3ournat of Anatomn aub Xpoitlogp. THE RETE MIRABILE OF THE NARWHAL. By HENRY SEASONWILSON, M.D.,Superintendent ofPracticalAnatomy, London Hospital. (PLATES XXII., XXIII.) THE specimen on which the following investigations were made was a fetal Narwhal, 641 inches long, and 481 inches in its greatest circumference. Its mother was captured in 1861 on the north side of Pond's Bay, Davis Strait, by Captain George Deuchars of the screw-steamer "Narwhal" of Dundee. The foetus was removed at the time of capture by Dr Robert Brown, who kindly permitted me to dissect the soft parts. Owing, however, to the decomposed state of the animal, I could only investigate the vascular system, and even that but imperfectly. My remarks will apply only to the thoracic rete mirabile, and are selected from the notes and drawings taken during my dis- sections when Demonstrator of Anatomy in the University of Edinburgh in 1862. It seems unfortunate that the term "rete mirabile" should have been applied so indiscriminately to all vascular plexiform arrangements, for, both anatomically and teleologically, when the various plexuses are compared with each other, a good deal of confusion and a greatwant of harmonyareatonce observable. I shall not enter here into that question. It will be best under- stood and more easily handled when we come to examine into the probable function of retia in general, and especially of the thoracic plexus in Cetacea. Suffice it to say that of all the retia mirabilia throughout the vertebrate, that in the thorax of the Cetacea resembles no other so-named plexus. John Hunter is the only author who has likened it to any other network, and VOL. XIV; D 378 DR HENRY SEASON WILSON. even that similitude is but trivial, for the spermatic arteries of the Bull resemble it only in the tortuosities of their branches. I will first state the facts evinced by dissection, deferring deduc- tions and other observations for subsequent attention. As the thoracic rete mirabile in the Narwhal is arterial, and its position is bythe side of the thoracic aorta, in order to under- stand the sources whence the vascular supply is derived, a brief enumeration of the modifications exhibited by the primary branches of the aortic arch and the thoracic aorta are necessary. Aortic arch.-The arch lay at first entirely concealed by the pulmonary artery. Beyond this only a narrow portion could be seen, owing to the close manner in which it was embraced by the lobes of the thymus (Plate XXII. A). The right lobe separated it from the vena cava anterior, whilst both lobes were interposed between the arch and the pneumo-gastric nerves, more especially on the left. By dissecting away the thymus, the aorta was seen to be continued forwards and to the right for a short distance, and thence to arch obliquely upwards towards the left of the spine. Close to its exit from the left ventricle the aorta supplied its coronary branches, each having its usual origin and distribution. From the convexity of the arch arose the innonlinate, the left common carotid, and the left posterior thoracic arteries. Theinnominate (Plate XXII.A,1) hadits originfrom thecom- mencement of the transverse portion of the arch, on a level with the head of the first rib, and greatly exceeded the others in size. It extended forwards towards the neck for about half an inch, and then divided into two large branches-the right posterior thoracic,andtherightcommoncarotid (PlateXXII.A,3, 2, 11,8). The right posterior thoracic in its origin and in the relations ofits first part resembled the right subclavian, but differed from it in its ultimate distribution. From its originitarchedhorizon- tally forwards and outwards towards the first rib. Reaching the latter at the distance of about three inches from the spine, it suddenly curved backwards, and continued along the inner aspect of the thoracic wall. In this course it crossed the first eight ribs and ended in the eighthintercostal space. Itwas thus divisible into two portions-the arch, situated in the neck, and the terminal portion related to the ribs, and intrathoracic in .379 THE RETE MIRABILE OF THE NARWHAL. position (PlateXXII.A). Forthe sakeof descriptionthese por- tions may be called the arch and the costal portion. The arch was much the shorter, and was cervical in position. It had the usual relation to the innominate vein, the pneumogastric, the branches of the sympathetic, and the phrenic nerves, as well as to the scalenus muscle. At first it was covered by the cervical part of the right thymic lobe. Only one branch, cervical (Plate XXII. A, 4) passed from this arch. It arose from the parent trunk, where that was crossed by the right innominate vein. This branch corresponded in its distribution with the"cervicalis ascendens" in man. In addition to its muscular branches it supplied offsets in great numbers to the cervical portion of the rete mirabile, and also formed large anastomoses with the anterior (superior), thyroid, and pharyngeal branches of the external carotid (Plate XXII. A). On the left side this branch came from the left internal mammary. The costal portion of the posterior thoracic ended, as above stated, in the eighth inter- costal space. In its course it lay, to a certain extent, embedded in the substance of the thoracic portion of the rete, running along the external margin of the rete, and keeping parallel to the spine. Immediately before its termination, however, it diverged fromthemesial line. Itsbrancheswereverynumerous, and were chieflydirected inwards towards the spine,or outwards from it. Towards the vertebral column it gave hundreds of long, delicate offsets,nearly equal in their calibre. These passed entirely to the rete, and a fuller account of their distribution will be more appropriate when considering that network. The branches passing outwards constituted groups which, with the exceptionofthefirst,occupiedtheintercostalspaces fromthethird to the eighth (Plate XXII. A, 16, 7). The first group was given off immediately before the artery entered the thorax, and was muchlarger than the others. It consisted of numerousbranches, directed forwards to the cervical muscles, and backwards to the first and second intercostal spaces. The other groups resembled intercostal vessels, and need not occupy time in minute descrip- tion. As the right common carotid artery has very little to do with the rete, supplying comparatively very few branches, and those simply to its cervical portion, I shall pass it over. The left common carotid (Plate XXII. A, 2') sprung from the 380 DR HENRY SEASON WILSON. aorticarch,ata distanceofaboutthreequartersofaninchfromthe innominate trunk. In size it was somewhat more than half the calibre of the latter. Its course was forwards, and slightly out- wards. Within the thorax it was related externally to the narrow anterior prolongation of the left thymic lobe, by which it was separated from the left pueumogastric, and internally it was in contact with the thyroid body. Its cervical portion need not be described. The left posterior thoracic (Plate XXII. A, 3'), in its place of origrin resembled the left subclavian in man; that is, it arose from the left aspect of the last portion of the aortic arch,directly opposite, but immediately preceding the ductus arteriosus. It was an inch and a half beyond the left common carotid, and about half its size. Its course was forwards to a level with the first rib and then backwards to end in the ninth intercostal space. The arch was intrathoracic and very short. It had the same relations to the sympathetic and pneumogastric as the corresponding subclavian in man. Its only branch was given off as it neared the first rib. This branch soon broke up into a number of offsets, and represented the first group of the costal portion of the right posterior thoracic (Plate XXII. A, 6'). The costal part of the left posterior thoracic extended one intercostal space beyond that of the same branch on the left side. It supplied the same innumerable branches inwards to the rete, and similar intercostalgroups outwards. The first group, already mentioned, resembled that on the right side in supplying the first and second intercostal spaces. It may here be added that these intercostal groups anastomosed freely with corresponding intercostal branches of the internal mammary arteries, and the last or terminal group also inosculated with those aortic inter- costals supplying the neighbouring intercostal spaces. Between the origins of the left common carotid and the left posterior thoracic, but closer to the latter, a very small fourth branch arose from the aortic arch. It was directed forwards to supply the apex of the left thymic lobe, and then ended in the left mass of the rete mirabile (Plate XXII. A). fThoracic aorta.-The thoracic aorta occupied the usual posi- tion along the spine asfar as the diaphragm. Its relations were similar to those in man, with this exception, that the rete THE RETE MIRABILE OF THE NARWHAL. 381 separated it from the spinal column. Its branches, seven in number, corresponded with the intercostal arteries, and arose from the vertebral aspect of the trunk, not in pairs, but as single offsets. These branches varied considerably in size. From the first to the fourth there was a gradual decrease, whilst the last three became greatly enlarged. The direction of the individual branches also differed. The first three passed slightly forwards to reach their destination, but the last two inclined backwards. Each terminated in two lateral branches at variable distances from its origin. These lateral divisions coursed along the transverse vertebral grooves and ended in branches which were destined to supply, some the rete mirabile alone, others partly the latter and partly the intercostal muscles. Their ramifica- tions in the intercostal spaces were peculiar in that thebranches of distribution passed off at acute angles from the trunks. As the same description does not answer for all, each must be examined separately. The first intercostal artery came off from the aorta at a level somewhat posterior to the fifth intercostal space, and, turning forwards, soon broke up into two divisions. One of these divisions continued onwards and subdivided into lateral branches to supply that portion of the rete corresponding with the two fourth intercostal spaces. The other division also became bifid, the resulting branches being spent in the mesh- work of the rete occupying the fifth interspaces. The second, third, and fourth intercostals branched almost directly into lateral divisions, and were lost in the rete under the sixth, seventh, and eighth interspaces respectively of the right and left sides. The fifth intercostal artery was a very large branch comparatively. Its lateral divisions were very unequal in size. That to the left far exceeded the other, and was in calibre the continuation of the parent trunk. It coursed along the trans- verse vertebral groove into the ninth intercostal space, lying above the constituent vessels of the rete. Having reached the space it subdivided into two unequal offsets. ,The smaller of these ran along the middle of the interspace and anastomosed with the ultimate twigs of the posterior thoracic and musculo- phrenic arteries, whilst the larger offset turned obliquely back- wards and followed the last rib. In its course along the rib it first supplied a considerable branch to the tenth interspace, and 382 DR HENRY SEASON WILSON. then terminated in offsets to the abdominal muscles on the one side, and the intercostals on the other. These terminal offsets of the left lateral division, as each passed along its intercostal space, supplied dorsal vessels which pierced the intercostal muscles and ended in the dorsal vertebral muscles, and while beneath the rete each, as well as the parent trunk, distributed numerous delicate twigs to that network. The right lateral division of the fifth intercostal was similar in size and distribu- tion to those of the preceding. It was lost in that part of the rete opposite the ninth interspace. The sixth intercostal branch was the largest of the last three. Its lateral divisions were proportionately as unequal in size. That to the left,.the smaller, continued entirely imbedded in the rete, as far as the intertransversales, through which it passed to ehd in the dorsal muscles of the spine. Whilst surrounded by the rete it gave to the latter a large supply of delicate, slender branches. The right lateral division subdivided into two branches opposite the head of the tenth rib. The smaller of these, after sending a dorsal muscular offset, coursed along the ninth rib and was lost, partly in distribution to the ninth and eighth interspaces, and partlyin anastomoses with the last ramifications of the posterior thoracic and musculo-phrenic arteries. The larger branch gave origin also to a dorsal muscular offset, and, continuing outwards for a short distance, bifurcated for the supply of the last two intercostal spaces. The whole of the right lateral division as well as part of its subdivision were covered by the rete to which they distributed innumerablevessels. The seventh of the inter- costal series of branches corresponded to a "last dorsal." Its course, being close to the posterior margin of the last rib, was in the abdominal wall. It was the smallest of the last three, and its lateral divisions were, like those of the others, unequal in size. That to the left, the larger, was directed above the rete mirabile as far as the posterior borderof thehead of the eleventh rib, along which it ramified, supplying the muscles in that region, and communicating with offsets from the fifth intercostal artery. When lying above the rete the latter received a rich supply of vessels, and when lying on the intertransversales it gave a dorsal muscular branch. The right lateral division was exceedingly small, and soon broke up into numerous twigs for THE RETE 'MIRABILE OF THE NARWHAL. 383 that portion of the rete corresponding with the tenth inter- costal space. Nowthat the main branches, whence the rete mirabile derives its vascular supply, have been described, the position of the principal part of the rete, that is, its thoracic portion, as well as the relations it bears to the thoracic vessels, will be easily understood. By the thoracic rete mirabile is understood that large mass of anastomosing vessels situated along the centre of the dorsalwall of the thorax. For convenience of description it may be said to consist of two elongated lateral masses, each lying athwart the ribs, and close to the side of the spine, the lateral masses being connected to each other by an intermediate central band, simi- larly constituted, and arching over the vertebral bodies. Each lateral mass measured 3 inches in breadth. They both pre- served their characteristic arrangement and structure from the base to the apex of the thorax, but beyond these limits they became altered and comparatively insignificant. Their exten- sion into the abdomen was scarcely perceptible, though their prolongation into the neck was more distinct and more readily traced. The intermediate band had the same extent in an antero-posterior direction, but was only 1A inch in breadth. The whole mass had thus a transverse measurement of about 7j inches. As the rete closely invested the structures over which it lay, it necessarily varied considerably in thickness. This was greatest nearest to the vertebral bodies where the anastomosing mass sunk deeply between the transverse pro- cesses and the heads of the ribs, and into the hollows of the vertebral grooves. It was least in the intermediate central band, which but thinly covered the bodies of thevertebrae. The parts of the thoracic wall concealed by the entire rete mirabile were the bodies of the dorsal vertebrae, the intervertebral fora- mina, and part of the ribs with the corresponding portion of their intercostal muscles. Though the meshwork closely in- vested these structures, fitting accurately into all theinequalities of their surfaces, yettheconnective tissue between them and the rete was of such a loose nature as to permit the entire mass to be reflected from them without much dissection. It was most firmly adherent over the transverse processes and heads of the 384 DR HENRY SEASON WILSON. ribs. The opposite aspect of the rete was almost entirely in contact with the pleura. The thoracic aorta, however, crossed lengthwise along the left of the central band, but the lateral masses were covered only by the pleurae. The free margins of the rete, well defined on each side, kept parallel to the spinal column, and were bounded along nearly the whole of their extent by the two posterior thoracic arteries. The three principal sources whence the rete mirabile derived the vessels entering into its composition were the posterior thoracic arteries on each side, and the thoracic aorta in the middle. The anastomotic vessels of the posterior thoracics branched in vast numbers from those portions of their trunks and of their intercostal offsets which were above the rete. As they emerged from these sources these branches presented one of the most beautiful of anatomical objects. The delicacy of their coats, the vast immensity of their numbers, their gently sinuous courses, their almost equal calibres, and their almost equally distanced origins at once marked them as peculiarly characteristic and uncommon. They soon became lost in intri- cate meanderings which, with care and patience, were gradually unravelled and their ramifications traced into the substance of the labyrinthine maze. These ramifications will be best under- stood by following as far as practicable any given branch. Quitting its parent trunk as a minute vessel it entered the rete with a sinuous course, retaining the same calibre throughout this extent. Once within the mass it lost its wavy undulations and wandered very tortuously. At the same time it became sometimes rapidly, sometimes gradually, widened into twice, thrice, and even four times its former size, and distributed here and there, at irregular intervals, secondary branches, having its own calibre, or occasionally rather smaller. Passing onwards in its intricate course, it branched on all sides until at last its individuality was lost in the complex network of the anastu- motic maze. Its secondary branches each presented a like distribution, and supplied tertiary offsets, which in their turn but repeated the same vascular intricacy, adding more and more to the general labyrinth. The interval between any two branches was occasionally very considerable, and in such cases the vessels wended about in a very tortuous manner, as if THE RETE MIRABILE OF THE NARWHNL. 385 profiting by their freedom from lateral ties. Nowhere did a secondary or a tertiary offset anastomose with like branches from a primary vessel. The other vascular source of the rete mirabile, the thoracic aorta, added its supply through the seven intercostal branches which have already been described. They entered the mass along its middle, and thus divided it into symmetrical halves. Though not presenting the same beautiful regularity in their origins, the vascular supply from these aortic branches was in every other respect similar to that-from each of the posterior thoracics. They entered the substance of the rete, and were there lost in the intricacies of the plexus. On unravelling the rete throughout its depth and to the extent of about the breadth of two or three intercostal spaces, the characters and arrangement of its constituent vessels were readily examined. In character and size they were divisible into three sets, easily distinguished from each other. For easier reference I have called them vasa maxima, vasa media, and vasa minima. The first set, vasa maxima, contained vessels of a large calibre, but possessing very thin walls. They were situated chiefly along the middle of the inferior aspect of the rete towards its posterior extent, and more particularly opposite the ninth and tenth intercostal spaces. Their walls were pecu- liarly puckered so as to give them an appearance somewhat like that of the large intestines. It was due to this puckering that these vessels became so curiously contorted when inflated, causing the whole superficial aspect of the rete to resemble a mass of cerebral convolutions in miniature. After loosening the vasa maxima from their connective tissue, the puckerings still remained, and seemed to be produced by a band or bands running more or less longitudinally in the coats of the vessels. The second set of vessels, vasa media, were smaller than the preceding, though their walls possessed a greater proportionate thickness. They were the most generally distributed through- out the mass, meandered in a very tortuous manner, and pre- sented puckered walls, though less distinctly marked than the preceding. The third set, vasa minima, consistedofvessels very much smaller than the preceding, but having proportionally the thickest walls. Their coats were entirely devoid of the pucker- ing so characteristic of the others, and they were generally 386 DR HENRY SEASON WITLSON. found deeply imbedded in the rete. In their course they were only slightly tortuous. They formed here and there distinctly isolated plexiform areas, easily distinguished from the rest by the straight, narrow, thread-like appearance of their constituent vessels. The anastomoses of each set were continuous with those of the others. The walls of the vasa maxima and media were lax, and the vessels appeared as if empty. This was doubtful in the case of the vasa minima. Occasionally vessels of a size intermediate to those specified were met with, but they were only exceptions. When compared with its great extent, the branches supplied by the rete mirabile to the structures beneath it were very few. These were chiefly muscular and osseous, and were most dis- tinctly seen on reflection of the central band from the bodies of the vertebrae, or of the lateral masses from the intercostal spaces. From their small and insignificant size, they probably but connected the vascular mass of the rete with nutrient vessels of the surrounding structures. They presented no puckering of their coats. In addition to the preceding, larger branches passed through the vertebral foramina into the spinal canal. They had puckered walls, and were evidently communi- cations between the rete and the vessels within the vertebral canal, or they extended a meshwork of the same character as the rete into that canal. In unravelling the rete mirabile many structures were found imbedded within its substance, or surrounded by its anasto- motic vascular plexus. These were-several veins, the first portion of the intercostal nerves, and the sympathetic ganglia with their branches. Some of the veins were large, all of them had very delicate walls. They seemed to pass towards the spine, and, while in the substance of the rete, they received many branches apparently derived from a very fine network situated within the interspaces and over the coats of the anasto- motic arteries. But they were much too small and delicate, and the parts much too decomposed for any satisfactory dissec- tion to be made with the scalpel. The intercostal nerves lay very deeply in the substance of the rete, and while thus covered each received its sympathetic root. The sympathetic ganglia were situated in the vertebral grooves close to the roots of the
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