The stomach can be divided into three anatomic (A)

and two functional regions (B)

A B
Fundus
Pylorus

Antrum Gastric pump Gastric reservoir

Corpus Phasic contractions Tonic contractions

Ehrlein Figure 1
 The relaxation of the gastric reservoir is mainly regulated
by reflexes. Three kinds of relaxation can be differentiated:
the receptive, adaptive and feedback-relaxation

1. Receptive
Mechanical relaxation Vagus
stimuli in the centre
pharynx Inhibitory
vagal fibre
3. Feedback 2. Adap tive (NANC-inhibition)
relaxation relax ation ACH
CCK NO + VIP et al.
Tension Relaxation of
Nutrients Nutrients
receptors gastric reservoir
Distension

Ehrlein Figure 2
The transport of digesta from the gastric reservoir
into the antral pump is caused by two mechanisms:
tonic contractions and peristaltic waves in the region
of the gastric corpus
Tonic
contraction

Pylorus

Accumulation Peristaltic wave

Proximal of chyme (Pump of the reservoir)
antrum
Backflow from antrum and flow from reservoir

Ehrlein Figure 3
 The function of the gastric pump can be differentiated into
three phases: A: phase of propulsion, B: phase of emptying,
C: phase of retropulsion and grinding
Phases A Phase of propulsion
ABC Contraction of proximal antrum (PA)
Pylorus

Proximal Propulsion of chyme into relaxing

antrum terminal antrum
+ duodenal contraction
PA
Middle B Phase of emptying
antrum
Contraction of middle antrum (MA)
Terminal
antrum MA Transpyloric and retrograde flow
closed + duodenal relaxation
Pylorus
open TA
C Phase of retropulsion
Contraction of terminal antrum (TA)
Duodenum
Jet-like back-flow with grinding
10 sec + duodenal contraction

Ehrlein Figure 4
 Liquids and small particles leave the stomach
more rapidly than large particles.
This discrimination is called „sieving function“
Phase of propulsion Phase of emptying Phase of retropulsion

Antrum

Bulge

Rapid flow of liquids with Emptying of liquids with Retropulsion of large

suspended small particles small particles whereas particles and clearing
and delayed flow of large large particles are retained of the terminal antrum
particles towards pylorus in the buldge of the terminal
antrum
Ehrlein Figure 5
 Grinding of solid particles is caused by a forceful
jet-like retropulsion through the small orifice of the
terminal antral contraction

Onset Late phase

of terminal antral of terminal antral
contraction contraction

Pylorus closing Pylorus closed

Ehrlein Figure 6
 Antro-duodenal co-ordination: Contractions of the proximal
duodenum cease during the phases of gastric emptying..
Phases of gastric emptying

Middle antrum

Terminal
antrum Antral
closed waves
Pylorus
open
9.9 3.5 6.6 3.5 9.9 sec

Proximal
duodenum 1 2 3 1 2 3 1 2 3 4 1

Lacking duodenal contractions

0 5 10 15 20 25 30 35 sec
Because of different frequencies between antral and duodenal contractions,
the duodenum can contract three to four times during an antral wave
Ehrlein
Figure 7
 Several factors of gastric and duodenal motility
co-operate and modulate gastric emptying:
A. Rapid emptying B. Delayed emptying

Pylorus
1a 6a
4 3 9 8

10 6b
5 1b
2 7

A. Rapid emptying is caused by tonic contractions of the reservoir (1a), deep

peristaltic waves along the gastric body (1b), deep constrictions of the antral
waves (2), a wide opening of the pylorus (3), a duodenal receptive relaxation (4)
and peristaltic duodenal contractions (5).
B. Delayed emptying due to feedback inhibition is caused by a prolonged
relaxation of the reservoir (6a), shallow peristaltic waves along the gastric body
( 6b), shallow antral waves (7), a small pyloric opening (8), a lacking duodenal
relaxation (9) and segmenting duodenal contractions (10).
Ehrlein Figure 8
 Balance between gastric reservoir and antral pump

Gastro-gastric reflexes
Enhanced and prolonged
relaxation of reservoir

Inhibitory
reflex Distension
Disten-
sion

Antral pump
switched on Excitatory
and intensified reflex

Ehrlein Figure 9
 Pyloric activity is modulated by antral inhibitory
and duodenal excitatory reflexes

Descending
inhibitory reflex
causing
pyloric relaxation

Ascending
excitatory reflex
causing
pyloric contractions
and increasing
pyloric tone
Contraction of
Duodenal stimuli middle antrum

Ehrlein Figure 10
 An additional function of the pyloric sphincter
is to prevent duodeno-gastric reflux

Antrum Pyloric closure

closed

Pylorus
open
Inhibition
Duod. bulb

Duodenum Stimulation

0.5 ml oleic acid + bile

into duodenum
Duodenal stimuli like oleic acid inhibit antral contractions, evoke
duodenal contractions, increase pyloric tone and elicit frequent pyloric
contractions
Ehrlein Figure 11
Solids and liquids of the gastric chyme are emptied with
different velocities.
Lag phase

100
Solids
Gastric volume (%)
80
Viscous
60 content

40
Liquid
20 content

0
0 20 40 60 80 100 120
Time (min)

Emptying of liquids is exponential, emptying of large solid particles only

begins after sufficient grinding (lag phase). Afterwards the viscous chyme
is mainly emptied in a linear fashion
Ehrlein Figure 12
 Nutrients in the gut activate a feedback control and
modulate gastric and duodenal motility

Non-caloric meal Nutrient meal

Feedback control causes
Reduced force of
Antrum antral contractions

closed Reduced
Pylorus pyloric opening

open
Reduced
Duodenal peristaltic waves
bulb
Enhanced
Middle
segmenting activity
Duodenum

Gastrointestinal motor patterns after a non-caloric and a nutrient meal

Ehrlein Figure 13
 The feedback regulation of gastric emptying is performed
by entero-gastric reflexes and release of intestinal hormones
Vagal
oric t center
ens ren +
Nutrients S ffe ers _
a ib + Inhibitory
Long chain fatty acids f
Amino acids vagal fibers
Dipeptids Stimulating cholinergic
ACH
Glucose vagal fibers
Osmolality NO, VIP et al.
Hydrochloric acid CCK
Enhanced
ACH
relaxation
and
Reduced opening storage
of pyloric sphincter Backflow
Reduced
contraction

It causes enhanced relaxation of the gastric reservoir, inhibition of the antral

pump, and reduced opening of the pyloric sphincter.
Ehrlein Figure 14
 Contractile patterns of the small intestine
Peristaltic Stationary Clusters
waves contractions of contractions
oral

aboral
1 minute 1 minute 1 minute

The most frequent patterns are peristaltic waves (dashed lines), stationary
contractions (arrows), and clusters of contractions, which occur either
stationary at an intestinal segment or slowly migrate aborally
Ehrlein Figure 15
 Phase III of the interdigestive motility
designated as ”migrating motor complex” (MMC)

Jejunal phase III (MMC)

oral 1 minute

aboral Aboral migration Velocity of the

of phase III peristaltic waves

Rectangles: strain gauge transducers, Data of dog.

Ehrlein Figure 16
Pathological contractile patterns of the proximal intestine
Antiperistaltic waves Aboral giant contractions
oral oral

0,2 Newton

Duodenum
Jejunum

aboral aboral
1 minute 1 minute

Alternating peristaltic (blue arrows) and antiperistaltic waves (red arrows).

Giant contractions sometimes originate as a cluster.
Ehrlein Figure 17
 Different kinds of contractile patterns
are caused by different kinds of excitation
Stationary Single
segmenting contractions peristaltic waves
Excitation Excitation

PP 1 1

PP
2
2

3 3
Time
course

Stationary segmenting contractions Single peristaltic waves are produced by

are produced by brief excitation of a short excitations of a long intestinal segment
short intestinal segment

Ehrlein
1, 2, 3 successive pacesetter potentials (PP) Figure 18
 Origin of clustered contractions
Stationary cluster Migrating cluster

1 1

2 2

3 3

Time
course
Stationary excitation Aboral migrating excitation

Clustered contractions are produced by a long lasting excitation of a short

intestinal segment. The cluster is stationary when the excitation remains at
the same segment. When the excitation slowly moves aborally the cluster of
contractions migrates along the intestine.
1, 2, 3 successive pacesetter potentials (PP)
Ehrlein Figure 19
 Central and peripheral control of contractile patterns

Vagal Vago-vagal reflexes

centre

Interneurons
Integrating circuits
Intestinal Contractile
Sensory neurons Motorneurons
wall patterns
Program circuits
Enteric nervous system
Peptide (CCK) Receptors
Intestinal
lumenl Glucose - Osmolality
Long chain fatty acids
Amino acids

Luminal stimuli elicit vago-vagal reflexes which activate integrating and

program circuits of the enteric nervous system. These activate specific
motorneurones responsible for specific contractile patterns.
Ehrlein Figure 20
Postprandial contractile patterns of the small intestine

oral

0,2 Newton

aboral

They are composed of stationary segmenting contractions (green arrows),

stationary and migrating clusters of contractions (red horizontal lines)
and single short peristaltic waves (dotted lines).
Ehrlein Figure 21
 The interdigestive motility consists of three phases
Interdigestive Cycles
III I Phases II III

Phase III Phase I Phase II

Stomach Contraction
Phase III of reservoir
Pylorus Sporadic
Forceful
Motor peristaltic waves
Duodenum peristaltic
Accumulation quiescence
waves Segmenting
of residues of stomach
contractions
of chyme Ab and duodenum
and single
Jejunum mi oral peristaltic waves
gra
Phase II ti o
n Motor
quiescence
Ileum Phase I
Phase III

The phase III of the migrating motor complex originates simultaneously

at the stomach and duodenum and migrates within 90 to 120 minutes
along the small intestine (dog)
Ehrlein Figure 22
Gastric phase III consisting of 1 - 3 forceful contractions
of the gastric reservoir and lumen occluding peristaltic
waves occurring at intervals of 2-3 min
Gastric phases III

Middle Antrum
0 mm
Pyloric diameter
6 mm
Duodenal bulb

Duodenum

1 min

A P Stomach is cleaned of residues

P P of chyme and secretions.

The antral waves are associated with a wide opening of the pylorus and
inhibition of duodenal contractions followed by duodenal peristaltic waves
occurring at maximal frequency.
Ehrlein Figure 23
 Phase III (MMC) of the small intestine
Intestinal phase III
Tim
e (abo
ut 2
oral 0 se
c)

Successsive
peristaltic waves

Chyme
Slow aboral
migration of
phase III
aboral

The peristaltic waves clean the intestinal segment from chyme which
accumulates aborally. Because the successive waves start and end
further aborally the phase III slowly migrates distally
Ehrlein Figure 24
 Ingestion of a meal suppresses the interdigestive
motility and induces a fed motor pattern

Phase III Fed motor pattern

Meal
Antrum

closed
Pylorus
open

Duodenum

5 min

Postprandial motility is characterised by a lower amplitude of the antral

waves occurring at maximal frequency, rhythmic pyloric opening and
closure and co-ordinated duodenal contractions occurring in sequence
with the antral waves

Ehrlein Figure 25
 Contractile patterns of the large intestine
Shallow peristaltic waves of caecum and colon

A backflow low propulsion

Shallow peristaltic waves at haustrated colon

B
small aboral flow

Colonic segmenting contractions migrating aborally

slow aboral propulsion
C
aboral migration

C: A special feature of the large intestine are multiple segmenting

contractions of long duration migrating aborally. They divide digesta
into boli pushing them slowly aborally. The motility tracings show a rise
of the baseline superimposed by phasic contractions
Ehrlein Figure 26
 Motility of the large intestine in pig
Ileum - Caecum - Colon
A Caecum B
C1 J1
Giant contractions
J2
C2
C1

C3 Colonic wave
Co1
1 min
Co2
Co3 Colon
Co3
Co2
Ileum 1 min
J1 A: Haustral movements of the caecum result in
Co1 J2 clustered contractions.
Caecum B: The ileum is emptied by giant contractions. They
occur either isolated or in co-ordination with
Distal C1 C2 C3 peristaltic waves of the caecum and colon.
colon Additional colonic waves originate at the beginning
Ehrlein of the colonic coil. Figure 27
 Motility of caecum and colon in sheep.
Caecum Colon
Spiral colon Giant
C4 contraction
Peristaltic C1
wave
C3
SC1
Co1
C2
SC2
C1 Co2
SC3
Co1 Co3
Ileum
Co1 C2 C3 C4 Co2 Co4
C1
Colon 1 min
Caecum 1 min
Co2 Co2 Co3
Caecal motility is characterised by peristaltic and
SC1 antiperistaltic waves. In the colon peristaltic waves
SC2
and giant contractions are the dominant feature. In
Co4 the spiral colon prolonged segmenting contractions
SC3
Spiral colon divide digesta into boli and push them distally.
Ehrlein Figure 28
 Motor patterns of the large intestine in rabbits
Colon Caecum
Giant contractions
J1
Co1
C1
Co2

C2
Co3
1 min
Co3 Co2 C3
J1
Ileum Co1
C1 C4
Colon

C2 C5
Caecum
1 min
C3 C4 C5
Caecal motility is characterised by peristaltic and antiperistaltic waves. Migrating
segmenting contractions are the dominant feature of the single haustrated colon.
Ehrlein Figure 29
 Colonic motor complexes (CMC’s) of the canine colon
A B
Colonic motor complex (CMC)
oral

Phasic
contractions

15 min 5 min

aboral

A: Slow paper speed. The CMC’s occur at all parts of the colon at intervals
of 20-30 min.
B: High paper speed. The CMC’s consist of a rise of the baseline super-
imposed of phasic contractions. The onset of the CMC‘s obviously differs
along the colon (indicated by lines).
Ehrlein Figure 30
 Retrograde giant contraction followed by vomiting
Retrograde
giant contraction Vomiting

Distal duod.
Prox. duod.
Bulbus
closed
Pylorus (P)
open
Antrum
1 2 34 5 1 min
Duodenum
P
P
P
Jejunum Stomach
1 2 3 4 5
(1) Normal segmenting contractions of the proximal jejunum
(2) Start of a retrograde giant contraction in proximal jejunum;
(3) Retropelled digesta reach the duodenum and (4) are forced across the
widely opened pylorus into the antrum; (5) The giant contraction proceeds
to the antrum, the chyme accumulates in the gastric reservoir.
Ehrlein Figure 31