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J. MARK BALDWIN, Johns Hopkins University
FRANK W. BANCROFT, University of California
LEWELLYS F. BARKER, University of Chicago
H. HEATH BAWDEN, Vassar College
ALBRECHT BETHE, University of Strassburg
G. E COGHILL, Pacific University
FRANK J. COLE, University of Liverpool
H. E. CRAMPTON, Columbia University
C. B. DAVENPORT, University of Chicago
WM. HARPER DAVIS, Lehigh University
HENRY H. DONALDSON, University of Chicago
LUDWIG EDINGER, Frankfurt a-M.

S. I. FRANZ, McLean Hospital, Waverley, Mass.
THOMAS H. HAINES, Ohio State University
A. VAN GEHUCHTEN, University of Louvain
R. G. HARRISON, Johns Hopkins University
C. F. HODGE, Clark University

S. J. HOLMES, University of Michigan
EDWIN B. HOLT, Harvard University
G. CARL HUBER, University of Michigan
JOSEPH JASTROW, University of Wisconsin
J. B. JOHNSTON, West Virginia University

B. F. KINGSBURY, Cornell University
FREDERIC S. LEE, Columbia University
JACQUES LOEB, University of California
E. P. LYON, St. Louis University

ADOLF MEYER, N. Y. State Pathological Inst.
THOS. H. MONTGOMERY, Jr., Univ. of Texas
WESLEY MILLS, McGill University

C. LLOYD MORGAN, University College, Bristol
T. H. MORGAN, Columbia University

A. D. MORRILL, Hamilton College
HUGO MUENSTERBERG, Harvard University
W. A. NAGEL, University of Berlin
G. H. PARKER, Harvard University
STEWART PATON, Johns Hopkins University
RAYMOND PEARL, University of Michigan
C. W. PRENTISS, Western Reserve University
C. S. SHERRINGTON, University of Liverpool
G. ELLIOT SMITH, Gov't. Medical School, Cairo
EDWARD L. THORNDIKE, Columbia University
JOHN B. WATSON, University of Chicago
W. M. WHEELER, Am. Museum of Nat. History
C. O. WHITMAN, University of Chicago

Published bi-monthly

DENISON UNIVERSITY, GRANVILLE, OHIO

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Comparative Neurology and Psychology

Volume XV

1905

Number 3

THE MORPHOLOGY OF THE VERTEBRATE HEAD FROM THE VIEWPOINT OF THE FUNCTIONAL DIVISIONS OF THE NERVOUS SYSTEM.1

By J. B. JOHNSTON.

With Plates I to IV.

TABLE OF CONTENTS.

5.

I.

2.

3.

Introduction.

a. Nature of the unsettled problems

b. Nerve components

C.

d.

Functional divisions of the nervous system

Bearing upon the subject of head morphology

Number and relations of mesodermic somites

Branchial apparatus and lateral musculature 4. Segmentation of the central nervous system

Typical sense organs of vertebrates

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The absence of nerve roots from one or more hind brain segments and

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The occipital region and the posterior limit of the head. The constitu

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1 Studies from the Zoological Laboratory of West Virginia University, No. 9.

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19. Relation of dorsal and ventral nerve roots to the myotomes

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a. Nature of the unsettled problems.

Shall we consider vertebrates as animals possessing a high degree of cephalization from their first appearace? The structure of their near relatives, Amphioxus and Ascidians, is against this view. The structural relations of vertebrates and invertebrates indicate that the ancestors of the vertebrates were segmented invertebrates in which the process of cephalization had not gone very far. Even within typical vertebrates evidence is not lacking that the special sense organs of the head were late to appear; that the branchial apparatus was at one time more extensive, reaching into what is now the trunk; that the nerves of the branchial region once had a more simple segmental arrangement; and that in the brain itself the several regions were once less highly specialized than at present. If Amphioxus be considered, the presence of true nephridia (41) in the head and the slight specialization in the head region seem to relate this "lowest vertebrate" with invertebrates rather far down the scale.

If, then, the ancestral vertebrate had only a slight head development, it is evident that the interpretation of the special organs of the head of typical vertebrates is to be reached by a study of their structure, function, and phylogenetic history, with a view to tracing them back to their unspecialized beginnings. When each organ has thus been followed back to its ancestral condition we shall have reduced the vertebrate head to terms-not of the trunk, but of a more simple condition which underlies both head and trunk. Such is the real problem of head morphology as the writer understands it.

The central difficulty in framing such a conception of the head is the matter of segmentation. Head specialization has

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