Embriology, Variations and Innervations of The Human Neck Muscles

HEAD & NECK CME
ABBREVIATION KEY
FGF ⫽ fibroblast growth factor
mRNA ⫽ messenger RNA is
synthesized from a DNA template;
during transcription it mediates
the transfer of genetic information
from the cell nucleus to ribosomes
Embryology, Variations, and in the cytoplasm, where it serves

as a template for protein synthesis
MYOD ⫽ a transcription factor
Innervations of the Human protein that plays a major role in
regulating muscle differentiation
Neck Muscles NOTCH ⫽ Notch signaling pathway

is a highly conserved cell signaling
system present in most
P.M. Som and J.T. Laitman multicellular organisms; there are
4 different notch receptors,
referred to as NOTCH1, NOTCH2,
NOTCH3, and NOTCH4; the notch
receptor is a single-pass
CME Credit transmembrane receptor protein
The American Society of Neuroradiology (ASNR) is accredited by the Accreditation Council for Continuing Medical Education and notch signaling is required in
(ACCME) to provide continuing medical education for physicians. The ASNR designates this enduring material for a maximum of one regulating cell polarity and cell–
AMA PRA Category one creditTM. Physicians should claim only the credit commensurate with the extent of their participation in the
cell communication
activity. To obtain credit for this activity, an online quiz must be successfully completed and submitted. ASNR members may access
this quiz at no charge by logging on to eCME at http://members.asnr.org. Nonmembers may pay a small fee to access the quiz and WNT ⫽ Wingless/int1 family of
obtain credit via http://members.asnr.org/ecme. secreted signaling molecules
Received December 10, 2015;

ABSTRACT accepted March 31, 2016.
The embryology of the muscles of the neck and face have been somewhat neglected in the From the Departments of
Radiology (P.M.S.), Otolaryngology
literature. In part, this reflects the fact that relatively little is actually known regarding the (P.M.S., J.T.L.), and Medical
development of these muscles. However, a picture has emerged that reveals that the neck Education (J.T.L.), Ichan School of
Medicine at Mount Sinai, New York,
muscles have branchial, myotomic, and mesenchymal origins. This review discussed what is New York.
presently known regarding the embryology of these muscles, with drawings to help the Please address correspondence
reader follow the text. It also discussed the variants of these muscles, which can cause to Peter M. Som, MD, Department
of Radiology, The Mount Sinai
confusion during surgery, and the embryology and variations of the nerves that innervate Hospital, One Gustave Levy Place,
these muscles. New York, NY 10029; e-mail:
Peter.Som@MSSM.edu
Learning Objective: The reader will understand the origins of the various muscles of the face http://dx.doi.org/10.3174/ng.3170206
and neck, and the origins of the nerves that innervate these muscles. Disclosures
Based on information received
from the authors, Neurographics
INTRODUCTION overview of our current understanding of has determined that there are no
Financial Disclosures or Conflicts
The detailed embryology of the neck mus- the embryology of the muscles of the neck, of Interest to report.
cles is not as well documented in the liter- with numerous illustrations to help clarify
ature as other areas of head and neck em- the text. The variations in these muscles
bryology. It was the embryology manuals and the variations and embryology of the
written by Keibel and Mall1,2 in 1910 and nerves that innervate these muscles will
1912 that likely describe in the most detail also be reviewed.
the development of the muscles of the neck,
and there are only a few additional detailed THE SOMITES AND MYOTOMES
reports.3,4 The facial muscle embryology Somitogenesis is regulated by a clock and
was discussed in an earlier review in this wavefront mechanism that is controlled by
series as was the laryngeal musculature and oscillating NOTCH and WNT signaling.
the tongue and pharyngeal musculature.5-7 This segmentation clock controls the
The muscles of the neck come from 3 main expression of segmentation cyclic genes,
sources, the branchial apparatus, myo- which provides a wave of fibroblast
tomes that arise from the somites, and growth factor (FGF) and of other proteins
myogenic cells from the paravertebral me- that has a ventral to dorsal gradient and
soderm. The following review presents an that establishes the rhythm of somitogen-
Neurographics 2017 May/June; 7(3):215–242; www.neurographics.org 兩 215

esis. At approximately 20 embryonic days, this wave causes the myotome have been defined based on these molecular
a compacting of the unsegmented paraxial mesenchyme biologic criteria.13 The primaxial domain refers to areas
that lies on either side of the neural tube into spherical that are composed only of somatic cells, and its myoblasts
epithelial somites that surround a central cavity that con- are the population of cells adjacent to the neural tube that is
tains free somitocele cells, often referred to as the core affected by neural tube signaling to generate muscle precur-
cells.8,9 These initial somites will eventually form bilaterally sors (Fig 4). These premuscle cells have limited migratory
paired epithelial blocks of paraxial mesenchymal meso- potential, and 1 group of these cells migrates onto the ven-
derm that develop along the head-to-tail axis of the devel- tral face of the adjacent developing vertebra and forms the
oping embryo and flank the notochord (Fig 1). These flexor muscles of the cervical and upper thoracic vertebra.
somites initiate from the ventral end of the embryo and The primaxial cells will also give rise to the extensor mus-
proceed dorsally. cles of the vertebra, which include the erector spinae, the
Shortly after they appear, the somites proceed to subdi- multifidus, the semispinalis and rotatores, and the splenius
vide into dorsal and ventral divisions. The ventromedial and suboccipital muscles.14,15 Likely arising from paraspi-
part of the somite undergoes an epithelial-to-mesenchymal nal mesenchyme are 3 additional premuscle masses. One
transformation, and these cells, together with the core cells, mass will develop into the suprahyoid and infrahyoid mus-
will form the sclerotome, which will develop into the verte- cles (strap muscles), 1 mass will become the scalene and
brae and ribs. After formation of the sclerotome, the re- geniohyoid muscles, and, finally, 1 mass will develop into
mainder of the somite consists of a dorsal epithelial layer, the levator scapulae and serratus anterior muscles.
which is the dermomyotome or the epithelial plate.10,11 The abaxial domain contains somatic cells that migrate
Initially, the dermatomyotome retains its epithelial struc- and mix with the lateral plate cells. The abaxial myotomes
ture and contains presumptive myogenic and dermal cells. are ventrolateral myoblasts that are farther from the neural
The dermatomyotome then gives rise to the more laterally axis and thus respond to signals from the adjacent lateral
and superficially placed dermatome, which will give rise to plate mesoderm and ectoderm. They give rise to a migratory
portions of the dermis of the back, and the more medially population of muscle precursors that stream out into the
positioned myotome, which contains dorsal edges of the body and form the muscles of the trunk and limbs. The
dermomyotome. boundary between the primaxial and abaxial domains is
Traditionally, each myotome then divides into a dorsal referred to as the lateral somatic frontier, and, as noted, the
epaxial division (epimere) and a ventral hypaxial (hypom- primaxial and abaxial domains are patterned indepen-
ere) division. This divisional nomenclature was originally dently.14,15 As such, the terms primaxial and abaxial are
based on the locations of the future adult muscles. At the not exactly equivalent to epaxial and hypaxial, and the
same time that the myotome divides, each developing spinal lateral somatic frontier does not separate hypaxial and ep-
nerve also divides into a ventral primary ramus that supplies axial structures (Fig 4).
the hyaxial division muscles and a dorsal primary ramus
that supplies the epaxial division muscles (Figs 2 and 3). MYOTOME TO SKELETAL MUSCLE
Thus, all of the future somitic muscles will be innervated The initial indication that myogenesis is occurring is the
by the spinal nerves. Between embryonic days 20 and 30 elongation of the nuclei and cell bodies of the mesenchymal
(the somite period of development), 38 pairs of somites will cells that will differentiate into premyoblasts, which, in
develop, and, by the end of the fifth embryonic week, there turn, will differentiate into myoblasts. Sonic hedgehog pro-
will be 42– 44 pairs of somites, although some of the most tein signaling from the ventral neural tube and notochord,
distal somites will regress, which usually leaves 35–38 and WNT and bone morphogenetic protein 4 signaling
somite pairs.10
from the overlying ectoderm regulate the initiation of myo-
genesis and the induction of the myotomes.10 The actual
THE SYNDETOME CELLS transformation of the mesenchymal cells into myogenically
As a result of close interactions between the myotome and committed cells is regulated by the MYOD family of tran-
the sclerotome, which involve both induction signaling and scription factors.16
regression, tendon progenitor cells develop in the interface The proliferating myogenic cells remain in the cell cycle
between the myotome and the sclerotome. These cells com- because of the action of growth factors such as FGF and
prise the syndetome, which will give rise to tendons. It is the transforming growth factor ␤. Because these myogenic reg-
close spatiotemporal regulation of these 3 cell populations ulatory factors accumulate within the cells, the synthesis of
that assures that the bones, tendons, and muscles will de- the cell cycle protein p21 is upregulated, and this irrevers-
velop into a functional musculoskeletal system (Fig 2).12 ibly removes the myoblasts from the cell cycle. Additional
growth factors cause the myoblasts to transcribe the mR-
DIFFERENT CELL POPULATIONS IN THE MYOTOME NAs necessary to produce the major contractile proteins
To more fully describe the changes in cell distribution, the actin and myosin. The myoblasts then start to fuse with
mixing of somatic and lateral plate cell populations, and other myoblasts to form the multinucleated myotubes or
the different patterning properties, 2 different domains of myofibers (Fig 5A). At this time, the regulatory muscle con-
216 兩 Neurographics 2017 May/June; 7(3):215–242; www.neurographics.org

Fig 1. A, Serial drawings of different embryos, illustrating the emergence of the somites and their increase in number as the embryo grows (modified
with permission from Hamilton WJ, Boyd JD, Mossman HW. Human Embryology. 2nd ed. Baltimore: Williams and Wilkins; 1952 [Figs 97–99]; and
Cochard LR. Netter’s Atlas of Human Embryology. Teterboro, NJ: Icon Learning Systems; 2002 [Fig 6.18, p 149]). B, Drawing, illustrating the devel-
opment of the somites and, in particular, the emergence of the myotomes from which the skeletal muscles will develop (modified with permission from
Ref 11 [Fig 8 –3, p 223]).

whereas the endomysium is formed by the external lamina
and reticular fibers. Eventually, the adult striated muscle is
formed (Fig. 5C). Further muscle-fiber growth is accom-
plished by a population of myogenic cells called satellite
cells, which are located between the muscle fibers and the
basal lamina that encases them.10,11,17
THE FACIAL MUSCLES

The embryology of the facial muscles has been discussed in
a previous review in this series as has the embryology of the
facial nerve.6,18 However, because 2 of these muscles qual-
ify as neck muscles, it seems appropriate to briefly review
this muscle group. These muscles include all of the muscles
supplied by the facial nerve, the nerve of the second bran-
chial (hyoid) arch. Between 27 and 32 embryonic days, the
second arch mesoderm immediately dorsal to the first bran-
Fig 2. A coronal drawing, illustrating the traditional separation of the
chial groove becomes increasingly attenuated with compact
somite into the sclerotome, the myotome, and the intervening syn-
detome; the emergence of the epaxial and hypaxial cells is also shown mesenchyme (Fig 6A). At approximately 32 embryonic
(modified with permission, from Ref 12). days, the main stem of the facial nerve extends into the
hyoid arch mesenchymal mass and the distal end of the
nerve has a brushlike configuration. Between 32 and 45
embryonic days, arising from this attenuated mesenchyme
are sheetlike laminas of premyoblasts, and early myoblasts
extend into the superficial part of the temporal, occipital,
cervical, and mandibular regions (Fig 6B). As the laminas
increase in size, they spread ventrally and dorsally into the
facial region as well as dorsally toward the shoulder to form
the platysma.1,19
The medial aspect of the premuscle mass also becomes
thickened to form the rudiment of the stylohyoid muscle,
the posterior belly of the digastric muscle, and the stapedius
muscle. As the muscle rudiments spreads out, the facial
nerve divides into several branches, which follow the wan-
derings of the muscle tissue. By 39 – 40 embryonic days, the
nerve stem distal to the chorda tympani and stapedius mus-
cle primordium divides into 3 branches, a thin one directed
Fig 3. Drawing, showing the division of the spinal nerve into ventral root medially sends nerves into the rudiments of the posterior
that will supply the hyaxial division of the myotomes and the dorsal root belly of the digastric muscle and stylohyoid muscles. A sec-
that will supply the epaxial division of the myotomes (modified with
ond, larger, branch, the posterior auricular nerve extends
permission from Netter FH. The CIBA Collection of Medical Illustrations.
Volume I, Nervous System Part I Anatomy and Physiology. CIBA USA; cranially, while the third main branch divides into the 5
1983 [section II, Plate 16]). major distal facial nerves branches (temporal, buccal, zygo-
matic, mandibular, and cervical) (Fig 7). Between 51 and 58
traction proteins troponin and tropomyosin are produced, days, the superficial muscles differentiate rapidly, and most
and aggregates of contractile subunit called sarcomeres are of them are composed of myoblasts. By 83 days, all of the
formed, each subunit composed of the thin myofibrils of muscles contain myotubules and are in their definitive po-
actin, tropomyosin, and troponin, and the thick myobrils of sitions, and, soon, the myotubules develop into young mus-
myosin (Fig 5B). cle fibers. Until term, the muscles increase in size and gain
The myofibrils extend the entire length of the muscle definitive attachments and the superficial muscles in the
fiber so that, when the myofibrils shorten, the entire muscle cervicomandibular and occipital regions differentiate ear-
unit contracts. The myofibrils are composed of smaller lier than those in the frontal and midfacial regions.1,19 The
structures called myofilaments, and, as the myotubes fill adult facial musculature is shown in Fig 8.
with myofibrils and myofilaments, their nuclei migrate to
the periphery. At this time, the myotube has differentiated THE MUSCLES INNERVATED BY THE TRIGEMINAL
into a muscle fiber cell or a myocyte. As the muscle fibers NERVE
develop, surrounding fibroblasts will form the perimysium The mesoderm of the mandibular process of the first bran-
and epimysium layers of the fibrous sheath of the muscle, chial arch gives rise to the muscles of mastication, which

Fig 4. Drawing, illustrating the newer concept of dividing the somite cells based on the signaling to which they respond; the primaxial cells will respond
to neural tube signaling, whereas the abaxial cells will respond to signaling from the lateral plate mesoderm and the ectoderm; the interface between
the primaxial and the lateral plate frontier (right side of diagram).
Fig 5. A, Drawing, showing the progression of development of skeletal muscle. B, Drawing, illustrating the contractile unit called the sarcomere and the
thick and thin myofilaments (modified from open access http://boundless.com/biology/textbooks). C, Drawing, illustrating the various segments and
their linings within a skeletal muscle (modified from open access http://www.easynotecards.com/print_cards/19549).
Fig 6. A, Sagittal drawing of an approximately 28 –30-day-old embryo, illustrating the initial mesenchymal concentration that will develop into the
facial muscle. B, Drawing, showing an approximately 47-day-old embryo and illustrating the emerging lamina that will populate the facial and calvarial
muscles (modified with permission from Ref 19 [Figs 1 and 6]).
include the temporalis, masseter, lateral pterygoid, and me- arch is filled with a uniformly closely packed mesenchyme,
dial pterygoid muscles, and, likely, the mylohyoid muscle most concentrated near the terminal end of the mandibular
(Fig 9). The tensor tympani and tensor veli palatini muscles nerve. This premuscle mass, with the mandibular nerve
also arise from this arch, as does the anterior belly of the within it, lies at approximately the middle of the arch. By 35
digastric muscle. In the 30-day-old embryo, the mandibular embryonic days, the premuscle mass has increased in size

Fig 7. Sagittal drawing of the facial nerve within the parotid gland, illustrating the major branches to the facial muscles (modified with permission from
Netter FH. Atlas of Human Anatomy. 5th ed. Philadelphia: Saunders Elsevier; 2011 [Plate 24]).
Fig 8. Frontal drawing of the face, showing the adult facial muscles partially cut away on the figure’s left side (modified with permission from Putz R,
Pabst R, eds. Sobotta Atlas of Human Anatomy. Vol 1. 13th ed. Philadelphia: Lippincott Williams and Wilkins; 2000 [Fig 139, p 75]).
but still has no differentiation into discrete muscles (Fig 10). ble, will develop into the mass for the medial and lateral
From its beginning, this premuscle mass is closely associ- pterygoid muscles.1
ated with the condensed mesenchyme that will be the man- By 40 embryonic days, the process of differentiation has
dible, and, with the differentiation of the proximal end of progressed, and the pterygoid muscles are now partially
the mandible, the premuscle mass partially divides into 3 separated by the Meckel cartilage. The coronoid process of
segments. The most cranial segment will become the tem- the mandible also partially separates the masseter from the
poralis muscle; the lateral segment will develop into the pterygoid masses. Meckel cartilage will eventually be in-
masseter muscle; and the inner segment, which is separated volved in the development of the sphenomandibular liga-
from the outer segment by the proximal end of the mandi- ment, the lingula of the mandible, and the upper portions of

Fig 9. A, Drawing, showing the skull and the muscles of mastication as seen from a left oblique view. B, A more sagittal view; the masseter muscle and
part of the mandibular ramus have been cut away to show the 2 heads of the lateral pterygoid muscle and part of the medial pterygoid muscle. C, The
drawing is viewed from behind and below the skull and shows the 2 heads of the lateral pterygoid muscle arising from the lateral aspect of the lateral
pterygoid plate and inserting onto the temporomandibular joint capsule and the condyle of the mandible. The medial pterygoid muscle arises from the
pterygoid fossa and inserts onto the inner aspect of the mandible near the lower ramus and angle (modified with permission from Netter FH. Atlas of
Human Anatomy. 5th ed. Philadelphia: Saunders Elsevier; 2011 [Plates 54 and 55]).
the incus and the malleus. Continued gradual differentia- and tensor veli palatini muscles can be recognized and are
tion of the muscles occurs with the maturation of the mem- connected with the pterygoid mass from which they likely
branous mandible and the adjacent skull to which the mus- arise. At this time, the tensor tympani muscle has already
cles of mastication will attached. At no time during gained its insertion to the malleus. The final development of
development are the muscles attached to Meckel cartilage, these muscles is related to the continued development of the
but they are always in relation with the membranous man- skull base, the Eustachian tube, and the soft palate.1
dible (Fig 11). By 45 embryonic days, the various muscles of
mastication are distinct but have only a slight resemblance THE TONGUE AND SUPRAHYOID MUSCLES
to their adult form. The temporalis muscle is quite small in In a previous review, the development of the palate, man-
proportion to the size of the head and only gradually does dible, and tongue were discussed, as was the separation of
the muscle extend over a much greater area of the calvaria. the tongue from the gums. The tongue has striated muscle
The masseter muscle is first attached only to the zygomatic derived from the occipital somites; cranial neural crest cell–
arch before it attaches to the mandible. derived mesenchyme; and a stratified squamous, nonkera-
The mylohyoid muscle apparently differentiates more rap- tinizing epithelium. The tongue will develop from contribu-
idly than the other muscles of mastication, and, by 35 embry- tions from the first 4 branchial arches in the ventral part of
onic days, it can be recognized by its nerve, the mylohyoid the foregut. All of the intrinsic tongue muscles are formed
branch from the inferior alveolar nerve of the mandibular from myoblasts that migrate from the occipital myotomes.
nerve. The mandibular nerve is the nerve of the muscles of The cranial nerve associated with the occipital somites (pos-
mastication. By 40 embryonic days, the mylohyoid muscle has totic myotomes) is the hypoglossal nerve (XII), which ac-
attained much of its adult morphology.1 As the mylohyoid companies the myoblasts as they travel a long-range migra-
muscles develop, they always leave the genial portion of the tion into the oral cavity and the tongue primordia. As a
symphysis menti free for the attachment of the genial mus- result of this migration, the tongue muscles are innervated
cles.20 Also, in the 40-day embryo, both the tensor tympani by the hypoglossal nerve. The cell migration pathway is first

Fig 10. Sagittal drawing of a 30 –35-day-old embryo, illustrating the development of the various premuscle masses. Note the continuous mesenchymal
muscle mass that extends from the tongue mass through the infrahyoid muscle mass (anterior neck muscles) and down to the diaphragmatic mass. The
nerves that will innervate these areas are also noted (modified with permission from Cochard LR. Netter’s Atlas of Human Embryology. Teterboro, NJ:
Icon Learning Systems; 2002 [Fig 9.10, p 225]).
Fig 11. A, Sagittal drawing of a 48 – 49-day-old embryo, illustrating the various premuscle masses and how they are developing compared with Fig 10
(modified with permission from Cochard LR. Netter’s Atlas of Human Embryology. Teterboro, NJ: Icon Learning Systems; 2002 [Fig 9.10, p 225]). B,
Drawing of a 50-day-old embryo and illustrating the continued progressive growth of the premuscle masses compared with Fig 11A. Note that the
sternocleidomastoid and trapezius masses are now clearly separated and the levator scapulae muscle mass is descending into the dorsal neck
(modified from Keibel F, Mall FP. Manual of Human Embryology. Philadelphia: JB Lippincott; 1912 [Fig 370]).
ventral and then cranial to eventually populate the already premuscle mass. Finally, the lingual mass merges dorsally with
formed endodermal portion of the tongue. the diaphragmatic premuscle mass. This lingual-infrahyoid-
In the 31-day-old embryo, the mesenchyme in the floor of diaphragmatic mesenchymal band is likely a primordial ven-
the mouth is similar to that present in the mandibular and tral visceral muscle complex and does not appear to be of
hyoid arches from which the musculature of those 2 arches direct myotomal origin (Figs 10 and 11). At this stage, the
will develop. In a 33-day-old embryo, the floor of the mouth individual muscle cannot be identified.
mesenchyme enlarges and thickens, and 2 bilateral masses de- In the 35-day-old embryo, each homogeneous lingual
velop within it. These bilateral premuscle masses extend from premuscle mass has split into 2 masses: a medial ventral
the region in which the mandible will later develop dorsally to mass for the geniohyoid and genioglossus muscles; and a
the hyoid region. From there, they become continuous with an dorsolateral mass for the hyoglossus, styloglossus, and
attenuated medial mesenchymal mass that is the infrahyoid chondroglossus muscles (the chondroglossus muscle is oc-

Fig 12. Drawing in the (A) sagittal view of the inner aspect of a 50 –54-day-old embryo, and (B) a 80 – 84-day-old embryo, illustrating the progressive
development of the membranous mandible about the Meckel cartilage (modified from model of jaw in Gray H. Anatomy of the Human Body. Philadel-
phia, Lea & Febiger; 1918).
casionally described as being part of the hyoglossus muscle, the hyoid anlagen. Over the dorsum and dorsolateral region
but it is separated from the hyoglossus muscle by the fibers of the tongue, the hyoglossus and styloglossus premuscles
of the genioglossus muscle). The chondroglossus muscle extend from the hyoid precartilage and styloid process, re-
arises from the medial side and the base of the lesser hyoid spectively, to the tip of the tongue. They lie dorsal and
cornu, and passes directly cranially to blend with the intrin- lateral to the radiating genioglossus muscle and their ori-
sic tongue muscle fibers that lie between the hyoglossus and gins. Although these genioglossus muscles are distinct, they
genioglossus muscles. The medial ventral mass extends are still close together and parallel. The hypoglossal nerve
from the region of the future symphysis menti to the prehy- now gives off branches to the geniohyoid muscle, then to
oid mass, and, dorsally and cranially, it expands into the the hyoglossus and styloglossus muscles, and then passes
tongue region. The mainstem of the hypoglossal nerve en- through the genioglossus muscle to its tip, and gives off
ters the mass dorsally and passes longitudinally nearly to its numerous lateral branches into this muscle. There is, at this
ventral margin. The dorsolateral mass extends from the stage, little interlacing of the tongue muscle, which is not yet
prehyoid region and the medial portion of the styloid pro- recognizable as the intrinsic muscles, which will eventually
cess into the dorsolateral region of the tongue for a short be the superior and inferior longitudinal, transverse, and
distance. A branch of the hypoglossal nerve enters its ven- vertical muscles (Figs 13 and 14 ).1
tral surface. The styloid process at this stage has an almost In the 45-day-old embryo, all the muscles are clearly
horizontal position and extends nearly to the midline. differentiated and increased in size. The origin of the
As the differentiation and development of these muscle styloglossus muscle has been carried to a more lateral
masses proceeds, the developing tongue gradually becomes position and enters the tongue at an angle to the hyoglos-
elevated above the mandibular arch. By 40 embryonic days, sus muscle. The hyoglossus muscle has spread out its
the differentiation of the mandibular arch has proceeded, attachment to the hyoid cartilage. The palatoglossus
and the Meckel cartilage is clearly present and partially muscle is now recognizable, extending from the laterally
enclosed by the membranous mandible (Fig 12). From the placed soft palate to the lateral surface of the tongue. The
membranous mandible, then arises the genioglossus mus- development of the intrinsic tongue muscles is most no-
cle, which extends in a fanlike manner into the tongue. Also ticeable at this stage, and the interlacing of the tongue
at this time, the geniohyoid muscle is present and extends to musculature is quite advanced. The palatoglossus mus-

Fig 13. A, Drawing from the front and below of the mandible, showing the mylohyoid muscles and their median raphe that extend from the mylohyoid
line on the lingual aspect of the mandible down to the hyoid bone. B, The anterior belly and portions of the posterior belly of the digastric muscles are
shown, with their tendon passing through the distal attachments of the stylohyoid muscles. C, The drawing is from above and behind the mandible and
illustrates the mylohyoid muscle’s attachment to the mylohyoid line of the mandible and the geniohyoid muscles, which lie immediately above the
mylohyoid muscles (modified with permission from Netter FH. Atlas of Human Anatomy. 5th ed. Philadelphia: Saunders Elsevier; 2011 [Plate 53]).
Fig 14. A, Sagittal drawing with the left mandible cutaway, showing the suprahyoid muscles extending from the mandible and hyoid bone to the styloid
process. B, The drawing is seen from lateral and below, and illustrates the more superficial suprahyoid muscles (modified with permission from Netter
FH. Atlas of Human Anatomy. 5th ed. Philadelphia: Saunders Elsevier; 2011 [Plates 53 and 59]). C, A coronal view of the tongue musculature illustrates
the 4 major intrinsic tongue muscles and the genioglossal muscle (modified with permission from Putz R, Pabst R, eds. Sobotta Atlas of Human
Anatomy. Vol 1. 13th ed. Philadelphia: Lippincott Williams and Wilkins; 2000 [Fig 198, p 109]).
cles arise from myoblasts associated with the branchial portion of the second branchial arch that gives rise to the
arches and, thus, are innervated by the branchial styloid process, the stylohyoid ligament, and the lesser
branches of the vagus nerve (X) (Fig 14).1 As the digastric cornu and upper body of the hyoid bone. Embryologic
muscle develops, it initially has only 1 belly, with a con- variations in the origin of this muscle include its attach-
striction of its continuous fibers where it passes the hyoid ment by fascicles that originate from the mandibular an-
primordium. The geniohyoid and genioglossus muscles gle, its attachment to the mandibular angle by a fibrous
basically have their adult morphology once they start to tract, or its attachment via muscle fascicles to a fibrous
develop, but they undergo proportional changes.20 tract that connects the Reichert cartilage to the mandible.
The muscle primordium then extends ventrally to insert
THE STYLOGLOSSUS MUSCLE on the posterolateral tongue primordium. In all cases, the
The styloglossus muscle originates in the Reichert carti- styloglossus muscle is innervated by the hypoglossal
lage, which is a cartilaginous formation in the cranial nerve (Fig 14).21

Fig 15. Drawing in the sagittal view of an approximately 38 – 42-day-old embryo, showing the various myotomes and noting that the occipital
myotomes do not extend as far ventral as the other myotomes (modified with permission from Cochard LR. Netter’s Atlas of Human Embryology.
Teterboro, NJ: Icon Learning Systems; 2002 [Fig 6.18, p 149]).
THE FLEXOR MUSCLES OF THE NECK

The occipital myotomes in the neck do not extend as far
ventrolaterally as do the cervical and thoracic myotomes
(Fig 15). The only complete ventral extension of the hyaxial
myotomes is between the transverse processes on the ven-
tral surface of the developing vertebral column in the cervi-
cal and upper thoracic regions. These premyoblast cells
form a flattened premuscle mass along the anterior surface
of these vertebrae, which will form the rudimentary flexor
prevertebral muscles, which are the longus capitis, longus
colli, and rectus capitis anterior muscles (Fig 16). As already
mentioned, the epaxial cells will give rise to the extensor
muscles of the vertebra.1,2,22
THE INFRAHYOID (STRAP) MUSCLES

By 45 embryonic days, densely packed mesenchyme forms a
paracervical premyoblast mass that likely arises directly Fig 16. Frontal drawing of the cervical and upper thoracic spine, illustrat-
from the mesoderm with, possibly, some contribution from ing the longus capitis, longus coli, and the rectus capitis anterior mus-
cles, all of which are derived from the hyaxial myotomes (modified with
the myotomes near the level of the third to fifth myotomes. permission from Netter FH. Atlas of Human Anatomy. 5th ed. Philadel-
This embryonic mesodermal premyoblastic mass will phia: Saunders Elsevier; 2011 [Plate 29]).
eventually form the infrahyoid muscles.1 By 47 embryonic
days, the neck muscles are beginning to differentiate and the The infrahyoid muscle mass soon consists of a distinct
infrahyoid premuscle mass is better developed than the band of premuscle tissue that extends on either side from
other premuscle masses that occur in the neck. Initially, as the base of the tongue dorsally and laterally toward the tips
noted above, at this early stage, the infrahyoid premuscle of the first ribs. The muscle masses are already supplied by
mass is continuous above with the tongue premuscle masses the ansa cervicalis from the ventral spinal nerve roots (Fig
and below with the diaphragmatic masses forming a lin- 17). Next, the muscle mass separates from the tongue mass,
gual-infrahyoid-diaphragmatic band (Figs 10 and 11). and, above the heart, the 2 lateral masses approach each

Fig 17. Drawing, in the sagittal view, of an approximately 38 –39-day-old embryo, illustrating the development of the ansa cervicalis, which will supply
the muscles of the anterior neck (modified with permission from Cochard LR. Netter’s Atlas of Human Embryology. Teterboro, NJ: Icon Learning
Systems; 2002 [Fig 5.5, p 118]).
other and nearly meet in the midline. However, as they muscle is innervated by branches of the lower 4 cervical
descend, they become widely separated by the heart and spinal nerves and the lateral muscular branches of the third
occupy a lateral position in the neck. As the heart starts and fourth cervical spinal nerves.24
to descend into the chest, the 2 muscle masses approach
each other from above downward, and, by 50 embryonic THE LEVATOR SCAPULAE AND SERRATUS ANTERIOR
days, the distinct identity of the infrahyoid muscles be- MUSCLES
gins (Fig 11B). Initially, the infrahyoid muscle primor- The levator scapulae muscle is a true posterolateral neck
dium divides into a superficial layer and a deep layer. The muscle that migrates dorsally to the scapula. It develops
deep layer will become the sternohyoid, thyrohyoid, and with the serratus anterior muscle, which migrates down
sternothyroid muscles, while the superficial layer will be- to the posterolateral thoracic region. These muscles arise
come the omohyoid muscle.23 By embryonic day 54, the from an additional premuscle mass that is in the region of
approximation of these masses is nearly complete, and the ventral ends of the lower cervical myotomes.1 By
there is a clearer distinction of the sternohyoid, sterno- approximately 47 embryonic days, the levator scapulae
thyroid, thyrohyoid, and omohyoid muscles, the latter and serratus anterior muscles are beginning to differen-
extending dorsally to reach the scapula (Fig 18). By 59 tiate from the mesenchyme, and soon this premuscle mass
fetal days, these muscles have almost attained their adult initially forms a column, without attachments to either
form and they lie parallel to each other near the midline the vertebrae or to the ribs over which it extends. This
and extend from the hyoid and thyroid cartilages to the column of cells extends from the cervical region to the
rudimentary halves of the sternum.1 thorax, and, by 50 embryonic days, the lower portions of
the levator scapulae and the serratus anterior muscles are
THE SCALENE MUSCLES well defined. By 54 embryonic days, the levator scapulae
The scalene muscles also arise from premuscle tissue, which has begun to resemble its adult form and its attachment
is most likely from the paraspinal mesenchyme, which is to the scapula is present (Fig 20).
ventral to the lower cervical myotomes and separate from
the infrahyoid muscles masses. By 50 embryonic days, the 3 THE TRAPEZIUS AND STERNOCLEIDOMASTOID
scalene muscles are fairly well differentiated, with their MUSCLES
adult attachments to the cervical vertebrae and the ribs (Fig The common rudiment of the trapezius and sternocleido-
19).1,2 The anterior scalene muscle is innervated by the mastoid muscles first appears in the 45-day embryo and
ventral branches of the fifth through eighth cervical spinal originates in the epipericardial ridge mesoderm of the bran-
nerves. The middle scalene muscle is innervated by branches chial arches. The embryonic ancestry of the trapezius and
of the third and fourth cervical spinal nerves and the lateral sternocleidomastoid muscles has been a topic of much dis-
muscular branches of these nerves. The posterior scalene cussion. Because they lie ventral to the 2 dorsal occipital

Fig 18. A, Frontal drawing of the neck, illustrating the infrahyoid (strap) muscles; the sternocleidomastoid muscles were removed to better see the
strap muscles; the most common position of the ansa cervicalis is shown in the left neck. B, The more superficial strap muscles have been partially cut
away to show the deeper strap muscles; more laterally, the scalene and trapezius muscles can also be seen. C, A lateral view of the neck with the
sternocleidomastoid and the inferior belly of the omohyoid muscle cutaway, illustrating the muscles of the floor of the posterior triangle of the neck,
the ansa cervicalis, and the accessary nerve with some of the connections to the cervical ventral spinal rami (modified with permission from Netter FH.
Atlas of Human Anatomy. 5th ed. Philadelphia: Saunders Elsevier; 2011 [Plates 28 and 31]).
myotomes and the 2 anterior cervical myotomes, they are in then gradually extends dorsally toward the arm bud pre-
closer proximity to the derivatives of the branchial arches muscle tissue, which, in the 47-day-old embryo, is near the
than to the myotomal muscles. Due to this proximity, the level of the fourth cervical myotome (Figs 10 and 11). It is
trapezius and sternocleidomastoid muscles are considered now when the dorsal end of this muscle mass starts to divide
by most researchers to be of branchial arch origin. How- into 2, one division for the trapezius muscle and one divi-
ever, other researchers see the muscles as myotomal deriv- sion for the sternocleidomastoid muscle. In the 49-day em-
atives based on their innervation arising from the separate bryo, the 2 dorsal aspects of these muscles are clearly sepa-
spinal root of the spinal accessory nerve (rather than from rated, with the trapezius muscle mass extending to near the
the cranial root that contains fibers bound for the pharynx level of the sixth cervical nerve but not yet attached to the
and the pharyngeal muscles) and the additional fact that no shoulder girdle and the sternocleidomastoid muscle mass
remnant of an aortic arch is in the area.25 extending toward the rudimentary clavicle, which is ante-
At this early stage, the common rudiment consists of rior to the first rib.
closely packed mesenchymal premyoblasts, which are indis- The trapezius soon becomes thicker and forms a colum-
tinguishable from the surrounding mesenchymal cells ex- nar mass, which extends from the occipital region dorsally
cept for their greater attenuation, and the presence of the toward the shoulder girdle and only slightly toward the
accessory nerve, which runs within the mass. Initially, spinous processes to which it is connected by a layer of
the ventral end of this muscle mass lies at the level where the fascia. By 55–56 embryonic days, the trapezius and sterno-
accessory nerve (XI) leaves the vagus nerve (X). The mass cleidomastoid muscles are fully separated and distinct

Fig 19. A, Frontal drawing of the cervical and upper thoracic spine, illustrating the scalene muscles and a more complete visualization of the rectus
capitis anterior muscle than shown in Fig 16; the scalene muscles have their origins from the anterior tubercles of the transverse processes of the
cervical spine (modified with permission from Netter FH. Atlas of Human Anatomy. 5th ed. Philadelphia: Saunders Elsevier; 2011 [Plate 29]). B, A lateral
view of the neck, with part of the sternocleidomastoid muscle cutaway, showing more clearly than in Fig 18C the muscles of the floor of the posterior
triangle of the neck (modified with permission from Putz R, Pabst R, eds. Sobotta Atlas of Human Anatomy, Volume 1, Head, Neck, Upper Limb. 13th ed.
Philadelphia: Lippincott Williams and Wilkins; 2000 [Fig 260, p 143]).
Fig 20. A, Drawing of the back, illustrating the origins and insertions of the normal levator scapulae muscles; the origins are from the posterior
tubercles of the first 4 cervical vertebrae, and the insertion is on the upper medial edge of the scapula. B, The drawing is from the back and illustrates
the levator scapulae (see also Fig 19B) and the trapezius muscles (modified with permission from Netter FH. Atlas of Human Anatomy. 5th ed.
Philadelphia: Saunders Elsevier; 2011 [Plate 168]).
throughout their lengths, while, at the same time, the devel- VARIATIONS IN THE MUSCLES OF THE NECK
oping arm bud and shoulder girdle have migrated dorsally. Although uncommon, a number of anatomic variations in
The trapezius is now attached to the spine of the scapula, the muscles of the neck have been reported, usually with an
the adjoining portion of the clavicle, and it has spread dor- overall individual incidence of ⬍3%.26 Most of these in-
sally to the sixth rib and dorsally toward the spinous pro- clude variations in the origins of the muscles, the insertions
cesses and the ligamentum nuchae. It is not until day 59 that or attachments of the muscles, and the presence of accessory
the trapezius acquires its adult extent. The splitting of the muscles. The following is a summary of some of these
trapezius into the divisions occasionally found in adults is a variations.
secondary process and not present in the early embryo.1
The sternocleidomastoid muscle also has progressively de- Muscles of Mastication
veloped by 54 embryonic days and extends from the mas- Variations that occur in the facial muscles were discussed in
toid process and occipital region dorsally to the clavicle. It a previous review in this series.6 Overall, variations in the
has started to split into 2, which will eventually attach to the muscles of mastication are uncommon. A pterygoideus pro-
manubrium and the sternum, and to the clavicle (Fig 21). prius muscle has been described as originating from the
The spinal accessory nerve innervates both the trapezius anterior infratemporal crest and extending vertically and
and sternocleidomastoid muscles. dorsally to insert onto the lateral pterygoid plate and then

sory mylohyoid muscle was noted to arise from the mylo-
hyoid line on the mandible and insert into the dorsal
portion of the mylohyoid raphe and the hyoid bone.32 The
mylohyoid muscle may be absent and replaced by the ante-
rior bellies of the digastric muscles or there may be no raphe
between the mylohyoid muscles, which results in a contin-
uous mylohyoid muscle (Fig 22B).33 Deficiencies in the my-
lohyoid muscle have also been reported with herniation of
any or all of the fat, the submental artery, or the sublingual
gland from the floor of the mouth. In 1 report, the sublin-
gual gland herniated in 72% of the cases of mylohyoid
muscles defects.34,35 The defect can also occur without any
herniation. The geniohyoid muscle can blend with the op-
posite muscle, and muscular slips to the genioglossus mus-
cle can occur.
Fig 21. Frontal drawing of the neck, illustrating the normal sternal and
clavicular heads of the sternocleidomastoid muscle (modified with per- Sternocleidomastoid
mission from Netter FH. Atlas of Human Anatomy. 5th ed. Philadelphia: Most of the reported neck muscle variations involve the
Saunders Elsevier; 2011 [Plate 27]).
sternocleidomastoid muscle. In addition to the 2 traditional
origins of the muscle to the sternum and clavicle (Fig 21), a
extending to the lateral pterygoid muscle. A temporalis mi-
third head was reported to originate from the investing
nor muscle has been described as having the origin of a
layer of fascia in the roof of the subclavian triangle near the
normal temporalis muscle but inserting into the mandibular
clavicle and that traversed obliquely upward and forward
notch rather than the coronoid process. An aberrant muscle
to fuse with the clavicular head.36 This may be the result of
band has been noted that extends from the anteromedial
splitting of the sixth arch mesoderm. In another report, a
margin of the temporalis muscle to interdigitate with the
third head of the sternocleidomastoid muscle was reported
lateral pterygoid muscle. A muscle has also been noted that
to arise from the capsule of the sternoclavicular joint and
originates from the anterior aspect of the infratemporal
crest of the greater sphenoid wing and extends toward the the superolateral border of the manubrium.37 There is a
inferior orbital fissure to insert onto the pyramidal process report of 4 heads of the sternocleidomastoid muscle, 2 each
of the palatine bone and then fuses with fibers of the dorsal on the clavicular and sternal origins. Each additional head
medial pterygoid muscle.27 joined the main muscle in the middle of the neck.38
There are numerous variations in the extent of the cla-
Suprahyoid Muscles vicular origin of the sternocleidomastoid muscle that have
In the literature, variations in the suprahyoid muscles are been noted. If the clavicular head is narrow, then the cla-
reported to occur, with incidences for the anterior bellies of vicular muscle origin is often narrow. Conversely, when the
the digastric muscles ranging from 5.9% to 53% of cases.28 clavicular head is broad, the sternocleidomastoid muscle
Accessory digastric and mylohyoid muscles have been de- origin may subdivide into several slips and comprise a
scribed. An accessory anterior digastric muscle belly was broad clavicular origin.26 As the trapezius and the sterno-
noted to arise from the intermediate digastric tendon on 1 cleidomastoid muscles arise from a common premuscle
side and insert into the opposite digastric fossa of the man- mass, it is not surprising that the adjoining margins of these
dible (Fig 22A). There can be 3 anterior bellies, 3 bellies muscles can be in contact or merged, or that there can be an
with a fourth fibrotic band, or 4 separate insertions of the absence of one or both muscles.26,39
anterior belly of the digastric muscle.28,29 Bilateral anterior
bellies of the digastric muscles have been observed with the Supernumerary Muscles
accessory muscles that originate in the digastric fossa but The supernumerary cleido-occipital muscle has been re-
that insert either into the hyoid bone or into the mylohyoid ported as a separate muscle from the sternocleidomastoid
raphe.30 Variations in the posterior digastric belly are less muscle. This cleido-occipital muscle may extend from the
common than in the anterior belly. There are reports of the mastoid tip to the medial clavicle or to the manubrium (Fig
posterior digastric belly arising from the styloid process or 23).39 Another muscle variation is the levator claviculae
being connected by slips to the middle or inferior constric- muscle, or the cleidocervical muscle, which arises from the
tor muscles. The digastric tendon may also pass ventral, or transverse processes of the upper cervical vertebra and in-
even rarely dorsal, to the stylohyoid muscle. serts onto the lateral aspect of the clavicle (Fig 24).26,40
An accessory mylohyoid muscle was described as arising There also has been a report of a rare hereditary disease
from the midline raphe of the mylohyoid muscle and then with shortening of the sternocleidomastoid and the exten-
having diverging muscle fibers that inserted into the inter- sor muscles of the neck, which limits neck motion.41 The
mediate tendon of the digastric muscle.31 Another acces- sternocleidomastoid muscle has been reported to be com-

Fig 22. A, Drawing, from below and in front of the mandible shows the normal mylohyoid and anterior bellies of the digastric muscles; however, there
is an accessory anterior belly of the digastric muscle (*), in this case, extending from the digastric fossa on the mandible to insert in the opposite
attachment of the normal anterior belly. B, There is an accessory mylohyoid muscle (*) extending from the mandible down to the opposite side of the
hyoid bone. These are just 2 examples of the variations noted in these muscles (modified with permission from Netter FH. Atlas of Human Anatomy. 5th
ed. Philadelphia: Saunders Elsevier; 2011 [Plate 53]).
Fig 23. Frontal drawing, illustrating the supernumerary cleido-occipital Fig 24. Frontal drawing, illustrating the levator claviculae or cleidocer-
muscle, extending from the mastoid tip to the medial clavicle. When vical muscle, which arises from the transverse processes of the upper
present, this muscle runs deep to the sternocleidomastoid muscle, and it cervical vertebra and inserts on the lateral aspect of the clavicle. This
may be uni- or bilateral (modified with permission from Netter FH. Atlas supernumerary muscle may be unilateral or bilateral (modified with per-
of Human Anatomy. 5th ed. Philadelphia: Saunders Elsevier; 2011 [Plate mission from Netter FH. Atlas of Human Anatomy. 5th ed. Philadelphia:
29]). Saunders Elsevier; 2011 [Plate 29]).
shoulder muscles and hand anomalies. New components of
posed of multiple layers, including the sternocleidooccipi- this syndrome have been reported and include the absence
tal, cleidomastoid, and sternomastoid muscles.36 There or hypoplasia of many muscles, including the trapezius.
may be no mastoid insertion but rather an insertion on the This may be secondary to the temporary interruption of the
occipital bone, the external auditory canal, or a lateral or subclavian artery and its branches in early embryogenesis
medial mastoid insertion. There may be no clavicular head because this artery supplies these muscles.43 Another ana-
or, as noted, a complete absence of the sternocleidomastoid tomic variation in the trapezius muscle has the lateral, up-
muscle.39 per three-fourths of the descending portion of the muscle
being separate from the remainder of the muscle and fusing
Trapezius with the main muscle above its midpoint attachment to the
With regard to the trapezius muscle, there has been a report clavicle or attaching to the clavicle by a separate tendon.44
of the familial absence of this muscle associated with the
absence of the pectoralis, supraspinatus, and serratus ante- Scalene Muscles
rior muscles.42 Poland syndrome is a rare unilateral congen- Scalene muscle variations include an accessory middle sca-
ital anomaly characterized by the absence of the pectoral lene muscle that can cause a thoracic outlet syndrome. This

Fig 25. A, Drawing in the left lateral view, illustrating the relationship of the 3 scalene muscles; an accessory scalene muscle is also shown extending
between the anterior and middle scalene muscles; as the brachial plexus and the subclavian artery pass between the anterior and posterior scalene
muscles, the accessory muscle can put pressure on these structures, causing symptoms (modified with permission from Putz R, Pabst R, eds. Sobotta
Atlas of Human Anatomy. Volume I, Head and Neck, Upper Limb. 13th ed. Philadelphia: Lippincott Williams and Wilkins; 2000 [Fig 260, p 143]). B, The
right brachial plexus passes normally between the anterior and the middle scalene muscles; on the left side, a variant is shown in which the brachial
plexus passes between the middle and posterior scalene muscles (modified with permission from Netter FH. Atlas of Human Anatomy. 5th ed.
Philadelphia: Saunders Elsevier; 2011 [Plate 29]).
muscle runs obliquely between the anterior and middle sca- superior angle of the scapula (Fig 26A).48 Another unusual
lene muscles and compresses the subclavian artery (Fig variation in this muscle had an accessory head that inserted
25).45 A scalenus minimus is a rare muscle that lies poste- via a flat aponeurotic band to the tendon of the rhomboi-
rior to the subclavian artery, underneath the inferior aspect deus major and the superior aspect of the serratus posterior
of the anterior scalene. There are 5 reported types of scalene superior muscle (Fig 26B).49
muscle variations that affect the upper brachial plexus
nerves: type I has a direct attachment of the anterior scalene The Infrahyoid Muscles
muscle to the perineurium of the upper plexus; type 2 is a Variations in the omohyoid muscle occur more com-
thin muscle bundle that connects the anterior and middle monly in the superior belly than in the inferior belly.
scalene muscles (this is a middle scalene muscle); type 3 is an
Duplications of the superior and inferior bellies have
abnormal development of the upper part of the anterior
been reported. In 1 case, the inferior belly of the omohy-
scalene muscle posterior to the C5 and C6 nerves, and thus
oid muscle inserted into the sternohyoid muscle.22 The
displaces the upper 2 nerves anteriorly; type 4 is a single
omohyoid muscle has been reported to have both bellies
mass of scalene muscle with the individual nerves that pen-
arise from the clavicle, with the absence of an intermedi-
etrate through the muscle mass, with separation of the mus-
ate tendon.50 The superior belly of the omohyoid may
cles occasionally occurring only at the point where the
be absent, with the inferior belly blending into the pos-
nerves traverse the muscles; and type 5 are strong fibrous
bands or ligaments that cross the cervical nerves vertically terior aspect of the sternocleidomastoid muscle. There
behind the anterior scalene muscle.46,47 The brachial plexus has also been a report of a cleidosternohyoid muscle and
can also pass between the middle and posterior scalene a cleidohyoid muscle. On 1 side, the cleidosternohyoid
muscles rather than between the anterior and middle sca- muscle originated from the clavicle and accompanied the
lene muscles (Fig 25B). The persistence of certain muscle inferior belly of the omohyoid muscle. On both sides, the
inclusions in the brachial plexus as well as groups of mus- muscle inserted into the sternohyoid muscle. It has been
cles that traverse elements of the plexus is related to the described as a distinct fifth infrahyoid muscle (Fig 27).51
original scalene premuscle mass being variously fragmented The cleidohyoid muscle runs from the hyoid bone to the
by the passage of the developing nerves as the limb bud clavicle and augments the sternocleidomastoid muscle.
develops.47 There have been reports of a cleidofascialis muscle,
which originates in the middle third of the clavicle and
Levator Scapulae Muscle inserts into the fascia colli, and a hyofascialis muscle,
The normal origin and attachments of the levator scapulae which originates from the hyoid and inserts into the
muscle are shown in Fig 20. There is a report of an anatomic omosternoclavicular fascia.23 There has been a report of
variation in the levator scapulae muscle, with the muscle an accessory belly of the sternothyroid muscle, with the
having a bifurcation at its midpoint that extends dorsally. muscle extending from the oblique line of the thyroid
Its medial band attached to the anterior aspect of the rhom- cartilage to the posterior surface of the sternothyroid
boideus major muscle while its lateral band was fixed to the muscle and the pretracheal layer of the cervical fascia.52

Fig 26. A, Drawing from a posterior view, illustrating an anatomic variation in the levator scapulae muscle, which has an accessory head that inserts into
a flat aponeurosis and then to the rhomboideus muscle (modified from Ref 49 [Fig 2]). B, Drawing of the upper back, showing, on the left side of the
drawing, the normal anatomy of the levator scapulae and rhomboid major muscles; on the right side of the drawing, the levator scapulae muscle has
a bifurcation at its midpoint, with the medial portion of the muscle attaching to the rhomboideus major, while the lateral portion attaches normally to
the scapula (modified with permission from Netter FH. Atlas of Human Anatomy. 5th ed. Philadelphia: Saunders Elsevier; 2011 [Plate 411]).
Fig 27. Frontal drawing, illustrating the cleidosternohyoid muscles on either side and the cleidohyoid muscle on the right side; these have been
considered as a fifth infrahyoid muscle (modified from Ref 51 [Fig 1]).
INNERVATION OF THE NECK MUSCLES dibular nerve (V3). The geniohyoid muscle is supplied by
This section reviews the nerves that innervate the the first cervical ventral ramus through the hypoglossal
above discussed muscles and the embryology of these nerve. The genioglossus, hyoglossus, styloglossus, palato-
nerves. glossus, and chondroglossus muscles are all innervated by
the hypoglossal nerve. The anterior belly of the digastric
The Suprahyoid Muscles muscle is supplied by the mylohyoid branch of the inferior
The mylohyoid muscle is innervated by the mylohyoid alveolar nerve, and the posterior digastric belly is supplied
branch of the inferior alveolar nerve, a branch of the man- by the facial nerve (Fig 28).53

Fig 28. Sagittal drawing as Fig 14A, illustrating the suprahyoid muscles that extend from the mandible and hyoid bone to the styloid process; the
muscles have been color coded to help identify the innervating nerves (modified with permission from Netter FH. Atlas of Human Anatomy. 5th ed.
Philadelphia: Saunders Elsevier; 2011 [Plates 53 and 59]).
The Hypoglossal Nerve nerve sheaths begins, adjoining axons can become more or
The hypoglossal nerve first appears at the end of the third less bound together and as the individual tongue and hyoid
embryonic week, at which time it consists of loose axonal muscles draw apart these nerves, they are led out into an
strands, which can be traced between the occipital myo- open plexus. The exact formation of these anastomoses
tomes that spring from the ground (basal) plate of the neu- must depend on the position of the axons at the time that
ral tube and extend a short distance into the mesenchyme. the sheaths are formed, which introduces a variability and
These rootlets are formed in 3 or 4 segmental groups and which might account for the different arrangements found.
develop in the same line with the ventral roots of the cervi- A further source of variation is presented by slight differ-
cal nerves. During the fourth embryonic week, they grow ences in the division line between the rootlets of the hypo-
forward and fuse in a common trunk. At the end of the first glossal nerve and the first cervical nerve.54
embryonic month, this trunk has passed around the nodose Thus, the descendens hypoglossi seems to develop simul-
ganglion and curves medially around the cervical sinus of taneously with the appearance of the anastomoses between
His to reach the anlage of the floor of the mouth (Fig 29A). that hypoglossal nerve and the upper cervical nerves. In the
At the point where the hypoglossal nerve bends upward early stages, the rootlets of the hypoglossal nerve have a
toward the anlage of the tongue, it is joined by fibers from close similarity to the ventral roots of the spinal nerves and
the first and second cervical nerves. A week later, its prin- are now generally considered as a cranial continuation of
cipal branches of distribution are present.54 them. That is, the hypoglossal nerve is derived from the
In the 34-day-old embryo, as the hypoglossal nerve fusion of 3 or 4 segmental spinal nerves, which, in the
crosses the nodose ganglion, it gives off the ramus descen- course of phylogenesis, have become enclosed in the cra-
dens hypoglossi. At the time, the descendens hypoglossi has nium. In a hypothetical ancestor, the segments of the nerve
the opportunity to communicate between the hypoglossal belonged to the spine and possess, in addition to ventral
nerve and the upper 2 or 3 cervical nerves that already exists roots, both dorsal roots and ganglia, the latter becoming
(Fig 29B). That is, the terminal axons of the cervical nerves subsequently reduced coincidently with the invasion of the
end in brushlike tufts in close contact with the hypoglossal vagal group into this region. Strong support for this view
nerve. The amount of interchange of axons cannot be accu- exists in the hoofed animals, in which there are persistent
rately predicted, but it is evident that the character of the dorsal roots and ganglia that belong to 1 or 2 of the more
descendens hypoglossi is dependent on the nature of the dorsal divisions of the hypoglossal nerve. Variations in the
contribution of axons from the cervical nerves. The course hypoglossal nerve are uncommon. However, there is a re-
in the development of this cervical anastomosis is as fol- port of an anastomosis between the 2 nerves situated just
lows: the axons of the hypoglossal and the upper cervical ventral to the hyoid bone region.53
nerves start out perpendicular to the neural tube. Due to the
curve of the neural tube, these axons come together like The Trigeminal Nerve
spokes in a wheel and then grow forward adjacent to each The largest sensory ganglion in the embryo is the trigeminal
other into the premuscle tissues. When the formation of the (Gasserian or semilunar) ganglion. It is connected both to

Fig 29. A, Sagittal drawing of a 4.5-week embryo, illustrating the common neural (ganglionic) crest for the vagus and accessory nerves, and their
shared multiple ganglia; also note the extension of the neural crest dorsally into the spinal ganglia that are connected by interganglionic neural crest
bridges; the hypoglossal nerve extends dorsally and ventrally around the nodose ganglion of the vagus nerve to then proceed to the developing
tongue musculature. B, In a 5-week embryo, the interganglionic bridges have disappeared in the spinal nerves as the dorsal cranial nerves mature; the
ansa cervicalis is also developing; initially, C1 joins the XII to form the superior root of the ansa cervicalis; C2 and C3 then join together to form the
inferior root and eventually the 2 roots connect to complete the ansa cervicalis (modified from Ref 54 [Fig 7 and pp 96 and 97]). C, A drawing based
on avian research, illustrating the various origins of the cell populations in the trigeminal ganglion (modified with permission from D’Amico-Martel A,
Noden DM. Contributions of placodal and neural crest cells to avian cranial peripheral ganglia. Am J Anat 1983;166:445– 68 [Fig 15, p 462]). D, Drawing
of the most common morphology of cervical plexus and its connections with the accessory, vagal, and hypoglossal nerves (modified with permission
from Netter FH. Atlas of Human Anatomy. 5th ed. Philadelphia: Saunders Elsevier; 2011 [Plate 32]).
the brain and the 3 peripheral trunks that extend into the comes fused into a single ganglionic mass. If this latter view
ophthalmic, maxillary, and mandibular regions. Centrally, is correct, then the semilunar ganglion arises partly from the
the ganglion is connected to the brain by a thick, short trunk neural crest and partly from the epibranchial ectoderm, and
that enters the brain opposite the first and second rhombic this composite origin morphologically corresponds with
grooves, which are transient furrows in the floor of the that found in lower vertebrates.1,55
future fourth ventricle. Fibers immediately form a flattened The trigeminal nerve is derived from the trigeminal pla-
longitudinal tract, part of which extends dorsally as the code, one of the epibranchial placodes. Initially, the trigem-
spinal tract and part of which extends ventrally and crani- inal ganglion may be bilobed, with an ophthalmic portion
ally to enter the cerebellar ridge. Although it generally is and a maxillary-mandibular portion. Soon, the latter por-
believed that the origin of the Gasserian ganglion is from a tion separates into distinct maxillary and mandibular divi-
single cellular neural crest mass, it has also been proposed sions. Based on avian and mouse research, it seems that the
that the Gasserian ganglion is formed from 3 separate pro- trigeminal ganglion is populated by placodal neurons, neu-
ganglia that are interconnected by a cellular lamina with 3 ral crest neurons, and neural crest satellite cells and
epibranchial proganglia, the entire group eventually be- Schwann cells (Fig 29C). The motor root of the trigeminal

Fig 30. On the top are illustrations of the development of the spinal cord from 5 embryonic weeks to 3 fetal months; the cross-sections extend from the
neural tube peripherally; note the progressive development of the intermediate (mantle) and marginal zones, which contain the gray and white
matter, respectively (modified with permission from Netter FH. Atlas of Human Anatomy. 5th ed. Philadelphia: Saunders Elsevier; 2011 [Plate 3.8]). The
bottom drawing that shows a cross-section of the spinal cord, illustrating the roof and floor plates, the sulcus limitans, which separates the alar
(sensory) and the basal (motor) portions of the cord (modified with permission from Carlson BM. Human Embryology and Developmental Biology. 4th
ed. Philadelphia: Mosby Elsevier; 2009 [Fig 11– 6, p 244]).
nerve develops from neuroblasts of the ventral zone of His, Overview of the Development of Spinal Nerves
in the hindbrain at the level of the Varolian bend. These By the end of the fourth embryonic week, the single-layered
neuroblasts gather together and form the trigeminal nucleus neuroepithelium of the neural tube has started to proliferate
that lies near the junction of the ventral and dorsal columns and produce the cells of the central nervous system. As the
and the ascending root of the trigeminal nerve. The fibers from wall becomes multilayered, the cells lose their clear outlines
these neuroblasts gather into a single stem that exits the brain and form a compact cellular syncytium, which is bound on
near the dorsolateral edge of the ventral zone.1,2 its outer and inner surfaces by an external and an internal
In the 34-day-old embryo, the ophthalmic division has limiting membrane, respectively. In the 34-day-old embryo,
extended ventrally and divided into the frontal and nasocil- the inner layer, or ependymal zone, has cells that abut the
iary nerves, the latter branch being located just dorsal to the internal limiting membrane, while their processes extend
optic stalk. The maxillary and mandibular divisions have outward. As these cells migrate peripherally, they establish
extended dorsally and then divided into their terminal the intermediate zone (previously called the mantle zone),
branches among the cells of the maxillary and the mandib- the precursor of the gray matter, and where most of the
ular processes, respectively. By the beginning of the sixth mature neurons will be situated. Fibers from the intermedi-
embryonic week, the chief branches of these divisions are ate layer then extend farther peripherally to establish the
clearly evident.1 marginal zone, initially a noncellular layer that is the pre-
cursor of the white matter (Fig 30).1,2
The Cervical Flexor Muscles The intermediate zone of the spinal cord and brain stem
The flexor muscle of the cervical and upper thoracic spine is then organized into a ventral pair of basal plates (motor)
are innervated by the ventral rami of the cervical spinal and, slightly later, a dorsal pair of alar plates (sensory).
nerves. Specifically, the longus capitis muscle is innervated Along the lateral border of the neural canal, the 2 plates are
by the ventral cervical spinal rami of C1–C3. The longus separated by the sulcus limitans, whereas dorsally and ven-
coli muscle is innervated by the ventral cervical spinal rami trally, the basal and alar plates are connected by the non-
of C2–C6. The rectus capitis anterior is supplied by a loop neurogenic roof plate and floor plate, respectively.10,11 In
between C1 and C2 (Figs 16 and 29D). the 27-day-old embryo, the ventral roots are already devel-

Fig 31. Serial drawings of spinal cord as seen from above and the left side. A, The ventral rootlet has already developed from the basal plate regions;
the dorsal spinal ganglia are interconnected by interganglionic neural crest bridges but not yet connected to the spinal cord. B, Fibers extended from
the dorsal root ganglia to the spinal cord alar plate regions via the dorsal roots, and the interganglionic bridges have disappeared. C, Fibers from the
dorsal root ganglia and dorsal roots extend to the ventral rootlets finally forming a complete spinal nerve (modified with permission from Ref 11 [Figs
10 – 4 and 10 –5, pp 304 and 305]).
oping as outgrowths of neuroblasts from the basal plates of the neural spinal crest is produced. The ventral ends of the
the spinal cord. After separating themselves from the syn- spinal ganglia then extend ventrally and join the already
cytium of the basal plate, these outgrowing fibers are assem- formed ventral roots (Fig 31).2,10,11
bled into rootlets. These primary processes may then course In the 34-day-old embryo, there is a clearly defined dor-
in the marginal zone of the neural tube or they may con- sal root, its sharply outlined ganglion, and a well-defined
verge and penetrate the marginal zone ventrally and later- ventral root that all join together to form the nerve trunk.
ally to form the ventral roots of the spinal nerves. Once At the same time, the dorsal and ventral roots give off
outside the spine, these rootlets are grouped into segmental lateral fibers that form the dorsal branches that will inner-
bundles, and, after being joined by the axons of the futures vate the cells that are forming the long muscles of the back
dorsal roots, they will constitute complete segmental spinal and the overlying skin. At the locations where the ventral
nerves (Fig 31). spinal rami leave the spinal cord, connecting neural loops
At the end of the fourth embryonic week, fibers start to may extend from one spinal nerve to another. In the cervical
extend back from the dorsal border of the neural crest spi- region, this leads to the development of superficial and deep
nal ganglia to attach to the spinal cord. These are the prim- nerve plexuses. The superficial plexus is formed by the
itive dorsal spinal roots, and they first appear in the cervical union of the lateral terminal branches, which form into
region and develop dorsally. They enter the marginal zone loops that supply cutaneous branches to the auricular, cer-
of the neural tube and eventually form a longitudinal bun- vical, and occipital regions. The deep cervical plexus will
dle that corresponds to the dorsolateral funiculus of the form the ansa cervicalis and the phrenic nerve.1,2 By 37
adult spinal cord. With the formation of these rootlets, days, the ventral cervical roots are well established. In most
there is a gradual disappearance of the dorsal bridges be- specimens, the ventral rami of the second and third cervical
tween the spinal ganglia, and a complete segmentation of nerves unite to form the inferior root of the ansa cervicalis.

Fig 32. A, Drawing of the most common morphology (approximately 80% of the time) of the ansa cervicalis (modified from https://en.wikipedia.org/
wiki/Ansa_cervicalis). B–F, Illustrate some of the different variations in the morphology of the ansa cervicalis. B, The vagus nerve contributes to the
superior root, and the C1–C3 rami are not interconnected. C, There is no contribution from the C3 ramus, and the C2 ramus inserts in the vagus nerve,
which then contributes to the innervation of the appropriate infrahyoid muscles. D, The rami of C1–C3 are not interconnected, the ramus of C3 forms
the inferior root, while the C2 ramus inserts into the vagus nerve, which then forms the superior root of the ansa cervicalis. E, the C1–C3 rami are not
interconnected, the C1 ramus runs with the vagus nerve rather than the hypoglossal nerve, and the rami of C2 and C3 independently contribute to the
inferior root. F, The ramus of C1 runs with the vagus nerve rather than the hypoglossal nerve; compare the various versions to Fig 29D (modified from
https://en.wikipedia.org/wiki/Ansa_cervicalis).
The superior root most often is formed by the first ventral either a uni- or bilateral absence of the ansa cervicalis,
rami uniting with the hypoglossal nerve. The 2 loops then which, in these cases, is replaced by a vagocervical complex
unite to form the complete ansa cervicalis.56 that is formed by the vagus nerve and the C1 and C2 ventral
rami from the cervical plexus. The vagus nerve can fuse
The Infrahyoid Muscles with the hypoglossal nerve immediately after exiting the
The infrahyoid muscles are innervated by the branches skull base and then either the vagus nerve can supply
of the ansa cervicalis, which is a neural loop normally the infrahyoid muscles and contribute to the formation of
formed by the ventral rami of the upper 3 cervical nerves. the superior root of the ansa cervicalis or the infrahyoid
The ansa cervicalis has a superior and an inferior root. The muscles can be innervated by C2 and C3 branches rather
superior root is formed by the ventral ramus of the first than the vagus nerve. In these cases, the C1 ramus joins
cervical (C1) spinal nerve. These fibers join the hypoglossal the hypoglossal nerve as usual. Another variant has the
nerve, and, after a short distance, some of these fibers de- superior root of the ansa cervicalis being formed by the
scend to form the superior root of the ansa cervicalis (de- C1 ventral ramus, which accompanies the vagus nerve
scendens hypoglossi). The remaining C1 fibers supply the instead of the hypoglossal nerve, and the C1 ramus joins
thyrohyoid and geniohyoid muscles, and a branch is given the inferior root from C2 and C3. There can be an absent
off from the superior root to supply the superior belly of the inferior root with C2 and C3 separately joining the su-
omohyoid muscle. The superior root then joins the inferior perior root. Some of these variations are shown in Fig
root (descendens cervicalis), which is formed by the ventral 32.57,60
rami of the second (C2) and third cervical (C3) spinal The inferior root is derived from the ventral rami of C2
nerves. Three branches then arise from the inferior loop of and C3 in 38% of cases, from C2, C3, and C4 in 10% of the
the ansa cervicalis to supply the remaining infrahyoid mus- cases, and from C2 alone in 12% of the cases. The inferior
cles (Fig 29). This configuration occurs in approximately root passes posterolaterally to the internal jugular vein in
80% of the cases.57-59 74% of people and anteromedially in 26% of the cases.60 In
There have been many anatomic variations reported for 81% of people, the ansa cervicalis is lateral to the carotid
the ansa cervicalis. To mention only a few, there can be sheath, and, in 15% of people, it passes between the inter-

Fig 33. A, Serial drawings, illustrating the progressive development of the neural tube from the neural plate and the separation of the neural crest cells
from the overlying ectoderm and the underlying neural tube; the neural crest cells then further divide into left- and right-sided masses that then extend
from cranial to dorsal into the spine. Compare with Fig 29A (modified with permission from Cochard LR. Netter’s Atlas of Human Embryology.
Teterboro, NJ: Icon Learning Systems; 2002 [Fig 3.2, p 53; Fig 3.8, p 59]). B, At 4 weeks, the separate origins of the superior and inferior ganglia of
cranial nerves IX and X are illustrated. C, Drawing of a 4.5-week embryo, showing the relationship of the accessory nerve (green) to the vagus nerve
(X) and their common neural (ganglionic) crest; also note the separate neural crest of the glossopharyngeal (IX) nerve. The superior and inferior
ganglia of IX and X are now clearly joined (B and C, modified with permission from Ref 65 [Fig 6, p 378]). D, A sagittal drawing, illustrating the ganglion
cell masses in the occipital region; note that the accessory nerve passes through the C1 ganglion (the most common variant); also note the numerous
ganglia of the cranial nerves X and XI (modified from Ref 54 [Fig 13, p 101]).
nal jugular vein and the common carotid artery. Rarely, the The Scalene Muscles
ansa passes behind and medial to the great vessels.61 The The scalene muscles are all innervated by ventral rami of the
precise knowledge of the position and anatomic relation- cervical spinal nerves, including the ansa cervicalis. Specif-
ships of the ansa cervicalis has become of great importance ically, the anterior scalene muscle is supplied by the ventral
to surgeons to avoid damage to these nerves. Recently, rami of C4 –C6. The middle scalene muscle is supplied by
transplantation of ansa cervicalis to the paraglottic space the ventral rami of C3–C8, and the posterior scalene muscle
has become the preferred procedure to reanimate the larynx is supplied by the ventral rami of C6 –C8.
after a recurrent nerve paralysis.62 With regard to the posi-
tion of the ansa cervicalis in the neck, although there, again, The Levator Scapulae and Anterior Serratus Muscles
is variation in the literature, 1 report noted that the ansa The levator scapulae muscle is innervated by the ventral
cervicalis loop was at the level of the hyoid bone in 5% of rami of the cervical spinal nerves C3–C4, and C5 via the
the cases, it was between the hyoid and the omohyoid mus- dorsal scapulae nerve (Fig 29). The serratus anterior muscle
cle in 87.5% of the cases, and it was below the omohyoid is innervated by the ventral rami of C5–C7 through the long
muscle in 7.5% of the cases (Fig 18).60 thoracic nerve.

Fig 34. Frontal drawing, illustrating the accessory nerve and its connections to the ventral cervical spinal rami and its innervation of the sternocleido-
mastoid muscle and the trapezius muscle; also see Fig 18C (modified with permission from Netter FH. Atlas of Human Anatomy. 5th ed. Philadelphia:
Saunders Elsevier; 2011 [Plate 126]).
Sternocleidomastoid and Trapezius Muscles

As noted previously, the spinal accessory nerve has histori-
cally been described as having a “cranial” root and “spinal”
root. The spinal root is often referred to as the “accessory”
nerve, and this component originates solely from the spinal
cord, with no connection to the so-called cranial root. The
definitional problems arise from our lack of knowledge re-
garding the derivatives and components of the dorsal bran-
chial arches, that is, what constitutes the fifth and sixth
branchial arches. This, in turn, precludes our full under-
standing of the embryology of the innervation of muscles
such as the trapezius and sternocleidomastoid. Indeed, a
redefinition of cranial nerve XI, by acknowledging these
inherent problems, seems warranted.63,64
In the early embryo, the accessory and vagus nerves do
not exist as 2 independent cranial nerves but rather as parts Fig 35. Diagrams of the primary anastomoses between the accessory
of a single structure, each possessing mixed motor and sen- nerve and the cervical plexus. A, In most cases, there is a nearly straight
neural connection between the nerves, B, Whereas, in only approxi-
sory root ganglia derived from the same neural (ganglion)
mately 8% of the cases, there is a true ansiform configuration (modified
crest (Fig 33). Unlike the combined neural crest of the 10th with permission from Caliot P, Bousquet V, Midy D, et al. A contribution
and 11th nerves, the neural crest of the glossopharyngeal to the study of the accessory nerve: surgical implications. Surg Radiol
nerve (IX) is separate, and, apparently, it is not influenced Anat 1989;11:11–15 [Fig 1, p 13]).

Table: Summary of the embryonic origins of the muscles and their innervations
Muscles Embryonic Origin Muscles Innervating Nerves
Muscles of somitic origin Extensor and flexor spinal muscles (epaxial or primaxial cells) Ventral and dorsal cervical rami
Extraocular muscles (preotic somites) III, IV, VI
Intrinsic tongue muscles (occipital somites) XII
Muscles of branchial origin Facial muscles, including the stylohyoid, posterior belly of the VII
digastric, and platysma muscles
Muscles of mastication, including the anterior belly of the V
digastric and the mylohyoid muscles
Stylopharyngeus muscle IX
Musculus uvulae, palatoglossus, palatopharyngeus, X
salpingopharyngeus, and the intrinsic and extrinsic laryngeal
muscles
Trapezius and sternocleidomastoid muscles (epipericardial ridge XI
(4th–6th arch)
Muscles of de novo Suprahyoid muscles, infrahyoid muscles (strap muscles), scalene Ansa cervicalis and ventral
mesenchymal origin muscles, levator scapulae, and serratus anterior muscles cervical rami
by the adjacent combined neural crest of the vagus and some rootlets emerging at their level of origin, whereas
accessory nerves. It should be noted that the upper ganglia others ascend for a variable distance within the spinal cord
(the superior ganglion of the glossopharyngeal nerve [Eh- before turning out of the cord. Usually the trunk of the
renritter ganglion] and the jugular ganglion of the vagus accessory nerve passes through the ganglion of C1 (Fig
nerve [X]) are derived from the neural crest. However, their 33C). In addition, the presence of ganglia along the trunk of
lower ganglia (the petrosal ganglion of cranial nerve IX and the accessory nerve indicates the presence of a sensory com-
the nodose ganglion of cranial nerve X) are derived from the ponent.65 Both the nucleus ambiguus and the accessory nu-
ectodermal epibranchial placodes and then eventually join cleus are derived from the same embryonic cell column. The
the upper ganglia (Fig 33).54 These different embryologic spinal rootlets form a trunk, which then ascends and enters
origins may explain the different nerve innervations of these the skull via the foramen magnum behind the vertebral
ganglia. The superior ganglion of cranial nerve IX is small artery. The nerve then turns upward and passes into the
and has no branches. It is the petrosal ganglion of cranial jugular foramen and exits the skull base, running down-
nerve IX that has the branches that supply the tympanic ward in a posterior lateral direction where it passes either
plexus, sensory fibers to the base of the tongue and phar- ventral (39.8%) or dorsal (57.4%) to the internal jugular
ynx, and secretomotor fibers to the parotid gland. The jug- vein or, rarely, through this vein. It runs to the deep surface
ular ganglion of cranial nerve X primarily supplies the pha- of the sternocleidomastoid muscle, where it forms an anas-
ryngeal plexus, whereas the nodose ganglion primarily tomosis with variable fibers of the C2–C4 ventral roots.
supplies the laryngeal nerves. In approximately 8% of cases, there is no contribution
The cranial component, or root, of cranial nerve XI orig- from the cervical branches and the muscle is solely inner-
inates in the ambiguus nucleus and exits the skull base via vated by the accessory nerve. When there are contributions
the jugular fossa where it unites for a short distance with the from the cervical nerves, the nerves involved are the follow-
spinal root (Fig. 34). It also connects to the jugular ganglion ing: C2–C3 (38.1%– 44%), C3 (8.8%–28%), C2 (44%–
of the vagus nerve. The cranial root then separates from the 53.1%), and, rarely, C3–C4 (3%), which form the ansa of
spinal root and immediately joins the vagus nerve superior Maubrac, although a true ansiform configuration is only
to the nodose ganglion of the vagus nerve. Some fibers will present in 8% of cases (Figs 29 and 35).66,67 On the deep
supply the pharyngeal and palatal muscles except the tensor surface of the sternocleidomastoid muscle, the nerve sends
veli palatini, which is supplied by the mandibular nerve of branches without penetrating the muscle in 45.9% of cases.
the trigeminal nerve, while other fibers will enter the recur- In 54.1% of cases, the nerve sends branches and penetrates
rent laryngeal nerve to supply the intrinsic laryngeal the muscle.66 The point of emergence from the posterior
muscles. border of the sternocleidomastoid muscle is variable, and
The spinal root arises from an elongated nucleus of mo- the nerve then crosses the posterior triangle inferiorly and
tor cells in the lateral aspect of the ventral horn, which laterally on the surface of the levator scapulae muscle, sep-
extends from the junction of the spinal cord and the me- arated from this muscle by the prevertebral layer of the deep
dulla to the fifth or sixth cervical segment, often referred to cervical fascia and adipose tissue.
as the spinal accessory nucleus (Figs 33 and 34). The in- In general, the nerve passes under the anterior border of
traspinal course of the accessory nerve is variable, with the trapezius muscle at a point at the junction of the supe-

rior two-thirds and inferior one-third of the border of the 12. Dubrulle J, Pourquie O. Welcome to syndetome: a new
trapezius. Approximately 3–5 cm above the clavicle, it then somitic compartment. Dev Cell 2003;4:611–12. 10.1016/
passes behind the anterior border of the trapezius muscle S1534-5807(03)00133-3
and may form a plexus on its deep surface, with contribu- 13. Burke AC, Nowicki JL. A new view of patterning domains in
tions from C3 and C4, or C4 alone. No plexus is present in the vertebrate mesoderm. Dev Cell 2003;4:159 – 65. 10.1016/
S1534-5807(03)00033-9
25% of the cases. The nerve finally enters the deep surface
14. Nowicki JL, Takimoto R, Burke AC. The lateral somitic
of the trapezius muscle. The function of these cervical
frontier: dorso-ventral aspects of anterio-posterior regional-
nerves is believed by some researchers to supply proprio- ization in avian embryos. Mech Dev 2003;120:227– 40.
ceptive fibers to the sternocleidomastoid muscle, while the 10.1016/S0925-4773(02)00415-X
accessory nerve supplies motor innervation. Other re- 15. Shearman RM, Burke AC. The lateral somitic frontier in on-
searchers believe that the cervical plexus may have some togeny and phylogeny. J Exp Zool B Mol Dev Evol 2009;312:
motor fibers and that they may supply both motor and 603–12
proprioceptive fibers to the lower two-thirds of the sterno- 16. Bailey P, Holowacz T, Lassar AB. The origin of skeletal mus-
cleidomastoid muscle in 75% of the cases.68 The embryonic cle stem cells in the embryo and the adult. Curr Opin Cell Biol
origins of the muscles discussed in this review and the 2001;13:679 – 89. 10.1016/S0955-0674(00)00271-4
nerves that innervate these muscles are summarized in the 17. Carlson B. Integumentary, skeletal and muscular systems. In:
Table. Carlson B, ed. Human Embryology and Developmental Biol-
ogy. 4th ed. China: Mosby Elsevier; 2009:200 – 07
18. Som PM, Curtin HD, Liu K, et al. Current embryology of the
CONCLUSION
temporal bone. Neurographics 2015. In press.
This review addressed the embryology of the muscles of the 19. Gasser RF. The development of the facial nerve in man.
face and neck, their innervations, and their variation. The Ann Otol Rhinol Laryngol 1967;76:37–56. 10.1177/
embryology of the nerves that supply these muscles was also 000348946707600103
addressed, as were variations in these nerves. The reader is 20. Radlanski RJ, Renz H, Tabatabai A. Prenatal development of
encouraged to use this review when needed to better under- the muscles in the floor of the mouth in human embryos and
stand how these nerves and muscles arise and what varia- fetuses from 6.9 to 76 mm CRL. Ann Anat 2001;183:511–18.
tions may exist. 10.1016/S0940-9602(01)80057-1
21. Mrida-Velasco JR, Rodriguez-Vazquez JF, de la Cuadra
Blanco C, et al. Origin of the styloglossus muscle in the human
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Embriology, Variations and Innervations of The Human Neck Muscles

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Embriology, Variations and Innervations of The Human Neck Muscles

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HEAD & NECK CME

Embryology, Variations, and in the cytoplasm, where it serves

Neck Muscles NOTCH ⫽ Notch signaling pathway

Received December 10, 2015;

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THE FACIAL MUSCLES

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THE FLEXOR MUSCLES OF THE NECK

THE INFRAHYOID (STRAP) MUSCLES

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Sternocleidomastoid and Trapezius Muscles

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