THE DIGESTIVE SYSTEM OF
VERTEBRATES
TOPIC: Motor activity of the
digestive tract

Figure This is a comparison of esophageal muscle
variations
in
birds and mammals. Dashed lines represent striated muscle and solid red
lines represent smooth muscle. The esophagus of some mammals, such as
Perrisodactyls and Lagomorphs, terminates in distinct, muscular
gastroesophageal sphincters. Although others , such as the birds,
primates, felines, canines, suiformes, ruminant, and rodents can
constrict the terminal segment of their esophagus, it lacks the
thickened muscular sphincter. (From Sellers & Stevens 1977)

Figure 6.1
Esophageal pressure events during bovine (A)
deglutition, (B) regurgitation, and (C) eructation.. Numbers represent
1) cervical esophagus 79 cm cranial to cardia, 2) thoracic esophagus 40
cm cranial to cardia, 3) thoracic esophagus 19 cm cranial to cardia,
and 4) pleural cavity. The peristaltic deglutition wave is accompanied
by no changes in respiratory cycle. However, regurgitation is
accompanied by a marked drop in pressure in the pleural cavity and
thoracic esophagus, followed by an antiperistaltic wave of esophageal
contraction. Eructation is recorded as a rise in intrathoracic pressure
of low amplitude and approximately one-second duration, due to
abdominal press. This is followed by a high pressure, short duration
wave of antiperistaltic esophageal contraction, often followed by
a fluid- clearing wave of deglutition. (From Sellers & Stevens
1960) < go
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Figure 6.2 Basal electrical rhythm of the
human stomach.
Slow
waves of partial depolarization of the circular muscle is initiated by
a pacemaker and passes over the distal
half of the stomach. These initiate contractions when accompanied by
spike potentials.
(Stevens 2001) < go
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Figure 6.3 Diagrammatic
sections of bovine reticulorumen
and
omasum. Structures and compartments of major importance are
numbered as follows: 1) cardia, 2) reticulo-omasal orifice, 3)
reticulum, 4) cranial sac of rumen, 5) dorsal sac of rumen, 6)
caudodorsal blind sac, 7) ventral sac of rumen, 8) caudoventral blind
sac, 9) ruminoreticular fold, 10) cranial pillar, 11) right
longitudinal pillar, 12) caudal pillar, 13) dorsal coronary pillar, 14)
ventral coronary pillar, 15 ) omasum, 16) omasal canal, 17) omasal
pillar, 18) omaso-abomasal orifice, 19) omasal lamina (leaf), 20)
abomasum. (From Sellers
& Stevens 1966.) < go
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Figure 6.4 A six-minute
recording of reticuloruminal
cycles and
associated events in the esophagus and omasum. Note the
relationship between the contractions of the rumen and omasal canal,
and the prolonged, acyclic contractions of the omasal body pressure
during primary contraction of the dorsal rumen and omasal canal.
Upper traces show eructation as a biphasic wave occurring during
secondary rumen contractions and the frequent waves of deglutition
associated with the swallowing of saliva. (From Sellers & Stevens
1966.) < go
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Figure 6.5 Movement of
digesta during primary and
secondary
contractions of the reticuloruminal cycles. (From Stevens & Hume
1995.) < go
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Figure 6.6 Movement of
digesta through the bovine
omasum.
Closed arrows show movement of digesta and open arrows show movement of
forestomach walls. Diagrammatic axial section (see Fig. 5.3) shows the
cranial reticulum (1), reticulo-omasal orifice (2), omasal leaf portion
of omasal body (3), omasal canal (4) and cranial abomasum (5). A:
All structures are relaxed during much of the cyclic contraction of the
forestomach. B: During the second reticular contraction, the
reticulo-omasal orifice and omasal canal are pulled ventrally,
producing a negative pressure in the canal and a closing and then
opening of the orifice, which results in aspiration of digesta from the
base of the reticulum. C: Primary contraction of the rumen is
associated with a primary contraction of the reticulo-omasal orifice
and omasal canal, forcing fluid and small digesta particles between the
leaves of the omasal body and into the abomasum. These events are
followed by relaxation of these structures (D), and repeated if the
forestomach undergoes a secondary contraction. E: At intervals
that vary and are unrelated to the cyclic contractions of the
forestomach, a wave of contraction passes over the omasal body,
releasing its contents into the abomasum. (From Stevens & Hume
1995.) < go
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Figure 6.7 Reticuloruminal
cycles during rumination in
cattle.
Numbers 1 and 2 mark first and second reticular contractions seen with
every cycle and the x marks the extra reticular cycle at the
regurgitation stage (R) of rumination. The remasticated bolus is
swallowed (D) just prior to the next reticulorumen cycle. The AP on the
reticular tracing is a registration of abdominal press at the time of
eructation (E). (From Stevens
& Sellers 1968.) < go
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Figure 6.8 Schematic
representation of the llama stomach. The
esophagus (1) enters the first compartment (A), which is partially
divided into cranial (3) and caudal (4) sacs by a pillar of muscle (7),
and separated from the second compartment (B) by a constriction (8).
Both compartments include regions of saccules (5,6,9) containing
cardiac glandular mucosa. A ventricular groove (2) runs along the
lesser curvature of the forestomach between the esophagus and the third
compartment (C). The initial four-fifths of the third compartment is
also lined with cardiac glandular mucosa (11,12). The terminal segment
(13) is lined with proper gastric mucosa and separated from a duodenal
ampulla (D), by a pyloric sphincter (14). (From Vallenas et al. 1971.) < go
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Figure 6.9 Left, lateral, longitudinal
section of the
llama
stomach showing the entrance of the esophagus (A), transverse pillar
(B) between the cranial and caudal sacs of the first compartment and
entrance to the second compartment (C). It also shows the openings to
the glandular saccules in the first compartment. (Modified from
Vallenas et al. 1971.)
< go
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Figure 6.10 Pressure
recording of cyclic contractions of
llama
forestomach. (From Vallenas
& Stevens 1971a.) < go
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Figure 6.11 Cyclic contraction of the
first compartment
of the
llama forestomach. A-D show contractions of pillar and
sacs. Cyclic eversion of caudal sac pouches during the three stages of
contraction is shown in drawings on the right. (From Vallenas &
Stevens 1971a.) < go
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Figure 6.12a Phases of
migrating myoelectric complexes (MMC) during interdigestive (fasting)
and feeding periods in carnivores and humans. This diagram shows
slow waves with no spike potentials during phase I, intermittent spike
potentials during phase II, and consistent spike potentials during
phase III. (From Hendrix 1987.)
< go
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Figure 6.12b Phases of
migrating myoelectric complexes (MMC) during interdigestive (fasting)
and feeding periods in carnivores and humans. This diagram shows
the effects of interdigestive periods on the MCC, which originate
in the stomach and lower esophageal sphincter (LES) and pass through
the small intestine. (From Hendrix
1987.) < go
to CD Chapter 6
Figure
6.12c Phases
of migrating myoelectric complexes (MMC) during
interdigestive (fasting) and feeding periods in carnivores and
humans. This diagram shows the effects of feeding on the
MCC, which originate in the stomach and lower esophageal sphincter
(LES) and pass through the small intestine. Feeding interrupts
the cycle and increases the duration of phase II. (From Hendrix
1987.) < go
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Figure 6.13 Relationship between
the velocity of propagation of myoelectric complexes and the length of
the small intestine. The hatched area shows the 95% confidence
limits. The median daily number of jejunal complexes is high in
ruminants and low in carnivores, because of the obliterating effect of
feeding in the latter species. (From Ruckebusch 1981.) < go
to CD Chapter 6
Figure 6.14 Relationship between digesta
flow and electrical slow waves and migrating spike bursts in the cat
colon. Slow waves (SW) appear to originate from a pacemaker midway
along the colon, spread toward the cecum, and tend to produce digesta
flow in the same direction. Migrating spike bursts (MSB) begin at a
variable position in the proximal colon and migrate toward the rectum.
These are accompanied by contractions, which tend to move digesta in
that direction. (From Christensen
et al. 1974) < go
to CD Chapter 6
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