BASIC SCIENCE
Physiology of swallowing
Jenny Walton
Priyamal Silva
Abstract
The mechanism behind normal swallowing is complex and multifacto-
rial. Due to the close proximity of the pathways of swallowing and respi-
ration, precise coordination between these functions is vital in order to
avoid entry of material into the airway and to ensure optimal health and
nutrition in general. Swallowing can be divided into three stages: oral,
pharyngeal and oesophageal and although initiation of the swallow is
often under voluntary control, swallowing is also triggered frequently
throughout the day as a reflex action due to the presence of saliva in
the oropharynx. Dysphagia is a symptom frequently encountered by cli-
nicians and its causes are vast and varied. A thorough understanding of
the physiology of swallowing remains necessary to conduct a full
assessment and instigate appropriate treatment for these patients in
whom dysphagia is often debilitating and may significantly affect their
quality of life. We present an account of the physiology of swallowing,
using clinical examples to illustrate certain aspects.
Keywords Anatomy; Deglutition; Dysphagia; Physiology; Swallowing
Introduction
The pharynx assists in the shared functions of respiration and
swallowing (deglutition); coordination between these functions
is of great importance. This article reviews the phases of the
normal swallow and describes relevant applied anatomy and
physiology to assist in the understanding of what is a complex
and multifactorial process. Examples of relevant pathology are
used where they illustrate the phases.
Phases of swallowing
Swallowing can be subdivided into three main phases: oral,
pharyngeal and oesophageal. The oral phase can be subdivided
into the preparatory and transit phases. The oral phase is under
voluntary control whereas the pharyngeal and oesophageal
phases are involuntary. It is estimated that humans normally
produce around 500 ml of saliva per day and although swal-
lowing is thought of as an active function related to eating and
drinking, it should be noted that much of this activity is occurring
without active stimulation or awareness.
The oral phase
Anatomy
The oral cavity extends from the soft tissues of the lips and
cheeks anteriorly to the anterior tonsillar pillars. Its functions
Jenny Walton MBChB MSc(dist) MRCS DOHNS is a Specialist Registrar on
the Oxford Rotation, UK. Conflicts of interest: none declared.
Priyamal Silva FRCS ORL-HNS is a Consultant in Otolaryngology at the
John Radcliffe and Churchill Hospitals, Oxford, UK. Conflicts of
interest: none declared.
SURGERY --:-
Please cite this article in press as: Walton J, Silva P, Physiology of swallowing, Surgery (2018), https://doi.org/10.1016/j.mpsur.2018.08.010
include mastication and formation of a bolus, phonation, artic-
ulation and ventilation. The oral cavity is in direct communica-
tion and continuity with the oropharynx, the boundaries of
which are the soft palate superiorly and the tip of the epiglottis
inferiorly.
The nasopharynx lies between the skull base superiorly and
the soft palate inferiorly. Its anterior boundary is the posterior
choana. Its function is concerned with respiration; in normal
swallowing, the nasopharynx is closed off by the elevation of the
soft palate and uvula (Table 1).
The oral preparatory phase
This phase is concerned with the formation of a bolus from
material placed into the oral cavity. This does not apply to liq-
uids, as these need no oral preparation. The key features of this
stage are coordinated, purposeful movements of the tongue,
mandible and lip/buccal musculature and the simultaneous
closure of the upper oesophageal sphincter (UOS) to prevent
premature passage of food; these movements are coordinated in
the cerebellum (Table 2).1,2
The oral transit phase
Tongue movement is the most important feature of this stage, as
in the preparatory phase. The shape and movement of the tongue
acts to seal the food bolus against the palate; the lateral aspects of
the tongue sit along the alveolar ridges each side, stabilising the
tongue and thereby enabling the central part to propel the bolus
posteriorly. This phase lasts approximately 1 second and is
prolonged with increasing viscosity of the bolus and with
increasing age.2 The transit and preparatory phases may be
bypassed by syringing liquid into the back of the mouth to
initiate the pharyngeal phase.
The oral phase is controlled by 3 cranial nerves: the trigeminal
nerve (CN V), which controls chewing, the facial nerve (CN VII),
which controls the buccal and lip musculature to assist in the
positioning of food within the mouth, and the hypoglossal nerve
(CN XII), which controls tongue movement.3
The swallowing reflex
Swallowing is a complex action; voluntary initiation of swal-
lowing is mediated in the cortex, but swallowing may also be
triggered as a reflex response to food/liquid in contact with
particular areas in the oral cavity or oropharynx, or simply by the
accumulation of saliva.1 This is mediated in the medulla, which
receives afferent impulses from the nucleus tractus solitarius and
the spinal trigeminal nucleus. Efferent impulses from the medulla
pass through the nucleus ambiguus, the hypoglossal nucleus and
the motor nuclei of the trigeminal and facial nerves, leading to
the actions involved in the pharyngeal phase.
The gag reflex
The gag reflex is mediated by the same nerves as the swallowing
reflex, the difference being the initial stimulus. The gag reflex
is triggered by the presence of a stimulus in the oropharynx
outside of normal voluntary swallowing and subsequent
muscle contraction leads to gagging, retching or even vomiting
(Figure 1).
1 Ó 2018 Published by Elsevier Ltd.
 BASIC SCIENCE

Muscles of the soft palate

Muscle Actions Innervation Vascular supply

Levator veli palatini Elevate and pull back the posterior part
of the palate to close off nasopharynx
during swallowing
Tensor veli palatini Tightens and depresses soft palate to
open the Eustachian tube
Palatoglossus Elevates root of tongue and medialises
palatoglossal arches to separate oral
cavity from oropharynx
Palatopharyngeus Pull pharynx up and forwards to
shorten it during swallowing
Musculus Uvulae Retracts and thickens middle part of
palate to help close off nasopharynx
Cranial accessory nerve via pharyngeal
plexus
Mandibular nerve via nerve to medial
pterygoid
Cranial accessory nerve via pharyngeal
plexus
Cranial accessory nerve via pharyngeal
plexus
Cranial accessory nerve via pharyngeal
plexus
Ascending palatine branch of facial
Greater palatine branch of maxillary
Ascending palatine branch of facial
Greater palatine branch of maxillary
Ascending palatine branch of facial
Ascending pharyngeal
Ascending palatine branch of facial
Greater palatine branch of maxillary
Ascending pharyngeal
Ascending palatine branch of facial
Descending palatine branch of
maxillary

Table 1

Components of the oral preparatory phase

Component of phase Muscles Function

Lip closure Orbicularis oris Keep food in the mouth

Contraction of buccal musculature Buccinator Prevent food entering the lateral sulci
Rotary motion of jaw Temporalis, masseter, medial pterygoid Chewing
Lateral rolling motion of tongue Intrinsic muscles, genioglossus Direct food towards teeth
Bulging of the soft palate anteriorly Palatoglossus, levator veli palatini Close off oropharynx and widen nasal airway

Table 2

The pharyngeal phase

Anatomy
The pharynx continues inferiorly as a tube surrounded by
muscular layers. The outermost layer consists of the three
pharyngeal constrictor muscles: muscle sheets situated posteriorly
with their fibres running obliquely. The inferior constrictor con-
sists of thyropharyngeus and cricopharyngeus, the latter of these
forming the primary component of the Upper Oesophageal
Sphincter (UOS). A deficiency occurs between these two parts: see
CAB 3. The inner layer consists of longitudinal muscles: palato-
pharyngeus, stylopharyngeus and salpingopharyngeus. Contrac-
tion of these muscles during swallowing acts to shorten and widen
the pharynx, facilitating passage of the bolus into the oesophagus.
The suprahyoid muscles: digastric, mylohyoid, stylohyoid and
geniohyoid, act to elevate the hyoid bone and larynx during
swallowing in order to protect the airway. This function is aided
by the epiglottis: a sheet of fibrocartilage sitting anterior to the
laryngeal inlet (Figure 2). The epiglottis is not essential clinically
and it is possible for a functional, safe swallow to be adopted if
the epiglottis is removed.

Figure 1 Nerves of the swallowing and gag reflexes. Image modified
from Standring S, ed. Gray’s Anatomy: the anatomical basis of clinical
practice. 40th edn. Elsevier. 2008. Fig 19.12 Swallowing and gag re-
flexes. p283.
Physiology
This phase is triggered by stimulation of the glossopharyngeal
(cranial nerve (CN) IX) and vagus (CN X) nerves by the presence
of the bolus, and is mediated in the medulla. Coordination

SURGERY --:- 2 Ó 2018 Published by Elsevier Ltd.

Please cite this article in press as: Walton J, Silva P, Physiology of swallowing, Surgery (2018), https://doi.org/10.1016/j.mpsur.2018.08.010
 BASIC SCIENCE
Figure 2 Posterior view of the larynx.
between this centre and the respiratory centre is vital due to the
need for respiration to cease for a fraction of a second during
airway closure.
As the bolus is propelled into the oropharynx, the soft palate
elevates, sealing off the nasopharynx and preventing nasal
regurgitation. Structural abnormalities in the soft palate can lead
to nasal regurgitation and hypernasal speech (Box 1). The
oropharynx is funnel shaped at this point to direct the bolus
inferiorly. Movement of the bolus through the pharynx occurs
sequentially, beginning at the superior constrictor muscle and
continuing inferiorly.
Contraction of the suprahyoid muscles elevates the hyoid
bone and larynx in an anterosuperior direction, away from the
cervical spine so that it is protected by its new position right
underneath the tongue base. The bolus can therefore safely pass
from the tongue base into the pharynx without entering the lar-
ynx. Laryngeal and hyoid elevation also acts to shorten and
widen the pharynx, creating a negative pressure in the
Clinical application: cleft palate
The anatomy of the palate sees levator palatini forming a sling, which
acts to elevate the soft palate during swallowing and exclude the
contents of the oral cavity from the nasopharynx. In a patient with
cleft palate, the fibres of levator palatini lie in a longitudinal orien-
tation, parallel to the cleft itself.4 The clinical consequence of this is
that total nasopharyngeal closure cannot be obtained during swal-
lowing and nasal regurgitation of milk during feeding in the neonate
is common. Following surgical repair, the act of re-orientating the
muscle fibres can consequently shorten the soft palate and lead to
incomplete closure during speech and swallowing; nasal regurgita-
tion during swallowing and perhaps more commonly escape of air
during speech (velopharyngeal insufficiency) can still occur.
Box 1
SURGERY --:-
Please cite this article in press as: Walton J, Silva P, Physiology of swallowing, Surgery (2018), https://doi.org/10.1016/j.mpsur.2018.08.010
laryngopharynx, which aids progress of the bolus. This move-
ment also occurs due to posteriorly directed pressure from the
tongue, inward pressure from contraction of the pharyngeal
constrictors and gravity (Box 2).3
Prevention of laryngeal penetration (entry of food/drink into
the laryngeal inlet) is avoided by several mechanisms.1 Firstly,
movement of the epiglottis acts as a diverter, directing liquid
material laterally via the piriform fossae into the oesophagus,
while solid material passes directly over the top. It is the com-
bination of pressure from the tongue base and active contraction
of the aryepiglottic muscles that causes the epiglottis to be
deflected posteriorly. It is felt that this is the least important
mechanism in airway protection during swallowing.2 Secondly,
the false vocal cords create a sphincter to protect the airway but
thirdly and most importantly, closure of the glottis by adduction
of the true cords seals the airway against entry of foreign mate-
rial. This closure is only maintained for a fraction of a second, as
it requires a complete pause in respiration. The muscles acting to
adduct the true cords are the lateral cricoarytenoid (closure of the
anterior part) and the posterior and oblique cricoarytenoids
(closure of the posterior aspect). Closure of the UOS and re-
establishment of normal respiration prevents air entry into the
stomach (Box 3).
If material does enter the larynx, the protective cough reflex
can be used to clear it from the airway. This reflex occurs due to
stimulation of laryngeal branches of the vagus nerve (CN X);
impulses travel via the trigeminal sensory nucleus to the nucleus
ambiguus and respiratory centres in the medulla. Stimulation of
intercostal and abdominal wall muscles cause coughing due to
pressure build up against an initially closed glottis.7
The oesophageal phase
Anatomy
The pharynx continues inferiorly into the oesophagus, a
muscular tube of about 25cm in length. The oesophagus in the
neck lies directly posterior to the trachea; behind the oesophagus
lie the prevertebral fascia, longus colli muscle and the cervical
vertebrae. The oesophagus can be divided into the upper two
thirds, which contain striated muscle, and the lower one-third,
which is smooth muscle. The lower segment forms the lower
oesophageal sphincter (LOS), a 3 cm long segment of oesophagus
extending just above and below the diaphraghm with a raised
Clinical application: pharyngeal pouch (Figure 3)
The location and orientation of the pharyngeal constrictors reveals a
deficiency in the posterior pharyngeal wall between the two parts of
the inferior constrictor. This deficiency is known as Killian’s dehis-
cence and is an area of potential weakness. Persistently high pres-
sures in the pharynx during swallowing can lead to a posteriorly
based protrusion of this weakness and is termed Zenker’s divertic-
ulum, or pharyngeal pouch. A pharyngeal pouch classically leads to
halitosis, regurgitation of food/fluid back into the oropharynx and
can even contribute to recurrent aspiration. Management of this
pouch is usually surgical, either endoscopically with staples or
externally via the neck.
Box 2
3 Ó 2018 Published by Elsevier Ltd.
 BASIC SCIENCE
Figure 3 Location of a pharyngeal pouch.
Clinical application: vocal cord palsy
A non-functioning vocal cord may occur due to many reasons:
congenital causes, trauma from surgery e.g. thyroidectomy or
cardiothoracic procedures, malignancy, neurologic conditions or may
be idiopathic. Paresis or paralysis of one or both vocal cords may be
due to injury to the superior (SLN) or recurrent laryngeal nerves (RLN)
and often the initial symptoms experienced by the patient include
significantly altered voice and difficulty breathing. However, an
important and debilitating consequence of vocal cord paralysis is to
the swallow. Both nerves provide sensation to the larynx: the internal
branch of the SLN supplies the laryngeal mucosa above the level of
the vocal folds and the folds themselves and the RLN supplies the
mucosa below, as well as the lining of the trachea. Damage to either
of these nerves has the potential to produce a partially insensate
larynx, increasing the risk of aspiration. The internal branch of the
SLN (ISLN) provides the laryngeal protective reflexes and damage can
therefore also result in dysphagia5; studies have shown an increased
incidence of laryngeal penetration and tracheal aspiration in the
anaesthetized larynx.6 In addition, patients with an anesthetized ISLN
may experience symptoms of globus and discomfort and feel the
need to produce a more forceful swallow to clear a bolus. The RLN
innervates all laryngeal muscles except cricothyroid; damage can
lead to an inability to fully close the glottis during swallowing.
Box 3
SURGERY --:-
Please cite this article in press as: Walton J, Silva P, Physiology of swallowing, Surgery (2018), https://doi.org/10.1016/j.mpsur.2018.08.010
intraluminal pressure created by circular muscle fibres and the
surrounding diaphraghm.
Physiology
When the pharyngeal phase is triggered, the UOS opens to allow
passage of the bolus into the oesophagus. The passage of food
past the UOS signals the start of the oesophageal phase. Suc-
cessful swallowing requires relaxation of the UOS at the correct
time with subsequent contraction to help prevent laryngophar-
yngeal reflux (Box 4). As soon as the bolus enters the oesoph-
agus, peristaltic waves propel the bolus rapidly onwards towards
the stomach and the LOS relaxes to allow unimpeded passage.
Peristalsis is initiated by Auerbach’s plexus, which is situated
between the muscle layers.
If the individual is upright at the time of swallowing, gravity
plays a role in the passage of liquids into the stomach however
the same is not the case for solid boluses; these always require
peristalsis to move the bolus.9 Peristaltic waves also act to clear
the remaining liquid bolus so passage is not entirely reliant on
gravity alone. During drinking, the oesophageal phase is not
triggered until the last swallow; then peristalsis will commence.
It takes a bolus around 5e6 seconds to travel the full length
of the oesophagus with normal peristalsis; this is a rate of around
3e4 cm/second.
Swallowing in the fetus and neonate
Anatomy of the larynx and pharynx in the neonate and young
child differs from that of the adult. The larynx is situated in a
more elevated position; often in close contact with the soft palate
in the neonate and it is often possible to visualize the epiglottis
when examining a child’s oropharynx without the use of in-
struments. After the first few months of life, the pharynx extends
and the larynx takes up its adult position lower in the neck7; this
may reduce the chance of aspiration.
Swallowing movements are first noticed in utero at around 11
weeks’ gestation.7 The fetus will swallow around 450 mls of am-
niotic fluid per day and will subsequently develop swallow-induced
peristaltic activity. Interestingly, aspiration does not occur in the
foetus under normal conditions and it would therefore seem that
the protective functions of the larynx are already established prior
to birth.10 At the time of birth, the neonate already possesses the
ability to coordinate sucking, swallowing and breathing and will
demonstrate this within the first couple of feeds.
Clinical application: laryngopharyngeal reflux
Laryngopharyngeal reflux (LPR) is the regurgitation of stomach con-
tents into the larynx or pharynx. It occurs due to failure of one of
more of the protective mechanisms designed to move food and
saliva onwards towards the stomach: the UOS and LOS, saliva pro-
duction, oesophageal peristalsis and gravity.8 The patient will often
experience excessive throat clearing, coughing, hoarseness, and
globus pharyngeus (the sensation of a lump in the throat). Man-
agement includes lifestyle modification, acid-suppression medication
and alginates, with surgery used uncommonly and of uncertain
benefit.
Box 4
4 Ó 2018 Published by Elsevier Ltd.
 BASIC SCIENCE
Investigating the abnormal swallow
Dysphagia is the term used to cover a number of symptoms
experienced by a patient whose normal swallowing mechanism
has failed.
Aspiration is the entry of food, liquid or saliva into the airway
below the level of the true vocal cords. In a patient with adequate
airway sensation, this will lead to coughing in an attempt to clear
the material. Patients who cannot clear this material either due to
inadequate sensation or weak cough reflex are at risk of aspiration
associated pneumonia.2 Material may enter the airway at several
points during the swallow. If tongue control is weakened; material
can fall backwards and enter the airway during the oral phase, as
laryngeal positioning and airway protection have not yet been
initiated. Secondly during the pharyngeal phase: if triggering of
airway protection is delayed, it is possible for material to enter the
airway. Altered neurology such as following a stroke, or any local
condition e.g. following treatment for head and neck cancer may
cause aspiration during either of these phases.
Bedside assessment of swallow
By careful observation, information can be gleaned regarding
movement of the lips, tongue, pharyngeal control, laryngeal
protection and respiratory control, although there is often
considerable variability between the findings of different tes-
ters.11,12 The advantage of such assessment is that the patient’s
swallow can often be optimized at the bedside. Flexible Endo-
scopic Evaluation of Swallowing (FEES) can help to assess pa-
tients with altered airway sensation: this involves direct
observation of the larynx using a nasendoscope whilst asking the
patient to swallow a coloured liquid, e.g. blue dye, or milk.
Although at the point of the pharyngeal swallow the tip of the
nasendoscope will become obscured due to movement of the
pharyngeal walls and passage of liquid, the view of the larynx is
quickly restored and laryngeal penetration will be easily visible.
Barium swallow
Radiologic images of the swallow are taken whilst the patient
swallows contrast material. This is commonly barium, although
water-soluble materials are available for use in any patient in
whom a perforation is suspected or those at high risk of aspira-
tion. Contrast material dilates the normally collapsed oesophagus
and allows visualisation of the anatomy and motility of the
oesophagus. Barium swallow is excellent for demonstrating
pharyngeal diverticulae, although strictures, webs and oesopha-
geal reflux may be better appreciated with the use of trans-nasal
oesophagoscopy (TNO) or oesophago-gastroduodenoscopy
(OGD) in order to directly visualise the area.
Videofluoroscopy
Videofluoroscopy (VF) allows dynamic, radiologic examination
of all the phases of swallowing. The aims are to identify: phases
of the swallow that are causing problems, consistencies that pose
the greatest challenge, and to find strategies to overcome these
difficulties. In addition to speech and language therapy and
radiology, members of the wider multidisciplinary team: otolar-
yngology, gastroenterology, neurology and psychiatry may also
be involved.
SURGERY --:-
Please cite this article in press as: Walton J, Silva P, Physiology of swallowing, Surgery (2018), https://doi.org/10.1016/j.mpsur.2018.08.010
The essential information gained from this procedure includes
direction of travel of the bolus and residual coating, presence of
laryngeal penetration or aspiration and patient reaction, and
coordination/timings of muscle movements.12 The test utilizes
different consistencies of material mixed with contrast; boluses
are given in increasing amounts to minimize risk of aspiration.
Visualisation of the mucosa and surrounding secretions is,
however, difficult.
Oesophagoscopy
Excellent direct views of the pharynx and larynx can be obtained
using a rigid oesophagoscope when the patient is under general
anaesthesia. This enables visualisation of any lesion in the upper
aerodigestive tract and allows biopsies to be taken. The disad-
vantage is that dynamic assessment of the vocal cords and the
swallow cannot take place.
Conclusion
The key component of a successful swallow is airway protection;
this requires precise coordination of the swallowing and
breathing centres in the cortex and brainstem leading to a com-
plex process of laryngeal manoeuvres and bolus movement. Such
a process has numerous steps at which problems can occur
leading to dysphagia and these can be investigated using the
methods described. A comprehensive understanding of relevant
anatomy and physiology will aid the clinician in this process and
allow appropriate subsequent management. A
REFERENCES
1 Leslie P, McHanwell S. Ch. 150. Physiology of swallowing. In:
Gleeson M, Browning GG, Burton MJ, et al., eds. Scott-Brown’s
otolaryngology, head and neck surgery. 7th edn, vol. 2. CRC
Press, 2008.
2 Logemann J. Ch. 90. Mechanisms of normal and abnormal swal-
lowing. In: Flint PW, Haughey BH, Lund VJ, et al., eds. Cummings
otolaryngology: head and neck surgery. 5th edn. vol. 1, 2010;
p1215e21.
3 Hennessy M, Goldenberg D. Surgical anatomy and physiology of
swallowing. Oper Tech Otolaryngol 2016; 27: 60e6.
4 Hoffman WY. Cleft lip & palate. In: Lalwani AK, ed. Current diag-
nosis and treatment: otolaryngology head and neck surgery. 3rd
edn. McGraw-Hill, 2012.
5 Scerrino G, Tudisca C, Bonventre S, et al. Swallowing disorders
after thyroidectomy: what we know and where we are. A sys-
tematic review. Int J Surg 2017; 41: 94e102.
6 Jafari S, Prince RA, Kim DY, Paydarfar D. Sensory regulation of
swallowing and airway protection: a role for the internal superior
laryngeal nerve in humans. J Physiol 2003; 550: 287e304.
7 Standring S, ed. Gray’s anatomy: the anatomical basis of clinical
practice. 40th edn. Elsevier, 2008.
8 Campagnolo AM, Priston J, Thoen RH, Medeiros T, Assunça ~o AR.
Laryngopharyngeal reflux: diagnosis, treatment, and latest
research. Int Arch Otorhinolaryngol 2014; 18: 184e91.
9 Goyal RK, Mashimo H. Physiology of oral, pharyngeal, and
esophageal motility. 2006. GI Motility online, https://www.nature.
com/gimo/contents/pt1/full/gimo1.html (accessed 14 May 2018).
5 Ó 2018 Published by Elsevier Ltd.
 BASIC SCIENCE

10 Jadcherla SR, Hogan WJ, Shaker R. Physiology and pathophys-
iology of glottic reflexes and pulmonary aspiration: from neonates
to adults. Semin Respir Crit Care Med 2010 Oct; 31: 554e60.
11 McCullough G, Wertz R, Rosenbek J, Mills R, Boss K, Ashford J.
Inter- and intrajudge reliability of a clinical examination of swal-
lowing in adults. Dysphagia 2000; 15: 58e67.
12 Hirst LJ. Ch. 151. Functional investigations of the upper gastroin-
testinal tract. In: Gleeson M, Browning GG, Burton MJ, et al., eds.
Scott-Brown’s otolaryngology, head and neck surgery. 7th edn, vol.
2. CRC Press, 2008.

SURGERY --:- 6 Ó 2018 Published by Elsevier Ltd.

Please cite this article in press as: Walton J, Silva P, Physiology of swallowing, Surgery (2018), https://doi.org/10.1016/j.mpsur.2018.08.010