The Importance of Being Asymmetric

The Physiology of Digesta Propulsion

on Earth and in Space

C.P. ARUN
Institute of Urology, University College London, London, United Kingdom

ABSTRACT: In the embryo, the mammalian gastrointestinal tract (GIT) is a

midline structure, but later becomes strikingly asymmetric. Such asymmetry
is in contrast to other organ systems that are essentially bilaterally symmetric.
Making a departure from the traditional straight tube model of the bowel, we
offer a more realistic model—a kinked collapsible conduit disposed as a con-
strained kinematic chain. We examine evidence for the importance of its asym-
metry to the unidirectional flow of digesta. A number of factors cooperate to
ensure a unidirectional flow. The anatomical factors must include (1) the shape
of the abdomen, the inverted truncated cone allowing several degrees of free-
dom of movement of the bowel allowing folds and twists; (2) the location of the
liver and the stomach under the diaphragm, providing for efficient force trans-
mission from the diaphragm (especially in the distended state in the case of the
stomach); (3) cranio–caudal gradients in length of the small bowel mesentery
and diameter of the bowel lumen. The physiological factors include (1) a delib-
erate conversion of ingested food into a non-Newtonian fluid with increasing
viscosity, (2) nonlinearity of the tube law, (3) respiratory excursions of the dia-
phragm, and (4) a “Law of the Intestine”. In microgravity, the bowel can be
expected to float and exhibit loss of polarity of propulsion of digesta, but this
can be compensated for by exercise (indirectly by increasing diaphragmatic
movement). The asymmetry of the GIT is an ingenious device to ensure a uni-
directional movement of digesta.
KEYWORDS: law of the intestine; peristalsis; digesta; low Reynolds number;
collapsible tube; tube law

INTRODUCTION

At the time of writing, human travel to space is a reality and the first space tourist
has safely returned to Earth. Thanks to the International Space Station (ISS), periods
of stay in excess of a year have been achieved. The fact that humans have conquered
such a harsh and forbidding environment as space is a testament not just to the
strength of the human spirit but also to the endurance and adaptability of our organ
systems. One such system whose value is often unappreciated is the gastrointestinal
tract (GIT).

Address for correspondence: C.P. Arun, Department of Urology, Colchester General Hos-
pital, Colchester CO4 5JL, England, U.K. Voice: 44-1206-742450; fax: 44-1206-742220.
arunpeter@yahoo.com

Ann. N.Y. Acad. Sci. 1027: 74–84 (2004). ©2004 New York Academy of Sciences.
doi: 10.1196/annals.1324.008
ARUN: PHYSIOLOGY OF DIGESTA PROPULSION 75

Mammals, like other animal life forms, need to extract nutrients from ingested
food. This ingested food then undergoes digestion, involving mechanical and chem-
ical processing to facilitate the later stage of nutrient absorption. As with other
organs, the principle of form tailor-made to function is in evidence in the gastrointes-
tinal tract. The type of food ingested and dietary habit involved determines the kind
of movement required in its processing. For example, in ruminants, undigested food
must be brought back into the mouth to be chewed with corresponding implications
for design and function.
In humans, the aboral movement of digesta is important to nutrition (flow in the
reverse direction can in certain conditions, be detrimental to health). This unidirec-
tional movement of content appears to be achieved by a combination of factors
involving the anatomy, physiology, soft tissue, and fluid mechanics of the GIT, its
content, and neighboring structures, all aided no doubt by gravity. Given the circum-
stances in which it occurs, the predominantly aboral movement of digesta appears to
be no small feat even in normal gravity. From classical physics, we know that a net
movement of a particle requires an asymmetric net force. In most systems around us,
we encounter asymmetry in the direction of the force imparted directly to the parti-
cle. The strategy of the gastrointestinal tract however, appears to involve asymmetry
in its disposition, such that forces applied from various directions assist wall move-
ment to produce a net aboral movement of content. An examination of how the asym-
metry in the lie of the GIT is important to its function would require the application
of skills from several fields of science: it is not surprising, therefore, that this issue
has not been addressed by previous workers. A brief outline of the relevant aspects
of anatomy of the GIT follows.
In humans, ingested food is transferred from the mouth to the esophagus. The
esophagus is a conduit fixed at the upper end, the pharynx, and beyond, the cardiac
sphincter, is continuous with the stomach. From the esophagus, boluses of food
reach the stomach. Inside the abdomen, the presence or absence of a mesentery (a
double fold of peritoneum that carries blood vessels and nerves) determines the
degree of mobility of the viscus. The stomach is a strong muscular receptacle con-
tinuous with the esophagus above and the small intestine below. In turn, the small
intestine (see FIGURE 1) is a 4–6-meter long tube separated for purposes of descrip-
tion into the fixed duodenum (except for the last inch or so) and the free jejunum and
ileum. The duodenum is attached proximally to the stomach and is continuous with
the jejunum and ileum. The jejunum and ileum have a 15-centimeter line of attach-
ment by the mesentery to the posterior body wall. Beyond the ileo-cecal valve, the
small intestine is continuous with the large intestine. The large intestine (see
FIGURE 2) has, in succession, a fixed ascending part, called the ascending colon; a
free part, the transverse colon with a mesentery (the transverse mesocolon); the fixed
descending colon; and a free part, the sigmoid colon, that is in continuity with the
fixed rectum and anal canal. The physiology of the processes involved in digestion
is outlined next.
In the mouth, food is mixed with saliva, chewed, and digestion of carbohydrates
is initiated. In the stomach, the majority of the water and all of the alcohol is
absorbed, converting the ingested boluses into chyme. In the small intestine, digesta
undergoes further chemical processing and more absorption of nutrient occurs. In the
large intestine, water is recovered and the unwanted material is prepared for egestion.
76 ANNALS NEW YORK ACADEMY OF SCIENCES

FIGURE 1. The lie of the small intestine. The small bowel is an asymmetrically
arranged collapsible conduit with a small fixed part, the duodenum (held down to the pos-
terior abdominal wall) and a long free part (invested by a mesentery that allows movement).
Its small line of attachment and the considerably longer free margin allows it to kink (twist
and fold) providing for mobile physiologic sphincters.

FIGURE 2. The lie of the large intestine. The large bowel is another asymmetrically
arranged collapsible conduit with alternating fixed (held down to the posterior abdominal
wall) and free parts (invested by a mesentery that allows movement). The junctions of the
fixed and free parts must function as fixed physiologic sphincters.
ARUN: PHYSIOLOGY OF DIGESTA PROPULSION 77

Since there is redundancy in both the small and large bowel (much more so in the
former) a much larger surface area is available for absorptive processes than a straight
tube would allow. Also, since it is a collapsible tube, the folds produce a series of cas-
cades with digesta undergoing “piecemeal digestion” in this “cascade reactor” as it
moves along.1
Generally, the processes of digestion and absorption are slow and the digesta has
to spend time in the lumen of the gut (termed residence time) in order to allow these
processes to occur. Nature’s solution to this need for long residence times appears to
involve altering the consistency of ingested food to a fluid that is intrinsically resis-
tant to quick movement. Common examples from every day life, of fluids that resist
rapid motion, termed non-Newtonian fluids, include treacle and gum. In contrast,
Newtonian fluids are obedient to Newton’s laws of motion and are best exemplified
by water (for the non-specialist, a useful aide memoire: Newtonian fluids fall with a
splash, Non-Newtonian fluids fall with a splat). In the stomach, a nearly Newtonian
fluid like milk undergoes processing involving precipitation of the protein and
absorption of much of the water, and ends up a non-Newtonian fluid. It must be
apparent from the above that digesta that leaves the stomach is essentially a complex
(as opposed to simple Newtonian) fluid.
The world we inhabit is vastly different from the world of such complex fluid
dynamics and to the majority of us, an unfamiliar one. Although the aboral flow of
digesta is natural and normal it is by no means simple. The mechanics of fluid flow
can be a trap for the unwary and the mere application of common sense can be
unhelpful. Certain phenomena can be frankly counterintuitive (e.g., the Brazil nut
phenomenon—after shaking a mixture of nuts and corn flakes, the heavier nuts are
found at the top). An easy experiment to try at home is to compare the ease with
which a candle flame can be blown out with efforts to try extinguishing the flame by
sucking air in. A brief introduction to the relevant aspects of fluid dynamics of rele-
vance to the propulsion of digesta is, therefore, in order here.
The behavior of fluids in flow or of bodies that are moving in fluid can be
described using a dimensionless parameter termed the Reynolds number (Re).
Named for the nineteenth century English engineer Osborne Reynolds, this number
is given by Re = ρud/µ, where ρ is density, u velocity, µ viscosity of the fluid, and d
the dimension of the channel or body. When the Reynolds number is high, the flow
is interpreted to be inertia driven; when low, to be dominated by viscous forces. To
illustrate, from the above equation, the flight of a rocket would involve very high
Reynolds numbers: the viscosity and density of the air are small, the body dimen-
sions large, and the velocity extremely high. In contrast, digesta has a high viscosity
and density, moves in a channel of small dimension, and has a very small velocity;
a very low Reynolds number, therefore, governs such flow. Such very low Reynolds
number regimes are not in the least uncommon; microorganisms (they far outnumber
mammals and insects on earth) inhabit such an environment.
As is evident from the above, digesta in transit in the gut involve the strange
world of what physicists would term low Reynolds number (LRN) flow. In such a
regime, in the memorable words of Purcell,2 “what you are doing at the moment is
entirely determined by the forces that are exerted on you at that moment, and by
nothing in the past.” In everyday terms, this implies that there is no such thing as low
Reynolds number football—a “kicked” ball would stop at the foot rather than move
78 ANNALS NEW YORK ACADEMY OF SCIENCES

independently toward the goal. Therefore, for net movement to be successful under
such conditions, the tactics employed have to be different. Attempts to move by a to-
and-fro undulatory movement of the body employed by most fish or a to-and-fro tail
waving laboratory based mechanical fish (see FIGURE 3) are unhelpful here. The pio-
neering work of Berg and Anderson3 pointed out that bacteria swim by a peculiar
helical action of their flagella. It was later shown that in fact, such asymmetric move-
ments are a must in order to move in low Reynolds number regimes.2 This is
because, under such conditions, solutions of the fluid flow equations are unique and
reversible.4 Purcell2 further proposed that for movement constrained to one plane,
at the least a two-link mechanism would be necessary for locomotion at low Rey-
nolds number. Following up on Purcell’s work, more recently, Becker et al.5 reiter-
ated the importance of local anisotropy to swimming motions at low Reynolds
number. Reversibility of fluid solutions in LRN regimes explains why a single-link
(FIG. 3) mechanical fish can happily swim in water but cannot make progress in corn
syrup.4 Thus, mere to-and-fro movement, the mechanism employed by scallops
swimming in water, is inadequate for treacle traversal. It is due to such consider-
ations that attempts to explain unidirectional digesta propulsion by bowel wall
movement alone come unstuck. Merely extrapolating from our understanding of
Newtonian fluid behavior is inadequate to explain the mechanism of digesta propul-
sion in the gut.
Humans, like other mammals, have bilateral symmetry and the majority of body
systems are more or less symmetric. For example, the left and right hand sides of the
face, hands, and feet are normally almost perfectly symmetric. The one organ system
that makes a gross departure from such symmetry happens to be the gastrointestinal
tract (GIT). Since, in nature, form is firmly wedded to function, it is tempting to
speculate that such asymmetry is indeed important to the function of the GIT. With
an ingenious but curious layout, the biomechanics and kinematics of the bowel, cou-
pled with LRN fluid dynamics constitute a problem of no mean magnitude for
researchers. It is not surprising therefore that previous workers have dealt with very
elementary models to try to explain gut propulsion. One such model termed the law

FIGURE 3. Fishy physics—the strange world of low Reynolds number flows. (A) The
mechanical laboratory swimming device “fish” can move in water by a to-and-fro move-
ment of its “tail”. In non-Newtonian fluids like corn syrup or treacle, however, locomotion
is not possible. (B) Microscopic beings like bacteria are able to achieve locomotion by
using helical movements of appendages, such as flagella, because such movements are non-
time-reversible.
ARUN: PHYSIOLOGY OF DIGESTA PROPULSION 79

of the intestine is a favorite with physiologists. In this paper, we examine evidence

for GIT asymmetry being of importance to the unidirectional flow of digesta. Since
this work is intended for a general scientific audience, various terms have been
explained and excessive use of formulæ and equations avoided.

REVIEW OF LITERATURE

The movement of the intestines and the motion of digesta has been of interest to
scientists for centuries. The model of Bayliss and Starling6—the Law of the Intes-
tine—that has held sway for more than a century, presupposed a straight collapsible
tube with waves of contraction that are directed distally. Interestingly, their experi-
ments also included observations on solids placed in the bowel lumen—content that
is quite unlike what the bowel is accustomed to conveying. Later work, both exper-
imental and theoretical, has also employed a straight tube to model the bowel.7–9
Such straight tube models are no doubt of relevance to propulsion in the esophagus
(food bolus—semi-solid) and ureter (urine—Newtonian content); for the non-New-
tonian fluid digesta traversing the loops of the small and large intestine, however,
applicability of the law of the intestine is doubtful.
More recently, doubts have been expressed if the law of the intestine is valid—
Hodgkiss,10 working on the avian small intestine model, and Spencer et al.,11 work-
ing on the guinea pig ileum, presented evidence against such a law. Work on human
tissue is not available as this paper to goes to the press, but observations on bowel
exposed at open operation reveals the bowel to be a writhing mass of tubes and no
preferred direction of wall contraction is apparent. Clearly, a travelling wave of con-
traction that does not continuously and consistently occlude the bowel lumen cannot
be credited with producing a unidirectional movement of digesta. Since waves of
contraction in the bowel are only intermittent, some form of valve mechanism must
exist to discourage a backward of flow of content between loops of bowel in the
interval periods.
Other works of relevance include the paper by Amidon et al.,12 which recognizes
the importance of gravity for the aboral propulsion of digesta proposing that buoy-
ancy, drag, and gravity are crucial factors in flow through the intestine. The mono-
graph of Stevens13 reviews the comparative anatomy and physiology of mammals
and includes a study of the lengths of the various parts of the digestive tract among
various species. The arrangement of the gut itself and its possible implications was
not studied in these works.
The first model of the small bowel as a constrained kinematic chain was that of
Arun1 who pointed out that the length of the mesentery increases from the jejunum
to the ileum and imposes a gradient that is antero-posterior in a supine position and
directed inferiorly in the erect posture (i.e., cranio-caudally in the erect and supine
positions). In the present work, we extend this model to help understand how bowel
action occurs on Earth and how it continues to be possible in space.
80 ANNALS NEW YORK ACADEMY OF SCIENCES

MATERIAL AND METHODS

The literature on gastrointestinal physiology relating to propulsion of digesta was

reviewed. The human GIT was examined from an anatomic, physiologic, and fluid
mechanics standpoint to look for mechanisms that are likely to assist the net unidi-
rectional propulsion of digesta.

RESULTS

By analyzing the GIT as a fluid propulsion system, we obtain interesting findings

and a coherent theory of digesta propulsion is made possible. Unlike machines
designed by humans, the functions of various biological organ systems are coupled.
Such coupling is prominent in the structure and function of the GIT.
The alimentary tract is a system of collapsible conduits in continuity that have
free and fixed portions. The distal small intestine is a kinematic chain held down at
the duodenal flexure and the ileo-cecal junction, packed into a small conical cavity
with play allowed by the length of its mesentery (see FIGURE 4) is kinked into form-
ing loops. At the kinks, the bowel can not only be folded upon itself, but also, due to
the significant anteroposterior diameter of the peritoneal cavity, it is capable of twist-
ing (see FIGURE 5). The ends of loops must restrict the flow of content until such time

FIGURE 4. Gradient imposed by the small intestinal mesentery and cascading flow in
the small intestine. Note the increasing length of the mesentery the cranio-caudal direction.
The direction of the gravity gradient is marked at the center of the diagram. The anterior
(A) and posterior (P) parts of the diagram are marked in circles. In the upright position (A)
and the prone position (B), the gradient offered by the length of the length of the mesentery
is retained. The direction of cascading flow of digesta is marked in (A). The increasing
diameter and thickness of bowel wall is depicted as well.
ARUN: PHYSIOLOGY OF DIGESTA PROPULSION 81

FIGURE 5. Digestion by numbers: the loopy dynamics of propulsion of digesta in the

small intestine. In the schematic diagram, the digesta moves from the duodeno–jejunal junc-
tion at the start of loop 1, loop to loop, and at the end of loop 6; that is, beyond the ileo–cecal
junction, joins the large intestine. The mesentery is depicted as a dotted line. Loops 1, 2, and
3 involve an anticlockwise spiral, whereas loops 4, 5, and 6 form a clockwise spiral.

when digestion and absorption is satisfactory. The alternation of fixed and free por-
tions allows the junctions between the fixed and free parts to function as physiologic
sphincters. The formation of loops is important to ensure an optimal residence time
for digestive processes. When work on the portion of digesta is complete, a local
reflex is initiated and wall movement provides a propulsive force for movement into
the next bowel loop. Thus, due to the nonlinearity of the tube law for collapsible
tubes,14 the ends of bowel loops function as mobile valves.
The motive force for the movement of digesta is no doubt provided in large part
by the contraction of the bowel wall. The mechanisms for implementing polarity
though, are several. These may be separated into anatomic and physiologic features.
Anatomic features of the GIT imposing polarity must include:
1. The shape of the abdomen a three-dimensional (3D) structure—an inverted trun-
cated cone. The muscular abdomino-thoracic diaphragm forms the base; sup-
porting the smaller lower end is the pelvic diaphragm. The mobility afforded by
a 3D enclosure allows movement of the bowel with several degrees of freedom
and content cascading from one loop to the next. This allows the junctions
between free and fixed parts of the bowel to act as fixed physiologic “sphinc-
ters”. As mentioned above, in the small intestine, kinks between successive
bowel loops may involve not only a fold but also a twist, allowing for several
such mobile physiologic sphincters (see FIG. 3).
2. The stomach is located directly under the diaphragm on the left side and corre-
spondingly, the liver lies under the right hemi-diaphragm. Such an arrangement
allows transmission of forces from the diaphragm to the intestines. The effi-
ciency of such force transmission, of course, increases when, after a meal, the
stomach is full of food.
3. The small intestinal mesentery increases in length from above downwards. This
tends to cause smaller loops to form (and be constrained to remain) above
(FIG. 5). This polarity persists, both in the erect as well as the prone position, but
is lost in the supine position or in a microgravity environment. It is interesting to
82 ANNALS NEW YORK ACADEMY OF SCIENCES

note that researchers simulate a microgravity environment in terrestrial laborato-

ries using the supine posture with a slight head down tilt.
4. The lumen of the bowel increases from above downwards. From the Young–
Laplace relation (P = T/R, where P is the pressure, T the wall tension or hoop
stress, and R the radius), and from force–tension relations for muscle, such an
arrangement imposes a gradient in the strength of contraction possible.
The physiologic factors imposing polarity must include:
1. Gradients in viscosity of the content. The viscosity is an important determinant
of the Reynolds number and the liability of a flow to reverse. The stomach
appears to deliberately produce a non-Newtonian fluid from ingested food by
precipitating protein and absorbing much of the water and alcohol content from
ingested food. From the stomach to the colon, the viscosity increases dispropor-
tionate to the density and this inevitably imposes an increasing gradient.
2. The nonlinearity of the tube law for collapsible tubes that allows kinked por-
tions (whether temporary or anatomical) and collapsed portions to function as
sphincters.
3. Respiratory excursions of the diaphragm—the importance of the diaphragm to
intra-abdominal physiology was recognized as early as the seventeenth century
by William Harvey who called it “the engine of the abdomen”.
4. Law of the intestine—aboral propagation of circular and longitudinal contractions.

DISCUSSION

The discovery of the circulation of blood by William Harvey (1578–1657) was a

watershed event in physiology and the medical sciences. It placed the science of
blood flow on a sound scientific basis and for coming up with such a radical idea,
rewarded its discoverer, at least initially, with much notoriety. The heart and the cir-
culation inspire much awe even in the mind of the layperson. Such interest is no
doubt, justified: the flow of blood at the time of exsanguinations can be terrifying and
if unchecked, the effect on life, drastic. Although the importance of the circulatory
system cannot be disputed, the mechanism of blood flow is fairly simple when com-
pared with that of the digesta.
In health, the propulsion of digesta is a languid affair and even if food is being
enjoyed at the table, the mechanism of what is involved is not likely to be pleasant
dinner time conversation. Given its complexity, it is not surprising that unravelling
the mechanisms of digesta propulsion has taken much longer. Just as bacteria and
spermatozoa employ asymmetric non-time-reversible movements in order to swim,
the strategy of the gastrointestinal tract appears to be to employ asymmetry in its lie
so as to ensure unidirectional movement of digesta. It is now possible to explain why,
in the stage of the embryo, the gut that starts life as a midline structure eventually
becomes starkly asymmetric. It appears that this asymmetry is a deliberate and inge-
nious device to ensure a unidirectional movement of digesta.
It appears that more than one feature is responsible for implementing polarity in
digesta flow. An aboral gradient is implemented, not only by gravity, but also by the
shape of the peritoneal cavity, its pressure–tension relations, and the influence of the
ARUN: PHYSIOLOGY OF DIGESTA PROPULSION 83

diaphragm and other abdominal muscles. This arrangement emphasizes the coupling
of one organ function to another that is a common theme in physiology. We formally
recognize that propulsion in collapsible tubes in physiology depends not only on
intrinsic wall contraction, termed peristalsis, but also by the self-explanatory neigh-
boring organ assistance (covenant of NOA15) phenomenon. In the case of the bowel,
NOA is provided by the movement of the diaphragm and the abdominal muscles. It
appears that we are rediscovering the contribution of the diaphragm to bowel func-
tion that was originally recognized by William Harvey when he credited the dia-
phragm with being the engine of the abdomen.
The present model involves several subsystems working in collaboration to effect
a unidirectional flow of digesta. It involves a multihierarchical multidimensional
system. To validate this model, were it ever to be attempted, some very complicated
experiments would have to be devised. Barium contrast studies of the small intestine
in a microgravity setting would be interesting but are unavailable in the indexed lit-
erature. It is not unreasonable to assume that in microgravity the bowel floats freely
inside the peritoneal cavity. This would to some extent upset the aboral gradient that
the length of the mesentery would normally impose. Whereas the advice for healthy
eating is to have a high roughage diet to avoid constipation, payload constraints dic-
tate that astronauts consume a low residue diet. Since the bowel wall requires disten-
sion in order to propel content, a low residue can be expected to be passed onward
with difficulty. Constipation is a well-known problem among astronauts and GI tran-
sit time is known to be reduced. Vigorous exercise that is required of astronauts in
order to avoid osteoporosis of the bones can indirectly help the propulsion of digesta
by increasing diaphragmatic movements and thereby bowel action. During periods
of prolonged residence or flight in space, problems of ileus may occur and the man-
agement of such problems carries its own challenges. Robot technology and telep-
resence technology not withstanding, time lags between Earth and space may
preclude safe surgery. The only safe option may be to have a couple of general sur-
geons on board for long duration flights in space.
The present work has brought home the realization that the gastrointestinal tract
is a system with an awe-inspiring complexity of mechanism. No more can we
explain gastrointestinal physiology in terms of itself (vitalism) or dismiss it as being
simple—as the astute fictional detective Sherlock Holmes would probably have
summed up for the perplexed doctor: “It is Alimentary, my dear Watson”. The above
analysis has demonstrated the vital importance of asymmetry to the unidirectional
propulsion of digesta. Since several mechanisms in the alimentary tract, both ana-
tomic and physiologic cooperate to impose polarity, we are afforded a more or less
unidirectional propulsion of digesta even if one or more factors, such as gravity, are
excluded. Without such asymmetry, let alone travel in space, human life on Earth as
we know it would not be possible. This paper is a taster of more work on the propul-
sion of gastrointestinal and urinary tract content. Our future papers will involve
numerical simulations including as many as possible of the several interacting sys-
tems from the above complex model.
84 ANNALS NEW YORK ACADEMY OF SCIENCES

ACKNOWLEDGMENT

I am grateful to Dr. Henry Oldwinkle, MBBS, House Officer at Colchester Gen-

eral Hospital for assessing the typescript for readability.

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