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ROGER WARWICK
Department of Anatomy, University of Manchester
seriously weakened also by Le Gros Clark (1926), who contrasted the absence
or undifferentiated nature of Perlia's nucleus in some primates, such as
Tarsius and the macaque, with its exuberance in certain other mammals,
like Sciurus and Tupaia, with no claim to binocular function.
Despite these damaging objections, the nucleus of convergence, as a
structural and functional entity, has remained a feature of standard
accounts and is propounded by many current monographs with little
or no apparent misgiving (vide, Duke-Elder (1932), Rea (1938), Walsh
(1947), Mettler (1948), Berens (1949), Wolff (1954), etc.). A few authorities,
such as Adler (1953), express doubts, but the concept continues, on the
whole, deeply engraven in neurological teaching. Indeed, Stutterheim
(J937, 1950) had no qualms in using it as a major keystone in a new
INTRINSIC MUSCLES
RECTUS MEDIALIS
OBLIQUUS (IN CONVERGENCE)
INFERIOR
RECTUS
INFERIOR
OBLIQUUS. L TROCHLEAR
INFERIOR NUCLEUS
RESULTS
Topographical Observations
The right and left somatic or main oculomotor nuclei of the macaque
are more compact and nearer each other than is so in the human mid-brain
(cf. figs. 4 and 7, see Plates at end), but they are nevertheless clearly
separated by a mid-line zone, the median raphe. This is a complex
arrangement of axons, dendrites, and neuroglia, which also contains nerve
cells) mostly multipolar and variable in size and in number.
At caudal levels of the oculomotor complex this intermediate zone is
wide dorsally in the macaque, chimpanzee and man. Hence, in transverse
sections, it has the appearance of a cuneiform mass, as if driven in between
the oculomotor nuclei from the direction of the aqueduct (Plate IX, fig. 2).
Situated centrally within this wide caudal part of the median raphe is a
dorsal unpaired group, the caudal central nucleus. This consists of multi-
polar neurones smaller than those of the chief oculomotor mass (cf. figs.
8 and 9), the cells being less compactly arranged in the chimpanzee and
man than in the monkey, where they form a highly distinctive feature
(fig. 2). This nucleus is usually accepted as Perlia's "diffuse nucleus of the
mid-line" (his original description referred to man). In its dorsal and
caudal position it certainly does not correspond with his better known
central nucleus (of convergence). In addition, it has been shown that the
caudal central nucleus innervates the Levatores in monkeys (Warwick,
1953fl). The same nucleus appears to be identical with the posterior dorso-
central nucleus of Panegrossi (Warwick, 1953b).
THE SO-CALLED NUCLEUS OF CONVERGENCE 95
EXPERIMENTAL OBSERVATIONS
The findings detailed below were derived from study of mid-brains of
95 monkeys submitted to various orbital and intracranial interruptions of
THE SO-CALLED NUCLEUS OF CONVERGENCE 99
the oculomotor nerve and its branches. These were examined in serial
preparations with special reference to the following questions:
(1) Are the nerve cells of a central nucleus of Perlia, where this
exists in the monkey, a radicular source of oculomotor fibres or
not?
(2) Do the neurones of such a nucleus show retrograde degenera-
tion following denervation of the Rectus medialis?
(3) If not situated in Perlia's nucleus, in part or whole, what is the
locus of the motor pool of the Rectus medialis?
(4) Does interference with the nerve supply of any other extra ocular
muscle produce observable changes in the central nucleus?
DISCUSSION
Binocular movements may be categorized for convenience as conjugate
deviation, convergence, and divergence, according to whether the visual
axes are moved in parallel, made to approach each other to intersect at a
near object, or moved apart again for more distant fixation. Confusion
sometimes arises, however, over the term divergence, which is sometimes
called "negative convergence." This usage seems misleading, for con-
vergence and divergence are evidently the same process in opposite phases,
both being equally exact and active in performance. Divergence is also
centre for convergence assumes a lesser role than its description suggests.
Precisely what those who have favoured this concept had in mind con-
cerning its functions must therefore be asked. The majority of recent
writers are vague upon this point, and the expression "nucleus for con-
vergence" appears to have been copied from text to text over many years
with little critical enquiry into its sources.
To postulate some sort of unpaired or median control of bilateral
structures in intimate association seems at first sight plausible, but it is
difficult to follow why this should be required for convergence and yet
not for other binocular co-ordination, though it is true that a centre for
divergence has been suggested (Holmes, 1921). A median brain-stem
centre has not been regarded as necessary in connexion with other motor
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found Perlia's nucleus absent from just this fraction of the human mid-
brains examined by him.
Evidently in anthropoids the parallel between the development of
Perlia's nucleus and the evolution of convergence urged by Brouwer (1918)
is untenable, and Le Gros Clark (1926), in a broader comparative series,
found other anomalies which cast grave doubts upon Brouwer's phylo-
genetic arguments. Indeed, one would have expected that his observa-
tions would in themselves have proved sufficient to dismiss Perlia's nucleus
as a centre for convergence. This concept was, however, already so far
accepted that neither then nor since has any progress been made by the
facts of comparative and topographical study, although they have now
accumulated for half a century in contradiction.
might help to clarify these obscurities, are not available. It would, for
example, be of interest to know whether a median oculomotor nucleus is
a common feature and whether it appears in the embryos of species which
do not display it after birth.
It is evident that the embryological data so far recorded, though
admittedly inadequate, lend no support to the concept under discussion,
tending rather to add to the gravamen of the anatomical denial.
Since the belief in Perlia's nucleus as a mediator of convergence fails
to carry conviction on purely anatomical grounds, the second criterion of
its validity—the function of its constituent nerve cells—must next be
considered.
In previous papers (Warwick, 1950, 1953a) it was demonstrated, in
FIG. 16.—Diagrams showing how the folding of the basal laminae of the human
embryonic mid-brain may produce an approximation of the somatic oculomotor
masses and a change in the position of the visceral, Edinger-Westphal columns
(E.-W.), from a lateral to dorsal position.
growth processes would account for the final position of the developing
Edinger-Westphal nuclei dorsal, or even dorso-medial, to the somatic
oculomotor masses if, as is usually stated, these visceral groups arise
further laterally in the basal larninse. Fusion of the oculomotor nuclei
across the mid-line may be in part a consequence of their approximation
and also of proliferation of their cells. Le Gros Clark (1926) has noted a
phylogenetic tendency to an increase in the number of oculomotor
neurones in higher forms. The growing axons of the Edinger-Westphal
nuclei have to pass ventrally to enter the third nerve, and in adult primate
material it can be seen that many do so by traversing the median raphe,
THE SO-CALLED NUCLEUS OF CONVERGENCE 109
forming the fibrse recta? of Bernheimer (1894) (fig. 17). If these axons do
not decussate, and evidence of this has already been recorded (Warwick,
1954), it is not difficult to visualize the two groups of fibres cleaving a
median nucleus from a junctional mass of nerve cells between the third
nerve nuclei. Moreover, in species whose right and left nuclei are more
closely juxtaposed, like the macaque, bands of ventrally running fibres
might equally well separate off the paramedian groups so characteristic of
this animal. It is also possible to explain in this manner, the inconstant
occurrence and loose arrangement of the paramedian groups noticed in the
human and chimpanzee series of this research, for in both forms the
median raphe is more spacious than in the macaque. These views are,
however, merely suggestions, and further embryological observation is re-
quired to elucidate such points.
Finally one may enquire whether some other centre exists, in the mid-
brain or elsewhere, for the integration of convergence. Although this
study affords no positive evidence concerning this, it is not out of place to
raise certain issues, for these have a direct bearing upon the concept under
110 ROGER WARWICK
ACKNOWLEDGMENTS
REFERENCES
ADLER, F. H. (1953) "Physiology of the Eye." 2nd edit. London, pp. 366 and 367.
BERENS, C. (1949) " The Eye and its Diseases." 2nd edit. Philadelphia and London.
pp. 69 and 806.
BERNHEIMER, S. (1894) "Das Wurzelgebiet des Oculomotorius biem Menschen.'
Wiesbaden.
, (1897) v. Graemes Arch. Ophthal., 4 4 , 481.
BROUWER, B. (1918) Z. ges. Neurol. Psychiat., 4 0 , 152.
BRUCE, A. (1889-90) Proc. roy. Soc. Edinb., 17, 168.
CLARK, W. E. L E GROS (1926) / . And., Lond., 60, 426.
COOPER, E. R. A. (1946) Brain, 69, 50.
CROSBY, E. C , and WOODBURNE, R. T. (1943) / . Comp. Neurol, 78, 441.
DUKE-ELDER, W. S. (1932) " Textbook of Ophthalmology." London, vol. I, pp
278-279.
EDINGER, L. (1900) " Vorlesungen fiber den Bau der nervosen Centralorgane des
Menschen und der Thiere." 6th edit. Leipzig.
THE SO-CALLED NUCLEUS OF CONVERGENCE H3
FOERSTER, O., GACEL, 0., and MAHONEY, W. (1936) Verh. dtsch. Ges. inn. Med., 48,
386.
HOLMES, G. (1921) Brit. J. Ophthal., 5, 241.
KNTES, M. (1891) Arch. Augenheilk., 23, 19.
LORENTE DE N6, R. (1947) /. cell. comp. Physiol., 29, 207.
MAJANO, N. (1903) Mschr. Psychiat. Neurol, 13, 291.
MANN, I. C. (1927) /. Anat., Lond., 61, 424.
METTLER, F. A. (1948) "Neuroanatomy." 2nd edit. London, p. 334.
VON MONAKOW, C. (1905) " Gehirnpathologie." Wien. p. 1040.
PANEGROSSI, G. (1898) Ric. Lab. Anat. norm. Univ., Roma, 6, 103. ;
, (1904) Mschr. Psychiat. Neurol, 16, 268 and 344.
PATON, L., and MANN, I. C. (1925) Trans. Ophthal. Soc. UX.., 45, 610.
PEARSON, A. A, (1944) /. comp. Neurol, 80, 47.
LEGENDS TO PLATES
PLATE IX
FIG. 2.—Transverse section of monkey'sjjculomotor complex in its lower (caudal)
third, showing the caudal central nucleus (C.C.N.). Cresyl fast violet, x 30.
FIG. 3.—Transverse section at upper (cranial) limit of monkey's oculomotor
complex, which is represented at this level only by the antero-median nucleus (para-
sympathetic component). Cresyl fast violet, x 30.
FIG. 4.—Typical transverse field at intermediate levels of oculomotor mass in
the monkey. Note loose paramedian groups of cells (P.M.) near mid-line and absence
of central nucleus of Perlia. Cresyl fast violet, x 30.
FIG. 5.—Transverse section through monkey's oculomotor nuclei showing ventrally
a central nucleus of Perlia (P.). The dorsal median mass (E.-W.) consists of the
Edinger-Westphal columns, joining to form the antero-median nucleus. Cresyl fast
violet, x 30.
114 ROGER WARWICK
PLATE X
FIG. 6.—Transverse section of chimpanzee's oculomotor complex in its upper
(cranial) half. Note breadth of median raphe, poorly developed Perlia's nucleus (P.),
Edingcr-Westphal columns (E.-W.), and cranial extension of caudal central nucleus
(C.C.N.). Cresyl fast violet, x 30.
Fie. 7.—Adult human oculomotor nuclei in transverse section at similar level
to figs. 4-6. Perlia's central nucleus (P.) is represented by a few cells in the ventral
part of the median raphe. The Edinger-Westphal nuclei (E.-W.) are well-developed.
Cresyl fast violet, x 20.
FIG. 8.—Typical multipolar nerve cells of the main (somatic) oculomotor nucleus.
Compare with figs. 9 and 10. Monkey. Cresyl fast violet x 550.
Fie. 9.—Nerve cells of the caudal central nucleus (u. fig. 2). Monkey. Cresyl fast
violet, x 550.
PLATE XI
Fie. 11.—Transverse section of chimpanzee's oculomotor nucleus near its upper
(cranial) extremity. Note absence of Perlia's nucleus. Dorsally the Edinger-Westphal
columns (E.-W.) are joining together. Cresyl fast violet x 30.
FIG. 12.—Transverse section of oculomotor complex in a monkey killed 10 days
after extirpation of the left Rectus superior. A nucleus of Perlia (P.) is present, and
retrograde degeneration has occurred in some of its nerve cells (v. fig. 14). Cresyl
fast violet, x 30.
FIG. 13.—Part of the field shown in fig. 5, from mid-brain of a monkey subjected
to extirpation of right Rectus medialis. Swelling, nuclear eccentricity, and
chromatolysis are seen in cells of the right oculomotor nucleus but not in the
adjoining nucleus of Perlia (P.). Cresyl fast violet, x 100.
FIG. 14.—'Higher magnification of some of the cells forming the nucleus of
Perlia shown in fig. 12. About half of these show the changes characteristic of
retrograde degeneration, following extirpation of one Rectus superior. Cresyl fast
violet, x 150.
PLATE IX
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FIG. 4 FIG. 5