DIGESTIVE PHYSIOLOGY

I. Morpho-functional particularities
of visceral smooth muscle

Organization in muscular layers:

- longitudinal → outside
- circular → inside
- obliques → deep (only at the stomach);

Between the longitudinal muscle layer and

the circular one is the myenteric nerve plexus
of Auerbach  intrinsic innervation;

Between muscle fibers and nerve endings

are established diffuse junctions  one
vesicle with NT (neurotransmitter) for several
muscle fibers. 2
• Morpho-functional organization of the syncytial type  muscle
fibers conduct excitation from one cell to another by connexons
- the excitation spreads in the cranio-caudal direction
- the contraction of the muscle tunica is unitary
• Presents functional automatism  Cajal's pacemaker cells
(Auerbach's plexus) is coupled to smooth muscle fiber by "gap"
type junctions
• The basal tone (myogenic mechanism)
 permanent state of contraction which
is based on resting potential fluctuations
of around -50 mV
• Contraction control  nervous and
hormonal mechanism
 Types of gastrointestinal tract
movements
Movements associated with
• Tonic movements: the contraction of
digestion and absorption
the digestive sphincters prevents the
premature passage of intestinal
contents from one segment to another

• Segmentation movements: mixing

Movements associated with • Peristaltic movements: propulsion

the "fasting" state
• Gastric movements: peristaltic
contractions of "hunger”

• Bowel movements: migration motor

complexes (MMCs) that underlie
regular peristaltic contractions of
complete gastrointestinal evacuation
 PERISTALTIC MOVEMENTS (peristalsis)

Are characteristic for smooth muscle syncytia of tube configuration

(bile ducts, glandular ducts, ureters, etc.);

Spread intestinal contents:

- over a distance of 5 to 10 cm
- step by step
- in the cranio-caudal direction

Are triggered by:

- mechanical distension of the digestive tract
- chemical and physical irritation of intestinal mucosal epithelium
- intense parasympathetic stimulation

Are coordinated by Auerbach's myenteric plexus  necessary for

effective peristalsis
 PERISTALTIC MOVEMENTS (peristalsis)

 Peristalsis is the progression of coordinated contraction of

involuntary circular muscles, which is preceded by a
simultaneous contraction of the longitudinal muscle and
relaxation of the circular muscle in the lining of the gut;
 The peristaltic movement comprises relaxation of circular
smooth muscles, then their contraction behind the chewed
material to keep it from moving backward, then longitudinal
contraction to push it forward;
 The peristaltic wave propels a ball of food (called a bolus
before being transformed into chyme in the stomach) along the
digestive tract;
 Peristalsis is generally directed caudal, that is, towards the anus.
This sense of direction might be attributable to the polarisation
of the myenteric plexus.
 PERISTALTIC MOVEMENTS (peristalsis)

 primary peristaltic wave: occurs when the bolus enters the

esophagus during swallowing, forces the bolus down the
esophagus and into the stomach in a wave lasting about 8–9
seconds. The wave travels down to the stomach even if the
bolus of food descends at a greater rate than the wave itself,
and continues even if for some reason the bolus gets stuck;
 secondary peristaltic wave: if the bolus gets stuck or moves
slower than the primary peristaltic wave (poor lubrication), then
stretch receptors in the esophageal lining are stimulated and a
local reflex response is triggered around the bolus, forcing it
further down the esophagus; these secondary waves continue
indefinitely until the bolus enters the stomach;
 tertiary peristalsis: is dysfunctional and involves irregular, diffuse,
simultaneous contractions (esophageal dysmotility).
 Starling's Law of the Intestine
explains peristalsis by stating that stimulation
of the gut produces excitation above and
relaxation below, in orderly sequence.

Distension of a digestive segment

Peristaltic reflex Anal targeting of

peristaltic movements
1. Contractile wave (Ach/SP), with
a length of 2-3 cm, located
proximal to the site of distension
+ LEGEND:
2. Receptive relaxation wave (NO/VIP), ACh = acetylcholine
SP = substance P
located caudal of the site of NO = nitric oxide
distension, in the direction of the VIP = vasoactive intestinal peptide
cranio-caudal front PPS = postsinaptic potential
 SEGMENTATION MOVEMENTS

CHARACTERISTICS:
Are series of other concentric alternating
contractions, followed by relaxation 
a new contraction occurs between the
two previous contractile parts;
Determine the fragmentation of the
intestine in a sequence of:
- mixing segments
- contractile segments of limited
bidirectional propulsion
Frequency of contraction movements is
characteristic for an intestinal segment
(local pacemaker);
Segmentation contractions cause
separation and mixing without pushing
materials further down the digestive
tract (unlike peristalsis).
The relation of the digestive system with
the others systems of the organism
 Two groups of organs compose the
digestive system

 Gastrointenstinal (GI)
tract or alimentary canal:
mouth, most of pharynx,
esophagus, stomach,
small intestine, and large
intestine
 Accessory digestive
organs: teeth, tongue,
salivary glands, liver,
gallbladder, and pancreas
 Functions of the digestive system

 Ingestion
 Secretion of water, acid, buffers,
and enzymes into lumen
 Mixing and propulsion
(motility)
 Digestion:
- mechanical digestion
(churns food)
- chemical digestion
(hydrolysis)
 Absorption – passing into blood
or lymph
 Defecation – elimination of
feces
Mouth, Pharynx, Esophagus, and Stomach
Layers – small intestine
Layers of small intestine

Copyright © 2008 Pearson Education, Inc., publishing as Benjamin Cummings.

Esophagus

 Muscular tube from pharynx

to stomach
 Upper 1/3—skeletal muscle
 Lower 2/3—smooth muscle
 Upper esophageal sphincter
 Skeletal muscle
 Between pharynx and
esophagus
 Lower esophageal sphincter
 Smooth muscle
 Between esophagus and
stomach
Pyloric Sphincter

Regulates the passage of the chyme between stomach

and small intestine
Small Intestine
Circulatory Route: Absorbed Material
Features of the Colon
Accessory Structures
The Pancreas
GI Organs and Functions
Innervation of the Digestive System
Innervation of the Digestive System
Innervation of the Digestive System
Innervation of the Digestive System
Innervation of the Digestive System
 II. Motor function
of the gastro-intestinal tract

 Mastication
 Deglutition
 Gastric motility
 Intestinal motility
 Defecation
1. MASTICATION

DEFINITION

 The process of chewing food, in preparation for

swallowing and digestion

 A motor act accomplished by the controlled activity

of the components of masticatory system which
are controlled by the neuromuscular mechanism.
 1. MASTICATION

DEFINITION: process of mechanical treatment of food and it’s

impregnation with saliva, in the mouth, called FOOD BOLUS
 dents
• Passive element  mandibule
 temporo-mandibulary artic.
 mm masticators (n.V)
• Active element  mm. of the tongue (n.XII)
 mm. oro-facials (n.VII)
Element Functional role
Dents  occlusion and trituration surface
Mm. masticators  provide mandibular movements
 provide the occlusion position of the mouth
Mm. of the tongue  propel food onto the crushing surface
Mm. oro-facials  lies saliva and food in the mouth
 CONTROL OF MASTICATION

MECHANISM: sequence of unconditioned lowering/raising reflexes

of the mandible, which is supplemented by voluntary control

Masticatory center of the pons Frontal motor area 4

(motor nucleus of the n.V) (voluntary control)
(-) () n.V Cortico – nuclear tract

Proprioceptiv  n.V  mandibulary movements

n.VII
stimuli  n.VII  oro-faciale coordination
n.IX
 n.XII  tongue coordination
Chewing reflex
(miotactic reflex)
Raising the mandibule
Taste stimuli
32
Réflexe linguo-maxillaire Lowering the mandibule
 MASTICATORY REFLEX
1. MYOTATIC reflex 2. Reverse MYOTATIC reflex

Voluntary opening of the mouth Contraction of the masseter muscle

Stretch of the masseter muscle Strech of Golgi tendon organ

Type I-a sensory fibers of the n.V Type I-b sensory fibers of the n.V
(+) (-)
Pons motor nucleus of the n.V Pons motor nucleus of the n.V

n.V somatic motor fibers n.V somatic motor fibers

Contraction of the masseter muscles Relaxation of the masseter muscles

RAISING OF THE MANDIBLE LOWERING OF THE

MANDIBLE
 FUNCTIONAL VALUE OF MASTICATIONS

1. Ensures GOOD DIGESTION of food

- digestive enzymes acting on the surface of mechanically and
chemically processed food particles

2. TROPHIC role for the mouth

- ensures the development of facial mass
- stimulates secretion, saliva
- provides self-cleaning mouth
- intervenes in the olfactory and gustatory perception

3. TROPHIC role for the gastrointestinal tract

- provides mechanical integrity of the gastrointestinal tract
- provides reflex stimulation of gastrointestinal secretion and
motility
COMPONENTS OF MASTICATION

 Dentition
 Periodontal supporting tissues
 Maxilla and Mandible
 Temporomandibular Joint
 Mandibular musculature
 Muscles of lips, cheeks, and tongue
 Involving soft tissue
 Supplying innervation and vascularization
MASTICATION CYCLE
 Chewing cycle: succession of unconditional reflexes of
lowering and lifting the mandible (jaw opening and closing in
the vertical plane), plus voluntary control.

 The chewing cycle consists of three phases:

1. Opening phase: the mouth is opened and the mandible is

lowered
2. Closing phase: the mandible is raised towards the maxilla
3. Occlusal phase (intercuspal phase): the mandible is
stationary and the teeth, from both upper and lower arches,
come in contact
 Most of the muscles of chewing are innervated by the
motor branch of the V-th cranial nerve
 The chewing process is controlled by nuclei in the brain
stem; stimulation of specific reticular areas in the brain
stem taste centers will cause rhythmical chewing
movements.
 Stimulation of areas in the hypothalamus, amygdala,
cerebral cortex near the sensory areas for taste and
smell can cause chewing.
FACTORS AFFECTING MASTICATION

• Saliva - facilitates mastication, moistens the food

particles, makes a bolus and assists swallowing

• Characteristics of the food - water and fat

percentage, food hardness - are sensed during
mastication and affects masticatory force, jaw muscle
activity, and mandibular jaw movements

• The movement of the jaw and thus the neuromuscular

control of chewing - role in the comminution of the food

• The integrity of the occlusion surfaces of the teeth

and of the muscles of mastication .
 SIGNIFICANCE OF MASTICATION

•Is the first step in digestion

• Increases the surface area of foods, allows more efficient

break down by enzymes

• The food is made softer and warmer and the enzymes in

saliva begin to break down carbohydrates.

• Lubrication and moistening of dry food by saliva, so that

bolus can be easily swallowed

• Appreciation of taste of food

 COMPONENTS OF MASTICATION

1. Passive component - maxillary dental system:

• teeth - provides a surface grinded involved in cutting,

crushing, grinding food and an occlusal surface in
premolars and molars

• mandible

• temporo-mandibular joint: a complex articulation with

limited capability of diarthrosis (free movement)
 COMPONENTS OF MASTICATION

 The teeth are designed for

chewing.
 The anterior teeth (incisors)
provide a strong cutting action
 The posterior teeth (molars)
provide a grinding action.
All the jaw muscles working
together can close the teeth with
a force as great as 55 pounds on
the incisors and 200 pounds on
the molars.
 COMPONENTS OF MASTICATION

2. Active component:

 masticatory muscles – provide jaw movements by

contraction and by their resting tone provide the occlusion;
 classified as muscles that raise the mandible: masseter
m., temporal and internal pterygoid muscles and muscles
that lower of the mandible: external pterygoid muscle,
external mylohyoid m., geniohyoid and anterior belly of
digastric
- tongue muscles: provide propulsion of food on grinded
surface
- oro-facial muscles: keep saliva and food in the mouth
Masticatory Muscles Non Masticatory Muscles

 Masseter muscle
 Digastric muscle
 Temporalis muscle
 Mylohyoid muscle
 Medial pterygoid
 Geniohyoid muscle
muscle
 Orbicularis Oris
 Lateral pterygoid
muscle
Muscles of Mastication
Muscles of Mastication
Non Masticatory Muscle
Action of muscles during masticatory
movements

 Opening (depressor jaw muscles)

mylohyoid / digastric / inferior lateral pterygoid

 Closing (elevator jaw muscles)

medial pterygoid / superficial masseter / tempolaris
 Tongue muscles

 Consist of 2 groups:
Intrinsic muscle: change in tongue shape
Extrinsic muscles (eg. Genioglossus): response for
protrusion and retrusion of the tongue; three major
muscles that anchor and move the tongue
 Innervated by cranial nerve XII (hypoglossal nerve)
 Complete tongue activity occurs in jaw movements,
respiration, speech, taste, mastication, swallowing, and
sucking.
Extrinsic Tongue Muscles

Figure 10.7c
Mandible - Chewing
 Most of the chewing process is caused by a
chewing reflex (trigeminal reflex)

 The presence of a bolus of food in the mouth initiates reflex

inhibition of the muscles of mastication, which allows the
lower jaw to drop.
 This drop in turn initiates a stretch reflex of the jaw muscles,
that leads to rebound contraction  automatically raises the
jaw to cause closure of the teeth.
 It also compresses the bolus against the linings of the mouth,
which inhibits the jaw muscles once again, allowing the jaw
to drop and rebound another time.
 This process is repeated again and again.
UNCONDITIONAL MASTICATORY REFLEX

 Receptors : tactile r., proprioceptive r., taste r.

 Afferent fibers:
- V cranial nerve (trigeminal n.) for tactile and proprioceptive
stimuli
- VII (facial n.) and IX (glossopharyngeal n.) cranial nerves for
taste stimuli
 Pons Masticatory center (ncl. n V)
 Efferent fibers:
- V cranial nerve for masseter m.
- VII cranial nerve for oro-facial muscles
- XII cranial nerve (hypoglossal nerve) for tongue muscles
 Efectors: masticatory muscle
 The trigeminal reflex of lifting the mandible:
- voluntary opening of the oral cavity  masseter muscle
extension  stimulation of the primary sensory
terminations of the neuro-muscular spindle - (type a) of
nerve V  afferents in sensitive nucleus of V-th nerve
(medulla oblongata)  ACTIVATION of the mastication
center in PONS (corresponds to nucleus of origin of
the V-th nerve)  motor efferents, trigeminal nerve 
CONTRACTION of the maseter muscle  lifting the
mandible.
 Reversed trigeminal reflex of lowering the mandible
- contraction of the maseter muscle  stretching of the Golgi
tendinous organ (sensitive terminations type Ib of Vthn.) 
afferents  sensitive nucleus of V-th nerve, from the medulla 
INHIBITION of the PONTIN mastication center 
RELAXATION of maseter m.  lowering of the mandible.
 Lingo-maxillary reflex, mandibular lowering reflex
- starting point: contact with food
- taste R from mucosa  afferents to VII-th nerve and IX-th nerve
 MEDULLA  PONTIN mastication nucleus
 Presence of food in the mouth initiates reflex inhibition of muscles of
mastication ,which allows lower jaw to drop (Reversed trigeminal reflex)

This inhibits the jaw muscles once

again, allowing the jaw to drop The drop in turn initiates a stretch reflex of
and rebound another time the jaw muscles that leads to rebound
contraction

This automatically raises the jaw to cause

closure of the teeth and compresses the bolus
again against the linings of the mouth (The
trigeminal reflex )
 2. DEGLUTITION
• It begins after mastication and
bolus formation.
• Swallowing reflex = sequence of
events that result in propulsion of
food from the mouth to the stomach.
• It can be initiated voluntarily, then
it is under reflex control.
• The bolus activates oropharyngeal
sensory receptors that initiate the
deglutition reflex.
• It is coordinated by the swallowing
center in medulla oblongata and
lower pons
• It involves 3 phases:
- oral,
- pharyngeal and
- esophageal phase.
 2. DEGLUTITION
DEFINITION: the mechanical process in 3 stages (buccal, pharyngeal
and esophageal) by which the food bolus is moved, from the mouth
through the pharynx and the esophagus, into the STOMACH
I. BUCCAL TIME (~0,3 sec) - voluntary
Tongue tip elevation • placeing the bolus on
the base of the tongue

• lifting of the anterior part of the tongue

Passage of the • contraction of the tongue muscle
oropharyngeal isthmus • extension of the oropharyngeal isthmus
• relaxation of the pillars of the soft palate
• soft palate lift

Pharyngeal aspiration
of food bolus •pharyngeal negative pressure
 I. BUCCAL (ORAL) - VOLUNTARY TIME

 Food, ready for swallowing is

voluntarily squeezed or rolled
posteriorly into the pharynx by
the pressure of the tongue,
upward and backward against
the hard palate.

 From there on, swallowing

becomes entirel (or almost
entirely) automatic and usually,
cannot be stopped.
II. PHARYNGEAL TIME (1-2 sec) – involuntary
BUCCAL CAVITY
 closure of the oropharyngeal isthmus
Closure of the oral  prolonged contraction of the tongue
route muscles
 contraction of the levator and tensor soft
Closure of the palate muscle
nasal passage
 lifting the larynx
Closure of the  lowering of the opening of the glottis
laryngeal passage  the opening of the glottis with the epiglottis
 contraction of the muscles of the pharynx
Opening of the  pressure that pushes the bowl is 70-100
esophageal cm H2O
pathway

OESOPHAGUS
 II. PHARYNGEAL STAGE - INVOLUNTARY

a. Soft-palate is pulled upward

closing off the nasopharynx
b. Palatoglossal and
palatopharyngeal arches are pulled
medially, forming a sagittal slit with
the fauces, through which food is
passed to posterior pharynx
c. Vocal cords close
d. Epiglottis swings backward over
larynx and larynx is pulled upward, to
close off the opening of the larynx
e. Upper esophageal sphincter
(UES) relaxes to that bolus can enter
the esophagus
• Tongue sweeps backward → bolus
into the oro-pharynx
• Elevation of soft palate and
contraction of the upper constrictor
muscle of the pharynx to close
nasopharynx.
• Epiglotis and vocal cords shut off
the larynx
• Reflex apnea
• Relaxation of UES
• Peristatic contraction begins in the
upper constrictor muscle → middle
and inferior constrictor muscle
Nervous control of pharyngeal stage
 Afferents
Ring of nerve plexus of pharyngeal opening, with greatest sensitivity
in the tonsillar pillar (receptors), via Trigeminal N, Glossopharyngeal
N, Vagus N.
 Nucleus
- Tractus Solitarius and Nucleus Solitarius - sensory nuclei in
the medulla oblongata
- Nucleus Ambiguus – motor nucleus in the medullary reticular
formation
 Efferent
To pharyngeal musculature and tongue via V th , IXth, Xth, XIIth
cranial nerves and even a few of superior cervical nerves.
 III. OESOPHAGEAL TIME (5 -7 sec) – involuntary
OESOPHAGUS

Transient relaxation
of the UES

Propagation of the
food bolus by
peristaltic waves

Transient receptive
relaxation of LES
and stomach
ESOPHAGEAL MANOMETRY
UES = upper esophageal sphincter
STOMACH LES = lower esophageal sphincter
PRIMARY PERISTALTIC WAVES
- are triggered by vagal mechanism
- begin with relaxation of the upper
esophageal sphincter (UES) determined
by food bolus
- end with receptive relaxation of the lower
esophageal sphincter (LES)

SECONDARY PERISTALTIC WAVES

- are triggered by the Auerbach myenteric
plexus
- begin at the site of esophageal over-
distension, when the primary peristaltic
wave CANNOT ensure bolus movement

TERTIARY PERISTALTIC WAVES

- Esophageal disfunction (iregular contractions)
UPPER ESOPHAGEAL SPHINCTER (UES)
- generates, by a tonic contraction, a pressure = 50
mm Hg,
- prevents the passage of air into the esophagus
(aerophagia),
- prevents the regurgitation of food and it aspiration
into the larynx.

LOWER ESOPHAGEAL SPHINCTER (LES)

- generates, by a tonic contraction, a pressure = 20
mm Hg,
- temporarily stops the progression of the food
bolus,
- prevents gastroesophageal reflux (reflux
esophagitis).
 III. ESOPHAGEAL STAGE - INVOLUNTARY

a. Peristalsis pushes the bolus

downward through the
esophagus.

b. Lower esophageal sphincter

relaxes and the bolus enters
the stomach.
c. It is usually closed to
prevent gastric reflux.
 DEGLUTITION (SWALLOWING) REGULATION

STIMULUS
Entry of bolus in the oropharyngeal region stimulates receptors of that
region

Afferent impulses pass via the glossopharyngeal n. - IX - fibers

to deglutition center
Center - at the floor of fourth ventricle in
EFFERENT IMPULSE medulla oblongata

Impulses travel through the IX and X nerves

to soft palate, pharynx and esophagus
RESPONSE

Upward movement of soft Upward movement of

palate larynx
Closes respiratory passage and food
Closes nasopharynx enters esophagus and hence into stomach
DEGLUTITION (SWALLOWING) REGULATION
 Swallowing reflex and regulation

 Reflexogenic areas - oral-pharyngeal isthmus

receptors excited by particles of saliva and wet
material
 Afferent fibers of cranial nerves V, IX and X
 Swallowing center - medulla
 Efferent fibers of cranial nerves XII, V, IX and X
 The areas in the medulla and lower pons that
control swallowing or deglutition are collectively
called swallowing or deglutition centers.
 Deglutition apnea is the inhibition of respiration
during act of deglutition in the pharyngeal stage,
lasting for 1 or 2 seconds.
 The centers of respiration and deglutition are
situated close by in the medulla
 The swallowing enter specifically inhibits the
respiratory center of medulla during this time,
halting respiration at any point in its cycle to allow
swallowing to proceed.
 SWALLOWING REGULATION
EXTRINSIC NERVOUS
MECHANISM
Reflex areas
nerf V, IX et X
Center of swallowing in the
MEDULLA
 n. V  mm. masseter
 n. VII  mm. of the mimics
 n. XII  mm. of the tongue
 n. IX  mm. pharyngeals
 n. X  mm. of oesophagus
INTRINSIC NERVOUS
MECHANISM
Auerbach myenteric plexus
 oropharyngeal isthmus stimulated by
particles of saliva and moist ingested
material
 mechanical distension of the
esophagus
 inhibits the respiratory center
 inhibits the masticatory center
 it triggers salivary, tear and
vasomotor reflexes and the
vasomotor reflexes associated
with swallowing a large bowl of
food
 coordinates the peristaltics of
the middle and lower
esophagus and the lower
esophageal sphincter
 1. ESOPHAGUS MOTILITY
•Innervation:
 afferent: sensory feedback to swallowing center
 efferent:
• vagal somatic motor neurons to striated muscle
• vagal visceral motoneurons to smooth muscle, terminating at neurons of myenteric
plexus

Structural and regulatory aspects:

• Upper third of the esophagus:
- circular and longitudinal muscle layers are striated
- innervation via cranial nerve
• Middle third:
- coexistence of skeletal and smooth muscle
- primary innervation from vagus nerve
- nerve input from neurons of myenteric plexus
• Lower third:
- smooth muscle, enteric nerve system (input from vagus nerve to
enteric nerve system)
 Vagal intrinsec fb.

Vagal extrinsec fb. VIP and NO

nicotinic R

Esophageal sphincters
• Upper esophageal sphincter (UES): prevents entry of air
• Lower esophageal sphincter (LES): zone of elevated resting pres. (~ 30 mm Hg),
prevents reflux of corrosive acidic stomach content.
LES tone is regulated by extrinsic and intrinsic nerves, hormones, neuromodulators.
Contraction: vagal (cholinergic ) excitatory nerve input (+) to nicotinic R ( i.e.
atropine insensitive) and sympathetic nerves (-adrenergic R).
Relaxation: primary peristalsis  inhibitory vagal nerve input (-) to circular muscle
of LES (neurotransmitters: VIP and NO) and reduced activity of vagal
excitatory fibers (cholinergic, nicotinic).
 Esophageal pressure profile

l
Intraluminal esophageal pressure profile

Pressure in the body of esophagus is negative,

reflecting intrathoracic pressure

pressure wave
during swallowing

0 mm Hg = ambient pressure