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( 105 ) V.—Distribution of the Cells in the Intermedio-Lateral Tract of The

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( 105 ) V.—Distribution of the Cells in the Intermedio-Lateral Tract of The ( 105 ) V.—Distribution of the Cells in the Intermedio-Lateral Tract of the Spinal Cord, By Alexander Bruce, M.A., M.D., F.R.C.P.E., F.K.S.E., Physician to the Edin- burgh Royal Infirmary. (With One Plate and Twenty-four Figures.) (Read June 5, 1905 ; MS. received November 16, 1905. Issued separately March 6, 1906.) The term intermedio-lateral tract was introduced in 1859 [Phil. Trans., 1859, p. 445) by LOCKHART CLARKE to designate a tract or column of nerve cells in the spinal cord, which he had previously described in 1851 [Phil. Trans., 1851, ii. p. 613) as occupying that portion of the lateral margin of the grey matter which is intermediate between the anterior and posterior cornua. According to CLARKE'S original account, the column in question was very transparent in appearance, and resembled somewhat the substantia gelatinosa of the posterior horn. It was found in the upper part of the lumbar enlargement, extended upwards through the dorsal region, where it distinctly increased in size, to the lower part of the cervical enlargement. Here it disappeared almost entirely. In the upper cervical region it was again seen, and could be traced upwards into the medulla oblongata, where, in the space immediately behind the central canal, it blended with its fellow of the opposite side. In the more complete account of the tract published in 1859 (p. 446), its component cells are described as in part oval, fusiform, pyriform, or triangular, and as being smaller and more uniform in size than those of the anterior cornua. In the mid-dorsal region, where they are least numerous, they are found only near the lateral margin of the grey matter, with the exception of some cells which lie among the white fibres beyond the margin of the grey substance. In the upper dorsal region the tract is larger, and not only projects further outwards into the lateral column of the white fibres, but also tapers inwards across the grey substance, almost to the front of Clarke's column. In the cervical enlargement it gradually disappears, although it seems to contain, in part at least, a few scattered cells resembling those of the intermedio-lateral tract of the dorsal region. In the upper cervical region, as already stated, it is again seen occupying a lateral horn similar to that found in the dorsal region. It is composed of the same kind of cells, and can be followed up into the medulla, where it is said to give origin to some of the fibres of the vagus and the spinal accessory. WALDEYER, GASKELL, SHERRINGTON, MOTT, and others have drawn attention to the probability that the dorsal vagus nucleus of the medulla belongs to the same system as the intermedio-lateral tract. GASKELL has shown that sympathetic fibres pass out with the second and third sacral roots, and SHERRINGTON, without giving a definite opinion on the subject, has suggested that the cells which appear in the sacral region as a lateral horn probably belong to the intermedio-lateral tract system. TRANS. ROY. SOC. EDIN., VOL. XLV. PART I. (NO. 5). 14 106 DR ALEXANDER BRUCE As the present paper concerns itself, however, purely with the tract as it is found in the dorsal, lower cervical, and upper lumbar regions, the question raised in the two previous paragraphs as regards its distribution in the upper cervical and lower sacral regions will not be further referred to. The majority of anatomists who have written on the subject since CLARKE'S original papers appeared have regarded the intermedio-lateral tract as being synonymous with the cells of the lateral horn. This view is not exactly in accordance with the definition or the description and figures of CLARKE, an examination of which shows that, as already stated, he was aware that the tract passed beyond the limits of the lateral horn, both backwards along the margin of the grey matter and also inwards towards the column of CLARKE (Phil. Trans., 1859, p. 446, PI. XX. figs. 2 and 4 ; also PL XXI. figs. 3, 5 and 6). WALDEYER, in his work on the Spinal Cord of the Gorilla (Abth. der Kb'nig. Akad. derWissenschaft, Berlin, 1898), has subjected the intermedio-lateral tract to a careful examination, and has done more than any other writer since CLARKE to advance our knowledge and broaden our views with regard to the character and distribution of its cells. He makes it clear that the cells of the tract are not limited to the lateral horn, but that also at the margin of the grey matter near the formatio reticularis there are found cells identical with those in the lateral horn. These are, in his opinion, therefore, component parts of the intermedio-lateral tract. He is further of opinion that the tract does not, as CLARKE thought, disappear in the cervical and lumbar enlargements, but that it is found throughout the whole length of the cord. While he admits that it may disappear as a lateral horn in the cervical enlargement, he maintains that it is continued upwards as the cells of the formatio reticularis. He is further in agreement with KRAUSE in thinking that the lateral horn of the dorsal region is not identical with the lateral projection of the anterior cornua in the cervical and lumbar enlargements, and that the cells of the intermedio-lateral tract are not mere transformations or modifica- tions (SCHWALBE, BRB, OBERSTEINER, QTJAIN) of the cells in the lateral part of the anterior cornua, but are of altogether independent origin and nature. SHERRINGTON (Journ. Physiol., 1892, p. 698), with reference to these conclusions of WALDEYER'S, says : " WALDEYER, in his description of the grouping of the cells in the cord of the gorilla, says that the lateral horn cells are found throughout the cord in all its segments, including those of the cervical and lumbar enlargements. He divides the ganglion cells of the cord into as many as fourteen definite groups, attaching to each a name, but treats the cells of the lateral reticular formation and those of the lateral horn as one group (Mittelzellen). When he says that the cells of the lateral horn are present in, for instance, the eighth and seventh cervical segments, his meaning is that there are cells in the reticular formation at that level which he considers are cells of the lateral horn, although that horn can no longer with certainty be recognised. But in regions where the lateral horn exists there are cells in the lateral reticular formation as well as in the lateral horn, and the assumption that the cells of the lateral reticular formation, although somewhat similar to the cells of the lateral horn, are identical with ON DISTRIBUTION OF THE CELLS IN THE INTERMEDIO-LATERAL TRACT. 107 them, or are their equivalents, is a somewhat insecure foundation on which to rest the statement that the cells of the lateral horn are present in a region where there is no other evidence of them beyond cells in the lateral reticular formation." SHERRINGTON has not been able to trace lateral horn cells higher than the middle of the superficial origin of the eighth cervical nerve. In Rhesus he has not found them above the surface origin of the first thoracic root. The lower extremity of the tract he finds to correspond to the surface origin of the fourth lumbar nerve. This would correspond to the third lumbar segment in man. As regards the longitudinal distribu- tion or grouping of the cells in the intermedio-lateral tract, WALDEYER notes, on p. 19, that they lie close together, and that they may be arranged in groups or clusters separated by interspaces in which there are no cells. " Beziiglich ihrer Anordnung ist zu sagen dass sie gewohnlich dicht zusammengedrangt liegen. Selbst wenn keine grossere Zahl dieser Zellen vorhanden ist liegen sie haufig zu zweien, dreien oder vieren nahe beisammen ; zwischen diesen einzelnen kleinen Gruppen konnen dann allerdings grossere Zwischenraume vorhanden sein. Jedenfalls bilden diese Zellen stets eine besondere Formation im Riickenmarke." This arrangement of the cells in groups or clusters has also been referred to by AINSLIE HOLLIS, in the Journal of Anatomy and Physiology, vol. xvii., 1883, p. 63 : " The (intermedio-lateral) tract consists of clusters of small (mostly pyriform) cells arranged linearly, and dissociated from each other, and from the column of sparsely scattered giant cells to which they are contiguous, by a delicate fibrillar stroma of synectic tissue. In the mid-dorsal region I have observed two adjacent columns of these cell-clusters." Again, at p. 520, referring to " the vesicular columns of CLARKE and the tractus intermedio-lateralis," he says that " in certain parts of the cord they are found closely congregated in cell-nests." MOTT (Brain, 1890, p. 444) says: "The cells of the intermedio-lateral tract are found throughout the dorsal region. The cells are bi-polar, and in vertical sections they are often seen to exist as little groups or nests of vesicular cells, from eight to twelve in number." ONUF and COLLINS (Sympathetic Nervous System, 1900, p. 140), in describing the cells of the lateral horn, say : " The group is represented by a very pure type in the lower dorsal region. In longitudinal sections it does not appear in the form of a continuous column, but segmented, in the form of cell-nests distributed at intervals." ARGTJTINSKY, a pupil of WALDEYER'S, in a valuable paper in vol. xlviii. of the Archiv fur mikroskopische Anatomie (1897), " Ueber eine regelmassige Gliederung in der grauen Substanz des Riickenmarks beim Neugeborenen und iiber die Mittelzellen," in which he for the first time describes a segmentation of the " Mittelzellen" of WALDEYER, refers, on p.
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