Gastric Digestion in Vivo

FO05CH06-Singh ARI 30 January 2014 9:4
ANNUAL
REVIEWS Further Gastric Digestion In Vivo
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Influence the Digestion Process
Gail M. Bornhorst1 and R. Paul Singh1,2
by Otterbein University on 10/01/14. For personal use only.
1
Department of Biological and Agricultural Engineering, University of California, Davis,
California 95616; email: gbornhorst@ucdavis.edu, rpsingh@ucdavis.edu
2
Riddet Institute, Massey University, Palmerston North 4442, New Zealand
Annu. Rev. Food Sci. Technol. 2014. 5:111–32 Keywords

First published online as a Review in Advance on food breakdown, gastric motility, gastric emptying, glycemic response,
January 2, 2014
digestibility
The Annual Review of Food Science and Technology is
online at http://food.annualreviews.org Abstract
This article’s doi: Food digestion is crucial for sustaining life. Although it has been examined for
10.1146/annurev-food-030713-092346
more than 300 years, the basic principles are not entirely understood. Antral
Copyright c 2014 by Annual Reviews. motility is well characterized, and current research is seeking to determine
All rights reserved
flow patterns generated by the stomach’s peristaltic contractions. The rate of
gastric emptying for solid and liquid meals has been determined according to
variations in meal composition, energy content, and subject characteristics.
The glycemic response has been measured for many carbohydrate foods
and is altered by factors such as amount of processing, particle size, and
starch structure. Similarly, ileal starch digestibility is altered by food and
starch properties. Even though many foods have been studied according to
their glycemic response, starch digestibility, and in vitro digestion kinetics,
the rate-determining processes and underlying mechanisms remain to be
established. The link between food properties, digestion processes, and final
health outcomes must be strengthened for functional food optimization.
111
1. INTRODUCTION
Gastric digestion is a crucial step in the absorption of energy and nutrients from foods. After
Gastric: related or mastication, the majority of the remaining food breakdown occurs in the gastric environment.
referring to the The stomach acts as a tank, mixer, grinder, and sieve (Meyer 1980) in a complex feedback control
stomach system that simultaneously breaks down ingested foods while mixing them with gastric acid and
GIT: gastrointestinal digestive enzymes. The rate of food breakdown in the stomach plays a key role in determining
tract other processes such as gastric emptying and nutrient absorption, but it is still not fully understood.
Many previous studies in the medical and nutrition fields have focused on topics such as gastric
motility, rate of gastric emptying, and absorption of nutrients into the blood.
Gastric motility, or the movement of the stomach walls, is characterized extensively in the
medical literature due to its use in diagnosing certain gastric diseases. Gastric motility parameters
will influence the rate of breakdown of food in the stomach, as they will change the forces,
pressures, and specific flow profiles encountered by food particles. Gastric emptying, or the rate
at which food leaves the stomach, is characterized in terms of both subject and food characteristics
(Benini et al. 1994, 1995; Hermansson & Sivertsson 1996; Kwiatek et al. 2009; Moore et al. 1984,
1990).
After food is emptied from the stomach, it will move into the small intestine, where the remain-
ing chemical breakdown will occur and the ingested nutrients will be absorbed for use in the body
(Borgström et al. 1957). Glycemic response, or the rate of glucose absorption into the blood after
meal consumption, is commonly measured in nutritional studies. Factors related to the specific
food consumed, such as initial food properties (Brand Miller et al. 1992, Panlasigui et al. 1991,
Ranawana et al. 2009), extent of food processing prior to cooking (Casiraghi et al. 1993), amount
of mastication (Ranawana et al. 2010b), and food physical form (i.e., ground or intact) (O’Dea
et al. 1980), influence food glycemic response.
The fundamental mechanisms causing the observed differences in gastric emptying and
glycemic response still require complete explanation. There is an imminent need for breakdown
and mixing kinetics to be determined in the gastric environment so that the food breakdown
process can be better understood and quantified. Current trends in the food industry are moving
toward designing innovative foods with unique health benefits, such as increased satiety, larger
nutrient availability, or decreased blood glucose response. However, to optimize functional food
design, it is essential to understand how the physical and chemical properties of foods influence
their breakdown in the stomach, a controlling factor in the health and digestive properties of
food products. Relationships must be developed between the levels of initial food processing, food
structure, and food composition and the food breakdown kinetics to be able to fully optimize
future food design. These future food products could provide many useful health benefits to con-
sumers, such as longer satiety (i.e., useful for weight management), more efficient energy release
(i.e., crucial for athletes), controlled glucose uptake (i.e., vital for diabetics), and optimized drug
and/or nutrient release and absorption.
2. DIGESTION OF FOODS
The process of digestion begins with the ingestion of a food product that is systematically broken
down via a multitude of mechanical and enzymatic processes to ensure that its components and
nutrients are fully available for absorption (see sidebar, Food Breakdown in the Stomach; also see
Figure 1). The gastrointestinal tract (GIT) begins at the mouth; goes through the esophagus,
stomach, and small and large intestines; and ends at the anus. It has a total length of 8–9 m
(Heuman et al. 1997).
112 Bornhorst · Singh

FOOD BREAKDOWN IN THE STOMACH
Gastric digestion is a complex process involving both physical and chemical food breakdown. Peristaltic mus-
cle contractions in the gastric antrum crush and fragment food particles. Gastric acid softens food particle tex-
ture, and digestive enzymes (pepsin, lipase) begin hydrolysis of nutrients so they can be absorbed once the
food reaches the small intestine. Food structure plays a crucial role in the food breakdown and nutrient release
processes.
2.1. Oral Digestion

The goal of oral digestion is to produce a mass of chewed food known as a bolus. The bolus
must have a sufficiently small particle size and be adequately lubricated with saliva so it can be
safely swallowed. Physiological variables, such as gender, personality type, and dentition status, as
well as food properties, such as food hardness, composition, and size of food portion, will affect
the chewing time, amount of saliva secreted, and final bolus particle size (Agrawal et al. 1997,
Bornhorst & Singh 2012, Dawes 1972, Dawes & Watanabe 1987, Gavião & van der Bilt 2004,
Gonzalez et al. 2004, Jalabert-Malbos et al. 2007, Lucas et al. 2004, Mioche et al. 2002, Peyron
et al. 2004, Prinz et al. 2007, Woda et al. 2006, Yurkstas 1965).
Organ Physical process Chemical process

Oral cavity (mouth) Mastication Enzymatic hydrolysis
Goal: Bolus formation Goal: α-amylase: Starch breakdown
Lingual lipase: Lipid breakdown
Esophagus Peristalsis
Goal: Bolus transport
Stomach Peristalsis Enzymatic hydrolysis

Goal: Continued food breakdown Goal: Pepsin: Protein breakdown
Lipase: Lipid breakdown
Acid hydrolysis
Goal: Gastric acid (pH ~2): Soften food
texture and break down structure
Small intestine Peristalsis Enzymatic hydrolysis

Goal: Digesta transport Goal: Lipase, phospholipase A: Lipid breakdown
Segmentation Amylase, amyloglucosidase: Starch breakdown
Goal: Mixing to facilitate absorption Trypsin, chymotrypsin, carboxypeptidase,
elastase: Protein breakdown
Diffusion (active/passive)
Goal: Absorption of nutrients
Large intestine (colon) Peristalsis Fermentation

Goal: Digesta transport Goal: Produces short-chain fatty acids
Segmentation and other by-products
Goal: Mixing to facilitate absorption
Diffusion (active/passive)
Goal: Absorption of fermentation
by-products and water
Figure 1
Summary of key physical and chemical processes that occur in the gastrointestinal tract (Heuman et al. 1997, Seidel & Long 2006).
www.annualreviews.org • Gastric Digestion and Food Structure 113

Esophagus
Cardia
Esophageal sphincter Fundus
Pyloric sphincter
Body
Pylorus
Duodenum
Antrum
Figure 2
Stomach anatomy showing approximate location of each region.
2.2. Esophageal Transit

Esophageal peristalsis is triggered once a bolus is swallowed. During esophageal transit, bolus
movement is also aided by the simple force of gravity, as long as the subject is not lying down
(Seidel & Long 2006).
2.3. Gastric Digestion

As the bolus passes through the lower esophageal sphincter and into the stomach, the proximal
stomach relaxes (Heuman et al. 1997, Sernka & Jacobson 1979), allowing the ingested bolus
to layer on top of the previously swallowed boluses (Schulze 2006). The stomach is primarily
comprised of the cardia, fundus, and antrum (Figure 2). The cardia is filled with mucin-secreting
cells. The fundus is lined with mucosa forming thick folds, known as rugae, where parietal cells
secrete HCl and chief cells secrete pepsinogen, an active precursor that is converted into the
proteolytic enzyme pepsin upon contact with acid. The antrum has a smooth surface filled mainly
with mucus-secreting cells. The pyloric sphincter forms the junction between the stomach and
the small intestine (Heuman et al. 1997, Seidel & Long 2006). Functionally, the stomach can be
separated into two regions: the fundus, or proximal stomach, comprising the cardia, fundus, and
body; and the antrum, or distal stomach, comprising the antrum and the pylorus. The proximal
stomach is thought to act as a food reservoir, where ingested food will remain before it moves
into the distal stomach. The distal stomach is thought to be the main location of the physical
breakdown of food in the stomach, where antral contraction waves (ACWs), or the peristaltic
contractions of the stomach wall, act to crush and grind food particles until they pass through the
pyloric sphincter into the small intestine (Heuman et al. 1997).
Peristalsis: sinusoidal
2.4. Small Intestinal Digestion
muscular contractions
used to mix and The small intestine extends from the pyloric sphincter to the ileocecal junction and is 6–7 m long.
transport food It ensures that the partially digested food, or chyme, is broken down into molecules small enough
throughout the
to be absorbed through the epithelial cells and carried into the bloodstream (Seidel & Long 2006).
gastrointestinal tract
The small intestine is separated into three sections. The duodenum secretes bicarbonate that will
ACW: antral
neutralize gastric acid and provide an appropriate pH for further enzymatic digestion to occur.
contraction wave
It receives secretions of bile and enzymes from the liver and pancreas, respectively (Figure 1).
The jejunum is approximately 2.5 m in length and extends from the end of the duodenum to

the ileum. The majority of nutrient absorption occurs in the jejunum. The ileum comprises the
remaining 3.5 m of the small intestine. Nutrient absorption is completed in the ileum, with
the distal 100 cm of the ileum being the only location where vitamin B12 and bile salts can be
MRI: magnetic
absorbed (Seidel & Long 2006). resonance imaging
Chyme is transported through the intestines by segmentation and peristaltic muscle contrac-
tions. Segmentation contractions aid in the mixing of chyme, allowing for a large amount of ma-
terial to contact the intestinal walls, which helps with nutrient absorption. Peristaltic contractions
result in a large forward movement, propelling chyme at a rate of 2–25 cm s−1 (Seidel & Long 2006).
2.5. Large Intestinal Fermentation

The large intestine extends from the ileocecal junction to the anal canal and is approximately 1.5 m
long. The movement and mixing of chyme occurs via segmentation and peristaltic contractions.
The large intestine is colonized by more than 1010 microorganisms/g of intestinal content, and
these anaerobic bacteria will ferment food particles that have not been completely digested. Only
short-chain fatty acids (by-products of the fermentation process) and water are absorbed (Seidel
& Long 2006).
3. GASTRIC DIGESTION PROCESS
3.1. Historical Perspective

Gastric digestion has been a topic of research for more than 300 years, with some of the earliest
documented experimentation conducted by Wepfer in 1679, who practiced vivisection on wolves,
dogs, and cats (Hertz 1910). It was not until 1833 that a detailed description of gastric digestion was
provided by William Beaumont, an army surgeon, who, over a period of multiple years, observed
the process of gastric digestion via a patient with a fistulated stomach. Almost 70 years later, Cannon
used a type of X-ray to examine the motility of the cat stomach. Cannon observed that the stomach
exhibits a type of receptive relaxation, that is, it will increase in volume to accommodate a large
meal. He also observed the movement of peristaltic waves through the antrum and was one of
the first scientists to cite the physiological differences in functionality of the proximal and distal
stomach (Cannon 1898, Cannon & Lieb 1911).
3.2. Gastric Motility

Gastric motility, or the movement of the stomach walls, is crucial for gastric digestion. The
frequency, duration, and intensity of ACWs will impact the rate and mechanism of physical food
breakdown and mixing in the stomach, given that they will change the forces, torques, pressures,
and flow profiles experienced by food particles. Gastric motility is commonly measured in the
medical field as a diagnostic measure. The classical methods to measure gastric motility require
the insertion of a barostat or manometer into the stomach and/or duodenum via the nose or mouth
(Collins et al. 1991; Desipio et al. 2007; Hebbard et al. 1995; Heddle et al. 1989, 1993; Szarka
& Camilleri 2009; Treacy et al. 1990). However, this testing procedure may alter normal gastric
function, leading to erroneous results. A newly emerging technique for monitoring gastric function
is magnetic resonance imaging (MRI), which has the capability to simultaneously measure gastric
volume, gastric motility, gastric secretions, and gastric emptying to help explain the relationship
between each of these parameters and overall gastric function (Ajaj et al. 2004; Hoad et al. 2007;
Marciani et al. 2001a,b,c; Szarka & Camilleri 2009). Another recently emerged technique is the
use of a wireless capsule, such as the commercially available SmartPillTM , to dynamically track the

force, torque, pressure, and/or pH in the GIT (Cassilly et al. 2008, Kloetzer et al. 2010, Laulicht
et al. 2010, Szarka & Camilleri 2009). The use of wireless capsules represents a noninvasive method
that is able to simultaneously measure multiple quantities; however, it still may require additional
validation with traditional measurements.
Table 1 provides examples of gastric motility measurements using the techniques described
above. Although there are small differences between studies, there is an overall consensus that
ACWs have an average speed of 1.5–3 mm s−1 and an average frequency of 2.6–3 waves min−1
for both solid and liquid meals. In spite of this, the effect of meal composition on gastric motility
is debated in the literature. Marciani et al. (2001c) showed no effect of meal composition (liquid,
liquid and solid, or high-viscous liquid and solid) on gastric motility, but Schwizer et al. (1996)
showed that dextrose concentration (10 versus 25%) changed the ACW frequency and depth.
However, meal composition was not always of high importance in the cited studies, as they were
generally done for diagnostic purposes. The available data suggest that food type, nutrient content,
and volume may affect gastric motility. Now that noninvasive techniques are available to monitor
motility parameters easily, research should be done using a wider variety of test meals with well-
documented food structures and composition.
3.3. Gastric Secretions

The stomach secretes an average of 2–3 L of gastric fluids each day (Seidel & Long 2006). Gastric
secretions are comprised of mucus, acid, enzymes, and intrinsic factor. Throughout the gastric
lumen, there is a gel-like mucous layer that protects the inner cells and musculature of the stomach.
Foveolar cells located in the superficial mucosal compartment of the stomach secrete mucus as
well as bicarbonate (HCO3 − ) ions (Heuman et al. 1997). The mucous layer is important for
protecting the gastric epithelium from its own acid secretions. The gastric fundus and body
contain gastric crypts filled with mucous neck cells (also secreting mucus to protect the gastric
epithelium), as well as parietal and chief cells. Parietal, or oxyntic cells, are triangular cells that
contain H+ -K+ -ATPase, which acts as a proton pump to exchange K+ for H+ , resulting in gastric
acid (HCl) secretion. In normal, healthy adults, average gastric acid secretion is 10 mEq H+ h−1
without stimulation, and can rise up to 30 mEq H+ h−1 with stimulation (Heuman et al. 1997).
Parietal cells also secrete intrinsic factor, a protein that binds to vitamin B12 and allows for its
absorption in the terminal ileum. Chief, or zymogenic, cells secrete the inactive precursor peptide
pepsinogen, which is converted into the active proteolytic enzyme pepsin upon contact with acid
by the removal of nine amino acids (Seidel & Long 2006). Chief cells also secrete gastric lipase,
which is responsible for 10–30% of dietary triglyceride hydrolysis (Gallier & Singh 2012a). The
pylorus is comprised mainly of mucus-secreting cells.
Some stomach cells also secrete hormones such as somatostatin, gastrin, cholecystokinin, and
glucagon that mediate the overall digestion process (Heuman et al. 1997, Seidel & Long 2006).
However, the specific feedback control mechanisms involving hormonal control of the digestion
process are outside the scope of this review.
3.4. Flow Patterns and Modeling in the Gastric Environment

As a result of the ACWs, there are flow patterns generated in the stomach, allowing for the
breakdown and mixing of food particles that are difficult to observe experimentally. Consequently,
various attempts have been made to computationally explore the flow patterns that are generated
in the stomach. Pal et al. (2004, 2007) developed a 2D computational fluid dynamic model of
the stomach, and Singh (2007) and Ferrua & Singh (2010) developed a 3D computational fluid

Table 1 Gastric motility measurements found in the literature

Technique Measurement Meal Values Obtained Notes Referencea
Manometry Intragastric 400 or 800 mL of Mean pressure increase The meal volume and (Kwiatek
pressure multinutrient from baseline (5.2 mm calorie content of a et al. 2009)
(Ensure R
) liquid Hg) after meal: 1.63 mm liquid meal influence
drink (varying Hg (400 mL meal) and intragastric pressure,
calories) 2.77 mm Hg (800 mL meal) but the magnitude of
the difference is quite
small
Solid-state Pyloric Liquid meal: Mean pyloric pressure: Solid and liquid meals (Desipio
manometry pressure Boost R
(240 kcal) 9.9 ± 1.8 mm Hg (solid had similar pyloric et al. 2007)
Solid meal: eggs, meal) and 12.8 ± 2.6 mm pressure, both during
toast, water Hg (liquid meal) and between ACWs
(345 kcal) Maximum pyloric pressure

(during ACWb ): 53.6 ±
8.4 mm Hg (solid meal) and
60.5 ± 8.9 mm Hg (liquid
meal)
MRI ACW Liquid meal: 500 Mean ACW frequency: All meal types resulted in (Marciani
frequency mL LBG solution 3.0 ± 0.2 waves min−1 similar ACW speeds et al. 2001c)
and speed (323 kcal) Mean ACW speed: and frequencies
Liquid/solid meal: 1.6 ± 0.2 mm s−1
500 mL LBG
solution with solid
agar gel beads
(323 kcal)
Viscous liquid/solid
meal: 500 mL
viscous LBG
solution with solid
agar gel beads
(323 kcal)
MRI ACW Liquid meal: enteral Mean ACW frequency: The frequency of ACWs (Kunz et al.
frequency and parenteral 2.9 ± 0.1 waves min−1 was similar for both 1999)
and speed feeding solutions (liquid and solid meal) meals, but the average
(523 kcal) Mean ACW speed: wave speed was
Solid meal: pancake 2.8 ± 0.1 mm s−1 (liquid different between the
prepared with meal) and 3.1 ± 0.1 mm solid and liquid meals
eggs, potatoes, s−1 (solid meal)
bacon, butter, and
water (523 kcal)
MRI ACW 500 mL of low- and Mean ACW frequency: The frequency and speed (Marciani
frequency high-viscosity 2.98 ± 0.03 waves min−1 of ACWs were similar et al. 2001a)
and speed; LBG meals with (low-viscosity meal) and for all meal types, but
force in addition of agar 2.92 ± 0.02 waves min−1 the high-viscosity meals
antrum gel beads of (high-viscosity meal) experienced higher
breaking strengths Mean ACW speed: 1.57 ± forces than the
from 0.15 to 0.90 0.04 mm s−1 (low-viscosity low-viscosity meals
N (323 kcal) meal) and 1.55 ± 0.03 mm
s−1 (high-viscosity meal)
Antral forces: ∼0.65 N
(Continued )
Table 1 (Continued)
Technique Measurement Meal Values Obtained Notes Referencea
MRI ACW 500 mL of 10% Mean ACW frequency: The ACWs did not (Kwiatek
frequency, liquid glucose 2.61 ± 0.03 waves min−1 cause liquids to empty et al. 2006)
speed, and (200 kcal) Mean ACW speed: (ACW propagation was
amplitude 2.7 ± 0.1 mm s−1 not related to liquid
Mean ACW amplitude: emptying)
7.0 ± 0.8 mm
MRI ACW 500 mL of 10% or Mean ACW frequency Both ACW frequency (Schwizer
frequency 25% dextrose (15–30 min after meal and depth decreased et al. 1996)
and depth solution consumption): 2.8 ± 0.6 when dextrose
waves min−1 (10% dextrose) concentration was
and 2.0 ± 0.9 waves min−1 increased
(25% dextrose)
Mean ACW depth
(% total): 40 ± 17% (10%
dextrose) and 34 ± 16%
(25% dextrose)
MRI ACW speed 400 mL of pudding Mean ACW speed Antral motility can be (Ajaj et al.
(15–30 min after meal characterized in normal 2004)
consumption): 2.4 mm s−1 and diseased subjects
using MRI
a
Gastric motility was not always the principal aim of the cited references; however, only the motility measurements are listed for comparison.
b
Abbreviations: ACW, antral contraction wave; LBG, locust bean gum; MRI, magnetic resonance imaging.
dynamic model of the stomach. These models were all developed using physiologically accurate
geometries and gastric motility parameters. These models have demonstrated various phenomena
that may occur during gastric digestion, such as the presence of eddies and a retropulsive jet in
the antrum (Ferrua & Singh 2010, Pal et al. 2004, Singh 2007). These models also show that flow
patterns in the stomach depend on the properties (viscosity, density, particle size) of the meal.
4. EXPERIMENTAL APPROACHES TO MONITOR DIGESTION IN VIVO
4.1. Measurements of Gastric Emptying

The rate of gastric emptying, or the rate at which food leaves the stomach, is influenced by
meal composition (Benini et al. 1994, Hertz 1910), meal nutrient content (Moore et al. 1984),
subject gender (Hermansson & Sivertsson 1996), and gastric motility (Camilleri et al. 1986, 1985),
among other factors (Figure 3). Methods used to measure gastric emptying include scintigraphy,
stable isotope breath testing, ultrasonography, the use of wireless motility capsules, and MRI
(Vantrappen 1994). Szarka & Camilleri (2009) present a detailed review of these methods.
4.2. Blood Glucose Response and the Glycemic Index

Blood glucose concentration is used in numerous studies as a noninvasive measure of the rate of
food digestion and absorption. Blood glucose levels are easily measured by taking small blood
samples from the finger with a capillary tube (Wolever et al. 1991). To determine the glycemic
response of a meal, blood glucose concentration is measured over a period of 2–3 h after meal
consumption and compared with baseline levels.

Subject
characteristics
Age
Gender
Activity level
Food properties
Composition
kCal
Physiological response Volume
Gastric secretions Structure
Hormonal response Processing
Food breakdown
Rheological properties
Particle size distribution
Gastric mixing
Figure 3
Representation of the food and subject factors that are interrelated in determining the gastric emptying rate
of a meal.
The glycemic index (GI) was developed to provide a means of comparison for the glycemic
response across many test foods. The GI is calculated as the area under the curve (AUC) of the
glucose response of a test food. This is compared to the AUC of a control (glucose solution or white
bread). Test meals contain 50 g of available carbohydrate ( Jenkins et al. 1981). The GI has become
a widely accepted measure of the glucose absorption in carbohydrate-based foods; international
GI tables are established for more than 150 food products (Foster-Powell et al. 2002).
More recently, the glycemic load (GL) was introduced. The GL is calculated similarly to the
GI; however, the portion of food fed to determine the GL represents a realistic serving of the food
under study instead of a 50-g portion (Foster-Powell et al. 2002). The GL may have more practical
implications for comparing the food portions that are actually consumed. However, due to the
differences in food size, comparisons may be hard to make between food products.
4.3. Ileostomates
Patients with diseases such as Crohn’s disease and ulcerative colitis may undergo ileostomy
surgery (McLeod et al. 1985). This is a procedure that removes inflamed tissue and attaches the
terminal ileum to the outside of the body, where a pouch will collect the effluent waste products.
Some ileostomates with a permanent need to use an external ileal pouch have volunteered to let
their ileal effluent waste be studied as an end product of the digestion process in the upper GIT.
Studies with ileostomates are quite unique in that, normally, ileal samples cannot be collected
easily from humans. Ileostomates have been used to study digestion and absorption of starches
(Botham et al. 1997, Chapman et al. 1985) and breakdown of almond cubes (Mandalari et al.
2008). However, because ileostomates do not represent a normal, healthy individual, applications
of these studies may be limited.
GI: glycemic index

4.4. Animal Models
AUC: area under the
It is not always possible to use a human model in studies of digestion, and/or it may be desired curve
to study the growth and nutrition of animals. Animals that are commonly used to study digestion
GL: glycemic load
include dogs (Dikeman et al. 2006, 2007a,b), chickens (Fuente et al. 1998, Short et al. 1996,
Weurding et al. 2001), rats (Holm et al. 1988, Rutherfurd et al. 2011, Zhao et al. 2003), and

pigs (Bornhorst et al. 2013b,c; Gregory et al. 1990; Kim et al. 2007; Rainbird & Low 1986). In
animal digestion trials, samples of chyme are taken from the GIT, which may be done using two
main techniques: cannulation and slaughter. In both techniques, an indigestible marker, such as
In vivo:
experimentation chromium oxide (Cr2 O3 ) or titanium dioxide (TiO2 ), must be added to the meal to determine the
conducted in a living proportion of the original meal that is collected at each location in the GIT (Bach Knudsen et al.
organism, such as an 2006, Short et al. 1996, van Bussel et al. 2010).
animal or human Cannulation involves the surgical insertion of a cannula into the wall of the GIT. The outer
In vitro: end of the cannula protrudes outside of the skin and contains a removable stopper, which allows
experimentation for samples to be extracted (Decuypere et al. 1977). The advantage of cannulation is that it allows
conducted in a
sampling from the same animal over a long time period. Also, different experimental treatments
laboratory setting
can be given to the same animals, reducing variability. However, cannulation requires surgery, and
the surgical modifications of the GIT may cause changes in gastrointestinal secretions and motility,
altering flow patterns and modifying the actual digestion process (Bach Knudsen et al. 2006).
In the slaughter technique, animals are sedated and euthanized at certain times after meal con-
sumption, and samples may be taken at any location(s) in the GIT. An advantage of the slaughter
technique is that it allows sampling from many locations from the same animal. The main disadvan-
tage is that it does not allow for repeated sampling from the same animal, which may increase the
number of animals needed in the trial to produce statistical significances (Bach Knudsen et al. 2006).
Both the cannulation and the slaughter techniques have their merits, and the type of samples
needed dictates which technique is selected. It has been demonstrated in pigs that the ileal and
fecal digestibility of nitrogen and amino acid from a meat and bone meal diet was not statistically
different between cannulated and slaughtered pigs, suggesting that either technique may be suitable
if only nutrient digestibility measurements are required (Donkoh et al. 1994).
5. EXPERIMENTAL APPROACHES TO MONITOR

DIGESTION IN VITRO
Due to the complex nature of the GIT, in vivo models of digestion provide the most accurate rep-
resentation of the intricacies of the digestion process. However, because of practical and logistical
constraints, it is not always possible to conduct studies in vivo. In these cases, in vitro, or laboratory-
scale, experiments are conducted. In vitro systems allow control of many factors that cannot be
controlled in vivo. This permits the isolation of key factors that are important in the digestion
process. Ultimately, in vitro models must be compared to in vivo models to establish their accuracy.
Hur et al. (2011) provide a detailed description of many in vitro digestion models used to
mimic human digestion (oral, gastric, and intestinal), as well as the enzymes used in each digestion
model. In addition, Bornhorst & Singh (2012) and Guerra et al. (2012) provide a more detailed
description of various in vitro oral and gastric digestion models. Following the description of in
vivo measurement methods for food gastric digestion, three main types of in vitro gastric digestion
models are briefly described.
5.1. Water Bath Incubation

The most simple and common in vitro gastric digestion model is a water bath incubation model.
Water bath incubation models may include any variety of digestive enzymes (amylases, proteases,
lipases), mucins, and salts depending on the specific food product under investigation and the
portion of interest in the GIT. Samples are commonly mixed with a simulated digestion fluid (with
appropriate pH and enzyme concentration) and incubated at 37◦ C for varying times, normally up
to 2–3 h (Hur et al. 2011). Some models employ agitation, or a shaking water bath, in an attempt

to mimic the hydrodynamic forces present in the GIT (Bornhorst & Singh 2013). However, this
agitation alone may not accurately imitate the physical forces present throughout the stomach
and intestines (Kamba et al. 2003). Water bath incubation models have an advantage in that they
HGS: human gastric
are simple to construct and maintain, allowing for many experiments to be conducted; however, simulator
care should be taken in the selection of specific pH and enzymatic conditions during digestion.
These models may be able to accurately simulate the chemical breakdown of certain foods and/or
nutrients, but their applicability to the digestion of whole food products may be limited due to
the absence of any physical and/or hydrodynamic forces.
5.2. Dynamic Mechanical Models

In an attempt to mimic both the physical and chemical conditions encountered in the GIT, several
groups have developed dynamic in vitro digestion models. These models not only incubate food
products with the appropriate concentration of acids and enzymes at 37◦ C but they also include
physical forces, such as peristaltic muscular contractions, to allow for both chemical and physical
breakdown (Bornhorst & Singh 2012). Such dynamic in vitro models include the TNO intestinal
model developed at the TNO Nutrition and Food Research Center (Zeist, Netherlands), which has
been used to study nutrient bioavailability and antimutagenic activity of food during digestion,
among other things (Krul et al. 2000, Verwei et al. 2003). The model gut, or dynamic gastric
model, was developed at the Institute of Food Research (Norwich, United Kingdom) and is
shown to accurately replicate the antral grinding forces observed in vivo (Vardakou et al. 2011).
In addition, the human gastric simulator (HGS), developed at the University of California, Davis
(Davis, California, USA), has been used to study the gastric digestion of rice and apples, and the
release of β-carotene from almond butter (Kong & Singh 2010, Roman et al. 2012). Dynamic
mechanical models of digestion have an advantage over other in vitro digestion approaches, as
they allow for examination of both physical and chemical breakdown of food products. However,
these models are more complex in their design and fabrication and require validation with in vivo
data prior to widespread use.
5.3. Cell Models

When nutrient bioavailability and uptake are concerned, some researchers have utilized cell mod-
els, most commonly a Caco-2 cell model. Caco-2 cells are found in the human epithelium and can
be used to monitor the uptake of certain nutrients, such as iron (Glahn et al. 1998). In cell models,
the food product of interest is generally subjected to simulated gastric digestion in a water bath–
type model, where it is mixed with enzymes at a low pH for 1–2 h. Following the gastric portion,
the pH may be adjusted and additional enzymes added to the partially digested food before it is
placed in a system with a dialysis membrane followed by a monolayer of Caco-2 cells. In this way,
the quantity of the nutrient of interest that is absorbed by the cells can be quantified over time as
a means to predict the cellular nutrient uptake (Glahn et al. 1998). The use of a Caco-2 cell model
may be advantageous when trying to predict the bioavailability of specific nutrients and to study
the underlying mechanisms in nutrient transport across the human epithelium (Bering et al. 2006).
6. FOOD PROPERTIES THAT INFLUENCE DIGESTION

The digestion of many food products has been studied in terms of the subject characteristics, food
physical form, and food processing method. The rate of digestion in vivo is mainly described by
the rate of gastric emptying, glycemic response, and ileal starch digestibility. In addition, in vitro

models have been utilized to study the digestive properties of many different food forms. Several
reviews are available regarding the digestibility of starch as influenced by the starch macro- and
microstructure (Parada & Aguilera 2011, Singh et al. 2010), as well as the starch digestion kinetics
Gastric emptying
rate: the rate at which (Dona et al. 2010); therefore, these relationships are not the focus of this review. This section
material leaves the focuses on in vivo studies that have been completed regarding the influence of food properties
stomach on their digestive characteristics as well as in vitro studies that have attempted to determine the
mechanisms behind the differences observed in vivo.
6.1. Gastric Emptying Rate of Foods

Gastric emptying is measured extensively in the medical and nutrition fields to determine its
relationship with individual and food-related parameters. Particularly in the medical literature,
the gastric emptying rate is quantified by the lag time, or the time it takes before a meal begins to
empty, and a t50 , or the time it takes for 50% of the meal to leave the stomach.
Subject characteristics, such as gender (Hermansson & Sivertsson 1996) and physical activity
(Moore et al. 1990), have been shown to play a role in the gastric emptying rate and must be
considered when conducting human trials. In addition, properties of the ingested meal such as
calorie content, meal volume, fat content, and acid concentration have all been shown to influence
the meal gastric emptying (Benini et al. 1994, 1995; Kwiatek et al. 2009; Moore et al. 1981, 1984).
In a study investigating the influence of meal size on solid particle gastric emptying, Collins
et al. (1996) fed healthy adult subjects a meal consisting of 150 mL of a 10% dextrose solution
with either 100 g (small) or 400 g (large) of a ground beef–chicken liver mixture and measured
the gastric emptying via scintigraphy. They found that the lag time of the solid emptying was
greater for the large meal (56 versus 31 min), but once the solid particles started emptying, the
particles from the large meal emptied almost twice as fast as those from the small meal. These
differences may have been due to the larger amount of particles that needed to be broken down in
the large meal, or the lag phase may have been longer in the large meal due to its larger volume and
caloric content (Kwiatek et al. 2009, Moore et al. 1981). Siegel et al. (1988) found that in a meal
containing solid and liquid components, the physical form of the solid component will influence
its lag phase. For example, they found that the lag phase of an egg meal was 31 min, and the lag
phase of a chicken liver meal was 62 min. These differences in gastric emptying may be caused by
a variety of food and subject factors (Figure 3).
Levels of food processing were shown to influence specific nutrient gastric emptying in pigs
after a meal of cooked brown or white (Calrose variety) rice. Bornhorst et al. (2013a) found that
the rate of emptying of dry matter and starch was similar after brown and white rice meals, but
brown rice meals had a slower rate of protein emptying (t1/2 = 309 min) compared to white rice
meals (t1/2 = 237 min). They hypothesized that the differences in protein emptying were due to
an accumulation of the bran layers from brown rice in the gastric antrum.
Meal macronutrient content may also play a role in the gastric emptying rate. Benini et al.
(1994) found that the total gastric emptying time of a fried pasta–based meal was significantly
delayed (317 min) compared to its boiled counterpart (227 min). The delayed emptying of the fried
meal, particularly in the later stages of digestion, could have been influenced by the ileal brake, a
phenomenon that occurs when a high level of fat reaches the jejunum and ileum, causing a decrease
in intestinal motility and the gastric emptying rate (Spiller et al. 1984, Van Citters & Lin 1999).
Various studies have examined the effect of organic acids and salts on the gastric emptying rate
and glycemic response to a meal. It has been suggested that hydrochloric acid, lactic acid (Lin et al.
1990), propionic acid (Liljeberg & Björck 1996), acetic acid (Liljeberg & Björck 1998), tartaric
acid, butyric acid, and other organic acids (Hunt & Knox 1969) all have a lowering effect on

the gastric emptying rate of a meal, and consequently on meal glycemic response. This response
may be part of an internal feedback control system in the stomach that neutralizes the acid and
consequently slows the gastric emptying rate.
Ileal digestibility:
amount of a certain
nutrient that has been
6.2. Food Properties Influencing Glycemic Index in Carbohydrate-Based Foods absorbed prior to
Numerous studies have been conducted in the nutrition field to examine the effect of different reaching the ileum
types of food products on their respective glucose response. Some of these studies have calculated a
GI for the food under study by following standard procedures (Wolever et al. 1991), whereas other
studies do not calculate a GI specifically, but measure the postprandial (after meal consumption)
glucose and insulin concentrations. These studies have shown that food structure, starch type,
starch amylose content, amount of moisture, amount of resistant starch, cooking/preparation
parameters, and fiber content, among many other factors, are important in determining food
glycemic response (Brand et al. 1985; Brand Miller et al. 1992; Casiraghi et al. 1993; Foster-
Powell et al. 2002; Goddard et al. 1984; Granfeldt et al. 1994; Hallfrisch & Behall 2000; Jenkins
et al. 1981, 1986, 1988; Josse et al. 2007; Kendall et al. 2011; Kristensen et al. 2010; Panlasigui et al.
1991; Parada & Aguilera 2011; Ranawana et al. 2009, 2010a,b, 2011; Read et al. 1986; Scazzina
et al. 2009; Vega-López et al. 2007).

In addition to compositional and cooking parameters, physical factors, such as particle size, have
also been shown to influence the glycemic response of rice. In comparing cooked brown and white
rice with their ground counterparts, O’Dea et al. (1980) found that ground rice elicited higher
glucose and insulin responses in the first hour after consumption compared to intact rice. There
was no significant difference observed between brown and white rice, suggesting that food particle
size is a key factor in determining glycemic response (O’Dea et al. 1980). Ranawana et al. (2010b)
also suggested that the rice particle size after mastication may play a role in the glycemic response.
6.3. Starch Digestibility from Animal Trials

Ileal starch digestibility is commonly quantified in animal trials as a measure of the starch avail-
ability for absorption during digestion. Pluske et al. (2007) reported in detail ileal digestibilities
of specific varieties of cooked rice. They found that pigs fed a medium-grain (Amaroo variety)
rice diet and a waxy rice (low amylose content) diet had high ileal digestibilities (96.2 and 99.1%,
respectively). However, the ileal starch digestibility was greatly reduced (88.6%) with a long-grain
(Doongara variety) rice diet. These differences were attributed to the higher amylose content of
the long-grain rice, resulting in a higher water-holding capacity, increased starch retrogradation,
and a larger resistant starch content (Pluske et al. 2007). Additionally, Bornhorst (2012) reported
that pigs fed a cooked medium-grain (Calrose variety) brown and white rice diet had higher ileal
digestibility from the white rice diet (98%) compared to the brown rice diet (95%).
Similar to studies of glycemic response in humans, many investigations of starch digestibility
have found that both physical form (i.e., particle size) and processing level (i.e., extrusion process-
ing) will influence ileal starch digestibility. Owsley et al. (1981) found that decreasing the particle
size of sorghum increased its ileal digestibility. This is similar to what was observed in another
study with rice, barley, sorghum, and oat diets (Solà-Oriol et al. 2007). However, the specific
starch source may determine whether the particle size influences the ileal starch digestibility, as
changes in wheat particle size did not influence ileal digestibility, whereas increasing rye particle
size decreased its starch digestibility (Bach Knudsen et al. 2006).
Animal trials have shown similar results to human trials examining the glycemic response
of processed foods; harsh processing treatments, such as extrusion, will increase the starch

digestibility in pig feed. Fadel et al. (1988) found that the ileal starch digestibility of raw barley
was 83.7% but an extruded barley meal was 96.9% digestible. Similarly, Sun et al. (2006) showed
that extrusion increased the ileal starch digestibility of barley, pea, potato starch, and wheat bran
diets, which was attributed to the gelatinization of raw potato starch, leaving it more susceptible
to hydrolysis by digestive enzymes.
6.4. Physical Property Changes During Gastric Digestion In Vivo

Many studies are published regarding the influence of food properties on meal gastric emptying,
glycemic response, and starch digestibility in vivo. A recent series of studies has examined the
physical and chemical properties of chyme during the gastric digestion process of cooked brown
and white rice in pigs. These studies examine how the food processing, structure, and composition
influence the actual breakdown process in the stomach, which will undoubtedly contribute to the
gastric emptying, glycemic response, and starch digestibility.
These studies have demonstrated that the presence of the bran layer in brown rice causes an
accumulation of fiber (Bornhorst et al. 2013c) and protein (Bornhorst et al. 2013a) in the gastric
antrum, which results in a slower gastric emptying rate of protein in brown rice compared to
white rice. In addition, they have also demonstrated that the gastric chyme behaves as a Herschel-
Bulkley fluid (Bornhorst et al. 2013b,c), exhibiting a positive yield stress followed by shear-thinning
behavior in both brown and white rice. The gastric chyme rheological and textural properties
were shown to decrease over a digestion period of 120–480 min, giving an indication of the rate of
breakdown of the meals. White rice showed a greater decrease in rheological properties, textural
characteristics, and particle size distribution compared to brown rice, indicating that it underwent a
more rapid breakdown process, presumably due to the decreased resistance to physical breakdown
in the absence of the bran layer and an increased rate of gastric secretions according to the rice
buffering capacity (Bornhorst et al. 2013a,b,c).
6.5. In Vitro Measurements of Food Breakdown During Gastric Digestion

To further elucidate the underlying mechanisms responsible for differences in food breakdown,
emptying, and absorption rates, many recent in vitro studies have been conducted. Although
a comprehensive review of these studies is outside the scope of this review, a few noteworthy
examples and recent reviews are given below.
Kong & Singh (2009) tested a variety of solid food products in a rotating turntable digestion
system to investigate the role of food structure as related to the mode of disintegration of individual
food particles during simulated gastric digestion. They found that the disintegration rates of solid
food particles of ham, fried dough, carrot, beef jerky, almond, and peanut followed a linear-
exponential profile. The disintegration profile shape varied between delayed-sigmoidal, sigmoidal,
and exponential, depending on food type. They showed that the rates of water absorption, texture
softening (from acid and enzymes), and surface erosion all play a role in the physical breakdown
of food particles during gastric digestion. In another study, Kong & Singh (2010) used the HGS
dynamic digestion model to demonstrate that the particle size distribution of rice was decreased
by the presence of peristaltic motion in the HGS compared to a shaking water bath model. This
indicates that when physical food breakdown is concerned, dynamic mechanical models may serve
to better describe the overall food digestion process.
Other in vitro studies have also demonstrated the influence of foods’ physical characteristics
and processing on their breakdown during digestion. A study examining the influence of physical
form on digestion kinetics was conducted with either hammer-milled or cryomilled sorghum of

various particle sizes (Mahasukhonthachat et al. 2010). This study showed that both the particle
size and milling process influenced the sorghum starch breakdown kinetics. These differences were
attributed to process-induced changes in water absorption, pasting, and water solubility properties
of sorghum. In a similar fashion as the research conducted on other carbohydrate-based food prod-
ucts, Bornhorst & Singh (2013) studied the disintegration kinetics of boluses prepared with differ-
ent types of bread during static or agitated incubation in gastric juice. They found that differences
in disintegration kinetics between bread types could be attributed to bread bolus cohesive forces,
which were caused by variations in bread structure, moisture content, and water holding capacity.
In an attempt to use natural food products as structuring agents and to optimize nutrient
bioavailability through processing, other researchers studied in vitro (through incubation and
Caco-2 cell models) and in vivo β-carotene and lycopene bioavailability and bioaccessibility.
They found that nutrient bioavailability could be increased by using an optimal combination of
food structuring and processing in a soup product with a tomato, carrot, and broccoli base (Van
Buggenhout et al. 2012).
In addition to food macrostructure, food microstructure will also play a role in nutrient break-
down and release. The specific structure of lipid droplets and emulsions may be altered to speed
up or delay lipid absorption; such topics are the subject of multiple recent reviews (Gallier & Singh
2012b, Golding & Wooster 2010, McClements et al. 2008).
7. SUMMARY
Despite numerous investigations, the underlying mechanisms and processes of digestion are
still not completely understood. Antral motility is extensively quantified, and current research
is ongoing to determine the specific flow patterns that are generated after different meals. The
rate of gastric emptying is quantified for meals of varying composition and calorie content, and
in different population groups. However, comprehensive studies that examine food properties,
antral motility, gastric chyme properties, and perhaps even hormonal responses after a meal are
required to fully explain the complicated feedback and control mechanisms employed in the body
during food digestion.
The glycemic response, often altered as a consequence of gastric emptying, is thoroughly
quantified for many carbohydrate-based food products. Factors such as starch structure, particle
size, and level of processing are all understood to alter the glycemic response to starchy foods.
However, to fully understand the starch digestion and breakdown process, the relationship to
the gastric emptying rate needs to be clarified. Also, because cooking was shown to be an im-
portant parameter in several studies, food preparation methods need standardization for proper
comparisons to be made.
Similarly, ileal starch digestibility is altered by starch physical form, meal particle size, and
grain processing. Although the ileal starch digestibilities for a variety of common pig diets have
been measured, chyme properties that may influence these digestibilities have not been extensively
quantified to allow for a mechanistic explanation of the starch absorption process.
Even though several studies have examined the relationships between many food structures,
compositions, and levels of food processing and their associated gastric emptying rate, glycemic
response, and starch digestibility, the underlying mechanisms responsible for the observed differ-
ences have yet to be fully explained. Instead of examining the end-point measurements (i.e., gastric
emptying, glycemic response, ileal digestibility), which have already been extensively quantified, a
mechanistic approach needs to be taken to fully understand the role of food physical and chemical
properties in food breakdown, and how these properties change during digestion. This type of
mechanistic approach has been used in several recent in vitro and in vivo studies; however, more

comprehensive trials must be completed. Only once the actual breakdown mechanisms are under-
stood and quantified will it be possible to optimize functional food design to allow for controlled
breakdown, nutrient release, and absorption characteristics.
SUMMARY POINTS
1. The digestion process begins in the oral cavity, where ingested food is formed into a bolus.
The bolus is swallowed and travels through the esophagus to the stomach, where food
physical and chemical breakdown is continued. Nutrient absorption occurs in the small
intestine, and unabsorbed nutrients are fermented in the large intestine prior to excretion.
2. Gastric digestion has been studied for more than 300 years, but additional information
is still needed about the influence of food properties on the digestion process.
3. Gastric motility has been extensively studied in the medical field by techniques such
as MRI, manometry, and the use of wireless capsules. There is an overall consensus
that ACWs have an average speed of 1.5–3 mm s−1 and an average frequency of 2.6–
3 waves min−1 . However, food has not been a key concern, and conflicting results exist
regarding the influence of meal characteristics on gastric motility.

4. Gastric digestion has commonly been quantified using end-point measurements such
as gastric emptying, blood glucose response, and ileal starch digestibility, neglecting the
underlying processes that occur in the stomach and how they influence certain end points
of interest.
5. The gastric emptying rate is influenced by both individual (consumer) characteristics
and food properties. However, the interaction between individual characteristics and
food properties is not yet clear.
6. Food structure, food processing, and cooking method are key factors influencing the GI
of starchy food products.
7. Future research is necessary to determine the connection between food properties, phys-
iological responses, and digestion end-point measurements to provide the knowledge
necessary to optimize functional food development.
DISCLOSURE STATEMENT
The authors are not aware of any affiliations, memberships, funding, or financial holdings that
might be perceived as affecting the objectivity of this review.
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FO05CH06-Singh ARI 30 January 2014 9:4

• Top downloaded articles

Annu. Rev. Food Sci. Technol. 2014. 5:111–32 Keywords

112 Bornhorst · Singh

FOOD BREAKDOWN IN THE STOMACH

2.1. Oral Digestion

Organ Physical process Chemical process

Stomach Peristalsis Enzymatic hydrolysis

Small intestine Peristalsis Enzymatic hydrolysis

Large intestine (colon) Peristalsis Fermentation

www.annualreviews.org • Gastric Digestion and Food Structure 113

2.2. Esophageal Transit

2.3. Gastric Digestion

114 Bornhorst · Singh

2.5. Large Intestinal Fermentation

3. GASTRIC DIGESTION PROCESS

3.1. Historical Perspective

3.2. Gastric Motility

www.annualreviews.org • Gastric Digestion and Food Structure 115

3.3. Gastric Secretions

3.4. Flow Patterns and Modeling in the Gastric Environment

116 Bornhorst · Singh

Table 1 Gastric motility measurements found in the literature

(345 kcal) Maximum pyloric pressure

4. EXPERIMENTAL APPROACHES TO MONITOR DIGESTION IN VIVO

4.1. Measurements of Gastric Emptying

4.2. Blood Glucose Response and the Glycemic Index

118 Bornhorst · Singh

GI: glycemic index

www.annualreviews.org • Gastric Digestion and Food Structure 119

5. EXPERIMENTAL APPROACHES TO MONITOR

5.1. Water Bath Incubation

120 Bornhorst · Singh

5.2. Dynamic Mechanical Models

5.3. Cell Models

6. FOOD PROPERTIES THAT INFLUENCE DIGESTION

www.annualreviews.org • Gastric Digestion and Food Structure 121

6.1. Gastric Emptying Rate of Foods

122 Bornhorst · Singh

et al. 2009; Vega-López et al. 2007).

6.3. Starch Digestibility from Animal Trials

www.annualreviews.org • Gastric Digestion and Food Structure 123

6.4. Physical Property Changes During Gastric Digestion In Vivo

6.5. In Vitro Measurements of Food Breakdown During Gastric Digestion

124 Bornhorst · Singh

2012b, Golding & Wooster 2010, McClements et al. 2008).

www.annualreviews.org • Gastric Digestion and Food Structure 125

regarding the inﬂuence of meal characteristics on gastric motility.

126 Bornhorst · Singh

www.annualreviews.org • Gastric Digestion and Food Structure 127

Lipid Technol. 24:271–73

128 Bornhorst · Singh

www.annualreviews.org • Gastric Digestion and Food Structure 129

ileostomy. Dis. Colon Rectum 28:152–54

properties obtained from different meat textures. Food Q. Prefer. 13:583–88

130 Bornhorst · Singh

Seidel E, Long M. 2006. Crash Course: Gastrointestinal System. Philadelphia: Elsevier

emptying. Gut 29:85–89 characterized by both a

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Contents Volume 5, 2014

From Tomato King to World Food Prize Laureate

Editor: Stephen E. Fienberg, Carnegie Mellon University