that the ligament was connected with muscles at the margin of the acetabulum. An investigation of the hip joint in a Horse and the curious New Zealand Lizard Hatteria (Sphenodon), lead me to regard the ligament in man as the divorced tendon of the pectineus muscle. Later I found muscle and ligament directly continuous in an Ostrich chick. The result was somewhat startling and incited me to trace out the ancestry of various ligaments in the body, as time and opportunity permitted. The results of the investigation were made known in a series of papers published in the Journal of Anatomy and Physiology, vol. xvii. (1883), vols, xviii., xix., and xx. (1886), under the title "The Nature of Ligaments." In this present treatise an attempt is made to write a systematic account of ligaments and fasciae generally, taking care to avoid tedious descriptions, but being con- tent to confine attention to morphology and ancestral history. It has been necessary to appeal again and again to Comparative Anatomy, and the student will feel, vi PREFACE. in studying even ligaments, the truth of Bacon's remark that, "No natural phenomenon can be studied in itself alone, but, to be understood, must be considered as it stands connected with all nature." Lastly, facts revealed by one series of dissections have led me to anticipate the existence of conditions in animals which I had not previously dissected. This recalled to my mind, with redoubled force, Miiller's statement, "Com- parative Anatomy in its complete form leads to such necessary consequences, that expressions may be found for organizations, which are like the expressions of an equation. If these expressions are found, the unknown quantities may, in a given case, be reckoned from the known." In order to meet the wants of the student of human anatomy, I have avoided overburdening the general account with facts purely comparative. These have for convenience been concentrated in one chapter. To my talented friends, Messrs. E. B. Osmond and W. E. Wynter, is due all the credit of the drawings from which the illustrations were taken. Mr. Osmond, in addition, transferred the drawings on to the blocks; I owe him more thanks than can be well expressed. My friend Mr. Daniel Thurston kindly assisted me in correct- ing for the press. 22 Gordon Street, London. January, 1887. CONTENTS. PAGE Introduction ........... i CHAPTER I. Regression of Muscles in its relation to Ligaments .... 8 CHAPTER II. Examples of Metamorphosis and Regression of Muscles from the Lower Vertebrates ......... 25 CHAPTER III. Migration of Muscles in relation to Ligaments . . . -35 CHAPTER IV. The Knee Joint and Ankle ........ 42 CHAPTER V. The Ligaments of the Pectoral and Pelvic Girdles . . . -58 CHAPTER VI. Ligaments in connection with the Appendicular Elements of the Skull and Vertebral Column ....... 75 CHAPTER VII. Ligaments of the Vertebral Column (continued) . . . .86 CHAPTER VIII. The Manus ........... 93 An Appendix containing a list of Mammals with a well developed Gleno-Humeral Ligament. ....... 100 Indix 105 LIST OF ILLUSTRATIONS. teres of FIG. i. Humerus of a foetal lion 2. The supra-condyloid process in man .... 3. The scapula of a two-toed sloth (Choloepus hoffmanni) 4. The epitrochleo-anconeus muscle of man . 5. The epitrochleo-anconeal fascia .... 6. The epitrochleo-anconeus muscle of an opossum 7. 8, 9. The variations of the human coraco-brachialis muscle 10. The lumbar fascia .... 11. The forelimb of a porpoise . 12. The bicipital loop in the thigh of a fowl 13. The bicipital loop in a Rhea 14. The upper end of the femur of an ostrich with ligamentum 15. The femoro-caudal muscle of Iguana tuberculata 16. Vertical section of the knee joint of a human embryo 17. The inferior anterior annular ligament of the ankle joint monkey ......... 18. The anterior annular ligaments of the ankle joint of a bird ig. The leg of a chick at the fifth day of incubation 20. The leg of a chick at the eighth day of incubation 21. The tibio-tarsus of an ostrich chick 22. A general view of the ligaments of the pectoral arch 23. The shoulder girdle of an ostrich chick , 24. The shoulder girdle of Ornithorhynchus 25. The gleno-humeral ligament of a beaver 26. The shoulder girdle of a bird with the subclavius muscle 27. The pelvis of a sloth 28. The human chondro-cranium and appendages 29. The transformation of the appendages 30. The conjugal ligaments of a seal 31. The conjugal ligaments of a human foetus 32. The constitution of an intervertebral disc . 33. A portion of the vertebral column of a skate 34. The intervertebral disc between the odontoid process and the body of the axis 35. The subnotochordal rod of a tadpole 36. The manus of the water tortoise 37. The carpus of a baboon .... 38. Rudimentary centrale of the embryonic carpus 39. Human scaphoids with fused centrals PAOE 5 5 6 10 10 10 12 21 26 29 29 40 44 47 49 50 5i 52 54 59 60 63 66 67 73 76 76 78 79 79 81 88 9i 94 94 96 96 LIGAMENTS THEIR NATURE AND MORPHOLOGY. INTRODUCTION. As our mother earth receives alike the remains of kings, philosophers and peasants, so Nature buries in fibrous tissue, structures which were in histological rank far superior to the ligamentous tissue which now hides them, so disguising their former nature and true significance that they become reduced to the level of structures whose duty- is menial indeed. Morphological problems of the greatest interest may centre around a single band of fibrous tissue, whilst in the immediate neighbourhood, ligaments abound of functional value, but of no interest to the morphologist. To sort out from among the enormous heap of structures known as ligaments, those which can boast a noble de- scent, and trace the history of their downfall is the object of the present treatise. Attention will also be devoted to those tracts of fibrous and tendinous tissues known as apo- neuroses. Joints and ligaments have long been a dry study with students, but the following pages will serve to shew that some of the most important processes of evolution may be inculcated by an earnest study of these apparently unin- teresting bands. The more important ligaments of the body are derived either from the metamorphosis and re- gression of muscles, or the degeneration of osseous and cartilaginous tissues. Each will therefore be separately considered. B LIGAMEXTS. [Introduction. Metamorphosis of Muscle. — Any one dissecting- an amphibian or reptile for the first time, must be impressed with the small amount of tendon entering into the composi- tion of the muscles. This becomes more marked when the muscular system of an amphibian is compared with one of the higher mammals. Muscle and tendon have too long- been regarded as distinct tissues ; the arrangement of the fibres in tendon strongly recalls those of the fasciculi in the belly of a muscle. It is very difficult to determine how the muscle and tendon are joined, or by what means the union is brought about, for the parts become insensibly blended. The position of tendons has an important bearing on this question : — As a rule they are situated at the extremities of muscles where contractile tissue has little opportunity of exerting itself advantageously, e.g., the long tendon of the bi- ceps at the shoulder, the elongated tendons of the flexors and extensors of the fingers and toes. When a tendon develops in the centre of a muscle, as in the digastricus of man, it is usually in a situation where muscular tissue would be of little avail. Again, when a muscle is so circumstanced that its contractile power can be brought into play throughout its whole length, it may remain muscular in structure from origin to termination, as in the case of the intrinsic and extrinsic muscles of the tongue. If either or both extremi- ties of a muscle fail to act to full advantage, the reason for the existence of such a tissue fades, and the ends degen- erate into tendons to play an important part nevertheless as passive agents. Physiologists agree that voluntary muscle fibre is to be regarded as one of the master- tissues of the animal body. To maintain such highly specialised structure in good condition the frequent exer- cise of its function is necessary. Parts frequently used are, as a rule, abundantly supplied with blood, for healthy performance of function depends on the organ being adequately nourished. If these premises be correct then the conclusions which must necessarily follow are these: — Introduction.] REGRESSIOX OF MUSCLE, 3 The ends of certain muscles are badly situated to exert their full contractile power, or on account of modification in the creature's habits, portions of, and in some cases whole muscles become rarely used or rended inoperative. Loss of function leads to diminished blood supply, decrease in the amount of nutrition ends in degradation of tissue- The converse of this leads to hypertrophy. The ends of muscles, when compared with the central portions, are passive; they need little blood supply, and become in consequence, metamorphosed into tendon. The active central parts are often called into play, are well nourished and increase in size and quality. A very good illustration of this presents itself in the biceps flexor cubiti. If the muscle to which the tendon belongs is one fre- quently exercised as in the case of the gastrocnemius and soleus, the attached tendon contains a very large amount of elastic tissue. Regression of Muscle. — When a muscle degenerates either from morphological or pathological reasons, the amount of contractile tissue is diminished, and the parts take on the character of fibrous tissue and become liga- ment. This change may conveniently be referred to as regression. These two processes metamorphosis and regression assisted by a singular change of position in muscles known as migration, which will be considered in detail later, have been most active in providing ligaments in the limbs. The view that tendon results from the metamorphosis of muscle receives great support from the fact that nerves terminate in them by developing end-organs something like those of muscle. These curious terminations have been especially studied by Golgi. They seem to be most numerous near the muscular insertion. They may form peculiar reticulated plates of primitive fibrillae, or end in structures recalling the end-bulbs found in the conjunctiva. The question of the relation of muscles, tendons, liga- ments and nerves has a practical bearing, inasmuch as it B 2 S LIGAMENTS. [Introduction . affords an anatomical explanation of the knee-jerk phe- nomenon so familiar to school-boys, which when used by the physician becomes an example of "philosophy in sport made science in earnest." It also serves to account for the intense pain caused by tension of a joint or the accumulation of pus under dense fascia. The curious and intractable contractions of palmar and plantar fasciae and other aponeurotic structures are more comprehensible if we regard them as arising from the metamorphosis and regression of muscles. Degeneration of osseous and cartilaginous tis- sues. — Ligaments arising by this method occur with greatest frequency in the axial skeleton and pectoral girdle. As a very large portion of what is to follow in the succeeding chapters is devoted to the consideration of this question, it will be unnecessary to spend much time in giving illustrations here, but it may be mentioned as a further inducement to those who are inclined to prosecute the further study of this question, that the inquiry will carry the investigator into some of the most fascinating regions of morphological science. As an example of the method by which ligaments arise from the degeneration of bone or cartilage, we may select the supra-condyloid foramen. In many carnivora, rodents, lemurs, and others, there exists above the internal condyle of the humerus, a bony tunnel known as the supra-condyloid foramen. This transmits as a rule, the median nerve and the brachial artery. In a young lion at birth as in the fcetal cat, the foramen is in part formed by an outgrowth from the diaphysis, as well as by an upgrowth from the epiphysis, as in fig. I. In man the most frequent condition of this foramen when existing, is to have the upper part of the ring formed by an osseous outgrowth from the humeral shaft, and named the supra-condyloid process. The lower part of the ring is completed by a band of fibrous tissue extending to the Introduction.] DEGENERATION. 5 internal condyle, and as a rule affords attachment to the pronator radii teres muscle. Fig. 2. Fig. i.— The humerus of a foetal lion, showing that the supra- condyloid foramen x is formed by the diaphysis and epiphysis. Comparing- the foramen when present in man with that of the lion, it will be seen that the portion of the ring- derived from the epiphysis is represented by fibrous tissue. Fig. 2. — The supra-condyloid process in man. (For full details the student should consult Struther's Anatomical and Physiological Observations, 1 85 4). * The observations of Baur seem to show that in the Armadillo the supra-condyloid foramen is complete in the shaft without the aid of the epiphysis. Morph. Jahrbuch, 1886. LIGAMENTS. [Introduction. The notch in the upper border of man's scapula which transmits the supra-scapular nerve and occasionally the artery, is usually converted into a foramen by a band of fibrous tissue. Frequently this ligament, known as the transverse, is replaced by bone. It is characteristic of sloths that this foramen is sur- rounded by bone, and its mode of formation is interesting. In the young ant-eater (I\/yn?iccophaga jubatd) the foramen is composed in part by the body of the scapula and com- pleted by the coracoid process, thus affording an interesting example in the limbs of a passage of a nerve between two centres of ossification, a condition of things almost con- stant in the skull. Therefore I am disposed to the view that the transverse ligament in man is the fibrous repre- sentative of this bony bridge constant in sloths, and that the occasional occurrence of a complete osseous foramen Fig. 3.— The scapula of a sloth, showing F, the osseous coraco-scapular (supra-scapular) foramen, C, coracoid. in this situation is not to be regarded as an ossification of the transverse ligament, but as a reversion to a former condition. As the suprascapular notch is formed in part by the scapula and coracoid process in man and other mammals, it would be far more convenient to refer to it by the name it bears in comparative anatomy — the coraco-scapular foramen, fig. 3. In man the tendon of the biceps plays in a groove at the upper end of the humerus, in him as in some carnivora, as the lion, tiger, and bear, this groove is arched over at its Introduction.] DEGEXERA TIOX. J upper end by a broad band of fibrous tissue, converting- it into a tunnel. In the mole as my friend Mr. Austin Freeman demonstrated to me, the tendon plays in a most perfect osseous tunnel. Among- other examples of the interchangeability of fibrous and osseous tissues, the following may be briefly referred to here : — In many carnivorous mammals and a few others, the tentorium cerebelli is replaced by a plate of bone. Among reptilians, some turtles present a curious ano- maly, for the temporal fascia, so conspicuous in man, is in these animals represented by a layer of stout bone. This condition is also found in a certain frog named Pelobates. The only known mammal in which this arrange- ment exists is a singular specimen of rodent from Africa named Lophiomys. The pillars of man's external abdominal ring, in that they consist chiefly of fibrous tissue, agree with the majority of mammals, yet in those singular groups, the Ornitho- delphia and Didelphia, the internal pillar is directly ossified to form the marsupial bone. On the other hand, bones in man which as a rule are well-formed, may be replaced by fibrous tissue. Thus the fibula has been represented by ligament, and numerous cases are on record of the shaft of the first rib in man being merely a fibrous band extending to the sternum. * Journal of Anatomy] and Physiology, vol. xx., p. 206. " On the Myology of the Forelimb of the Mole." CHAPTER I. Metamorphosis and Regression of Muscles in Relation to Ligaments. The principal arguments in this chapter concerning- the nature of some of the ligaments of man's body, depend on the circumstance that muscles under certain conditions degenerate into fibrous tissue, which may become utilised as ligament if occasion demands. It will be well, therefore, to commence the discussion by the consideration of a few indisputable instances. Professor Watson mentions the occurrence of the curvatores coccygis muscles in man ; they arose from the anterior surface of the sacrum between the third and fourth sacral foramina, also from the anterior surface of the body of the fifth sacral vertebra. They were inserted into the front surface of the body of the second, third, and fourth coccygeal vertebrae. The Professor then goes on to relate that Albinus, in his Histories Musculorum, p. 336, had noted this muscle in three subjects, but that in others they were replaced by ligament- ous or tendinous fibres. The exact statement is: " Inveni cumintribus; in alio imperfectionem et degenerantium; in aliis non musculo, sed ligamento simile." The -extract is quoted here as affording excellent testimony to the degeneration of muscles into ligamentous tissue, from so famous an anatomist as Albinus (1734). The bands of fibrous tissue passing from the lower sacral vertebrae to the coccyx, and known as the posterior sacro- coccygeal ligaments, represent in man the extensor coccygis of anthropomorphae, or extensor of the caudal vertebrae of other animals. In man the place of these ligaments is occasionally occupied by muscles, the attach- ments of which are thus described by Hyrtl : — * Journal of Anatomy and Physiology, vol. xiv., p. 407. Chap, i.j METAMORPHOSIS OF MUSCLE. "The muscles are found between the hinder surface of the sacral vertebrae and the last piece of the coccyx. They are a pair of tendinous muscles, repetition of a muscle which appears in most mammals — sacro-coccygeus posticus seu extensor coccygis" (Lehrbuch der Anatomie, 14M Aujiage, 487). The preceding cases serve as excellent illustrations of the rules to be followed in forming" conclusions as to whether any particular band of fibrous tissue represents a degen- erated muscle : — 1. It must correspond with its presumed representative in origin and insertion. 2. The muscle should occasionally reappear as an anomaly in the human subject and replace the fibrous tissue. 3. Failing Rule 2, the muscle should assume a functional importance in other animals. If Rules 1 and 2 be satisfied, the assumption that a given fibrous band represents the muscle is reasonable ; if the third rule be sustained, the evidence is then very strong, and becomes almost a certainty. When Rules 1 and 2 are not satisfied, the notion of re- presentation should be entertained with caution. Thus — (1) The place of the curvator coccygis is occa- sionally occupied by a fibrous band. (2) This band is occasionally replaced by the curvator as a functional muscle; it is then regarded as an anomalous structure. (3) The curvatores coccygis, as seen in man, are repre- sentatives in him of the depressores caudae present in very many of the lower mammals. These rules are well illustrated by the epitrochleo-anconeus, an occasional muscle in man, which arises from the back part of the internal condyle of the humerus and is inserted into the inner side of the olecranon process. It forms the tunnel for the ulnar nerve, and when not represented by muscle a piece of fascia occupies its place, this fascia may be regarded as .the degenerated muscle. Gruber has described the epitrochleo-anconeus as being the most frequent muscular abnormality in the body. IO LIGAMENTS. [Chap. i. The chief points in its anatomy are represented in the accompanying- drawings, fig". 4 represents the muscle as it occurs in man, fig. 5 shows it replaced by a tract of fibrous tissue — the usual condition. Fig-. 6 represents the elbow- JUl. joint of a Virginian opossum, in whom this muscle con- stantly occurs. UJl. E.A.muscIe Fig. 6. The epitrochleo-anconeus exists in a very large number of mammals. * For further details consult J. C. Galton, Journal of Anatomy and Physiology, vol. ix., p. 169. Chap, i.] C0RAC0-BRACH1ALIS. II A muscular slip in the sole of the foot of man, known as Wood's muscle, is a case in point. It arises from the external tuberosity of the calcaneum, and is inserted into the prominence at the base of the fifth metatarsal bone in common with the outer margin of the plantar fascia. The muscle is occasionally referred to as the abductor ossis metatarsi quint 7. According- to most authorities it is present once in every two subjects, but my observations shew it to be even of such frequent occurrence that I am in the habit of teaching - that the first layer in the sole of the foot con- sists of four muscles, thus regarding Wood's muscle as of normal occurrence. When not present in a functional con- dition a stout band of fibrous tissue occupies its place. The muscle was present in the feet of two chimpanzees which I had the opportunity of dissecting- ; and it occurs in many other quadrumana. Like the preceding- muscles it well exemplifies the neces- sary rules, thus it may be present as a functional muscle, or represented by fibrous tissue, while it is a well and normally developed muscle in lower mammals. The Coraco-Brachialis Muscle. For the true history of this interesting- muscle we are indebted to the labours of Prof. Wood. This anatomist has pointed out that the coraco-brachialis in mammals has for the most part a triple constitution. The muscle arises from the tip of the coracoid process of the scapula in company with the short head of the biceps, with which it is united for some distance, and is inserted into the inner border of the humerus near its middle. Some of the higher fibres are attached to a fibrous loop extending from the coracoid process to the lesser tuberosity of the humerus. Viewed in the light afforded by a study of the * Journal of Anatomy and Physiology, vol. i., p. 44, 1867. 12 LIGAMENTS. [Chap. i. variations this muscle is subject to in the human body, and information gained from comparative myology, the coraco- brachialis presents three parts. 1. The portion represented by the fibrous loop may develop as a distinct muscle attached to a lesser tuberosity of the humerus and known as the rotator humeri. 2. The middle portion is the cot aco-brachialis of ordinary human anatomy. N Fig. 7. — The usual arrangement of the muscle, C, fibrous ligament representing coraco-brachialis sup., C", the medius, and C", the longus, in part ligament, N, the musculo-cutaneous nerve. 3. The third piece may exist as a muscular belly extend- ing from the common tendon to the internal condyle of the humerus. It is rare to find this piece fully developed, its situation being marked out by the fibrous band which commonly extends from the lower part of the muscle to the elbow, and is familiar as the internal brachial ligament of Struthers. Figs. 7, 8 and 9. The first may be called the coraco-brachialis superior or Chap, i.] CORA CO-BRA CHIALIS. 13 brevis, the second is the coraco-brachialis proprius or medius, and the third coraco-brachialis longus. This arrangement Fig. 8.— C C" C"\ as in fig. 7. recalls in a very striking- manner the disposition of the adductor brevis, longus, and magnus, in the thigh. Fig. 8. Fig. 9. — C, functional upper segment, C'' and C" as in fig. 7. A great deal of variation exists among mammals as to the development of the various parts of this muscle. In i 4 LIGAMENTS, [Chap. i. some the rotator humeri alone is developed, in others only the middle segment. A few have the second and third portions as a well developed muscle extending- to the in- ternal condyle. For further details the student should consult Wood's admirable paper before mentioned. The Interosseous Membrane of the Forearm and Leg. Passing - between the radius and ulna is a flat fibrous membrane, known as the interosseous membrane of the forearm. Its fibres are directed for the most part obliquely downwards from the radius to the ulna, the membrane commences a little below the bicipital tubercle. On the posterior aspect small bundles of fibrous tissue intersect the direction of the main fasciculi at right angles. Lying" on the membrane are the anterior interosseous nerve and arterv. Careful dissection shows that small bands of tissue, intimately connected with the membrane, arch over and thus enclose the nerve and artery in a fibrous tunnel. In the leg a similar interosseous membrane exists, the fibres of which pass more or less horizontally from the fibula to the tibia and are intersected by fibres, passing- obliquely between the two bones in different directions. The anterior tibial nerve and artery bear the same relation to the membrane as is the case in the forelimb. In the iguana, gecko, menopoma, sphenodon. chameleon, and the like, a muscle exists, the fibres of which arise from the fibula to be inserted into the tibia, thus occupying the position of the interosseous membrane so common among mammals. The muscle in question has been named the peroneo-tibial and it exists in a well-developed form in the hind leg of the wombat. The anatomical relations of the muscle may be studied most completely among the lizards. If the interosseous membranes in the forearm and leg of man were not originally derived from a peroneo-tibial Chap, i.] PALMAR AND PLANTAR FASCIA. 15 muscle, they are at least the morphological representatives of that structure. As the muscle occurs within the mam- malian circle, it is not unreasonable to regard these mem- branes as degenerated muscles. If this view be accepted, then the anterior interosseous arteries and nerves are to be regarded as those originally ministering to this re- markable muscle. The fibrous tunnel in which they lie is formed as a consequence of the degeneration of the muscle which they once supplied. It is quite possible that the popliteus is a remnant of the original peroneo-tibial muscle, also the rotator fibulae of the gibbon. The Palmar and Plantar Fascia. The palmar and plantar fasciae are structures so well known to the student of human anatomy, that it would be tedious even to recapitulate the details connected with their anatomy. It is sufficient for the purposes of morphology to remember that the palmar fascia is connected superiorly with the anterior annular ligament, whilst the middle, strongest portion of the fascia, is directly continuous with the tendon of the palmaris longus muscle. This muscle, so far as man is concerned, presents itself normally as a feeble structure; its importance may be estimated from the fact that it is frequently absent without in the least im- pairing the utility of the limb. It is calculated that this muscle is wanting in ten per cent, of all bodies examined. Usually it consists of a definite belly with a rounded tendon, occasionally two bellies are present with a ten- dinous intersection ; the muscle may end in the fascia of the forearm, or may be wholly represented by ligament. In many animals, e.g., the armadillo and aard vark, the palmaris longus is large and important, muscular throughout its whole extent, sending slips to all the 1 6 LIGAMENTS. [Chap. i. digits, and serving- as one of the chief agents in flexing them. That the palmar fascia arises from the degeneration of the distal end of the palmaris longus muscle, admits of no doubt, for it occasionally happens that portions of the fascia reassume their lost character and present themselves as slips of muscle. This occurs most frequently in connection with the strips of fascia going to the thumb. There is good reason to believe that the abductor pollicis and abductor minimi digiti are derived from the distal end of this once extensive muscular sheet, of which the sole representatives, in a muscular form besides them, are the belly of the palmaris longus and the palmaris brevis muscles. The plantar fascia. — The disposition of this fibrous sheet in the sole of the foot exactly repeats that seen in the hand, and the same arguments will be used to show that in this case the fascia results from the regression of the plantaris muscle. This muscle, like the palmaris longus, is vestigial in man, and like such structures in general is liable to all kinds of variations. Normally it is inserted into the inner side of the tendo Achillis, but it may join the deep fascia, the internal annular ligament, or plantar fascia, occasionally it is absent. These vagaries alone are sufficient to indicate that the muscle has an interesting ancestry. Turning to lower forms we find that in the armadillo the plantaris muscle, instead of being, as in man and many mammals, chiefly represented by a long, thin, and straggling tendon, is larger than the gastrocnemius. The muscle arises as usual from the back part of the external condyle of the femur, it then expands into a large fleshy belly extending the whole length of the calf; at the heel it forms a tendon which glides in a well-formed groove on the back of the os calcis, and spreads out as the plantar fascia, slips of which pass to the hallux, second, and third toes, extending even to the terminal phalanges. As many of the short muscles belonging to the thumb Chap, i.] PLANTAR LIGAMENTS. 17 and fifth digit may have arisen from portions of the original palmaris longus, so may some of the short muscles in the first layer of the sole be considered as detached portions of the plantaris. An exception must be made in the case of the abductor hallucis, which is prob- ably a migratory muscle, and will be considered later. For an excellent account of the plantaris and palmaris longus muscles in Orycteropus, the student should consult Mr. J. C. Galton's account of the myology of this animal in the Transactions of the Linnean Society, vol. xxvi., p. 567. The Long and Short Plantar Ligaments. ( Calcaneo- Cuboid). If the hind leg of a horse be examined, a strong ligament passing from the posterior surface of the os calcis to the cuboid and head of the rudimentary external metatarsal bone may be readily dissected. This fibrous band is known as the calcaneo-metatarsal ligament. Near the origin of this ligament from the calcaneum, the tendo Achillis or combined insertion of the gastrocnemius and soleus muscles is attached. In the fcetal horse and foetal deer, the tendo Achillis and the calcaneo-metatarsal ligament form one continuous structure. It would be most illogical to say that because the tendon and ligament were one structure in a limb so peculiarly modified as that of the horse, the same source of the ligament could be maintained for the long and short plantar ligaments of man's foot, unless other evidence exists. If a hiuTian foetus even as late as the sixth month of intra-uterine life be examined by a vertical section carried through the tendo Achillis, os calcis and cuboid, the ligament and tendon will be seen to form one continuous band, and the attachment of the tendon to the under surface of the os calcis is of the loosest kind. After the seventh month of intra-uterine life the tendon is c i8 LIGAMENTS. [Chap. i. gradually divided by extra growth occurring at the posterior extremity of the os calcis, the inferior portion becoming" the long- and short plantar ligaments, the other gaining a permanent attachment to the lower border of the os calcis. If in the adult foot, the origin of the muscles composing the third lay< r of the sole of the foot be dissected, they will be found, with the exception of the transverse pedis, 10 be thoroughly incorporated with the plantar ligaments. When th _■ corresponding- parts are dissected in the foot of a seal the anatomist will quickly come to the conclusion that the adductor pollicis, flexor brevis pollicis, flexor minimi digiti, long plantar ligaments, tendo Achillis, gastrocnemius, and soleus muscles, have been derived from a muscular layer which in lower primitive forms was a continuous muscular stratum. The conclusion is inevitable, that the long and short plantar ligaments are degenerated muscular tissue. The Deep Fascia of the Limbs. On comparing the limbs of a lizard or a bird with that of man or any of the higher mammals, we shall be struck with the fact that in the latter forms the muscles are invested by strong aponeuroses known as the deep fasciae, and separated one from the other by fibrous intermuscular septa. Not so in the Sauropsida (birds, lizards, croco- diles). Careful inquiry, by means of dissection will force an impartial worker in the end to the conclusion, that the deep fasciae and the intermuscular septa are in reality the result of the metamorphosis and regression of muscular tissue, at least so far as the fore- and hind-limbs of most mammals are concerned. An attempt is made on page 31 to show that there is good ground for believing that the broad band of fascia continuous with the biceps temoris tendon and extending Chap, i.] DEEP FASCIA 19 thence to the external annular ligament of the ankle may be traced to a large and functional muscle in lizards, crocodiles, and birds. It is commonly recognised that the thickening of the fascia lata known as the ilio-tibial band is the degenerate representative of that portion of the tensor fasciae femoris muscle which in some animals extends to the knee. There can be little doubt that the larger portion of the fascia lata on the posterior aspect of the limb is derived from degeneration of the distal portion of the gluteus maximus muscle. The glutei muscles in birds are very numerous, as many as four are present in some species. In others some of these muscles are entirely represented by tracts of fibrous tissue or fascia. If the dissector wishes to find the external cutaneous nerve in the thigh he has to seek it in a tunnel formed in the fascia lata. When dealing with the interosseous mem- brane of the forearm it was shown that the artery and nerve were enclosed in a tunnel in consequence of the degeneration of the muscle to which they ministered. Comparative anatomy supports the view that the external cutaneous nerve of man lies in a tunnel in consequence of the muscle which it supplied, having undergone regression, and burying the nerve and artery in fibrous tissue, for, as in the case of the interosseous membrane, the nerve is always accompanied by an artery. The same line of argument is true of the deep fascia of the arm. I have seen an instance in which the bicipital expansion was replaced on both sides by a thin sheet of muscular tissue, much broader than the fan-shaped arrange- ment of fibrous tissue which it replaced. In the ensuing section evidence is forthcoming that the same line of argu- ment is applicable to the fasciae of the trunk especially those in the loins. C2 20 LIGAMENTS. [Chap. i. The Lumbar Aponeurosis. Under this name human anatomists have found it conve- nient to describe three stout layers of fascia ensheathing the erector spinae and quadratus lumborum muscles. The superficial layer springs from the spines of the lumbar vertebrae and the posterior part of the iliac crest. After closely investing the erector spinse, it joins with the middle layer of the aponeurosis at the outer border of that muscle. The middle layer springs, by strong fibrous bundles, from the tips of the transverse processes of the lumbar vertebrae ; and, passing between the erector spinae and quadratus lumborum muscles, blends with the posterior layer, as already described, and forms a strong tendon of origin for the transversalis abdominis muscle. The anterior layer of the lumbar aponeurosis is attached to the front of the transverse processes of the lumbar vertebrae, covers the anterior surface of the quadratus lumborum muscle, and blends with the middle layer of fascia. Thus the anterior layer blends with the middle at the outer border of the quadratus lumborum, whereas the posterior and middle layers of the aponeurosis blend at the outer border of the erector spinae muscle. Careful inquiry into the comparative anatomy of the posterior layer of the aponeurosis makes it evident that it arises from the metamorphosis of the lower fibres of the latissimus dorsi muscle ; for in many animals this fascia is wholly replaced by muscle-fibres continuous with the latissimus dorsi sheet. The same mode of origin may be maintained for the anterior aponeurosis of the external oblique, internal oblique, and transversalis muscles, also the resplendent tendon of the serrati postici superior and inferior, and the rhomboids. Even the central tendon of the diaphragm has arisen from the metamorphosis of its muscular fibres ; for in the porpoise there is no trace of Chap, i.] LUMBAR APONEUROSIS. 21 tendon, but it is muscular in structure throughout the whole of its extent. The middle layer has a very curious origin. If a recent subject be chosen, and the structures of the loin be re- Fig. io. — To show the mode of origin of the posterior and middle layers of the lumbar aponeurosis from the latissimus dorsi and de- generate levatores costarum muscles respectively. moved down to the quadratus lumborum muscle, care being taken not to injure its investing fascia, there will be seen arising from the tips of the transverse 22 LIGAMENTS. [Chap. i. processes of the three upper lumbar vertebrae a fan- shaped arrangement of fine fasciculi composed of fibrous tissue, but more elastic in character than the rest of the aponeurosis, and strongly recalling in appearance the nacreous-looking fibres of origin of the serratus posticus inferior muscle. These fan-shaped masses have a direction downwards and outwards, the apices being attached to the tip of the transverse process exactly like a levator costae muscle, and there can be little doubt that the thickenings in the middle layer of the lumbar fascia are the degenerate remains of a set of muscles in continuation of the levatores costarum series. These fan-shaped collections of tissue in the middle layer of the transversalis aponeurosis are by no means occasional, but may be demonstrated in any dissect- ing-room subject with ordinary care. In bodies possessing a lumbar rib. however small, the fan-shaped ligament attached to the twelfth dorsal vertebra is replaced by a functional muscle, as shown in fig. 10. The anterior layer is in all probability derived from the regression of that portion of the transversalis muscle lying nearest the spine. An examination of the pleuro-peritoneal cavity of lizards leads me to believe that in a typical condition the abdo- minal and thoracic parietes may be conceived as possess- ing in their lateral and ventral aspects three muscular strata. 1. The external oblique sheet intersected in a part of its course by the ribs, giving rise to the external intercostal muscles. 2. An internal oblique stratum, also partially interrupted by the ribs forming the internal intercostal muscles. 3. The transversalis sheet immediately subjacent to the serous lining of the pleuro-peritoneal cavity. Mammals are distinguished from lower vertebrata in the possession of a complete diaphragm. This singular struc- ture probably represents that portion of the transversalis sheet which covers in lizards the inner aspect of the ribs. Chap, i.] LUMBAR APONEUROSIS. 23 The thickenings known as the ligamentum arcuatum in- ternum and externum, are to be regarded as tendinous intersections between the thoracic and abdominal segments. The original extension of this muscular stratum into the thorax is indicated by the occurrence of the sub- or infra- costal muscles which are found most frequently near the angles of the ribs on their inner aspect. There is good ground for believing that the longus colli muscle of man is a derivative of this same transversalis sheet, for in such animals as Jfenopoma, Cryptobranchus, etc., a prolongation extends forwards to the basis cranii. The origin of the three layers of the lumbar aponeurosis may be briefly stated thus. (a) The posterior layer results from the metamorphosis of the lower fasciculi of the latissimus dorsi muscle. (&) The middle layer contains degenerate elements of the lower representatives of the leva/ores costarwn series of muscles, one of which forms the ilio-lumbar ligament, and possibly this sheet of fascia is made up entirely in this way. (c) The anterior layer is the result of the metamorphosis of the vertebral portion of the transversalis muscle. In that man possesses in the posterior and anterior regions of the trunk, (for there can be little doubt that the aponeuroses of the obliqui and transversalis muscles are the result of the metamorphosis of the fasciculi of these muscles,) large sheets of tissue composed of minute glisten- ing tendons, he contrasts markedly with amphibians, lizards, and ophidians, in whom these aponeurotic tracts are represented by sheets of well developed muscular tissue. Absent Muscles. In the Journal of Anatomy and Physiology, vol. xix., p. 333, Professor Sir W. Turner notes a case of absence of the extensor carpi ulnaris muscle. On careful dissection, 24 LIGAMENTS. [Chap. i. however, a narrow band could be detected in the deep surface of the aponeurosis of the forearm, somewhat stronger and more opaque than the rest, extending- longi- tudinally down the limb, parallel with the ulna. Above the styloid process the band became isolated from the fascia and entered the groove on the back of the ulna in which the tendon of the ex. carpi ulnaris ought to have been lodged ; finally it became attached to the base of the fifth metacarpal bone. Turner regards this as an example of the conversion of a muscle into fibrous tissue, and refers to a similar case recorded by Gruber.° The interest of the case is this : when a muscle is found wanting in a subject in which normally it exists, search should be made for its fibrous representative. Doubtless many instances of muscular variations and absence of muscles receive explanation in this way. Thus, Professor Flower and Dr. Murief, in their account of the dissection of a bushwoman, state that there was no abductor ossis metatarsi quinti, but a strong tendinous band stretched from the plantar surface of the os calcis, close to the origin of the abductor minimi digiti to the head of the fifth metatarsal bone. In an old woman, Mr. Macdonald Brown found the muscles of the thenar eminence replaced by bands of fibrous tissue. In the six-banded armadillo, Mr. Galton found the short flexors in the sole of the foot similarly represented. Every anatomist of experience must also have come across examples of this not infrequent form of replacement in the human subject. * Virchow's Archiv, bd. xcix., p. 478. f Journal of Anatomy and Physiology, vol. i., p. 204, 1867. CHAPTER II. Examples of Metamorphosis and Regression of Muscles from the Lower Vertebrates. Instances of the transformation of muscles into ligaments are almost as frequent in the lower vertebrates as in man. It may not be uninteresting- to devote a chapter to the con- sideration of some striking examples chosen from the horse, whales, and birds. William Percivall in his excellent work on the Anatomy of the Horse, 1858, adds a foot note to his accountof the sus- pensory ligament of the fetlock, stating that Bourgelat regarded it as a tendon, but Girard as a muscle. Percivall describes it thus: — ''The suspensory ligament (so-called, I imagine, because the sesamoid bones seem to be suspended by it) is, perhaps, the strongest in the whole body, and is remarkable for its high degree of elastic property. It takes root, superiorily, in a projection at the upper and back part of the cannon, whence it passes, enclosed within a cellular sheath, between the splint bones, filling up their interspace. Opposite to about the termina- tions: of these small bones, it splits into two divisions, which, diverging in their descent, become implanted into the lateral and posterior parts of the sesamoid bones, and into the fibro-cartilaginous substance uniting them. From the places of implantation, two lateral slips are continued from it downward and forward to join the extensor tendon. Between the suspensory ligament and the joint, enveloped in adipose membrane, are some large bursa mucosa. In composition and texture, this ligament possesses peculiari- ties : it has a sanguineous tinge interiorly, which is not perceptible in other ligaments or in tendons ; and its fibres, which are very coarse, are disposed in layers. But its chief peculiarity consists in its exhibiting an inter-tex- 26 LIGAMENTS. [Chap. II. ture of delicate, pinky, fleshy fibres, which appear to be the uniting medium of the ligamentous fasciculi." The most conclusive evidence on this question is furnished by Prof. Cunningham in a paper on "The Fig. ii. — The forelimb of a Porpoise. The muscle of the forearm and manus are represented by fibrous tissue. Development of the Suspensory Ligament of the Fetlock in the fcetal Horse, Ox, Roe-deer and Sambre-deer," he shows most conclusively that this ligament is formed by the transformation of the short flexor muscle of the middle digit into fibrous tissue. The corresponding structure in * Journal of Anatomy and Physiology, vol. xviii., p. i. Chap, ii.] THE BICIPITAL LOOP. 27 the ox, sheep and camel arises by the coalescence and fibrous metamorphosis of the short flexor muscles of the middle and annular digits. In the forelimb of the porpoise represented in fig". 1 1 the muscles are no longer present, but tracts of fibrous tissue occupy corresponding positions. Professor Struthers has pointed out that in the Narwhal the finger muscles are present morphologically, but histo- logically they are represented by fibrous tissue. A similar but not >o extreme a condition may be studied in the wing of the Penguin. f The Bicipital Loop in the Thigh of Birds. Let anyone take the trouble to dissect the thigh of a fowl from the outer side, then a most interesting arrangement of myological structures will reward his labour, but the one to which especial attention is invited, is the singular loop which transmits the tendon of the biceps muscle and a branch of the great sciatic nerve. The great point of interest in connection with the bicipital loop is the fact that it is an exclusively avian character, at least so far as recent forms are concerned, and that with very few exceptions, among them a Steganopod {Phtzthon), no known bird lacks this sling. On reflecting the skin, the most superficial muscle is the tensor fascia or gluteus primus as it is termed by some. If this muscle be turned up, or removed, the deep structures will be exposed ; the most conspicuous is the biceps, which has the following attachments: — It arises from the upper- three-fourths of the post-acetabular ridge (a slight bony elevation which separates the post-acetabular area from * "On some points in the Anatomy of a Great Fin Whale," Journal of Anatomy and Physiology, 1871. f See especially Watson's " Report on the Spheniscidas," Zoology of the Voyage of H. M. S. " Challenger:' 28 LIGAMENTS. [Chap. ii. the external lateral surface of the ischium). Just in front of the tensor fasciae the fibres converge to form a rounded tendon, which on the outer side of the popliteal region is bent sharply downwards by passing- through a tendinous sling or loop; it is inserted nearly half way down the fibula to a bony prominence on its outer side. The loop in question is attached to the lower end of the femur; the outer limb is slightly connected with the head of the gas- trocnemius, which is situated to its outer side. A nerve to the leg and foot, derived from the great sciatic, accompanies the tendon through this sling (fig. 12). This tendinous loop, studied in the fowl and similar birds, is curious and puzzling, and evidence will be adduced in support of the view, that it is derived from the metamorphosis of the muscular fibres of the outer head of the gastrocnemius muscle. Before doing so, however, it will be necessary to state that the muscle, here termed biceps, has not the same anatomical relationship as the muscle so named in the leg of mammals, indeed, to my mind it is certainly not homologous with that structure, and there is little utility in discussing the morphology of muscles in so highly special- ised a form as that of the fowl as compared with man. In the thigh of the Rhea {Rhea Americana), the Emu (Dromceus Novce-hollandice), Ramphastos loco and many other avian forms, the loop through which the biceps tendon passes is modified in this way: — The inner limb is a tendon of great strength and of considerable thickness, but the outer limb of the loop is constituted by the muscular and ten- dinous fibres of that head of the gastrocnemius muscle which is attached to the external femoral condyle ; hence one half of the loop is tendinous, the other muscular (fig. 13). This arrangement clearly affords the clue by which we may seek to interpret the origin of so singular a pulley, for clearly it is the result of the metamorphosis of muscle fibres. In the lizard, Iguana tuberculata, if the dissection be com- menced from the outer side, as in the bird, a prominent muscle, referred to by Mivart as the ilio-peroneal, is easily Chap, ii.] THE BICIPITAL LOOP. 29 recognised. It has the following- attachments and rela- tions: — It arises from the posterior part of the outer side Fig. 12. — Thigh of a bird (Tinamou) dissected from the outer side to show the tendinous loop which transmits the tendon of the biceps and a nerve, the latter is represented by X. B. Biceps. G. Gastrocnemius. of the ilium, covered by the posterior portion of the tendinous origin of the gluteus maximus, and even a little overlapped by the gluteus medius. It is inserted by a llill Fig. 13. — The bicipital loop in the thigh of an Emu. One half is formed by the gastrocnemius, the other half is tendinous, the nerve is represented by X . strong tendon, which dips between a muscle called peroneus pri?nus by Mivart, and the external head of the gastrocnemius, into the outer side of the fibula near its summit. * Mivart's admirable account of the " Myology of Iguana tubercu- lata" (Proc. Zool. Soc, 1867), has been to me of the utmost utility in my dissections. 30 LIGAMENTS. [Chap. ii. It will be necessary, before discussing the ilio-peroneal muscle, to take into consideration the muscles named peroneus primus and the gastrocnemius. The first muscle arises by a strong tendon from the summit of the outer side of the external condyle of the femur. It becomes tendinous below the outer malleolus; a portion of this expansion gains an insertion into the peroneal border of the fifth metatarsal bone, a little above its middle. On the posterior aspect of the ankle-joint it forms a broad aponeurosis, and joins with the expansion of the gastroc- nemius. The gastrocnemius consists of two distinct parts ; one arises from the internal condyle of the femur, with a slight attachment to the head of the tibia near the insertion of the semi-membranosus and semi-tendinosus. Passing downwards it becomes aponeurotic near the ankle-joint, and blends with a broad tract of tissue — the plantar fascia. The second portion arises from the femur, just above the external condyle ; passing down the leg it becomes apo- neurotic near the ankle-joint, and blends with the plantar fascia. In very many birds the gastrocnemius arises by three heads, an inner one from the tendon of the rectus, and cnemial crest of the tibia ; the middle head from between the condyles of the femur, and an outer one arising from the external condyle of that bone. Between these two heads — the middle and the external — passes the biceps tendon. On comparing the fowl's leg muscles with those of Iofuana, it becomes evident that the muscle marked peroneus primus in the Iguana is really the equivalent of the outer head of the fowl's gastrocnemius, and that the bird's biceps cruris is really the representative of the ilio- peroneal of the Iguana; they agree in position and especially in their relation to the great sciatic nerve. Admitting this to be so, it will give us the steps or stages by which the bicipital sling has become fashioned. They may be briefly summarised as follows: — Chap, ii.] THE BICIPITAL LOOP. 3* 1. In its original condition the so-called biceps passed between two muscles ; one the third head of gas- trocnemius or peroneus primus, and the other the outer head of gastrocnemius. 2. The two muscles tend to fuse near the knee-joint, and enclose the distal end of the biceps. 3. The leg becomes bent at a more or less acute angle with the thigh ; this leads the biceps to drag on the muscle-fibres of the gastrocnemius through which it passes. The muscular tissue in relation with the tendon of the biceps thus piays a passive part, and becomes metamorphosed into tendon. 4. The inner limb first separates from the parent muscle; the outer head may do so and form a complete tendinous sling ; differentiation may not proceed so far, the outer head remaining in many birds per- manently attached to the outer portion of the gastrocnemius. The question at once arises, Have the facts any bearings on the myology of the mammalian limb? My answer is this. If the biceps femoris in man be carefully traced, the insertion is not by any means confined to the head of the fibula, but its fibres spread into a broad aponeurotic ex- pansion covering the outer side of the leg. The relative size of the biceps in man, as Mivart points out, is much inferior to its possible development. In some forms, e.g., the Agouti (Dasyprocta agouti)^ the muscle combines with the tensor fasciae and gluteus maximus to form an almost continuous sheet of muscle, extending from the pelvis over the outer side of the leg to the ankle-joint. I was very much interested in the biceps of a rhinoceros, in whom the muscle, after being inserted into the fibula, was prolonged by means of an enormously thick and broad tendon to the ankle-joint. This and similar facts induces me to venture the following explanation. The muscle in Iguana called peroneus primus, that in birds known as outer or third head of gastrocnemius, the broad muscular expansion in the 32 LIGAMENTS. [Chap. ii. Agouti, the enormous tendon of the biceps femoris in the Rhinoceros, and its broad fibrous expansion in man, must all be regarded morphologically as one and the same structure. The nerve which originally supplied this muscle, lies in part of its course beneath the fascia, and in part superficial to it, and is named musculo-cutaneous. The Wing of a Bird. A bird's wing abounds in examples of the conversion of muscular tissue into tendon. In this place we cannot do more than notice a few of the more important ones. In most birds the flexor carpi ulnaris is a fairly well developed muscle, but in the Vulture and in the Rail, Rallus aquaticus, Razor bill, Aha torda, and others, this muscle is replaced by a strong, thick and elastic tendon. Mr. D'Arcy Thompson in a paper " On the Nature and Action of Certain Ligaments," which is worthy careful perusal, points out that ligaments of this kind, though degenerate muscles, continue to perform the function of the muscle they represent. Tendons of this kind are developed from muscles which pass over two joints and which have a more or less ligamentous action. f This is well seen in the case of the bird's flexor carpi ulnaris, for when the elbow is flexed the tendon serves automatically to flex the wrist joint. Mr. Thompson writes that he " knows of no case in which the muscles have become purely ligamentous, it is indisputable that the stronger the wing the weaker these muscles, and the longer are their tendons in proportion to the fleshy bellies." As a matter of fact the flexor carpi ulnaris is purely ligamentous in many birds. The observations of Prof. Thompson are confirmatory • Journal of Anatomy and Physiology, vol. xviii., p. 406. t See especially Cleland, Ibid., vol. i., p. 85. Chap, ii.] WING OF A BIRD. 33 of the opinion regarding tendons of muscles frequently used, as stated on page 2. Every student of human anatomy must have experienced a certain amount of curiosity when he dissected for the first time the plantaris muscle ; this strange structure sinks into insignificance when compared with the celebrated ambiens of the bird's leg, or the tendon of the femoro-caudal in the lacertilia. Of all strange muscles, the one known as the expansor secundariorum (Garrod) in the bird's wing stands pre-eminent. It is a small triangular muscle arising from the quills of the last few secondary remiges at the elbow. Its remarkably long and slender tendon, which frequently traverses a fibrous pulley on the axillary margin of the teres muscle, runs up the arm side by side with the axillary vessels and nerves, to be inserted in the thorax into the middle of a tendon which runs from the inner side of the middle of the scapular element of the scapulo-coracoid articulation, to near the middle of the thoracic border of the sterno-coracoid articulation, at right angles to it when the fore-limb is extended. In the ducks and geese, among the Anseres, the tendons under consideration, when they enter the thorax, run towards one another and join, (after having expanded out), in the middle line in front of the oesophagus, and behind the trachea. Not the least interesting feature in this muscle is the im- portance Garrod attached to it for the aid it afforded him in his Classification of Birds. For all that relates to this extraordinary muscle, I must refer the reader to Garrod's memoirs: — " On the Anatomy of Chaufia derdiana" (Proc. Zool. Soc, 1876), " The Anatomy of Passerine Birds" .(Hid., 1874, 1877, and 1878), and to the volume of his Collected Scientific Papers, 1 88 1 . My investigations into the morphology of this tendon in- duce me to believe that it is the representative in the bird's wing of the coraco-brachialis longus of mammals, and the long brachial ligament of man. See page 13. D 34 LIGAMENTS. [Chap. ii. Thompson in the paper already alluded to, insists that as a general rule when a muscle by reason of it passing over two joints is capable of being replaced partly or wholly by ligament without its efficiency being lost, then ligamentous change will set in. The best example is furnished by the flexor sublimis muscle of the mole. In this remarkable mammal there lies in the front of the forearm an enormous ligament, thicker than the ulna. It divides in the palm to be inserted into the five digits. Above it is attached to a depression in front of the internal supra-condyloid ridge and process of the humerus, it represents the flexor sublimis muscle metamorphosed completely into a tendon. For full details concerning the mechanism of this arrangement, the reader must refer to the original paper. The heart furnishes another illustration. The valves of the auriculo-ventricular orifices are muscular in the foetus, but later metamorphose into tissue, ligamentous in its essential characters. The right ventricle of a bird's heart contains a muscular valve, and this is the case even in the mammalian circle, e.g., in the echidna. The conclusion, however, seems to be this ; when a muscle undergoes metamorphosis into tendon, it is because its action is needed frequently and forcibly. It is of course unnecessary to remind the reader that it is essential to bear well in mind the distinction between tendinous metamor- phosis and fibrous regression, the latter being the immediate result of disuse. CHAPTER III. The Migration of Muscles in Relation to the Formation of Ligaments. The term " migration of muscles " may be denned as: — The changing of the situation of a muscle by alteration of its origin, or insertion, or both. The lateral ligaments of the knee joint furnish some excellent examples, not only of the process, but also of its results. In adult men the adductor magnus muscle terminates at a tubercle on the internal femoral condyle ; from near this spot the internal lateral ligament of the knee joint takes origin. In two young orangs this muscle was found directly continuous with the ligament. In the human foetus at the fourth month of intra-uterine life the proportions of the various parts of the limbs to one another, are more or less anthropomorphous. At this date the adductor magnus muscle is inserted into the head of the tibia. Later migration occurs, the muscle becoming attached to the condyle of the femur, whilst the distal portion of the tendon becomes the internal lateral ligament. The External Lateral Ligament of the Knee Joint. Every student engaged in dissecting a knee joint must have had his attention arrested by the beautiful, rounded, cord-like conditions of the external lateral ligament. The notion of it being, or rather that at some time it had been, the tendon of a muscle seems irresistible, and this suspicion D 2 36 LIGAMENTS. [Chap. hi. is strengthened on finding occasionally, in young- subjects, a synovial sheath investing it. After dissecting this joint in a large number of animals, the muscles to which it could possibly have belonged were limited to two, — the peroneus longus and the extensor longus digitorum pedis muscles. The external lateral ligament is in reality the tendon of the peroneus longus, which has become gradually separated from the belly of the muscle, and formed a new attachment to the head of the fibula. The extensor longus digitorum muscle may be safely excluded for this reason : in many animals this muscle takes origin from the external condyle of the femur, e.g., hyrax, pig, horse, hare, and the orang among apes, and in the ostrich among the birds, and at the same time a well-developed external lateral ligament may be found quite apart from and in no way connected with this muscle. On the other hand, the peroneus longus muscle may often be found connected with the ligament, as in the opossum, in the domestic ox, and, what is more pertinent to this ques- tion, in the gibbon {Hylobates leucucus), the tendon of the peroneus longus arises from the external condyle of the femur, and replaces the ligament, the tendon itself having no attachment to the head of the fibula. This arrange- ment leaves the fibula free to submit to the action of the well-developed rotator fibulse which is present in this ape. Even in man the muscular fibres of the peroneus longus often have a considerable origin from the external lateral ligament. But by far the most important contribution to our know- ledge of migration is afforded by Dr. Ruge of Heidelberg, who satisfactorily traces out the history and ancestry of the extensor brevis digitorum muscle. He shows that in the Monotremata this muscle belongs to the peroneal group, and arises entirely from the fibula. Ascending the scale * " Untersuchung iiber die Extensorengruppe am Unterschenkel [und Fiisse der Saugethiere," Morph. Jahrbxich for 1878. Chap, hi.] LIGAMENT OF KNEE-JOINT. 37 of mammalian forms we find the muscle passing down tendon by tendon until it reaches the condition presented in the foot of man. The koala or native bear, as it is called in New Zealand, presents an intermediate condition, for we find two tendons on the dorsum of the foot, whilst the remainder still retain an attachment to the fibula. In very many mammals and even quadrumana we find two muscles arising" from the fibula, and gaining the dor- sum of the foot by passing behind the external malleolus. They are known as the peroneus quarti metatarsi, and peroneus quinti metatarsi muscles. They are really the representatives of the slips from the ex. brevis digitorum muscle to the fourth toe and the peroneus tertius muscle. The peroneus tertius muscle is regarded as a muscle exclusively human, but it is simply the most external slip of the short extensor of the digits, which has failed to migrate when its companions passed downwards from the leg to the dorsum of the foot. Dr. Hans Gadow° in an interesting paper on " Com- parative Myology," gives some examples of the process, and states that as a rule the origin of a muscle is more subject to variation than the insertion. My own obser- vations lead me to concur with this statement, and the history of the foot muscles serves as a striking instance. I am also of opinion that the abductor hallucis muscle is an emigrant, having descended from the tibia to its present position on the os calcis. For arguments in support of this, the student anxious for a full discussion of the matter should consult the reference. f The interest this singular process has for us is, that as the muscles migrate they leave some part of their structure to be transformed into ligament, and there is satisfactory reason to believe that the external and internal lateral * Journal of Anatomy and Physiology, vol. xvi., 1882. f Especially, Ibid., vol. xix., p. 42. 38 LIGAMENTS. [Chap. hi. ligaments of the ankle joint have arisen in this manner. Before quitting- the foot it may be interesting - to refer to the excellent observations made by Ruge on a remarkable change of position he has detected in the case of the dorsal interosseus muscles of the foot. In the early embryo the dorsal interossei are plantar in position and the metatarsals are in contact with each other, gradually the metatarsals separate and the interossei make their way between them eventually appearing on the dorsum of the foot. In their earliest stage these muscles possess one head, it is only when in their new position that they become bi- penniform. It is also important to remember that the varying posi- tion these muscles pass through, indicate stages which are permanent in lower vertebrates. Thus, the first stage is characteristic of the majority of mammals and may be con- veniently studied in the dog. In others, the second stage is represented ; whilst the third stage is probably never so completely attained in any mammal as in man. To those anxious to follow up this inquiry I would recommend them to study Ruge's paper, " Processes in the Development of the Muscles of the Human Foot," Morpho- logische Jahrbuch, 1878, and Cunningham's admirable monograph on the muscles of the foot, contained in the Zoology 0/ the Challenger Expedition, part xvi., 188 1, vol. v., " Marsupialia," also a paper " On the Intrinsic Muscles of the Human Foot," published in the Journal of Anatomy and Physiology, vol. xiii., p. i., 1878. The Ligamentum Teres. The round ligament of the hip joint has long been an anatomical puzzle, and many opinions have been held concerning its nature. It is, however, best regarded as the divorced tendon of a muscle, and here arguments will be Chap, hi.] LIGAMENT UM TERES. 39 raised in order to show that in all probability it belonged to the pectineus, but has become separated from it in con- sequence of skeletal modifications. The ligamentum teres is a structure fairly constant in Mammalia, but is wanting in the Seal, Elephant, Rhinoceros, Walrus, Ornithorhynchus, Echidna, Sea-otter, (Enhydra marina) (of this form I have had the good fortune to dissect the joints in two specimens), Sloth, Orang, Walrus, Hyrax, and Pangolin. The two last mentioned animals are additions to the list which I have been enabled to make, in consequence of dissecting two specimens of each. In birds the ligament is said to be present with only one exception — the cassowary, — but in a specimen of Casuarius appendiculaia dissected by me, the ligamentum teres was present in both hip joints. In reptiles possessing limbs, and in amphibians, a band representing the round ligament is universally present. It is in the horse that we first get the glimpse of the true nature of the ligament, for in this animal it consists of two parts, one hidden within the joint termed the cotyloid portion, the other passes out of the cavity to join the linea alba at its junction with the pubes, hence it is termed the pubio-femoral portion. From this band the pectineus takes origin. In the ostrich the ligamentum teres has a true tendinous structure. It is dense and strong, contains a large quantity of yellow elastic tissue arranged in fasciculi as in the tendon of a muscle. In this bird it may measure three-fifths of an inch in transverse section. Many birds possess in their thigh a very extraordinary muscle known as the ambiens, full of interest on account of its remarkable course, variability, and relations. When fully developed it has the following attachments. It arises from the tip of the short anteriorly directed spine which is situated just above the anterior border of the acetabulum, and runs along the inner side of the thigh to the inner side 4 o LIGAMENTS. [Chap. hi. of the knee, where it is covered by the sartorius, which is above it in the former part of its course. Its thin tendon then crosses the knee, running in the substance of the fascial extensor tendon, just in front of the patella, to the outer side, where it joins the fibres of origin of the flexor perforatus digitorum. In the adult ostrich a section carried through the ace- tabulum so as to divide the ambiens at its point of origin, and the ligamentum teres, will show that the two latter are connected by fibrous tissue. In the winter of 1883, I was fortunate enough to obtain an ostrich chick, and a dissection of its hip joint showed clearly enough the ligament and a small muscular slip, parallel with the ambiens, in direct continuity, as shown in fig. 14. The actual specimen is preserved in the museum Fig. 14. — The femur of an Ostrich chick with the pectineus muscle and ligamentum teres in continuity. of Middlesex Hospital. The ambiens and the muscular slips mentioned above, are the representatives in birds of the mammalian pectineus. The hip-joint of a sphenodon was then examined. In this very curious lizard, as in lacertilia generally, the hip joint is of very simple character, and the muscle corresponding to the ambiens of birds and the pectineus of mammals arises by two heads, one from the lateral spine of the pubes, the other lies inside the capsule and gains an attachment to the head of the femur, thus corresponding in its relation with the joint, to the ligamentum teres of mammals and birds. Chap, hi.] LIGAMENTUM TERES. 41 The varying relation of muscle and ligament may be arranged in a tabular form thus : — 9/)/ // (Tendon of ambiens (pectineus) passes in- " c { side the capsule to the head of the femur. 0j jL . (Ligamentum teres and ambiens muscle Struthio . \ ,. ., I directly continuous. „ (Ligament in two parts, one continuous with 1 U ' \ the pectineus outside the joint. jr fLigamentum teres, a fibrous band carrying ( blood vessels to the head of the femur. There is no ligament in the body which can boast such an extensive literature, or has exercised more the ingenuity of physiologists and surgeons than the one we have been considering. Teleologists like Paleyf have been enraptured with this structure, and anatomists have ascribed to it wonderful mechanical resistance and uses. Alas ! in this, as in so many like cases, morphology demands for it a low level and determines it to be a vestigial and practically useless ligament. In this sense teleology is as poetry, but mor- phology as plain history. * For all that relates to this strange muscle consult Garrod's remark- able paper in Proc. Zool.Soc, 1873^.626. "On certain muscles in the Thigh of Birds, etc." t See especially his Natural Theology, page 55. CHAPTER IV. The Knee Joint and Ankle. The general relations of the fibro-cartilages of the knee joint in man are familiar to every student of human anatomy. It must sound somewhat startling to him, how- ever, to learn that they are metamorphosed muscles and tendons. The history of the mode by which they have acquired so remarkable a position is replete with interest. The interarticular fibro-cartilages. — If one of the tailed batrachians be examined, and Menobranchus lateralis will serve as an excellent example, we shall find the knee joint deficient in crucial ligaments and interarticular fibro- cartilages, so that this animal forms a very good starting point. On looking closely into the myology of these amphibians, a curious fact at once becomes evident; in mammals, and in man especially, the muscles destined for the pes take origin, with very few exceptions, from the tibia and fibula; in these animals they arise almost exclusively from the femur. This is significant, and clearly indicates that it is in the forms filling up the gap between Urodele batra- chians and the lowest mammals that we must seek for an explanation of the change. In our first ascensive step we are encountered by the frogs, who startle us with the richness of the musculature of their limbs, for in many respects myological differentia- tion in them surpasses that which pertains in man. On examining the knee joint of a frog, it will be found to possess interarticular fibro-cartilages and crucial liga- ments, but these structures differ in a very important manner from those of mammals. In the first place, the interarticular fibro-cartilages are directly continuous with the tendons of muscles; the inner Chap, iv.] KNEE JOINT AND ANKLE. 43 cartilage is in direct continuity with a muscular mass corresponding to the semi-membranosus. This muscle is usually described as being inserted into the back of the tibia, but the main mass will be found inseparably asso- ciated with the cartilage. The external fibro-cartilage is not well formed, and seems rather to be a continuation of the inner one than as representing a discrete element of the knee joint, such as we find in higher forms. The question at once suggests itself — Why should so much difference prevail in the anatomy of this joint in animals so closely related as the tailed and tailless ba- trachians ? Dr. Gadow,° in his instructive paper on "Comparative Myology" puts the matter very beautifully thus : — " We must remember that the change of an aquatic ani- mal, which used its limbs simply like paddles, into a semi- aquatic and partly terrestrial, and therefore crawling, creeping and running creature, involved a complete change of its muscles, and at a later period of its bony framework. In swimming animals, such as ceratodus and menobranchus, the longitudinal axis of the whole limb is nearly straight, whilst a terrestrial life necessitates the bending of a limb at several angles, and the development of more or less complicated joints." It is this flexion which explains how tendons outside the joint in animals whose limbs are nearly straight, get drawn into it when the limb is acutely bent. When the leg is extended, the anterior surfaces of the femur and tibia are in the same plane ; when the leg is flexed, the broad upper extremity of the tibia is brought into contact with the distal part of the posterior surface of the femoral shaft, and immediately in relation with the ten- dons of origin of the muscles destined for the foot, and in many cases with muscular tissue. Therefore in menobranchus, the axis of whose hind limb * Journal of Anatomy and Physiology, vol. xvi., p. 501. 44 LIGAMENTS. [Chap. iv. is nearly straight, the tendons are excluded from the joint, and arise from the femur. In the frogs, whose limbs are flexed, the tendons of origin of many of the leg muscles in- truding upon the joint get drawn between the opposed surfaces of the tibia and femur, constituting fibro-carti- lages, crucial or lateral ligaments, as the case may be. The lizards furnish the strongest possible evidence that the interarticular fibro-cartilages arise in this way. The iguana may be selected as a type. In the thigh we find a very remarkable muscle known as the femoro-caudal. It is exceedingly large and arises from the infero-lateral aspect of the caudal ver- tebrae : it is inserted by a large, broad, and strong Fig. 15. — The femur, with the femoro-caudal muscle, F., of an iguana. Its long tendon X passes downwards to the interarticular cartilage of the knee-joint. The cut ends of the muscles are those of the biceps, semi-membranosus, etc. tendon into the base of the trochanter on its extensor aspect. A little space before its insertion the muscle gives off, at right angles to its lower border, a long, thin, and delicate tendon, which passes down the thigh, on the inner side of the great sciatic nerve, to the popliteal re- Chap iv.] KNEE JOINT AND ANKLE. 45 gion, where it passes between the fibula and tibia to blend with the outer part of the interarticular fibro- cartilage of the knee joint. Fig - . 15. This would seem to shew that in lizards, at least, the in- terarticular cartilages are the modified tendons of muscles which have become interarticular, during- the modifications the joint has passed through in its evolution. For further information concerning the morphology of this extraor- dinary muscle, consult this reference. An extended survey of the relation of these parts in ani- mals from the lizards onward to man, leaves little doubt that the fibro-cartilages in the human knee joint are sur- vivals of the distal termination of the femoro-caudal and associated muscles. The crucial ligaments. It is fairly conclusive that the muscles in the region of the knee joint have been affected by very considerable disturbance For example, in primi- tive forms, like lepidosiren, the muscular layers, pass from the trunk and invest the limbs without any transverse segmentation occurring. In frogs, lizards, and crocodiles, a gap occurs in the muscular sheet, the proximal segments attaching themselves to the tibia, the distal muscles arising from the lower end of the femur. In this way the quadri- ceps extensor, sartorius, gracilis, the hamstring group, and the adductor magnus, become inserted into the tibia ; whilst the extensors and flexors of the digits, the peroneal group, gastrocnemius, plantaris, tibialis anticus and posti- cus, arise from the femur. Many of the muscles have shifted their insertion from the tibia and formed new attachments to the femur, e.g. the adductor magnus. On the other hand, many left their acquired origin from the femur and descended to new at- tachments on the tibia and fibula (migration). Hence, it is necessary to decide whether a given ligament of the knee joint, had its origin from one of the thigh muscles shifting * Journal of Anatomy and Physiology, vol. xx., p. 39. 46 LIGAMENTS. [Chap, iv, its insertion, or a leg muscle migrating to the tibia or fibula from the femur. Both these processes have been in operation, and in Chapter II. evidence was adduced to support the view that the b>t^aal lateral ligament was originally the tendon of the peroneus longus, which muscle has migrated from the femur to the fibula. Whilst the internal lateral ligament results from the adductor magnus muscle withdrawing its point of insertion from the tibia to the femur. There can be little doubt that the crucial ligaments have a similar origin, but it is not quite clear to which muscles they originally belonged. It is possible to limit the mus- cles to three or four to which they may have been attached. The long extensor of the toes and the tibialis anticus mus- cle arise in many animals from the femur when crucial ligaments are present, so that these may be excluded satis- factorily enough. The muscles on which the lot falls are the peroneo tibial, tibialis posticus, flexor longus and the extensor brevis digitorum. See table on page 48. The Synovial Membranes. — In the beaver the ar- rangement of the synovial membranes of the knee joint is after this fashion. There is a distinct synovial pouch for the patella and the patellar surface of the femur. The cavity for the articular surfaces of the femur and tibia is divided into two parts by the crucial ligaments and a septum of connective tissue, thus making three synovial membranes in all : one for the patella, and one for each femoral condyle and the corresponding articular surface on the head of the tibia. In man, however, this septum disappears, save a few strong fibres in the middle line, which pass from the lower border of the patella to the intercondyloid notch immedi- ately below the patellar facet. This band of tissue, with its lateral fringes, adipose and fibrous in structure, is denominated in human anatomy the ligamentum mucosum and the ligamenta alaria. In reality they are simply vestiges of the sac-wall of the bursa, originally developed between the patella and the femoral condyles. Chap, iv.] KNEE JOINT AND ANKLE. 47 Examine a human foetus at the third month, by dividing the knee joint vertically, the synovial membrane will be found arranged as follows : — The proper cavity ends on a level with the upper border of the articular cartilage of the femur. Immediately above this, and lying behind the quadriceps extensor muscle, is a large bursa, whose sac communicates with that of the knee joint proper by a small opening in its lower part. After birth, however, the diaphragm between the two cavities quickly disappears ; the synovial membrane of the joint and the bursa blend to form one common cavity. Thus it follows that the large cul-de-sac of synovial membrane extending under the extensor muscles ot the thigh is due to secondary causes, Fig. 16. — Vertical section through the knee joint of a human foetus at the third month. F. Femur; P. Patella; T. Tibia. I. Quadriceps bursa. II. Patella bursa. III. Knee joint proper. it has also been shown that this has occurred in connection with the patella, so that the knee joint, as seen in man, is made up of its own synovial membrane reinforced by two very considerable bursae. Viewed in full light, the evolution of the structures con- cerned in man's knee joint may be considered as resulting from the changes produced by the regression and migration of muscles to provide it with ligaments, the metamorphosis of tendon to furnish it with interarticular fibro-cartilages, and lastly the annexation of bursae to extend the dimen- sions of its synovial territory. 43 LIGAMENTS. [Chap. iv. For convenience, the origin of the various structures of the knee joint are arranged in a tabular form : — Internal External Internal — External Anterior Posterior Lateral Ligaments. Tendon of the adductor magnus muscle. Tendon of the peroneus longus. Fibro-Cartilages. j Modifications of the femoro-caudal, biceps and ( semi-membranous muscles. Crucials. ( There are no certain facts as to which muscle ( it originally belonged. Tibialis posticus muscle. (?) Synovial Membrane. (Original bursa between the quadriceps extensor The great cul-de-sac . \ . . . & y and the femur. Middle portion . . The original patellar bursa. Lig. mucosum and (The remains of the original sac-wall of the alaria . . . ( patellar bursa. The Anterior Annular Ligament of the Ankle Joint. The structures included under the above title are two in number, one is placed at the lower end of the leg, and is simply a transverse band of fibrous tissue passing from the tibia to the fibula and serving to retain the vertical portion of the extensor tendons. Although it is a structure very general, from amphibians to man, it has apparently no morphological significance. The other ligament is opposite the bend of the ankle joint, and is attached to the os calcis by its two extremities which are in contact where they arise from the bone, so Chap, iv.] LIGAMENTS OF ANKLE JOINT. 49 that they form an exceedingly narrow loop, which just admits of the passage of the extensor longus digitorum tendon; from the inner limb of the pulley a fibrous cord passes beneath the extensor proprius hallucis and tibialis anticus muscles, to be attached to the internal malleolus, blending- with the tibial attachment of the superior liga- ment as shown in fig-. 17. By studying the relations of this structure in a large number of mammalian forms, it turns out that the loop which encircles the extensor longus digitorum is the essential portion of the ligament, but the fibrous cord passing to the internal malleolus is entirely secondary, and exhibits an extreme vagary in the way it disports itself with regard to the extensor hallucis and tibialis anticus muscles. F.L.D. Fig. 17. — The inferior anterior annular ligament of the ankle joint of a monkey. F.L.D., extensor longus digitorum muscle. It is impossible to deal satisfactorily with the annular ligament of the ankle joint without taking into considera- tion the condition these ligaments present in birds, and as the inquiry has led me deeply into the subject of the devel- opment of the tarsus in the feathered tribe, and certain facts of some interest have come under notice, it will be necessary to go systematically into the matter. The Avian Tarsus and Annular Ligaments. The tibia, or more correctly tibio- tarsus, of a bird is a bone very characteristic of this class. It is a cylindrical bone, usually about half as long again as the femur. The So LIGAMENTS. [Chap. iv. proximal extremity presents two somewhat concave articu- lar surfaces for the femoral condyles; in front it is pro- duced into a prominent cnemial crest or process, which in a few cases (struthio and rhea) is ossified as an epiphysis. In the Dinosauria this process may be variously subdivided. The distal end of the bone is terminated by a trochlear or pulley-like articular surface. Not unfrequently there is an oblique bar of bone on the anterior face, just above the trochlea ; this osseous loop transmits some of the extensor tendons (fig. 18). The true composition of this bone will Fig. 18. — The ligaments of the ankle-joint of a fowl. I. The super- ficial; II. The deep ligament. be considered when dealing- with its development. The fibula of birds is usually imperfect, and presents itself as a slender osseous rod attached to the outer side of the tibio- tarsus. Its proximal end articulates with the outer con. dyle of the femur being- received in the fibular fossa; the distal end gradually thins and fuses with the tibio-tarsus. Although as a rule the fibula is shorter in the adult bird than the tibia, yet it may be of the same length as in some of the penguins. The segment beyond the tibio-tarsus is a compound of certain bones of the tarsus and metatarsus, hence it may be correctly referred to as the tarso-metatarsus ; the various metatarsals which compose it, fuse together to form a single bone. As the proximal part of the tarsus becomes confluent with the tibia, and the distal portion fuses with the meta- Chap, iv.] THE A VI AN TARSUS. 5i tarsus, it thus comes about that the joint between the tibio-tarsus and the tarso-metatarsus is in fact a mesotarsal articulation, really corresponding with the medio-tarsal joint of man, which exists between the os calcis and astragalus as its proximal elements, and the scaphoid and cuboid as the distal bones. The above statements are warranted by an appeal to the development of the parts in birds. Gegenbaur Fig. 19. — Leg of a chick at the fifth day of incubation (after Miss Johnson). Fe. Femur ; T. Tibia ; F. Fibula. The numerals refer to the digits. ( Uniersuchungen zur Vergleichenden Anatomie der Wibelihiere, Erstes Heft, " Carpus und Tarsus ") proved that the distal end of the tibia, which in the young bird is separated from the main bone by a suture, is not an epiphysis, for the cartilage in which it ossifies is separate ; this individuality in the cartilaginous state distinguishes it as an element, an epiphysis being a distinct ossification in a continuous cartilage. In order that the true nature of the parts shall be cor- rectly comprehended, it will be safer to give in brief outline e 2 52 LIGAMENTS. [Chap. iv. the development of the bird's leg, which has been worked out by Gegenbaur, Rosenberg-, Baur, and Miss Johnson. In the limb of a chick, five days old, the tissue is con- densed axially into a single mass, and the skeleton of the Fig. 20.— The leg of a Chick at the eight day of incubation, (after Johnson). Fe. Femur ; T. Tibia ; F. Fibula ; A. Astragalus (tibiale) ; C. Os calcis (fibulare) ; Ta. Tarsalia. The numerals refer to the digits. limb is produced by the subsequent elongation i ■ d seg- mentation of the mass. * " Development of the Pelvic Girdle in the Chick," Qitar. your. Mic. Science, vol. xxiii. (This paper contains numerous references of value). Chap, iv.] THE AVIAN TARSUS. 53 At this stage the limbs present the condition represented in fig-. 19. All the chief elements may be recognised, though they are completely continuous. The tarsus is continuous with the tibia and fibula above and with the metatarsals below. Five metatarsals are present. At the eighth day all the elements of the tarsus are at their most distinct and independent stage, though they are still united with one another, as well as with the tibia, fibula, and the metatarsals, by the condensed tissue of the groundwork of the tarsus (fig. 20). Later, the distal and proximal parts of the tarsus become separated, and the two proximal elements fuse together. Next, the proximal parts fuse with the tibia, which has grown more than the fibula, so that the latter no longer reaches the tarsus. The posterior lower edge of the tibia first becomes continuous with the proximal tarsal cartilage* while the anterior face of the latter gives off an upward process, the so-called "ascending process" of the astra- galus, which fits into a groove in the tibia, and remains lor a long time separate from it. At about the same time the distal part of the tarsus fuses with the metatarsals, first with the second, next with the fourth, and lastly with the third. All these processes take place while the tarsus is still cartilaginous. In the chick two ossific nuclei later make their appearance in the proximal piece of cartilage, the inner one representing the astragalus, the outer and smaller piece the os calcis. The distal piece of cartilage represents the tarsalia. In the majority of birds the astragalus sends up a long osseous " spike," which fits into a grove on the anterior surface of the tibia, as shown in fig. 21, and is commonly referred to as the ascending process of the astragalus. This spike has been shown by Morse to have an inde- * Anniversary Memoirs of the Boston Society of Natural History, i88o > Morse, On the Identity of the so-called Ascending Process of the Astragalus, with the Intermedium. 54 LIGAMENTS. [Chap. iv. pendent ossific centre, and he considers it to correspond with the intermedium of the typical tarsus. The ques- tion is one so full of interest, and as I wish to give an account of some observations of my own on the subject, it is well to refer in detail to Morse's paper. It appears that Professor Wyman sent a note to Morse, to the effect that he had found in the tarsus of an embryo heron, a long style-shaped bone, broadest at its distal Fig. 21. — The distal end of the tibio-tarsus of an Ostrich chick. A. Astragalus ; C. Os calcis ; I. The so-called ascending spike of the astragalus (intermedium) ; F. Fibula. extremity, lying in front of and at the distal end of the tibia, which he believed to represent the so-called ascend- ing process of the astragalus, but which had an independ- ent centre of ossification, and remained free from the other tarsal bones until the young had left the egg. Morse had previously shown that as the proximal series of bones become united, the fibula diminishes in size proportionally with the rapid increase of the tibia, becoming finally a splint-like bone with its attenuated distal end far removed from the tarsus ; the tibia, on the contrary, enlarges, so that its distal extremity equals in transverse diameter the two proximal tarsal bones. Chap, iv.] THE AVIAN TARSUS. 55 The intermedium, while occupying- its proper position between the tibiale and fibulare, and finally uniting- with them, becomes displaced, so to speak, by standing in front of the tibia. As proving the correctness of this assumption, it may be mentioned that the bone may be found occupying its true position in the tarsal series, and between the distal extremity of the tibia and fibula in the embryos of those birds in whom it is represented. An examination of the tarsus in the embryos of the Tern (Sterna hirundo), Petrel (Procellaria pelagica), Sea Pigeon (Una grylle), Eider Duck (Somateria mollesstma), Herring Gull (Larus argentatus), and in the Great Auk (Aptenodytes pennanti), showed that in these birds the intermedium is present as a separate bone. It lies at first in a line with the proximal row of tarsal bones, opposite the space be- tween the tibia and fibula. As the tibiale and fibulare coalesce the intermedium is crowded upwards and out- wards so as to occupy a position in front of the tibia. It is the last bone to unite with the confluent tibiale and fibulare. Since reading Morse's paper, I have had an opportunity of investigating the condition of the ascending process of the astragalus in a cygnet of Cygnus mutis, and in an ostrich and an emu chick. In the cygnet the "bony spike " was present as a separate ossification, exactly as Morse figures it. In the the case of Struthio and Dromoeus the process was just in the act of uniting with the astra- g-alus; on making a longitudinal section through the lower end of the tibio-tarsus the line of separation was clearly visible. These three specimens have convinced me that the ascending process of the astragalus is an independent element. I felt it was absolutely necessary to enter minutely into the details of the ossification of the bird's tarsus for two reasons: — (i) To show that the bony bridge on the lower end of the tibio-tarsus is an "adventitious" ossification, (2) to prove its relationship with the tarsus— these two con- ditions being essential to establish its identity with the in- ferior annular ligament of the mammalian ankle joint. 56 LIGAMENTS, [Chap. iv. In order to show that a great deal of misconception pre- vails as to the real nature of the oblique bar of bone, I will make the following- extract from Mr. Parker's paper, "On the Osteology of Balaniceps rex" (Trans. Zool. Soc, vol. iv. ) : — '• The inferior or distal end of the tibia is developed from a distinct osseous centre in young birds, which piece forms all the articular parts and sends upwards a wedge- shaped process in front, the seat of the ossification which makes the large, wide, oblique, tendon-like brid.^ Morse finds that this oblique tendon-like bridge in the heron and many other birds has no relation with the in- termedium, and I am convinced, from the examination of a very large number of birds, that the pulley in question is really the homologue of the loop which in mammals binds down the tendon of the extensor longus digitorum, and has its main attachment to the proximal row of tarsal bones. It is. therefore, an adventitious ossification and not an intrinsic element of the bird's tarsus. So far as the anterior annular ligaments of the bird's leg is concerned, it must be clear to any one who has fol- lowed the preceding account that the two structures differ very considerably in their mode of origin and in their rela- tion to each other. It must also be very evident that the upper ligament, which involves all the extensor tendons, is the homologue of the superior ligament of the mammalian ankle joint, but the deep or inferior band, ossified in some birds, which transmits the extensor longus digitorum ten- don, is the avian representative of the loop attached to the os calcis in most mammals, including man. It may per- haps be advisable to state that the fact of it often under- going ossification in birds is no objection to it being con- sidered as the homologue of a purely fibrous structure (and. so far as one knows, always so in mammals) for in the West African Lemur iPerodicticus potto) the anterior annular ligament of the wrist joint (which arises as the result of regression of certain portions of the short muscles of the hand > contains an osseous nodule, thus completing Chap, iv.] THE AVIAN TARSUS. 5 7 a bony tunnel for the flexor tendons, comparable in a certain measure to the osseous loop at the lower end of the bird's tibio tarsus. Admitting- then that the ligaments in the legs of birds and mammals are homologous, it necessarily follows that the question be asked, How do these structures arise, and what is their nature ? A full discussion of the mode of evolution of these two ligaments, would lead us far beyond the scope and inten- tion of this little work, the enthusiastic student is referred to the author's original paper dealing with this particular question. The principles involved are of great interest. Briefly the ancestry of the anterior annular ligaments may be stated thus : — i. The upper or vertical band arises from thickening of fibrous tissue the result of long continued strain. 2. The lower one represents an old attachment of the extensor longus digitorum muscle as seen in Iguana and Parson's Chameleon. Lastly as far as my dissections have extended, an anterior annular ligament is a structure peculiar to birds and mammals. * journal of Anatomy and Physiology, vol. xx., " Nature of Ligaments, part iv." Consult also Mivart, "Myology of Parson's Chameleon," Proceedings of the Zoological Society, 1870. CHAPTER V. The Ligaments of the Pectoral Arch. The term "pectoral arch " or "shoulder girdle " is applied to those elements of the skeleton which serve the purpose of attaching the anterior limbs to the trunk. The compo- nent parts of this arch or girdle in the adult, consist of the clavicles and scapulae with their connecting ligaments. It will be necessary to briefly recapitulate the leading characters of the ligaments, in order that we may fully understand their significance when dealing with the devel- opment of the parts and their morphology. i. The Ligaments at the Sternal (Ventral) End of the Clavicle. (a) The interclavicular. This is a T-shaped band of dense fascia uniting the clavicles across the median line and sending a vertical slip downwards to be attached to the top of the sternum. (b) Capsular ligaments. {c) An inter articular fibro- cartilage, interposed between the sternum and the clavicle. Occasionally a cartilagin- ous, or an osseous nodule may be found in the lower end of these ligaments, named the epi-sternal bone. 2. The Ligaments at the Acromial End of the Clavicle. (a) The capsular, an insignificant structure. (b) An interarticular cartilage, this is occasionally absent. 3. The clavicle is connected with the costal cartilage of the first rib by the rhomboid ligament, and with the coracoid process of the scapula by means of the conoid and trapezoid ligaments. 4. Closely connected with the preceding group are two ligaments passing from the scapula to the humerus; one Chap, v.] PECTORAL ARCH. 59 the coraco- humeral lying outside the capsule of the shoulder joint, the other the gleno-humeral which may be inside the joint, but in man is usually situated between the capsule and the synovial membrane. 5. The coraco- acromial ligament is a very strong- band of fascia passing from the outer side of the coracoid to the acromion process. 6. The cost 0- coracoid ligament, a dense band of tissue marking the limit of the costo-coracoid membrane and running from the tip of the coracoid to the costal cartilage of the first rib, and thence to the sternum. Frequently it contains cartilage nodules. It is stronger and more pronounced in many of the quadrumana than in man, but in him it is frequently very conspicuous. Fig. 22.— A general view of the clavicular ligaments and associated structures. I. The interclavicular ligament ; II. Interarticular fibro- cartilage; III. Rhomboid ligament; IV. The subclavius muscle; V. The conoid and trapezoid ; VI. The gleno-humeral ligament ; VII. The costo- coracoid membrane and ligament; VIII. The minor pectoral muscle; Its tendon as the coraco-humeral band. The dotted line indicates the situation of the coraco-acromial ligament. The ligaments of the arch when considered morpho- logically readily fall into two groups ; viz., those derived from its intrinsic elements, and those which arise from muscular regression or metamorphosis. To the first group belong the interclavicular ligamen^ the interarticular fibro-cartilages at each end of the 6o LIGAMENTS. [Chap. v. clavicle, the coraco-acromial and the costo-coracoid ligament. Those which constitute the second group will be considered later. The mode of origin of the ligaments derived from intrinsic elements of the arch are so involved in the ontogeny of the various bones and their cartilaginous pre- cursors, that the development of the clavicle and scapula must be detailed. Before doing so, however, it will be advisable to give an account of what may be considered the typical or generalised form of the cartilaginous arch. This consists essentially of a bar of cartilage which may be divided into three parts: — i. A dorsal segment repre- senting the scapula. 2. A ventral bar, the coracoid, forming with the scapula a socket for the reception of the proximal bone of the limb. 3. A rod or bar situated anteriorily to the coracoid, and named in consequence the precoracoid. These two segments agree in the circum- stance that they meet the sternum anteriorily and the scapula posteriorily, but the precoracoid is, as a rule, excluded from the glenoid fossa. Fig. 23. The bar of Fig. 23 — The shoulder-girdle of an ostrich chick. S. Scapula ; Sc. Supra-scapula cartilage; C. Coracoid; P. Precoracoid; E. Epicora- coid ; G. Glenoid fossa. cartilage which unites the ventral ends of the coracoid and precoracoid is termed the epicoracoid. From such a shoulder-girdle as this, we may derive any form, from chelonians onwards to man, by the two pro- cesses of addition and suppression, the suppressed parts usually being represented by fibrous tissue. This type form is also valuable, when we wish to com- pare homologous parts of the pectoral with the pelvic girdle, for although in the adult the differences are Chap, v.] OSSIFICA TION OF CLA VICLE. 6 1 considerable, yet in the cartilaginous state they are very similar. From the type-form, man's primitive pectoral arch differs only in this particular. The continuity of the coracoid and precoracoid bars is interrupted. This defect in the pre- coracoid is due to the intrusion of the clavicle as a secondary element in the arch. In the case of the coracoid it is a consequence of partial suppression. The ossification of the clavicle. — This bone is the first in the skeleton to ossify. It commences about the sixth week of intrauterine life by a deposition of ossific matter, at a spot corresponding- to the middle of the shaft of the clavicle, it may be detected before the formation of the precoracoid cartilage, consequently it must be regarded as a membrane bone. Later, the extremities of the clavicle invade the cartilage and serve as an example of a bone originally arising in membrane becoming secondarily engrafted on cartilage and assuming a composite charac- ter. This is interesting, for it serves to explain what would otherwise be singular, viz., the occasional trans- mission through the middle of the clavicular shaft of some twigs of the descending cervical nerves, and at times a cephalo-jugular vein. There can be little doubt that these represent nerves and veins which should pass be- tween the clavicle and precoracoid bar in forms where the latter exist as distinct entities. When these elements fuse to form man's composite clavicle, the nerves and veins are involved in the ossification. The clavicle is completed by an additional ossification at the sternal end. The nucleus is usually detected about the eighteenth year, and unites with the shaft at twenty- two. The interarticular fibro-cartilages at the sternal and acromial ends of the clavicle, arise from the regression of the inner and outer extremities respectively of the cartila- ginous precoracoid. The nodule of bone known as epi- sternal, is developed in this remnant of the precoracoid bar. 62 LIGAMENTS. [Chap. v. The ossification of the scapula. — This bone is pre- formed in cartilage. The primary centre appears about the third month of intrauterine life near the base of the spine, and quickly extends itself throughout the cartilage, so that at birth we only find a thin strip of cartilage along the vertebral border; the glenoid fossa, the coracoid and acromion processes are yet unossified. In the first year after birth a nucleus appears for the coracoid, and from this centre ossific matter extends into the glenoid fossa and as far backwards as the notch in the upper border of the scapula ; ossification only extends a short way into the coracoid bar, the chief portion being represented by the costo-coracoid ligament. At the fifteenth year the coracoid becomes ankylosed to the scapula and two nuclei appear for the acromion ; the supra-scapular cartilage ossifies from two centres, one opposite the spine, the other at the inferior angle. A thin lamina is added later to the glenoid fossa, and by the twenty-fifth year the ossification of the scapula is complete. The most remarkable feature in the development of this bone is the early appearance of the coracoid centre, and the long persistence of the strip of cartilage at the vertebral border. With regard to the coracoid, many look upon its nucleus as a primary one, even then it differs from the usual course of events, for its appearance is late, so that considering it from either point of view its de- velopment is anomalous and suggestive. This opinion is confirmed when the part is studied morphologically. On comparing the scapula of a human fcetus at the seventh month with the type-form, we shall at once see that the strip of cartilage along its vertebral border is homologous with the supra-scapular cartilage, which in batrachians persists throughout life, merely becoming im- pregnated with lime-salts. There is substantial agreement in that the glenoid fossa is formed by the confluence of the coracoid and the body of the scapula. The coracoid differs from the type-form, in the fact that it is repre- Chap, v.] OSSIFICATION OF SCAPULA. 63 sented by bone in only the scapular third, the sternal portion being - replaced by fibrous tissue. In order to elucidate the nature of the interclavicular ligament, we must study Omithorhynchus. The complex pectoral arch of this low mammal is shown in fig - . 24, where S represents the scapula, tipped with its thin streak of supra-scapular cartilage, C is the coracoid, which in no group of mammals other than the monotremata is prolonged as bone to the sternum. The precoracoid is represented by an osseous nodule resting on the sternum anterior to the coracoid, and as it articulates with the latter it contains also the epicoracoid element. The clavicles, C", are supported at their inner extremities by a large T-shaped bone, of membranous origin, the inter- clavicle. It is certainly remarkable that in order to find a similar skeletal element we must descend to lizards. Fig. 24. — The shoulder girdle of Omithorhynchus. S. Scapula ; C. Coracoid ; C". Clavicle ; E. Epicoracoid ; X. The interclavicle. A general survey of the pectoral girdle teaches us that fundamentally it consists of a dorsally placed scapula and two ventral segments, coracoid and precoracoid. The scapula and coracoid together form a glenoid fossa. The scapula is in relation with coracoid and pre-cora- coid, whilst the two latter are united ventrally by a 64 LIGAMENTS. [Chap. v. bar of cartilage, the epicoracoid. In the majority of types the precoracoid becomes divided in its centre by the clavicle, and its extremities are invaded by the growing- bone. Its median extremity is in man represented by the interarticular fibro-cartilage between the clavicle and sternum, occasionally a little cartilage or osseous nodule is present in them (the episternal bones of Breschet). The scapular remnant of the precoracoid finds a representative in the interarticular fibro-cartilage between the acromion and clavicle (Parker's confusing mesoscapular segment). The coracoid in man is a stunted process, which early fuses with the scapula, but its prolonged condition in frogs, birds and monotremes, is represented by the long costo-coracoid ligament running in the free border of the membrane of that name, and presenting chondral particles at its sternal attachment. The interclavicular ligament is the fibrous representative of the bone of that name in lizards and in monotremes. The coraco-acromial ligament. — This is a dense band of fascia passing from the acromion to the coracoid process, and is supposed by teleologists to have been de- signed to prevent upward dislocation of the head of the humerus. It has, however, a far more interesting signifi- cance. In the scapula of the sloth the acromion process sends a long hook-like piece of bone to meet the cora- coid. In the foramen triosseum of very many birds the scapula joins the coracoid above as well as below the foramen. In man and other mammals this portion of the acromion is represented by the coraco-acromial ligament. The view is supported by the fact that in the foetus of many mammals a tract of cartilage may be detected in the same situation. The disposition of these parts in the sloth is repre- sented in fig. 3. The nature of the transverse ligament is discussed on page 6. The following ligaments connected with the pectoral girdle remain for consideration, viz., the rhomboid, the Chap, v.] THE GLENO-HUMERAL. 65 conoid and trapezoid, the gleno humeral and the coraco- humeral. The rhomboid ligament. — The costo- clavicular or rhomboid ligament is attached to the cartilage of the first rib near the sternal end, and passes obliquely upwards, backwards and outwards to a rough impression on the under surface of the clavicle near its inner extremity. The coraco-clavicular ligament. — This ligament consists of two parts, an inner fasciculus termed the conoid, and an outer the trapezoid ligament. The conoid fasciculus is attached by its base to the conoid tubercle of the clavicle, whilst the apex is inserted into the root of the coracoid process. The trapezoid portion passes from the upper surface of the coracoid to a ridge on the under aspect of the outer extremity ol the clavicle. Be- tween the two ligaments a bursa may frequently be de- tected. The gleno-humeral ligament. — This ligament is usually described as an accessory band, springing from the edge of the glenoid fossa at the root of the coracoid process, and passing downwards to the lesser tuberosity of the humerus. It runs more or less parallel with the long tendon of the biceps, and may be identified in the usual conditions of the parts as a bulging into the joint, separated from the synovial cavity by the thin serous membrane only. Occasionally the ligament is completely surrounded by synovial membrane and is as prominent a structure in the joint as the biceps tendon ; indeed it then exists as a veritable lig amentum teres. This condition is far more frequent in the foetus than in the adult. It also exists as a strong and well defined ligament normally in a very large number of mammals. We shall return to this subsequently, the general character of the ligament is shewn in fig. 25 from the beaver, Castor canadensis. The coraco -humeral ligament. — This is a strong wide band passing from the root and outer border of the coracoid process to the neck of the humerus above the F 66 LIGAMENTS. [Chap. v. great tuberosity, it is intimately associated with the capsule of the joint. The morphology of the subclavius muscle. — In man the subclavius muscle arises from the first costal arch at the junction of the rib with its costal cartilage, close to the costo-clavicular (rhomboid) ligament. The fibres pass outwards and upwards arranged in a prismatic manner to be inserted in a groove on the under surface of the clavicle extending as far as the interval between the conoid and trapezoid ligaments lying beneath the costo-coracoid mem- brane. Fig. 25. — The shoulder joint of a Beaver {Castor canadensis), to show the large size of, G. The gleno-humeral ligament; S. Scapula; H. Humerus; B. Biceps. Its most important variations are the following : — 1. It may be inserted into the root of the coracoid process instead of the clavicle. 2. The muscle may divide its insertion, one part passing to the clavicle, the other ending in the root of the coracoid process. In some instances a few fibres may be inserted into the conoid ligament. 3. Occasionally it is attached to the coracoid process, and a bursa exists between the tendon and the coracoid, whilst a prolongation may pass from this tendon to the humerus. Other variations are Chap, v.] SUBCLAVIUS MUSCLE. 67 described, but these three are alone of importance at present. Leaving- man for the present and descending- to the amphibia we shall find that the interesting- form Mew bronchus lateralis presents about the shoulder a very distinct muscle, having the following- attachments: — It arises from the ventral surface of the long- precoracoid cartilage and passes backwards to be inserted into the head of the humerus, between the deltoid, pectoralis major and supraspinatus muscles. St. George Mivart in his paper on the myology of this creature, identifies the muscle as the subclavius. Whether it deserves this name or that of epicoraco- humeral is a matter Fig. 26.— The subclavius of a bird. H. Humerus. S. Subclavius. T. Its tendon. C. Clavicle. Co. Coracoid. of little moment, but its situation and points of attach- ments are, for the purpose of the argument, of great importance. It is probable that this muscle is the ancestor of the subclavius of human myology. If the shoulder joint of a carinate bird be dissected, a muscle will be found arising from the keel and median portions of the body of the sternum, as a broad fiat sheet, which passing forward ends as a strong tendon, in order to traverse the foramen formed by the union of scapula, coracoid and clavicle; emerging from the osseous ring it is inserted into the dorsal surface of the head of^the humerus, in the immediate neighbourhood of the attach- ment of the pectoralis major muscle. The tendon after F 2 68 LIGAMENTS. [Chap. v. threading - the foramen triossium lies within the capsule of the joint. This muscle enjoys a variety of names, e.g., pec t oralis secundus, levator humeri, and subclavius. Fig". 26. Prof. Rolleston proved most conclusively that this muscle, whose main function consists in raising the wing, is homologous with the mammalian subclavius and not with the pectoralis minor. Having discussed the morphology of this muscle, an attempt will now be made to render it responsible for the rhomboid, coraco-clavicular and gleno-humeral ligaments. In many mammals the subclavius arises from the sides of the sternum and the costal cartilage of the first rib, replac- ing" the rhomboid ligament, and it is reasonable to regard the rhomboid ligament as arising from the regression of the sternal fibres of the subclavius. We must now consider the relation of the subclavius muscle to the coraco-clavicular ligaments. When describ- ing the variations to which that muscle was liable, we noted that it may be inserted into the coracoid process, or partly into the coracoid and clavicle, and in some instances into the fibres of the conoid ligament; under these con- ditions it seems the most probable explanation that these ligaments arise from regression of a portion of the sub- clavius. If their relation to the gleno-humeral be ex- amined, it will be found that the two sets come into relation at the base of the coracoid process. Seeing" that the gleno-humeral ligament is closely associated with the coraco-clavicular set, and these are intimately connected with the subclavius muscle, the inference would be that the gleno-humeral ligament is the divorced tendon of that muscle. This is further supported by a case reported by Walshamy in which the subclavius was inserted into the coracoid process, and a tendinous expansion passed thence to the capsule of the shoulder joint. Between the tendon and the coracoid a bursa existed. * Linnean Soc. Trans., vol. xxvi., 1868. f St. Bartholomew's Hospital Reports, vol. xvi. Chap, v.] SUBCLAVIUS MUSCLE. 69 Previously the only gleno-humeral known, besides that of man, was the structure so commonly referred to as "the ligamentum teres in the shoulder joint of the frog-." If the view which would regard the gleno-humeral band as originally belonging to the subclavius is correct, a gleno- humeral ligament ought to be present with tolerable frequency, especially as the subclavius is so constant in mammals and in other forms above fishes. The examina- tion of a very large number of mammals has shewn that a gleno-humeral, as well-developed as that represented in Fig. 25, is present very frequently, in many instances it is as thick as the tendon of the biceps. In this condition it exists in many Quadrumana, Lemuridae, and Insectivora, in all Rodentia, and many of the Marsupialia. At the end of the book a tabulated list of the various mammals is given in whom the ligament has been detected. It is curious that those mammals which have no liga- mentum teres in the hip joint have not a gleno-humeral ligament in the shoulder :— The Seal, Elephant, Rhinoceros, Walrus, Ornithorhynchus, Echidna, Sea-otter (Enhydra marina) (of this form I have had the good fortune to dissect the joints in two specimens), Sloth, Orang, Hyrax, and Pangolin. 1. Every mammal in whom I have up to the present time found a gleno-humeral ligament, likewise possesses a ligamentum teres in the hip. 2. Those mammals in whom a ligamentum teres is absent, also lack a gleno-humeral band. 3. In those forms in whom the gleno-humeral is well developed, the ligamentum teres of the hip is also very thick and strong. The reasons for regarding the gleno-humeral ligament (or gleno-humeral band when it merely appears as a thickening in the capsule) as the divorced tendon of the subclavius muscle may be briefly enumerated. 1. All amphibians which possess the equivalent of the subclavius — the epicoraco- humeral of Mivart — in a /O LIGAMENTS. [Chap, v. well-developed form, lack the gleno-humeral liga- ment. 2 In birds the subclavius muscle (under the name of levator humeri) reaches its maximum of development, and passes through the shoulder joint as a tendon to be inserted into the humerus; no gleno-humeral ligament is present. 3. In mammals the subclavius is a very constant muscle, and a gleno-humeral ligament is present in a very- large number of them ; when not represented as a free ligament it may often be detected blended with the capsule of the shoulder joint, lying betrceeii it and the synovial membrane. Viewed in the full bearings of this speculation the history of the subclavius is as instructive a muscle as could well be. Commencing with menobranchus we see it as a muscle arising from the precoracoid, and at its insertion envelop- ing the outer aspect of the head of the humerus, being muscular throughout. Next we see it luxuriating in full perfection in flying birds, its distal end metamorphosed into tendon, per- forming the laborious and important function of raising the wing. Lastly, in man it becomes reduced to almost insignificant proportions lying as a small second rate muscle under the clavicle and representing in its retirement the middle portion only of the bird's levator humeri, its proximal end degenerated into a uniting band to connect the clavicle with the first costal arch, whilst its outer end is represented by the coraco-clavicular ligaments, and the small insig- nificant band, so far as function is concerned, known as the gleno-humeral ligament. In some cases in man the muscle may abort. The coraco-humeral must now be disposed of. In man it is usual for the pectoralis minor muscle to be in- serted into the upper surface of the coracoid process near its outer border ; a bursa frequently intervening between Chap, v.] 1HE CORACO-HUMERAL. 71 the tendon and the coracoid. Frequently it happens that the tendon glides over the coracoid process and gains an insertion into the great tuberosity of the humerus. In very many monkeys this is the normal condition. This induced some anatomists to believe that the gleno-humeral ligament was the tendon of the pectoralis minor muscle. Their view of the matter is invalidated on two points. 1. The tendon of the pectoralis minor when it takes this course lies out- side the capsule, whereas the gleno-humeral ligament is situated between the capsule and the synovial membrane. 2. The tendon of the levator humeri is inside the capsule, and this muscle of the bird is homologous with the sub- clavius. Lastly, the relation of the pectoralis minor to the capsule in certain cases occurring in man and monkeys, induces me to believe that the coraco-humeral ligament is the tendon of that muscle transformed into a fibrous band. The ancestry of the various ligaments of the pectoral arch may be arranged in two categories : — Those repre- senting skeletal elements and those derived from muscles. 1st Group. The ventral end of precoracoid . The interarticular fibro-cartilage between clavicle and sternum and the so-called episternals. The scapula end of ,, . . Acromial interarticular fibro-car- tilage. Coracoid Coracoid and costo-coracoid liga- ment. Coracoid extension of acromion . Coraco-acromial ligament. Coraco-scapular bridge, (sloth, Transverse ligament, etc.) Interclavicular bone, (lizards and Interclavicular ligament, monotremes) 2nd Group. Sternal end of subclavius . . . Rhomboid ligament. Humeral ,, ,, ... Conoid, trapezoid and gleno- humeral ligaments. Humeral end of pect. minor . . Coraco-humeral ligament. 72 LIGAMENTS. [Chap. v. The Ligaments of the Pelvic Girdle. As in the case of the shoulder girdle so with regard to the pelvis, a study of the hard parts is not sufficient to enable us to comprehend in its fulness their morphology. The various ligaments in connection therewith must be con- sidered. For this purpose the ancestry of the following structures will be detailed. The great and small sacro-sciatic ligaments, the triangu- lar ligament of the urethra, and the sub-pubic ligament. The great and small sacro-sciatic ligaments. — When the tendons of origin of the hamstring muscles are traced to the tuberosity of the ischium, they may by a very little dissection be seen to pass onwards into the great sacro- sciatic ligament, and by pulling on the muscles the ligament is tightened. When traction is made on the biceps the coccyx can be made to move on the sacrum. In fact the ligament may not unfairly be described as a tendinous ex- pansion of these muscles. This relation of the ligament to the hamstring muscles was first pointed out by Mr. Morris and the relation of the parts are shewn in a figure. Among the Ruminantia this ligament is replaced by a broad, strong tendon. The lesser sacro-sciatic ligament has a very different his- tory. This band of fibrous tissue passes from the spine of the ischium to the side of the sacrum. In the sloth we find the spine of the ischium extending to the sides of the pseudo- sacral vertebra, and bounding the great sciatic foramen. Fig. 27. In the armadillos this is an extensive piece of bone. Except among the edentata including the extinct mastodon and megatherium in mammals this bone is re- placed by fibrous tissue. This ligament should be referred to as the ischio-sacral ; those vertebrae into which it is inserted are pseudo-sacral. * Anatomy of the Joints of Man, 1S79. Chap, v.] TRIANGULAR LIGAMENT. 73 The triangular and sub-pubic ligaments. —These are somewhat puzzling structures and the explanation now offered to account for them, la^ks that amount of almost absolute proof which has been brought to bear upon so many of the ligaments which have been dealt with in this work. The matter is, however, not devoid of interest, and its consideration will serve to show how true is the remark ''that morphology is the very soul of anatomy." Fig. 27. — The pelvis of a sloth, Bradvpus didactylus, the small sacro- sciatic (ischio-sacral) ligament replaced by bone. (After Albrecht). In 1883, Professor Albrecht among his many contribu- tions to morphological anatomy, published a paper on the Pelvisternum of Edentata and endeavoured, but unsuccess- fully, to shew that there is an arrangement of parts in some animals, whereby anatomical relations similar to those of the sternum, in the anterior pare of the trunk, are repro- duced in the pelvis. * De Bulletin dc V Academic royale de Belgique, t. vi., nos. 9-10. 74 LIGAMENTS. [Chap. v p Albrecht in this paper draws attention to the existence of an independent ossicle in the pelvis of the sloths and armadillos, indicated in fig - . 27 by the letter X, which he names the pelvisternum. The existence of this ossicle admits of no doubt, I was able at once on reading- Albrecht's observations to confirm them in a chlamydophorous and three armadillos. In 1886 Dr. J. R. Anderson showed that Prof. Owen knew of the presence of this ossicle in the Potoroo, Hypsiprymnus, and he has himself detected it in the beaver and in phacochcerus. Anderson then cautiously pointed out that it is quite possible that the triangular ligament of the human urethra may be the fibrous representative of this bone. It is difficult to imagine this ossicle as representing anything more than the sub-pubic ligament, for the triangular liga- ment is a broad tract of tissue as compared with this bone. Beyond mentioning these views I shall not make any further observation, for the whole subject requires investi- gation with an abundant supply of material. * Proceedings of the Zoological Society, 1886, " On the Pelvisternum." CHAPTER VI. The Ligaments of the Appendicular Elements of the Skull and of the Vertebral Column. The appendicular elements of the skull are represented in the early embryo by three rods of cartilage named from before backwards the Eustachian, Meckelian and hyoidean bars. A great deal of interest surrounds these cartilages, for the last one is the undoubted representative of the branchial cartilage of a fish, the Meckelian bar is homo- logous with the lower jaw of certain fish whilst the anterior one represents the pterygo-quadrate bar of Elasmobranchs. They are diagrammatically shewn in the accompanying drawing fig. 28. In the course of development they undergo a most remarkable metamorphosis which will be explained on reference to fig. 29. The Eustachian bar becomes ossified at its distal end to form the internal pterygoid plate, which later fuses with the ala of the sphenoid and eventually with the lingulum. The middle segment remains as the cartilaginous portion of the Eustachian tube, the posterior and smallest piece degenerates into ligament and remains permanently as a connecting bond between the cartilage and the malleus. The Meckelian bar is transformed thus : — Its distal end is ossified and gives origin to all that portion of the inferior maxilla which lies anteriorly to the mental foramen, round the outer side of the bar, nuclei, fi\ e in number, are deposited to form the inferior maxilla; their names are the dentary, angular, coronoid, condyle and splenial. For a long time the cartilage occupies a depression in the bone later recog- nised as the mylo-hyoid groove, but eventually it disap- pears leaving a fibrous band known as the long internal lateral ligament of the jaw, extending from the inferior 76 LIGAMENTS. [Chap. vi. dental foramen to the spine of the sphenoid bone. The proximal segment becomes the malleus. The hyoidean bar is somewhat complicated. Its distal end fuses with the fellow of the opposite side and with the remains of a fourth arch to form the hyoid bone ; from the Fig. 28. — The Chondro-cranium of a human foetus at the third month with its associated cartilaginous bars. FXP. Fronto-nasal plate; E # Eustachian bar; If. Meckel's cartilage ; S. Styloid cartilage ; H. Hyoid bone. Fig. 29.— The later modification of the bars. P. Pterygoid bone ; E. Eustachian tube ; M. Malleus; I. Incus; T. Tympano-hyal ; S. Styloid cartilage; L\ Long internal lateral ligament of jaw; L. Stylo-hyoid ligament. lesser cornu to the styloid process it is, as a rule, repre- sented by a ligament, the stylo-hyoid. Above this point we get the styloid process, but for some months after birth it remains as cartilage, movable upon a nodule of Chap, vi.] VERTEBRAL COLUMX. 77 bone, discovered and named by Prof. Flower the tympano- hyal. Above, the cartilage is terminated by the incus, which is continuous with the tympano-hyal by means of a fibrous band. The Ligaments of the Vertebral Column. The ligaments which bind the vertebrae tog-ether may be divided thus: — 1. Those common to all the vertebrae. 2. Those which are confined to two vertebrae. To the first group belong the anterior and posterior common ligaments, and the supra-spinous ligament. The second group claims the intervertebral discs with the ligamentum conjugale costarum, the interspinous and inter- transverse ligaments, and the ligamentum subflavum. The anterior common ligament. — This consists of a longitudinal band of dense fibres placed on the anterior aspect (ventral) of the vertebral column, extending from the under surface of the basi-occipital bone, with a slight break at the atlas, to near the middle of the sacrum ; here it becomes scarcely distinguishable from the periosteum, but as it nears the coccygeal vertebrae it again becomes prominent, and ends at the tip of the coccyx. The super- ficial fibres extend over several vertebras (four or five), the intermediate ones pass over two, whilst the deepest fibres only connect adjacent vertebrae. The posterior common ligament Lying on the dorsal surfaces of the centra of the vertebrae, and extending from the upper surface of the basi-occipital bone to the lower end of the sacrum, is a fairly broad fibrous band. The upper part of the ligament, that which lies between the occipital bone and the axis, is often designated the posterior occipito-axial ligament. The more important features of the posterior common ligament are these: — In the neck it extends quite across the bodies of the vertebrae, in the yS LIGAMENTS. [Chap. vi. thoracic and lumbar regions it is broader opposite the intervertebral discs than at the middle of the bodies of the vertebrae; between these narrower portions and the ver- tebral centra is some loose connective tissue containing - a venous plexus; the ligament is only connected with the spine at the intervertebral discs and the corresponding- margins of the vertebrae, presenting throughout its course a series of attached and unattached segments. The supra-spinous ligament. — This consists of bundles of longitudinal fibres which form a continuous cord from the seventh cervical vertebra to the sacrum. It serves to connect the tips of the spinous processes together. Fig. 30. — The intervertebral disc of a seal with the anterior and posterior conjugal ligament. X' and X". We must now consider the second group : — The intervertebral discs. — These are plates composed of fibrous tissue arranged in laminae externally, with a pulpy nucleus internally. They are situated between the centra of the vertebrae beginning at the axis and ending in the adult at the sacrum, but in the young child they extend even to the coccyx. The outer or circumferential portion of each disc is composed of laminae of white fibro-cartilage arranged concentrically, which run obliquely between the vertebrae and are firmly attached to both. The direction of the fibres is reversed in successive layers. The centre of the disc is made up of pulpy substance which when exam- ined microscopically is seen to be composed of fine fibrous Chap, vi.] INTERVERTEBRAL DISCS. 79 tissue containing- cells. The ventral edge of each disc is firmly attached to the anterior common, and the dorsal bolder to the posterior common ligament. In many ani- mals, notably the seal, a very strong rounded band glides Fig. 31. — Ligamentum conjugate costarum, anterius and posterius, of a foetus at the seventh month. across the dorsal aspect of the disc, as shewn in fig. 30. This serves to connect the heads of the corresponding ribs and the ligament is provided with a synovial membrane. A band also passes along the anterior face of the disc, and may Fig. 32. — An intervertebral disc of an adult shewing the lig. conj. cost. post, incorporated with the disc. Its ends as the interosseous of the ribs. The anterior one represents the middle fasciculus of the stel- late ligament ; N. remains of the notocord. be named the anterior conjugal ligament. In the human foetus similar bands can be dissected from the dorsal and ventral aspect of the discs. The posterior band is continu- 5u LIGAMENTS. [Chap. vi. ous with the interosseous ligament of the rib-head, whilst the anterior becomes later the middle fasciculus of the well known stellate ligament of the costo-vertebral articula- tion. After birth the intermediate portion of these ligaments become incorporated with the intervertebral discs. The ac- companying figures represent the ligaments in a foetus at the seventh month and their fate in the adult. Figs. 31 and 32. The ligamentum conjugale costarum was first described by Mayer* of Bonn in 1S34 in various mammals. It has escaped the attention of most anatomists. Prof. Humphry in his well known work on the skeleton merely refers to Mayer's observation in a foot note ; Cleland unaware of Mayer's paper described the ligament in a seal in the Edinburgh New Philosophical Journal, 1858. These researches were unknown to me, when in 1S82 I noticed the ligament whilst dissecting a seal. Mr. Henry Morris then drew my attention to Cleland's papers. Later I examined the con- dition of this structure in a large number of mammals, including the horse, lion, otter, fox, dog, weasel, squirrel, rabbit, calf, sheep, badger, pig, ape, etc. The ligament is best developed in that part of the thoracic region of the spine where the ribs articulate with two vertebral centra. f The credit of drawing attention to the existence of this ligament in man rests with me, if we exclude the transverse ligament of the atlas, to be considered in the succeeding pages. It is very remarkable, but Hunter knew of this con- jugal ligament. Mr. Henry Morris requested the admirable assistant at the College of Surgeons, William Pearson, to prepare some dissections of this structure and having his attention thus drawn to the matter, Pearson shewed me an old preparation labelled Hmiterian. It is the dissection of two thoracic vertebrae of a horse so arranged as to shew the posterior ligamentum conjugale costarum. * Meckel's Arch. f For further details the student may consult journal of Anatomy and Physiology, vol. xviii., p. 225. Chap, vi.] LIGAMENTA SUBFLAVA. 8i The interspinous and intertransverse ligaments. — These are fibrous cords resulting from the regression of muscles of the same name, occupying the spaces their names indicate, but best studied as muscles in the cervical region. The ligamenta subflava.— Called also ligamenta intercruralia, they are plates of yellow elastic tissue alter- nating with the laminae of the vertebrae from the axis to the middle of the sacrum. They are so arranged as to be attached to the ventral surface of the laminae of the anterior vertebra, but are attached to the dorsal surface of the posterior laminae. A great deal of light seems to be shed on the nature of the ligamenta subflava by studying the vertebral column of a cartilaginous fish. In the skate these ligaments are Fig. 33 — A portion of the vertebral column of a skate shewing the interneural plates, N. represented by pieces of cartilage hexagonal in shape, known as the interneural plates. Fig. 33. The laminae are represented by a cartilaginous pedicle known as the neural processes, these only exceptionally join the neural spines. In the skate the ventral root of a spinal nerve leaves by a foramen in the neural process, whilst the dorsal root traverse the interneural plate. The elementary nature of the present book prevents me pursuing the argument to the fullest extent, but it may be mentioned that in man we have no alternation of the roots of the spinal nerves as in some of the cartilaginous fishes, G 82 LIGAMENTS. [Chap. vi. nevertheless, as in them, man's spine consists of segments composite in character, thus : — A centrum, with pedicles, laminae and a spinous process, succeeded by a disc composed of notochordal remains, and two plates of yellow elastic cartilage— the ligamenta subflava. The researches of American paleontologists, especially Cope's, go to support the view that the intervertebral disc may with good reason be regarded as representing a sup- pressed segment. In many of the remarkable batrachian forms, now extinct, unearthed in the Permian formation of Texas, the vertebral column possessed intercentra rivalling in size the true vertebral centra, and helping to form part of the bony boundaries of the neural canal. The first real attempt to attach a morphological value to a spinal ligament was that of Rathke, in his famous Entwicklung der Schildkroten, 1848, where, in a masterly piece of anatomical reasoning, he shews most conclusively that the os odontoideum of the Turtle is really the repre- sentative of the body of the atlas, and that the ligamentum suspensorium, a thin fibrous band passing from that process to the margin of the foramen magnum, is the remains of the notochordal sheath. Rathke had previously promulgated this doctrine in his Entwicklungsgeschichte der Natter, so far as the odontoid process is concerned. Cuvier, in his Recherches sur les ossemens fossiles, had previously identified the true homology of this bone by a study of its condition in the Turtle and in the Matamata Terrapin. Bergmann confirmed the state- ments of Rathke, and now the matter rests beyond all disputation after the investigations made into the subject as regards mammals by J. Miiller, Robin, and Hasse. Then, the fact that the nucleus pulposus in the middle of the invertebral discs is the remains, in part, of the noto- chord, lends additional interest to the matter. For this last satisfactory advance we are indebted to Luschka- It is a curious fact that, though in adult mammals, we Chap, vi.] LIGAMENTA SUBFLAVA. 83 find a definite ligamentum suspensorium attached only to the odontoid process, yet in birds, as Jiiger* has shown in Das Wirbclkcrpcrgehnk der Vogel, we find a suspensory ligament in many other parts of the column. In its most typical form the arrangement may be thus described: — Between the centra of contiguous vertebrae is a fibro- cartilaginous ring, which is continued inwards in the form of a disc which has free anterior and posterior faces, this portion is called the meniscus; this thins towards the centre, which is always occupied by a hole. The meniscus, when perfect, divides the space between the centra of the opposed vertebra into two synovial cavities. The central perforation in the meniscus is traversed by a ligament, which in the chick contains the remains of the intervertebral portion of the notochord. In- some instances the meniscus is reduced to a mere rudiment, in other cases it is united with the vertebral bodies, whilst in the caudal region, in its relation to the bodies of the vertebrae, it resembles an ordinary intervertebral disc. There seems to be no rule by which one could predict the presence or absence of this disc. In two specimens of Rhea [Rhea Americana) dissected by me, the discs and suspensory ligaments were absent, the vertebral centra coming into direct apposition. In the emu the meniscus is alone repre- sented, whilst in the pigeon, fowl, and similar birds the meniscus and suspensory ligaments are present in a perfect condition ; when carefully dissected, the neck vertebrae and the discs, alternating and threaded as it were by the sus- pensory ligaments, present a striking appearance. If the cervical vertebrae of a child be macerated until the intervertebral discs drop away, it will be found that the vertebral bodies will in many cases remain attached to each other by the thickened sheath surrounding the remains of the notochord; in this way it strikingly resembles the avian suspensory ligament. * Sitzungberichte der Wiener Akademie, 1858. 2 G 84 LIGAMENTS. [Chap. vi. In the skeleton of a modern bird not the least among the peculiarities of its bones is the shape of the articular surfaces of the pre-sacral vertebrae. Choosing a typical one from the middle of the cervical series, we find that the anterior articular surface of the centrum is slightly exca- vated from above downwards and convex from side to side: the posterior is the reverse of this, being convex from above downwards and concave from side to side. The anterior aspect resembles the curious surface on the trapezium for the metacarpal bone of man's pollex. The origin of this curious mode of articulation among the vertebrae of birds has long been a mystery, especially as whole series of extinct avian forms from the tertiary deposits present the same peculiarity. Prof. O. C. Marsh has, however, described a remark- able small Cretaceous bird named Ichthyornis, which had great powers of flight and bi-concave vertebrae as in fish, amphibians and a few reptiles. The third cervical vertebra of Ichthyornis when viewed in vertical section is, in its anterior articulation moderately convex, transversely it is strongly concave, thus presenting a near approach to the saddle-like articulation. No other vertebrae in lchthyor?iis present this character. The specialisation occurs at the bend of the neck and greatly facilitates motion in the vertical plane. This is the dominant motion in the neck of the modern bird ; any- thing that facilitates this motion would itself tend directly to produce the modification. Such modification of the vertebrae would naturally appear first where the neck had most motion, viz. in the anterior cervical region and would gradually extend down the neck ; and so on to the sacrum if the flexion continued. In modern birds behind the axis we find no exception to the saddle articulation in the whole cervical series. In a • ■• The Vertebrae of Recent Birds." The American Journal of Science and Arts, vol. xvii., p. 266. Chap, vi.] L1GAMENTA SUBFLAVA. 85 few birds especially that remarkable parrot, Stringops, the penguins, the terns, and a few others, one or more ver- tebrae are opisthoccelian, or imperfectly bi-concave. In the caudal series of recent birds the vertebrae preserve their orginal bi-concave structure. Thus Bi-Concave Vertebra are the rule in fish and am- phibians. From this form we derive : — The Plane Ver- tebra of mammals. The Cup-and-Ball variety of reptiles which are Procizlian in serpents and Opisthocalian in Dino- saurs and Saddle-shaped in Birds. CHAPTER VII. The Ligaments of the Vertebral Column (continued). The Atlas and Axis. The ligaments in connection with the first two cervical vertebrae are of sufficient importance to require separate mention, especially as they must be taken into considera- tion in questions connected with the morphology of these two extraordinary elements of the vertebral column. The ligaments to be stu ied are : — i. The transverse. 2. The check ligaments. 3. The ligamentum suspensorium. 4. The lateral occipito-atlantal ligaments. 5. The intervertebral disc between the odontoid process and the body of the atlas. It will also be necessary to extend the discussion to the anterior arch of the atlas, for an endeavour will be made to demonstrate that this part of the atlas may be considered as an ossified anterior conjugate ligament, and is in its nature comparable to the intercentrum of batrachians and reptilians. The transverse and check ligaments. — The trans- verse ligament divides the ring of the atlas into two unequal parts. The anterior space lodges the odontoid process, whilst the posterior serves for the transmission of the spinal cord. The ligament which is dense and strong derives firm attachments from the tubercle on the inner aspect of each lateral mass of the atlas. Where it glides around the constricted portion of the neck of the odontoid process it has a synovial covering. The check ligaments arise from the tubercles on the inner margin of the ex-occipitals. and passing transversely, Chap, vii.] INTERVERTEBRAL DISC. 87 meet tog-ether at the tip of the odontoid process into which they are inserted. Many text-books of anatomy figure them as though passing- obliquely to the odontoid process from above downwards, but they maintain a direction strictly parallel with that of the transverse liga- ment. In two instances I have seen the check ligaments form a continuous band so that it seemed as if there were twO transverse ligaments. There can be little doubt that these two bands are repetitions of the ligamentum conjugale costarum posterius which is described on page 79. This view of the matter also occurred to Cleland in his paper before referred to. To enter into a full discussion of the relationship would be to enter step by step into the question of the morphology of the atlas and axis, and is beyond the scope of this treatise. The student anxious for further information on this head may consult with advantag-e the references fur- nished on pages 89 and 90. The suspensory ligament.— Functionally of no value, but morphologically intensely interesting is the ligamentum suspensorium dentis. It passes from the summit of the odontoid process and joins, at the margin of the foramen magnum, the basi-occipital bone. Its true nature was de- tected by Rathke when he shewed that it was derived from the sheath of the notochord as it passed onwards to the base of the skull. The lateral occipito-atlantal ligaments are bands of fibrous tissue passing from the lateral mass of the atlas to the jugular process of the occipital bone. They run parallel with the rectus capitis lateralis muscle. The last named muscle is simply a continuation of the intertransver- sales series, and the lateral cccipito-atlantal ligament results from the conversion of a posterior rectus capitis lateralis muscle into fibrous tissue, thus becoming homologous with an intertransverse ligament. The intervertebral disc between the odontoid process and the body of the axis.— Cunningham has 88 LIGAMENTS. [Chap. vii. recently shewn that if, in the adult, an axis be bisected, a lenticular disc of cartilage of the shape and situation shewn in fig. 34. It is on all sides surrounded by bone so as only to be brought into view by means of sections. It is best marked in young subjects and gradually becomes smaller as life advances, but only disappears in extreme old age. The cartilage is of interest as it affords additional support to the view, that the odontoid process is the displaced body of the atlas. The evidence mav be arranged in a table for con- venience. 1. The notochord passes through the process on its way from the axis to the basis cranii. Fig. 34. — Transverse section of the body of the axis of a youth, aged 19. to shew the intervertebral disc between the odontoid process and the body of the axis. 2. Between the odontoid process and axis there is a •• swelling" of the notochord in the early embyro as in other intervertebral regions. 3. This '•swelling*' is later indicated by an intervertebral disc hidden in the bone, but persistent even in old age. 4. The odontoid process has independent centres. 5. It persists as an independent ossicle in chelonians, * For further information regarding this cartilage, and its comparative significance consult author's paper in the Proceedings of the Zoological Society, 1S86, " On the intervertebral disc between the odontoid process and the axis."' Chap, vii.] ARCH OF THE ATLAS. 89 lizards, and crocodiles. In Ornithorhynchus it remains dis- tinct for a very long time. 6. An epiphysial plate develops between it and the body of the axis in man and many mammals. 7. The neck of the odontoid process is embraced by a "conjugal *' ligament — the transverse. The bilateral ossific nuclei for the main portion of the mammalian odontoid process has given rise to a great deal of speculation among morphologists. Cope° has found that in an extinct batrachian, exhumed from the Permian formation of Texas, the vertebral bodies com- posed of two lateral pieces. The skeleton belonged to an animal named Eryopsmegalocephala of the order Ganocephala. This seems to indicate that in the possession of two lateral nuclei, the odontoid process retains a primitive charac- ter. The anterior arch of the atlas.— In the seal, as represented in fig. 30, the rib-heads of opposite sides are united by a ligament passing round the dorsal aspect of the intervertebral disc, known as the ligamentum con- jugale costarum In addition they are united by a fibrous band passing round the anterior aspect of the disc, known as the ligamentum conjugate costarum anterius. The fate of these ligaments has been fully described on page 79, and evidence has also been adduced to prove that the transverse ligament of the atlas, represents the posterior conjugal ligament of the ribs. I shall now attempt to show the anterior conjugal band has, in the case of the atlas, become ossified. In the first place it is necessary to mention the anterior portion of the lateral mass of the atlas agrees in its nature with the corresponding part of the transverse processes of the remaining cervical vertebrae, and is a rudimentary rib. The most conclusive arguments on this head are furnished by Macalister.f * Proc. American Philosoph. Soc, vol. xix., p. 51, 1SS0-S2. f Journal of Anatomy and Physiology, vol. iii.. p. 54. 90 LIGAMENTS. [Chap. vii. Lying 1 between the vertebrae in the lumbar region of a mole on the ventral aspect, are some small ossicles known as hypapophyses or sub-vertebral wedge-bones. In the remarkable lizard, sphenodon. these ossicles or intercentra form a continuous chain in the cervical, thoracic and lumbar regions of the spine. A careful examination of these bones in the mole, convinced me that they must be regarded as ossifications of the lig. conjugate costarum anterius. According to this view, the anterior arch of the atlas is really an intercentrum fused with the two rudi- mentary ribs, which constitute the anterior boundary of the lateral mass of the atlas of man." The subnotochordal rod. — In all Ichthyopsida (fish and amphibians), at a period slightly subsequent to the formation of the notochord, there appears a rod-like thickening, split off from the dorsal wall of the alimentary canal. This singular body is known to embryologists as the subnotochordal rod. Its situation and appearance are drawn by Gcette, in sections of Bombinator igneus. The rod lies between the notochord and alimentary canal, and is of almost the same longitudinal extent as these two structures. It was first detected by Gcette, who gave an account of it in Schultze's Archiv. fur Mik. Ana/., 1869. In his ela- borate account of the Entwicklungsgeschichte der Unke {Bombinator igneus), 1 875, he refers to it under the name of der Axenstrang des Darmblattes, and suggests that later it becomes a lymphatic trunk (page 775). The subnoto- chordal rod had been observed independently by Prof. Semper, and Balfour has given a description of it in his account of the development of Elasmobranch fishes. It is now known to be present in the embryos of Elasmo- branchs, Ganoids, the Teleostei, Dipnoi, and in the Amphibia, in all of whom it develops fundamentally in the same way. Fig. 35. Although it has not yet been found in a fully developed * Mivart, Elementary Anatomy, p. 217. Humphrey, The Human Skeleton. Chap, vii.] SUBNOTOCHORDAL ROD. form in any amniotic vertebrate, nevertheless a structure, which in all probability is a rudiment of the subnoto- chordal rod, was detected by Balfour and Marshall as a thickening- of the hypoblast in the neck of chicks. Salensky has further shown that in the sturgeon (Acci- penser) the subnotochordal rod becomes in the adult the subvertebral ligament, an observation which, according to Balfour, confirms an earlier suggestion made by Bridge. This point is important, and it will be interesting to endeavour to extend this view to vertebrata in general, and show that the anterior common ligament of the spine results, in some forms at least, from the degeneration of the subnotochordal Fig. 35. -Transverse section of a tadpole showing the subnotochordal rod. Later this becomes the anterior common ligament. rod, in the same way that ligamentous tissue in various parts of the adult column is the vestige of the embryonic notochord. I have made a series of observations on tad- poles of the common frog (Rana temporaria), and have been able to satisfy myself that in them, after the disap- pearance of the subnotochordal rod, a band of tissue can be seen in the sections replacing that structure. Further, as chondrification proceeds in the tissue lying between the notochord and the subnotochordal rod, the latter can be seen to flatten out over the bodies of the future vertebrae and finally assumes a fibrous appearance; having degene- rated into fibrous tissue, which grows with the vertebral column, it becomes in adults, the subvertebral ligament. 92 LIGAMENTS. [Chap. vii. It may be argued that, even admitting - this to be true, so far as amphibians or fish are concerned, yet it cannot apply to vertebrate forms above that group, in whom, with the exception of the chick, a subnotochordal rod has not yet been demonstrated. However, its existence in fish and amphibia is indisputable, therefore we may assume, unless later on its presence may be detected in higher forms, that the process has suffered abbreviation, and that the anterior ligament appears in these forms in obedience to the great principle — heredity. In support of the notion that this anterior common ligament may represent the subnoto- chordal rod of Sauropsida, without actually existing in the peculiar condition it presents in this class, I would draw attention to the fact that although Comparative Anatomy shows incontestably that the various structures about the knee joint — ligaments and fibro-cartilages — are in reality the disguised remnants of the tendons of muscles, yet during the development of the human embryo, as Barth has shown in an interesting article (" Die Entwicklungs- geschichte der Menschen," Morph. Jahr., Bd. iv. s. 403, 1878), the various ligamentous elements of the joint arise as thickenings of the connective tissue. In this case, as in so many others, embryological testimony will not always satisfy our demands when we ask for ancestral history, because the process has become abbreviated ; hence we are driven to Comparative Anatomy. CHAPTER VIII. The Manus. The skeleton of the manus consists of three distinct segments, carpus, metacarpus, and phalanges. The carpus. — This consists, as a rule, of eight small, and for the most part cubical bones arranged in two rows, a proximal and a distal, of four each. These bones com- mencing from the radial side of the proximal row are named, scaphoid, semilunar, cuneform and pisiform. Those of the distal row are recognised, following the same order, as trapezium, trapezoides, os magnum and unciform. Occasionally the number of the bones composing the carpus is increased to nine or even more. In order, there- fore, to fully comprehend the significance of these variations in the number of the carpal elements, as well as to com- pare it with the various members of man's class, it is essential first to consider some generalised form. It will, therefore, be expedient to examine the carpus of the water tortoise, Chelydra serpentina, for this presents, as Gegenbaur shewed, an excellent type-form to serve as a standard among most of the vertebrata. Fig. 36. In the carpus of this chelonian we find three bones occu- pying the proximal row, the preaxial or outer one articu- lating with the radius is known as the radiale, the inner or post-axial articulates with the ulna, and is termed the ulnare, between these bones and in contact partly with the ulna and radius is a third bone, named from its position the intermedium. The distal row is composed of five bones, each supporting a metacarpal. These, the carpalia, are numbered 1, 2, 3, 4, and 5 from the radial side. Inter- polated between the two rows of the carpus is a third element named from its position the os centrale. 94 LIGAMENTS. [Chap. viii. If we compare the carpus of man with this type-form, it is easy to see that the scapoid is the equivalent of the radiale, the cuneform of the ulnare, and the semilunar Fig. 36. — Manus of the Water Tortoise, Chelydra serpentina. R. Radius; U. Ulna; R. Radiale; /. Intermedium; U. Ulnare; C. Os Centrale; I-V. Carpalia. (After Gegenbaur). fulfils the characters required of an intermedium. Of the second row the carpalia, 1, 2, and 3, are represented by the trapezium, trapezoides, os magnum, whilst the unciform is Fig. 37.— Dorsal view of the carpus of a Baboon to shew the os cen- trale, C. the homologue of the fourth and fifth. The chief difference is that in the adult carpus the os centrale is wanting. Chap, viii.] THE CARPUS. 95 If, however, we examine the carpus of other mammals, e.g., baboons, macaques, orangs, lemurs, gibbons, and moles, we find an os centrale wedged in between the trapezium, trapezoides, os magnum and scaphoid. Fig. 37. In man also, an additional ossicle exists occasionally in the carpus at a spot corresponding to the position occupied by the centrale in those mammals normally possessing this bone.° The occurrence of this bone as an element in the carpus attracted considerable attention when Henke, Reyher and Rosenberg drew attention to its temporary existence in in the human manus. Rosenberg shewed that the "cen- trale" may be detected as a cartilage nodule in the carpus of a human embryo at the second month of intra- uterine life ; it is well marked during the third, but disap- pears during the fourth month of embryonic life. The existence of a cartilage between the trapezoides, os magnum and scaphoid in the foetus of man at the third month admits of no doubt whatever, and that it occasionally persists and forms a conspicuous element in the carpus is also indisputable. Opinions are, however, divided as to what becomes of the cartilaginous nodule in the usual course of development. The early investigations of Rosenberg seemed to shew that it underwent suppression, Kolliker also holds this view. Henke and Reyher, how- ever, are of opinion that the nodule fuses with the cartila- ginous scaphoid ; the scaphoid bone according to this view would be a compound of the radiale and centrale. Professor Gruber of St. Petersburg! has investigated an enormous number of human hands and scaphoid bones, (the total number of scaphoids in fresh hands, skeletons, or as separate bones amount 105129) has come to the conclusion from these macroscopic researches that the scaphoid may in exceptional cases, be compounded of a * Wenzel Gruber, Archiv. Anat. Phys., 1869, p. 331 ; also, Beobacht, aus der Mensch. und Vergleich Anatomie, Heft iv., Berlin, 1883. f " The Os Centrale in the Human Carpus," Journal of Anatomy and Physiology, vol. xviii., p. 119. 9 6 LIGAMENTS. [Chap. viii. radiale and centrale, but that as a rule it consists of a radiale only. Gruber also believes that scaphoid bones with a paral- lelogrammatic form, and those of a biscuit shape, which Fig. 38.— Manus of an early human foetus shewing the cartilaginous centrale. (After Leboucq). possess a tooth, or small process on, or near, the dorsal ulnar angle, are compounded of a radiale and centrale, whilst all other scaphoids are formed solely of the radiale. In support of this view Gruber also points out reasons Fig. 39.— Scaphoid bones of the human carpus shewing fusion of ossa centralia with them. for regarding- the scaphoid in the gorilla and chimpanzee, as being a compound of radiale and centrale ; whilst in certain mammals normally possessing a distinct centrale, it may, in old animals, fuse with the scaphoid, whilst in certain mammals the scaphoid is normally thus com- Chap, viii.] THE CARPUS. 97 pounded. Prof. Flower has found in the carpus of a dog six weeks old, the centrale as a separate element. In this mammal the scaphoid and semilunar bone fuse to form a scapho-lunar, and Flower's observation shows that it is of triple constitution. Prof. Leboucq of Gand has published some observa- tions on the morphology of the carpus, which induce him to believe that the os centrale fuses with the scaphoid. His valuable research published in Bull, de V Acad, royale des Sciences de Belgique, 3rd series, t. iv., no. 8, 1882, is illustrated with a very extensive series of draw- ings. In sections of the human carpus a small ligament passing from the scaphoid to the neck of the os magnum, will be occasionally detected; it is deemed so insignificant that only a few text-books of anatomy deign to consider or figure the little band. In 1885 I made an investigation into the development of the carpus with the hope of detecting the ancestry of this band of tissue. The relations of the ligament closely correspond to the position of the centrale, for it passes from the scaphoid to be attached to the neck of the os magnum. The result of the inquiry serves to show that this ligament, when present, is the remains of the cartila- ginous nodule, for whenever the cartilage nodule repre- senting the os centrale is present, the ligament cannot be detected, but in many cases, by no means in all, the spot previously occupied by the cartilage is filled up by fibrous tissue, which may persist through life, but in the largest number of instances disappears entirely. Thus the os centrale is represented in the carpus of the human embryo by a distinct cartilaginous nodule, which may undergo suppression, fuse with the scaphoid, or be- come represented by ligament. Finally, in very rare cases it may persist as an independent element. * Journal of Anatomy and Physiology, vol. vi., p. 62, 1872. H 98 LIGAMENTS. [Chap. viii. Weidersheim has observed in several specimens of Axolotl, that the os centrale has been double, and in some instance three ossa centralia were present. There were also two centralia in the tarsus. The duplicity of the os centrale in the carpus of Axolotl is of some interest, for it appears that in the carpus of the insectivorous mammal, Centetes Madagascariensis, there are two distinct ossa centralia. Leboucqf mentions that he had found in a foetus twenty- five millimetres in the length, the os centrale divided into two parts. One occupied the position of the normal cartilage representing- this bone, it was in part fused with the scaphoid. The other part was, on the palmar aspect, in contact with the scaphoid and the distal extremity of the semilunar. The "os centrale" must therefore be counted as one of the primitive elements of the carpus, and is of equal value with the remaining bones. The triangular fibro -cartilage in the wrist. — This has been made the subject of some interesting investiga- tions by Professor Leboucq, also contained in the paper " On the Morphology of the Carpus " before referred to. This observer finds that in the human foetus, from the third to the fourth month, there exists in this structure a small elliptical cartilaginous nodule, which at that period of development is in close relation with the pisiform bone. Until quite recently it has been customary to regard the pisiform as a sesamoid bone developed in the tendon of the flexor carpi ulnaris muscle. Recently, owing mainly to the comparative researches of Gegenbaur and Wiedersheim, doubts have been thrown over this view of its nature. According to Leboucq, the cartilage for the pisiform appears before the tendon of the flexor carpi * Morph. Jahrbuch, vol. vii., § 581, 1880. " Ueber die Vermehrung des Os Centrale im Carpus und Tarsus des Axolotls." f " Sur la Morphologie du Carpe et du Tarse." Anatotnischer Anzeiger, Jena, no. i., p. 17, 1886. Chap, vin.] FIBRO-CARTILAGE. 99 ulnaris, therefore it cannot be a sesamoid. He then points out that the mode of development of the interarticular fibro-cartilage and the pisiform ossicle induces him to' believe that these two structures are part of a complex apparatus, of which the pisiform bone is a separate portion. Leboucq also figures the carpus of a Gibbon (Hylobates leuctscus) preserved in the Museum of the Uni- versity of Gand, in which there is a cartilage nodule occupying the position of the interarticular fibro-cartilage and articulating by its distal end with the pisiform bone; this interarticular nodule he believes represents the tem- porary cartilage nodule in the human foetus. Although Leboucq's conjectures regarding the meaning of this cartilaginous nodule are rather far-fetched, it is nevertheless interesting to find, as in so many other in- stances, that this interarticular fibro-cartilage is the rem- nant of a structure which, in primitive forms, was of great functional value. APPENDIX. Containing a list of mammals which have been examined to determine the presence or absence of the gleno -humeral ligament. In all the mammals enumerated in the accompanying- list, the ligament was present and as well developed as represented in the beaver on page 69. The numeral refers to the number of each species specially examined. Quad rum ana. — Cercopithecus cynosutus, 8; C. lalandii, J ; C.callitrichus, 2; C.mona; C. erxlebeni ; C. campbelli ; C. me ti- tans ; C.albigena. Macacus sinicus, 20; M. cynomologus, 10; M. cyclopis; M. nemesirinus, 6. Theropithecus gelada. Cynocephalus hamadryas ; Cyn. anubis, 2. Chrysothrix sciurea, 5. Cebus fatuellus ; Ceb. capucinus, 3; Ceb. albifrons. Ateles paniscus ; A. ater, 2. Hapale jacchus, 4; H. penicillata, 2. Cercocebus albigena (in this species the ligament is thin). Semnopithecus leucoprymus, Xyctipithecus vociferans. Lemurid^:. — Lemur cat la, 3. Galago allem, 3; G. maholi, 2. Pcrodicticus potto, 2. Chirogaleus coquereli. Microcebus smith'. In Chiromys madagascarensis it was present but very thin. Insectivora. — Talpa europcea, 3. Erinaceus europczus. Tupaia peguana, 2. Centetes madagascaiensis. Microgale longicauda. Sorex vulgaris. Rodentia. — Pteromys magnificus. Arctomys monax. Cynomys ludovicianus, 3. Castor canadensis, 8. Mvoxus glis ; M. dry as. Fiber zibet hie us. Dipus cegypticus ; D. hirtipes. Myopotamus coypus, 6. Hystrix cristata. Erithrizon dorsatus. Sphingutus prehensilis. Ccelogenys paca. Dasyprocta agouti. Sciurus hypopyrrhus. Xtrus getulus. Sciuropterus fimbriatus. Ger- billus indicus. Cricetomys gambianus. Mus barbarus. Rhizomys badius. Octodon cumingi. Ccrodon rupestris. Arvicola amphi- bius. Mus musculus ; Mus decumanus. Lepus cuniculus. APPENDIX. 101 In some rodents the ligament is very large, and occupies a groove in the cartilage covering the head of the humerus^ Marsupialia. — Didelphys cancrivora ; D. virginiana, 2; D. opossum ; D. philander, 2 ; D. viverrina. Phalangista vulpina, 3. Hypsiprymnus gaimardi. Phascolomys wombat. Cuscus maculatus, 2. Belideus breviceps. Edentata. — Chlamydophorus truncatus. The gleno-humeral ligament is absent in the mammals mentioned in the lists which follow, but in many a thick- ened band was detected in the capsule which may have represented it. Quadrumana. — Chimpanzee (Anthropopilhecus troglodytes), 3. Orang-outang (Simia satyrus), 2. Gibbon (Hylobates leuciscus), 2. Macacus inuus (in this form a thick band may be seen in the capsule). Chiroptera. — Pteropus poliocephalus. Desmodus rufous. Plecotus auritus. Hyraces. — Hyrax capensis, 2. Ungulata. — Rhinoceros {Rhinoceros sumatrensis). Tapirus americanus. Cariacus virginianus ; C. ru/us. Pudu humulus. Tragulus meminno. Equus. Bos taurus ; B. indicus. Oreas canna. Llama huanacos ; L. peruana. Sus scrofa. Porcula sal- vania. Balirussa alfurus. Slrepsiceros kudu. Capra hircus. Ovis aries ; O. bland/ordi ; 0. burrhel. Moschus moschiferus . Tragulus, species incerta. Hydropotes inermis. Muntjac sclateri. Hippopotamus amphibius. Oryx beisa. Cephalophus mergens ; C. pygmcEus ; C. hydophylex. Cetacea. — Phocaina communis. Carnivora. — Felis leo ; F. tigiris. Cam's familiaris. Her- pestes ichneumon ; H. griseus. Lycaon pictus. Lutra vulgaris. Nasua rufa. N. narica. Phoca vitulina. Enhydra marina, 2. The Common Cat. Felis Javanensis, 2. Paradoxurus typus. P. trivirigatus. Ursus syriacus Very many other species of Carnivora and Ungulata 102 APPENDIX. have been examined with a negative result, but at first I did not make note of these cases, therefore, the species cannot be indicated individually. Although in most of the Carnivora the gleno-humeral ligament is absent, yet it turns up in that singular form the Two-spotted Paradoxure (Nandinia linotatta) ; it is also present in the Himalayan Bear (Ursus tibetanus). Edentata. — Bradypus tridactylus, 3. Tatusia peba ; T. hybrida ; T. sexcinctus ; T. villosus. Myrmecophaga jubata, 2. Tamandua tetradactyla, 2. Mam's didactyla, 2. Orycieropus cap.nsis. Marsupialia. — Perameles lagotis. Phascolarctos cinereus, 3. Ttlacropus rufus ; M. giganticus ; M. melanops. Halmaturus bennettu ; H. ualabatus ; H. derbia?ius. Moxotremata. — Echidna hystrix. Ornithorhynchus para- doxus. * It was absent in a specimen subsequently dissected. REFERENCE LIST OF AUTHORS. Albinus, 8 Albrecht, 73 Anderson, 74 Balfour, 90 Barth, 92 Baur, 5, 52 Bourgelat, 25 Bridge, 91 Brown, Macdonald, 24 Cleland, 80 Cope, 82, 89 Cunningham, 26, 38, 87 Cuvier, 82 Flower, 24, 77, 97 Freeman, 27 Gadow, 37, 43 Galton, J. C, 10, 17, 24 Garrod, 33, 41 Gegenbaur, 51, 93 Girard, 25 Gcette, go Golgi, 3 Gruber, g, 10, 24, 95 Hasse, 82 Henke, 95 Humphry, 80 Hunter, 80 Hyrtl, 8 Jager, 83 Johnson, 51, 52 Kolliker, 95 Leboucq, g7 Luschka, 82 Macalister, gi Marsh, 84 Marshall, 91 Mayer, 80 Mivart, 29, 31, 57, 90 Morris, 72, 80 Muller, 82 Murie, 24 Owen, 74 Paley, 41 Parker, 64 Percival, 25 Rathke, 82, 87 Reyher, 95 Robin, 82 Rolleston, 68 Rosenberg, 52, 95 Ruge, 36 Salensky, 91 Semper, go Struthers, 5, 12, 27 Thompson, 32 Turner, 23 Walsham, 68 Watson, 12, 27 Wood, 11, 14 Wyman, 54 INDEX. Aard vark, 15 Agouti, 31 Alca torda, 32 Amphibia, 2, go Ankle-joint, 42, 48 Anteater, 6 Aponeurosis, abdominal, 23 lumbar, 20 Articulation, meso-tarsal, 51 Astragalus of birds, 53 ascending process of, 54 Atlas, 89 Auriculo-ventricular valve, 54 Avian metatarsus, 53 tarsus, 49 Axis, ligaments of, 86 intervertebral disc of, 78 ossification of, 89 Band, gleno-humeral, 58, 65 ilio-tibial, 19 Beaver, 46 Bicipital loop of birds, 27 emu, 28 Bombinator igneus, go Brady pus didactylus, 73 Carpus, morphology of, 98 Casuarius appendiculata, 39 Chelydra serpentina, 93 Chimpanzee, 11 Chondro-cranium, 76 Clavicle, ligaments of, 58 ossification of, 61 Costo-coracoid membrane, 58 Coraco-scapula foramen, 6 Cretaceous birds, 82 Cryptobranchus, 23 Cygnus mutis, 55 Degeneration of bone, 4 cartilage, 4 muscle, 3 Diaphragm, 22 Dinosauria, 50, 85 Dipnoi, 90 Disc intervertebral, 78 DramcBus novce-hollandia, 28 Echidna, 34, 3 g, 6g Edentata, 72 Eider duck, 55 Elasmobranchs, 75, go Elephant, 3g Emu, 28 End-organs in tendon, 3 Enhydra marina, 3g Episternal bones, 64 Eryops megalocephala, 89 Eustachian cartilage, 75 Expansor secundariorum, 33 Fascia lata, ig palmar, 15 plantar, 16 temporal, 7 Femoro-caudal, 44 Fibro-cartilage of clavicle, 58 knee, 42 wrist, g8 Fin of porpoise, 26 Foramen coraco-scapular, 6 supra-condyloid, 5 triossium, 64 I io6 INDEX. Ganocephala, 89 Gecko, 14 Gibbon, 36 Great Auk, 55 Gull, 55 Horse, 39, 69 Hylobates leuciscus, 96 Hypsiprymnus, 74 Hyrax, 36, 69 Ichthyornis, 85 Iguana tuberculata , 14, 28, 44 Ilio-tibial band, 19 Insectivora, 69 Intermedium, 55 Koala, 37 Lemuridas, 69 Ligament, alaria, 46 annular of ankle, 49 of wrist, 56 anterior of spine, 77 arcuatum, 23 atlantal, 87 calcaneo-cuboid, 17 check, 86 conjugal, 79 coraco-acromial, 58, 64 coraco-clavicular, 65 coraco-humeral, 65, 70 costa-coracoid, 58 crucial, 45 ex-latera! of knee, 35 gleno-humeral, 58, 65 interclavicular, 58 internal brachial, 12 interspinous, 81 intertransverse, 81 intervertebral, 78 ischio-sacral, 73 mucosum, 46 occipito-atlantal, 87 of pectoral arch, 58 Ligament, plantar, 17 post, of spine, 77 rhomboid, 65 sacro-sciatic, 72 subflava, 81 supraspinous, 78 suspensory of axis, 87 of horse, 25 of deer, 25 teres, 38 transverse, 86 Lophiomys, 7 Lumbar aponeurosis, 20 Manus, 93 of baboon, 93 tortoise, 93 Marsupial bone, 7 Marsupialia, 38, 6g Mastodon, 72 Matamata terrapin, 82 Meckle's cartilage, 75 Megatherium, 72 Membrane, interosseous, 14 synovial, of knee, 46 Menobranchus, 67 Menopoma, 23 Muscle, abduct, metatarsi, 11 absence of, 23 ambiens, 39 biceps of birds, 27 coraco-brachialis, 11 curvator coccygis, 8 degeneration of, 3 epicoraco-humeral, 67 epitrochleo-anconeal, 9 extensor coccygis, 9 digitorum, 36 femoro-caudal, 44 ilio-peroneal, 29 interosseous, 14, 19 latissimus dorsi, 20 levator costarum, 22 humeri, 86 INDEX. 107 Muscle, metamorphosis of, 2, 25 migration of, 3 obliqui, 23 palmaris longus, 14 peroneus longus, 36 primus, 36 quartus, 37 quintus, 37 tertius, 37 peroneo-tibialis, 14 plantaris, 15 regression of, 3, 25 rotator humeri, 12 subclavius, 66 transversalis, 22 Narwhal, 27 Nerve, external cutaneous, 19 interosseous, 19 great sciatic, 27 Odontoid process, 82 Orang, 36, 39, 69 Ornithorhynchus, 39, 63, 69, 89 Orycteropus, 17 Ostrich, 36, 39 Os centrale, 94 intermedium, 54, 94 magnum, 95 odontoideum, 82 scaphoid, 95 trapezium, 95 Pangolin, 39, 69 Parson, chameleon, 57 Pelobates, 7 Penguin, 50, 55 Petrel, 55 Porpoise, 27 Potto, 56 Potoroo, 74 Quadrumana, 69 Rallus aquaticus, 32 Razor bill, 32 Rhamphastos loco, 28 Rhea, 28, 50, 55, 83 Rhinoceros, 32, 39, 69 „ Rod, subnotochordal, 91 Sauropsida, 92 Scapula, ossification of, 62 of sloth, 6 Seal, 69 Sea otter, 3g, 69 pigeon, 55 Skate, vertebrae of, 81 Skull, appendicular elements of, 75 Sloth, 6, 39, 6g Sphenodon, 14, 90 Struthio, 50, 55, 60 Sturgeon, 91 Subnotochordal rod, gi Subvertebral wedge bones, 90 Supra-condyloid foramen, 5 of cat, 5 of lion, 5 of man, 5 formation of, 4 Teleostei, go Tendo-Achillis, 17 Tendon, end-organs in, 3 metamorphosis of, 2 migration, 2, 35 relation to muscle, 3 use of, 32 Tern, 55 Tentorium cerebelli, 7 Tibio-tarsus, 4g of ostrich, 54 Vertebra atlas, 86 axis, 86 opisthoccelian, 85 procoelian, 85 saddle shaped, 85 Vulture, 32 Walrus, 3g, 6g Wombat, 14 Printed by H. 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Pharmaceutical Chemist ; Examiner to the Pharmaceutical Society ; Teacher of Pharmacy and Demonstrator of Materia Medica to University College Hospital. ELEMENTS OF MATERIA MEDICA AND PHAR- MACY. Crown 8vo, 8s. 6d. [Just published. HENEAGE GIBBES, m.d. Lecturer oh Physiology and on Normal and Morbid Histology in the Medical School of Westminster Hospital ; etc. PRACTICAL HISTOLOGY AND PATHOLOGY. Third Edition, revised and enlarged, crown 8vo, 6s. C. A. GORDON, m.d., c.b. Deputy Inspector General of Hospitals, Army Medical Department. REMARKS ON ARMY SURGEONS AND THEIR WORKS. Demy 8vo, 5s. W. R. GOWERS, M.D., F.R.C.P., M.RX.S. Physician to University College Hospital, &c. DIAGRAMS FOR THE RECORD OF PHYSICAL SIGNS. In books of 12 sets of figures, is. Ditto, unbound, is. J. B. GRESSWELL, m.r.c.v.s. Provincial Veterinary Surgeon to the Royal Agricultural Society. VETERINARY PHARMACOLOGY AND THERAPEU- TICS. With an Index of Diseases and Remedies. Fcap. 8vo, 5s. SAMUEL D. GROSS, m.d., ll.d., d.c.l. oxon. Professor of Surgery in the Jefferson Medical College of Philadelphia. PRACTICAL TREATISE ON THE DISEASES, IN- JURIES, AND MALFORMATIONS OF THE URINARY BLADDER, THE PROSTATE GLAND, AND THE URETHRA. Third Edition, revised and edited by S. W. GROSS, A.M., M.D., Surgeon to the Philadelphia Hospital. Illustrated by 170 engravings, 8vo, 18s. Catalogue of Works Published by H. K. Lewis. SAMUEL W. GROSS, a.m., m.d. Surgeon to, and Lecturer on Clinical Surgery in, the Jefferson Medical College Hospital, and the Philadelphia Hospital, &c. A PRACTICAL TREATISE ON TUMOURS OF THE MAMMARY GLAND : embracing their Histology, Pathology, Dia- gnosis, and Treatment. With Illustrations, 8vo, ios. 6d. WILLIAM A. HAMMOND, m.d. Professor of Mental and Nervous Diseases in the Medical Department of the University of the City of New York, &c. I. A TREATISE ON THE DISEASES OF THE NERVOUS SYSTEM. Seventh edition, with 112 Illustrations, large 8vo, 25s. II. A TREATISE ON INSANITY. Large 8vo, 25s. in. SPIRITUALISM AND ALLIED CAUSES AND CON- DITIONS OF NERVOUS DERANGEMENT. With Illustrations, post 8vo, 8s. 6d. ALEXANDER HARVEY, m.a„ m.d. Emeritus Professor of Materia Medica in the University of Aberdeen; Consulting Physician to the Aberdeen Royal Infirmary, &c. I. FIRST LINES OF THERAPEUTICS ; as based on the Modes and the Processes of Healing, as occurring Spontaneously in Dis- ease ; and on the Modes and the Processes of Dying, as resulting Naturally from Disease. In a series of Lectures. Post 8vo, 5s. 11. ON THE FCETUS IN UTERO AS INOCULATING THE MATERNAL WITH THE PECULIARITIES OF THE PATER- NAL ORGANISM. In a series of Essays now first collected. Fcap. 8vo, 4s. 6d. ALEXANDER HARVEY, m.d. Emeritus Professor of Materia Medica in the University of Aberdeen, &c. AND ALEXANDER DYCE DAVIDSON, m.d., f.r.s. edin. Regius Professor of Materia Medica in the University of Aberdeen. SYLLABUS OF MATERIA MEDICA FOR THE USE OF STUDENTS, TEACHERS AND PRACTITIONERS Based on the relative values of articles and preparations in the British Pharmacopoeia (1885). Seventh Edition, 32mo, is. 6d. Catalogue of Works Published by H. K. Lewis. 9 K. M. HEANLEY. Matron of Boston Cottage Hospital. MANUAL OF URINE TESTING-. Compile4 for the use of Matrons, Nurses, and Probationers. Post 8vo, is. 6d, GRAILY HEWITT, m.d. Professor of Midwifery and Diseases of Women in University College, Obstetrical Physician to University College Hospital, &c. OUTLINES OF PICTORIAL DIAGNOSIS OF DIS- EASES OF WOMEN. Folio, 6s. BERKELEY HILL, m.b. lond., f.r.c.s. Professor of Clinical Surgery in University College ; Surgeon to University College Hospital and to the Lock Hospital. THE ESSENTIALS OF BANDAGING. For Managing Fractures and Dislocations ; for administering Ether and Chloroform ; and for using other Surgical Apparatus. Fifth Edition, revised and much enlarged, with Illustrations, fcap. 8vo, 5s. BERKELEY HILL, m.b. lond., f.r.c.s. Professor of Clinical Surgery in University College ; Surgeon to University College Hospital and to the Loch Hospital. AND ARTHUR COOPER, l.r.cp., m.r.c.s. Surgeon to the Westminster General Dispensary. SYPHILIS AND LOCAL CONTAGIOUS DISORDERS. Second Edition, entirely re-written, royal 8vo, 18s. 11. THE STUDENT'S MANUAL OF VENEREAL DIS- EASES. Being a Concise Description of those Affections and of their Treatment. Fourth Edition, post 8vo, 2s. 6d. HINTS TO CANDIDATES FOR COMMISSIONS IN THE PUBLIC MEDICAL SERVICES, WITH EXAMINATION QUESTIONS, VOCABULARY OF HINDUSTANI MEDICAL TERMS, ETC. 8vo, 2s. 10 Catalogue of Works Published by H. K. Lewis. SIR W. JENNER, Bart., m.d. Physician in Ordinary to H.M. the Queen, and to H.R.H. the Prince oj Wales. THE PRACTICAL MEDICINE OF TO-DAY: Two Addresses delivered before the British Medical Association, and the Epidemiological Society, (1869). Small 8% ? o, is. 6d. C. M. JESSOP, m.r.c.p. Associate of King's College, London ; Brigade Surgeon H.M. British Forces. ASIATIC CHOLERA, being a Report on an Outbreak of Epidemic Cholera in 1876 at a Camp near Murree in India. With map, demy 8vo, 2s. 6d. GEORGE LINDSAY JOHNSON, m.a., m.b., b.c. cantab. Clinical Assistant, late House Surgeon and Chlor of ormist, Royal Westminster Ophthalmic Hospital, &c. A NEW METHOD OF TREATING CHRONIC GLAU- COMA, based on Recent Researches into its Pathology. With Ulus- trations and coloured frontispiece, demy 8vo, 3s. 6d. RUSTOMJEE NASERWANJEE KHORY, m.d. brux. Member of the Royal College of Physicians. THE PRINCIPLES AND PRACTICE OF MEDICINE. Second Edition, revised and much enlarged, 2 vols., large Svo, 28s. NORMAN W. KINGSLEY, m.d.s., d.d.s. President of the Board of Censors of the State of New Ytrk ; Member of the American Academy of Dental Science, &c. A TREATISE ON ORAL DEFORMITIES AS A BRANCH OF MECHANICAL SURGERY. With over 350 Illustra- tions, 8vo, 16s. E. A. KIRBY, M.D., M.R.C.6. ENG. Late Physician to the City Dispensary. I. A PHARMACOPOEIA OF SELECTED REMEDIES, WITH THERAPEUTIC ANNOTATIONS, Notes on Alimentation in Disease, Air, Massage, Electricity and other Supplementary Remedial Agents, and a Clinical Index ; arranged as a Handbook for Prescribers. Sixth Edition, enlarged and revised, demy 410, 7s. II. ON THE VALUE OF PHOSPHORUS AS A REMEDY FOR LOSS OF NERVE POWER. Fifth Edition, 8vo, 2s. 6d. Catalogue of Works Published by H. K. Lewis. 11 J. WICKHAM LEGG, f.r.c.p. Assistant Physician to Saint Bartholomew's Hospital, and Lecturer on Pathological Anatomy in the Medical School. ON THE BILE, JAUNDICE, AND BILIOUS DiSEASES. With Illustrations in chromo-lithography, 719 pages, roy. 8vo, 25s. A GUIDE TO THE EXAMINATION OP THE URINE: intended chiefly for Clinical Clerks and Students. Sixth Edition, revised and enlarged, with Illustrations, fcap. 8vo, 2s. 6d. III. A TREATISE ON HAEMOPHILIA, SOMETIMES CALLED THE HEREDITARY HEMORRHAGIC DIATHESIS. Fcap. 4to, 7s. 6d. DR. GEORGE LEWIN. Professor at the Fr. IVilh. University, and Surgeon-in-Chief of the Syphilitic Wards and Skin Disease Wards of the Chariti Hospital, Berlin. THE TREATMENT OP SYPHILIS WITH SUBCUTA- NEOUS SUBLIMATE INJECTIONS. Translated by Dr. Carl Prcegle, and Dr. E. H. Gale, late Surgeon United States Army, Small 8vo, 7s. LEWIS'S PRACTICAL SERIES. Under this title Mr. Lewis is publishing a Series of Monographs, em- bracing the various branches of Medicine and Surgery. The volumes are written by well-known Hospital Physicians and Sur- geons, recognized as authorities in the subjects of which they treat. The works are intended to be of a thoroughly practical nature, calculated to meet the requirements of the general practitioner, and to present the most recent information in a compact and readable form ; the volumes will be handsomely got up, and issued at low prices, varying with the size of the works. HANDBOOK OF DISEASES OF THE EAR FOR THE USE OF STUDENTS AND PRACTITIONERS. By URBAN PRITCHARD, M.D. Edin., F.R.C.S. Eng., Professor of Aural Surgerv at King's College, London ; Aural Surgeon to King's College Hospital ; Senior Surgeon to the Royal Ear Hospital. With Illustrations, crown 8vo, 4s. 6d. [Sow ready. A PRACTICAL TREATISE ON DISEASES OF THE KIDNEYS AND URINARY DERANGEMENTS. By CHARLES HENRY RALFE, M.A., M.D. Cantab, Fellow of the Royal College of Physicians, London; Assistant Physician to the London Hospital ; Examiner in Medicine to the University of Durham, etc., etc. With Illustrations, crown 8vo,ios. 6d. [Now leady. DENTAL SURGERY FOR GENERAL PRACTITIONERS AND STUDENTS OF MEDICINE. Bv ASHLEY W. BARRETT, M.B. Lond., M.R.C.S., L.D.S. Dental Surgeon to, and Lecturer on Dental Surgery and Pathology in the Medical School of, the London Hospital. With Illustrations, cr. 8vo, 3s. [Sow ready. BODILY DEFORMITIES AND THEIR TREATMENT: A HANDBOOK OF PRACTICAL ORTHOPEDICS. By H. A. REEVES, F.R.C.S. Edin., Senior Assistant Surgeon and Teacher of Practical Surgery at the London Hospital; Surgeon to the Royal Orthopaedic Hospital, &c. With numerous Illustrations, cr. 8vo, 8s. 6d. l Noxv read y- FEVERS AND THEIR TREATMENT. By ALEXANDER COLLIE, M.D. Aberd., Member of the Royal College of Physicians of London ; Medical Superintendent of the Eastern Hospitals ; Secre- tary of the Epidemiological Society for Germany and Russia. Illustrated with Coloured Plates, crown 8vo. i In the P resSt Further volumes will be announced in due course. *** Prospectus of the Series with specimen pages, &c, on application. 12 Catalogue of Works Published by H. K. Lewis. LEWIS'S POCKET MEDICAL VOCABULARY. Over 200 pp., 321T10, roan, 3s. 6d. J. S. LOMBARD, m.d. Formerly Assistant Professor 0/ Physiology in Harvard College. I. EXPERIMENTAL RESEARCHES ON THE REGIONAL TEMPERATURE OF THE HEAD, under Conditions of Rest, In- tellectual Activity and Emotion. With Illustrations, 8vo, 8s. II. ON THE NORMAL TEMPERATURE OF THE HEAD. 8vo, 5s. WILLIAM THOMPSON LUSK, a.m., m.d. Prtfesser of Obstetrics and Diseases of Women in the Bellevne Hospital Medical College, &c. THE SCIENCE AND ART OF MIDWIFERY. Third Edition, revised and enlarged, with numerous Illustrations, 8vo, 18s. JOHN MACPHERSON, m.d. Inspector-General of Hospitals H.M. Bengal Army {Retired). Author of " Cholera in its Home," &c. I. ANNALS OF CHOLERA FROM THE EARLIEST PERIODS TO THE YEAR 1817. With a map. Demy 8vo, 7s. 6d. 11. BATH, CONTREXEVILLE, AND THE LIME SUL- PHATED WATERS. Crown 8vo, 2s. 6d. DR. V. MAGNAN. Physician to St. Ann Asylum, Paris; Laureate of the Institute. ON ALCOHOLISM, the Various Forms of Alcoholic Delirium and their Treatment. Translated by W. S. Greenfield, M.D., M.R.C.P. 8vo, 7s. 6d. A. COWLEY MALLEY, b.a., m.b., b.ch. t.c.d. PHOTO-MICROGRAPHY; including a description of the Wet Collodion and Gelatino-Bromide Processes, together with the best methods of Mounting and Preparing Microscopic Objects for Photo- Micrography. Second Edition, with Photographs and Illustrations, crown 8vo, 7s. 6d. Catalogue of Works Published by H. K. Lewis. 18 PATRICK MANSON, m.d., cm. Amoy, China. THE FILARIA SANGUINIS HOMINIS ; AND CER- TAIN NEW FORMS OF PARASITIC DISEASE IN INDIA, CHINA, AND WARM COUNTRIES. Illustrated with Plates and Charts. 8vo, ios. 6d. PROFESSOR MARTIN. MARTIN'S ATLAS OF OBSTETRICS AND GYNJECO- LOGY. Edited by A. Martin, Docent in the University of Berlin. Translated and edited with additions by Fancourt Barnes, M.D., M.R.C.P., Physician to the Chelsea Hospital for Women ; Obstetric Physician to the Great Northern Hospital ; and to the Royal Maternity Charity of London, &c. Medium 4to, Morocco half bound, 31s. 6d. nett. WILLIAM MARTINDALE, f.c.s. Late Examiner of the Pharmaceutical Society, and late Teacher of Pharmacy and Demon- strator of Materia Medica at University College. AND W. WYNN WESTCOTT, m.b. lond. Deputy Coroner for Central Middlesex. THE EXTRA PHARMACOPOEIA with the additions in- troduced into the British Pharmacopoeia, 18S5, and Medical References, and a Therapeutic Index of Diseases and Symptoms. Fourth Edition, revised with numerous additions, limp roan, med. 24mo, 7s. [Now ready. WILLIAM MARTINDALE, f.c.s. COCA, COCAINE, AND ITS SALTS : their History, Medical and Economic Uses, and Medicinal Preparations. Fcap.8vo, 2s. MATERIA MEDICA LABELS. Adapted for Public and Private Collections. Compiled from the British Pharmacopoeia of 1885. The Labels are arranged in Two Divisions : — DiviBion I. — Comprises, with few exceptions, Substances of Organ- ized Structure, obtained from the Vegetable and Animal King- doms. Division II.— Comprises Chemical Materia Medica, including Alco- hols, Alkaloids, Sugars, and Neutral Bodies. On plain paper, ios. 6d. nett. On gummed paper, 12s. 6d. nett. %* Specimens of the Labels, of which there are over 450, will be sent on application. 14 Catalogue of Works Published by H. K. Lewis. S. E. MAUNSELL, l.r.c.s.i. Surgeon-Major, Medical Staff. NOTES OF MEDICAL EXPERIENCES IN INDIA PRINCIPALLY WITH REFERENCE TO DISEASES OF THE EYE. With Map, post Svo, 3s. 6d. J. F. MEIGS, m.d. Consulting Physician to the Children's Hospital, Philadelphia. AND W. PEPPER, m.d. Lecturer on Clinical Medicine in the University of Pennsylvania. A PRACTICAL TREATISE ON THE DISEASES OF CHILDREN. Seventh Edition, revised and enlarged, roy. 8vo, 28s. Wm. JULIUS MICKLE, m.d., m.r.c.p. lond. Medical Superintendent, Grove Hall Asylum, London, &c. GENERAL PARALYSIS OF THE INSANE. Second Edition, enlarged and rewritten, 8vo, 14s. MIDDLESEX HOSPITAL, REPORTS OF THE MEDI- CAL, SURGICAL, AND PATHOLOGICAL REGISTRARS FOR 1883 ; and 1SS4. Demy Svo, 2s. 6d. nett each volume. KENNETH W. MILLICAN, b.a. cantab., m.r.c.s. THE EVOLUTION OF MORBID GERMS : A Contribu- bution to Transcendental Pathology. Cr. Svo, 3s. 6d. STEPHEN MONCKTON, m.d., f.r.c.p. THE METAPHYSICAL ASPECT OF NATURAL HIS- TORY. (A Lecture). Second Edition, with Illustrations, crown 8vo, as. Catalogue of Works Published by H. K. Lewis. 15 E. A. MORSHEAD, m.r.c.s., l.r.c.p. Assistant to the Professor of Medicine in University College, London. TABLES OF THE PHYSIOLOGICAL ACTION OF DRUGS. Fcap. 8vo, is. A. STANFORD MORTON, m.b., f.r.c.s. ed. Surgeon to the Royal South London Ophthalmic Hospital. REFRACTION OF THE EYE: Its Diagnosis, and the Correction of its Errors. Third Edition, with Illustrations, small 8vo. [Just ready. WILLIAM MURRELL, m.d., f.r.c.p. Lecturer on Materia Medica and Therapeutics at Westminster Hospital ; Examiner in Materia Medica and Therapeutics in the University of Edinburgh, and to the Royal College of Physicians of London. I. MASSAGE AS A MODE OF TREATMENT. Crown 8vo, 3s. 6d. [Just published. ii. WHAT TO DO IN CASES OF POISONING. Fifth Edition, royal 32mo, 3s. 6d. [Just published. in. NITRO- GLYCERINE AS A REMEDY FOR ANGINA PECTORIS. Crown 8vo, 3s. 6d. WILLIAM NEWMAN, m.d. lokd., f.r.c.s. Surgeon to the Stamford Infirmary. SURGICAL CASES: Mainly from the Wards of the Stamford, Rutland, and General Infirmary. 8vo, paper boards, 4s. 6d. DR. FELIX von NIEMEYER. Late Professor of Pathology and Therapeutics ; Director of the Medical Clinic of the University of Tubingen. TEXT-BOOK OF PRACTICAL MEDICINE, WITH PARTICULAR REFERENCE TO PHYSIOLOGY AND PATHO- LOGICAL ANATOMY. Translated from the Eighth German Edition, by special permission of the Author, by George H. Humphrey, M.D., and Charles E. Hackley, M.D. Revised Edition, 2 vols., large 8vo, 36s. 16 Catalogue of Works Published by H. K. Lewis. C. F. OLDHAM, m.r.c.s., l.r.c.p. Surgeon H.M. Indian Forces ; late in Medical charge of the Dalhousie Sanitarium. WHAT IS MALARIA ? and why is it most intense in hot climates ? An explanation of the Nature and Cause of the so- called Marsh Poison, with the Principles to be observed for the Preser- vation of Health in Tropical Climates and Malarious Districts. Demy 8vo, 7s. 6d. G. OLIVER, m.d., m.r.cp. THE HARROGATE WATERS : Data Chemical and Therapeu- tical, with notes on the Climate of Harrogate. Addressed to the Medical Profession. Crown 8vo, with Map of the Wells, 35. 6d. 11. ON BEDSIDE URINE TESTING: a Clinical Guide to the Observation of Urine in the course of Work. Third Edition, revised and enlarged, fcap. 8vo, 3s. 6d. SAMUEL OSBORN, f.r.c.s., Assistant-Surgeon to the Hospital for Women ; Surgeon Royal Naval Artillery Volunteers, I. AMBULANCE LECTURES: FIRST AID. With Illus- trations, fcap. 8vo, is. 6d. *" II. AMBULANCE LECTURES : NURSING. With Illustrations, fcap. 8vo, is. 6d. ROBERT W. PARKER. Surgeon to the East London Hospital for Children, and to the Grosvenor Hospital for Women and Children. TRACHEOTOMY IN LARYNGEAL DIPHTHERIA, AFTER TREATMENT AND COMPLICATIONS. Second Edition. With Illustrations, 8vo, 5s. JOHN S. PARRY, m.d. Obstetrician to the Philadelphia Hospital, Vice-President of the Obstetrical and Pathologi- cal Societies of Philadelphia, &c. EXTRA-UTERINE PREGNANCY ; Its Causes, Species, Pathological Anatomy, Clinical History, Diagnosis, Prognosis and Treatment. 8vo, 8s. E. RANDOLPH PEASLEE, m.d., ll.d. Late Professor of Gynecology in the Medical Department of Dartmouth College ; President of the New York Academy of Medicine, &c, &c. OVARIAN TUMOURS : Their Pathology, Diagnosis, and Treatment, especially by Ovariotomy. Illustrations, roy. 8vo, 16s. Catalogue of Works Published by H. K. Lewis. 17 G. V. POORE, M.D., F.R.C.P. Professor of Medical Jurisprudence, University College; Assistant Physician to, and Physi- cian in charge of the Throat Department of, University College Hospital. LECTURES ON THE PHYSICAL EXAMINATION OF THE MOUTH AND THROAT. With an Appendix of Cases. 8vo, 3s. 6d. R. DOUGLAS POWELL, m.d., f.r.c.p., m.r.c.s. Physician to the Middlesex Hospital, and Physician to the Hospital for Consumption and Diseases of the Chest at Brompton. DISEASES OF THE LUNGS AND PLEURAE, INCLUD- ING CONSUMPTION. Third Edition, entirely rewritten and en- larged. With coloured plates and wood engravings, 8vo, 16s. [Just published. URBAN PRITCHARD, m.d. edin., f.r.c.s. eng. Professor of Aural Surgery at King's College, London ; Aural Surgeon to King's College Hospital ; Senior Surgeon to the Royal Ear Hospital. HANDBOOK OF DISEASES OF THE EAR FOR THE USE OF STUDENTS AND PRACTITIONERS. With Illustrations, crown 8vo, 4s. 6d. [Now ready. Lewis's Practical Series) . CHARLES W. PURDY, m.d. (queen's univ.) Professor of Genito-Urinary and Renal Diseases in the Chicago Polyclinic, &c, &c. BRIGHT'S DISEASE AND THE ALLIED AFFECTIONS OF THE KIDNEYS. With Illustrations, large 8vo, 8s. 6d. CHARLES HENRY RALFE, m.a., m.d. cantab. Fellow of the Royal College of Physicians, London ; Assistant Physician to the London Hos- pital ; Examiner in Medicine to the University of Durham, etc., etc. A PRACTICAL TREATISE ON DISEASES OF THE KIDNEYS AND URINARY DERANGEMENTS. With Illustrations, crown 8vo, 10s. 6d. [Now ready. Lewis's Prattical Series] , AMBROSE L. RANNEY, a.m., m.d. Adjunct Professor of Anatomy in the University of New York, etc. THE APPLIED ANATOMY OF THE NERVOUS SYS- TEM, being a study of this portion of the Human Body from a stand- point of its general interest and practical utility, designed for use as a Text-book and a Work of Reference. With 179 Illustrations, 8vo, 20s. 18 Catalogue of Works Published by H. K. Leivis. H. A. REEVES, f.r.c.s. ed. Senior Assistant Surgeon and Teacher of Practical Surgery at the London Hospital; Surgeon to the Roy ai Orthopaedic Hospital, &c. BODILY DEFORMITIES AND THEIR TREATMENT: A Handbook of Practical Orthopaedics. With numerous Illustrations, crown 8vo, 8s. 6d. [Now ready. Lewis's Practical Series] . RALPH RICHARDSON, m.a., m.d. Fellow of the College of Physicians, Edinburgh. ON THE NATURE OF LIFE : An Introductory Chap- ter to Pathology. Second Edition, revised and enlarged. Fcap. 4to, ios. 6d. W. RICHARDSON, m.a., m.d., m.r.c.p. REMARKS ON DIABETES, ESPECIALLY IN REFER- ENCE TO TREATMENT. Demy 8vo, 4s. 6d. SYDNEY RINGER, m.d., f.r.s. Professor of the Principles and Practice of Medicine in University College; Physician to, and Professor of Clinical Medicine in, University College Hospital. I. A HANDBOOK OF THERAPEUTICS. Eleventh Edition, thoroughly revised, 8vo, 15s. [Now ready. II. ON THE TEMPERATURE OF THE BODY AS A MEANS OF DIAGNOSIS AND PROGNOSIS IN PHTHISIS. Second Edition, small 8vo, 2s. 6d. FREDERICK T. ROBERTS, m.d., b.sc, f.r.c.p. Examiner in Medicine at the Royal College of Surgeons ; Professor of Therapeutics in University College ; Physician to University College Hospital ; Physician to Brompton Consumption Hospital, &c. I. A HANDBOOK OF THE THEORY AND PRACTICE OF MEDICINE. Sixth Edition, with Illustrations, in one volume, large 8vo, revised and enlarged to over 1000 pages, 21s. [Just published. %* Copies may also be had bound in two volumes cloth for is. 6d. extra. II. NOTES ON MATERIA MEDICA AND PHARMACY. Fcap. 8vo, 7s. 6d. Catalogue of Works Published by H. K. Lewis. 19 R. LAWTON ROBERTS, m.d., m.r.c.s. ILLUSTRATED LECTURES ON AMBULANCE WORK. Second Edition, copiously Illustrated, crown 8vo. 2s. 6d. [jf ust published. A. R. ROBINSON, m.b., l.r.c.p., and l.r.c.s. edin. Professor of Dermatology at : >h Polyclinic. A MANUAL OF DERMATOLOGY. With 88 Illustrations, large 8vo, 21s. ROBSON ROOSE, m.d. Fellow 0/ the Royal College of Physicians in Edinburgh. GOUT, AND ITS RELATIONS TO DISEASES OF THE LIVER AND KIDNEYS. Third Edition, crown 8vo, 3s. 6d. {just ready. D. B. St. JOHN ROOSA, m.a., m.d. Professor of Diseases of the Eye and Ear in the University of the City of New York ; Surgeon to the Manhattan Eye and Ear Hospital ; Consulting Surgeon to the Brooklyn Eye and Ear Hospital, &c, &c. A PRACTICAL TREATISE ON THE DISEASES OF THE EAR, including the Anatomy of the Organ. Sixth Edition, Illustrated by wood engravings and chromo-lithographs, large 8vo, 25s. J, BURDON SANDERSON, m.d., ll.d., f.r.s. Jodrell Professor of Physiology in University College, London. UNIVERSITY COLLEGE COURSE OF PRACTICAL EXERCISES IN PHYSIOLOGY. With the co-operation of F. J. M. Page, B.Sc, F.C.S. ; W. North, B.A., F.C.S., and Aug. Waller, M.D. Demy 8vo, 3s. 6d. W. H. O. SANKEY, m.d. lond., f.r.c.p. Late Lecturer on Mental Diseases, University College and School of Medicine for Women, London; Formerly Medical Superintendent (Female Department) of Hanwell Asylum ; President of Medico-Psychological Society, &c. LECTURES ON MENTAL DISEASE. Second Edition, with coloured plates, 8vo, 12s. 6d. ALDER SMITH, m.b. lond., f.r.c.s. Resident Medical Officer, Christ's Hospital, London. RINGWORM : Its Diagnosis and Treatment. Third Edit., rewritten and enlarged. With Illustrations, fcap. 8vo, 5s. 6d. 20 Catalogue of Works Published by H. K. Lewis. J. LEWIS SMITH, m.d. Physician to the New York Infants' Hospital ; Clinical Lecturer on Diseases of Children in Bellevue Hospital Medical College. A TREATISE ON THE DISEASES OF INFANCY AND CHILDHOOD. Fifth Edition, with Illustrations, large 8vo, 21s. FRANCIS W. SMITH, m.b., b.s. THE SALINE WATERS OF LEAMINGTON. Second Edit., with Illustrations, crown 8vo, is. nett. JAMES STARTIN, m.b., m.r.c.s. Surgeon and Joint Lecturer to St. John's Hospital for Diseases of the Skin. LECTURES ON THE PARASITIC DISEASES OF THE SKIN. VEGETOID AND ANIMAL. With Illustrations, crown 8vo, 2s. 6d. LEWIS A. STIMSON, b.a., m.d. Surgeon to the Presbyterian and Bellevue Hospitals ; Professor of Clinical Surgery in the Medical Faculty of the University of the City of New York, &c. A MANUAL OF OPERATIVE SURGERY. With three hundred and forty-two Illustrations. Second Edit., post 8vo, 10s. 6d. [Just published. HENRY R. SWANZY, a.m., m.b., f.r.c.s.i. Examiner in Ophthalmic Surgery at the Royal College of Surgeons, Ireland ; Surgeon to the National Eye and Ear Infirmary, Dublin ; Ophthalmic Surgeon at the Adelaide Hospital, Dublin. 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