Afferent Projections to Pharynx and Soft
Palate Motoneurons: A Light and
Electron Microscopical Tracing Study in
the Cat
JOSE BOERS,
1
ANTOINETTE C. HULSHOFF,
2
HENK DE WEERD,
1
LEONORA J. MOUTON,
1
RUTGER KUIPERS,
1
AND GERT HOLSTEGE
1
*
1
Department of Anatomy and Embryology, Faculty of Medicine, University of Groningen,
9700 AD Groningen, The Netherlands
2
Department of Otorhinolaryngology, University Hospital Groningen, 9700 RB Groningen,
The Netherlands
ABSTRACT
Pharynx and soft palate are muscles for respiration, vocalization, swallowing, and
vomiting. In cat, motoneurons innervating pharynx/soft palate are located in the dorsal group
of the nucleus ambiguus (dgNA) in the medulla oblongata. In cat, dgNA is the only part of
nucleus ambiguus that can be distinguished as a separate cell group, which makes it possible
to study its afferent input. In two cats, WGA-HRP injections in dgNA and surrounding
tegmentum resulted in retrogradely labeled cells at several levels of the neuraxis. In 170
cases anterograde tracers were injected in areas in which the cells of origin were identified.
Results demonstrate that dgNA afferents originate from the tegmentum dorsolateral to the
superior olivary complex, medullary ventromedial tegmentum, caudal raphe nuclei, medul-
lary lateral tegmental field, nucleus retroambiguus (NRA), and adjoining tegmentum, ex-
tending into the first cervical segment of the spinal cord. In order to determine whether
periaqueductal gray (PAG) and parabrachial nuclei (PB) make synaptic contacts with dgNA,
ultrastructural studies combined anterograde tracing from PAG, PB, and NRA with retro-
grade tracing of pharyngeal and soft palate motoneurons. The results showed that PB, but
not PAG, projects to the dgNA and that NRA afferent synapses are three times as numerous
as those from PB. The morphology of PB and NRA synapses is consistent with excitatory
input. In conclusion, pharyngeal and soft palate motoneurons receive their afferents almost
exclusively from the pontine and medullary tegmentum and first cervical spinal segment. J.
Comp. Neurol. 486:18 –38, 2005. © 2005 Wiley-Liss, Inc.
Indexing terms: nucleus ambiguus; vocalization; respiration; parabrachial nuclei; periaqueductal
gray; nucleus retroambiguus
The pharynx and soft palate play a crucial role in res-
piration (Lunteren and Dick, 1996), swallowing (Doty and
Bosma, 1956; Miller, 1972), vomiting (Gre ´lot and Miller,
1997), and vocalization (Thoms and Ju ¨ rgens, 1987; Saka-
moto et al., 1996). Both groups of muscles are innervated
by motoneurons located in the so-called dorsal group of the
nucleus ambiguus (dgNA; Lawn, 1966; Yoshida et al.,
1981; Holstege et al., 1983; Davis and Nail, 1984). In cat,
the nucleus ambiguus (NA) is located in the medullary
lateral tegmental field, extending from just caudal to the
facial nucleus rostrally to the level of the obex caudally
(Holstege et al., 1983; Davis and Nail, 1984). At most
levels the NA is difficult to recognize, because in each
transverse section only a few scattered NA motoneurons
are present. In contrast, the dgNA is the only part of NA
that can be recognized as a distinct nucleus in Nissl-
stained sections. Despite the great importance of pharynx
*Correspondence to: Gert Holstege, Dept. Anatomy and Embryology,
Univ. Groningen, A. Deusinglaan 1, 9700 AD Groningen, The Netherlands.
E-mail: g.holstege@med.rug.nl
Received 4 June 2004; Revised 4 October 2004; Accepted 4 December
2004
DOI 10.1002/cne.20530
Published online in Wiley InterScience (www.interscience.wiley.com).
THE JOURNAL OF COMPARATIVE NEUROLOGY 486:18 –38 (2005)
© 2005 WILEY-LISS, INC.