Resistant Starch and HealthA Evans, Tate and Lyle, Hoffman Estates, IL, USA

ã 2015 Elsevier Inc. All rights reserved.

Topic Highlights

• What is resistant starch?

• Resistant starch intake; measurement and consumption

• Effect of resistant starch intake on gut health

• Metabolic effects of resistant starch intake

Learning Objectives

• To get a basic understanding of what resistant starch is and

where it is found

• To learn about how its consumption has been linked to a

variety of potential beneficial health outcomes in the area

of gut health and various metabolic effects such as blood

glucose, insulin, and lipid levels

What Is Resistant Starch?

Starch is the major source of carbohydrates in the human diet.

Starch is present in many different fruits, vegetables, roots, and

grains. Starch and starch derivatives are a nutritive, abundant,

and economical food source. Starch can be consumed unpro-

cessed in the form of raw fruits and vegetables or in the form of

more shelf-stable processed foods. Food starches contribute to

the characteristic viscosity, texture, mouthfeel, and consistency

of many food products.

In the human body, starch is digested by a-amylases. First,

salivary a-amylase in the mouth catalyzes the hydrolysis of

amylose to maltose, maltotriose, and maltotetraose and the

hydrolysis of amylopectin to the same products plus two

a-limit dextrins. The partially digested starch then passes into

the stomach. After some resident time in the low pH environ-

ment of the stomach, the partially hydrolyzed starch then

passes into the small intestine where it is neutralized. The

majority of hydrolysis of the starch is then accomplished by

pancreatic a-amylase that is secreted from the pancreatic duct

via a multiple attack mechanism. a-Amylases hydrolyze the

starch into small mono-, di-, and oligosaccharides that then

need to be further broken down to be absorbed as glucose. Two

other enzymes are necessary to convert the hydrolysis products

of the a-amylases into glucose, which can be actively trans-

ported across the small intestine membrane. These two

enzymes are the brush-border glucogenic enzymes maltase–

glucoamylase and sucrase–isomaltase. The resulting D-glucose

is then actively transported across the luminal membrane of

the small intestine and passes into the blood via the sodium–

glucose cotransporter, which is located at the luminal surface

of enterocytes.

Several factors influence the rate and extent of starch diges-

tion. The main hydrolysis of starch is performed by the

a-amylases. Starch digestion by a-amylases requires a series of

steps. First, the enzymes diffuse into the starch matrix of the

food. In the second step, the enzymes bind to the substrate,

and finally, the enzymes cleave the a-1,4-glycosidic linkages ofthe substrate (starch). The structure of the food matrix and the

structure of the starch itself influence the kinetics of the amy-

lase hydrolysis.

Resistant starch (RS) has been defined as “the starch and

products of starch digestion that are not absorbed in the small

intestine of healthy individuals.” Based on the source of the

enzyme resistance, RS has been classified into five different

types (Table 1).

In type 2, type 3, and type 5 RS, the enzyme resistance is due

to the physical structure of the starch molecules. Type 2 RS is

found in raw starch granules and the enzyme resistance is due

to the natural organization of the starch within the starch

granules. Significant amounts of type 2 RS can be found in

green banana, potato, and high-amylose maize.

In type 3 RS, the enzyme resistance is due to the physical

structure of starch chains that have undergone some type of

restructuring due to retrogradation or heat treatment. Different

factors, such as amylose/amylopectin ratio, chain length, lipid

content, and processing conditions, have been shown to influ-

ence the amount and quality of type 3 RS. Annealing and heat

moisture treatment are two types of heat treatments that are

often used to create RS. Both of these treatments, as well as

gelatinization, either fully or partially melt crystalline structure

present in the native starch granules. After heat treatment, linear

amylose molecules and linear regions of amylopectin molecules

can organize into a mix of amorphous and crystalline areas with

varying degrees of enzyme resistance. Formation of the crystal-

line structures usually takes place above the glass transition

temperature and below the melting temperature, and any com-

ponents present that influence the glass transition temperature

can therefore be expected to influence the formation (yield and

quality) of the formed type 3 RS. The amylose content of starch

has been positively correlated with RS yield and the formation

of type 3 RS is strongly related to the crystallization of amylose.

The amount of RS formed is also dependent on the water

content and temperature used during the heat treatment.

Water does act as a plasticizer in the system and a minimum

water content is necessary to achieve the chain mobility needed

to form crystalline structure resistant to enzyme digestion. At

high-starch concentrations, the starch chains interact more eas-

ily, leading to increased crystal and RS formation. The presence

of lipids has been shown to decrease the formation of type 3 RS

due to formation of amylose–lipid complexes (type 5 RS).

The enzyme resistance in type 5 RS is due to the molecular

structure of amylose–lipid complexes that canbe either present in

the native starch or formed by controlled reactions using non-

granular starch and lipids to form the resistant amylose–lipid

complex.

The enzyme resistance in type 4 RS is due to chemical

modification of the starch. Chemical modification of

Encyclopedia of Food Grains http://dx.doi.org/10.1016/B978-0-12-394437-5.00097-8 1

starch creates chain irregularities or branches in the starch

chains. Cross-linking of starch covalently links two starch

chains together, in effect creating a branch point on both

chains. Chemical substitution introduces a bulky side group

to the starch chains. The introduction of these chemical groups

may create a steric hindrance to one or more human digestive

enzymes.

RS Intake

Quantification of RS

To determine the effect of RS intake on health, it is important

to determine the amount of RS consumed as accurately as

possible. Ideally, RS consumption would be determined

in vivo by quantification of undigested starch in healthy indi-

viduals. This is, however, very difficult to do. Several in vivo

approaches have been used for measurement of RS. These

in vivo methods include the hydrogen breath test, direct collec-

tion of ileal effluent from ileostomy patients, and direct col-

lection of the ileal effluent from healthy subjects using a long

triple lumen tube.

The measurement of hydrogen breath to determine the

amount of RS in a diet is based on the concept of fermentation

of dietary fibers including RS. When RS enters the colon, it may

be fermented by the colon microbiota that leads to the pro-

duction of short-chain fatty acids (SCFAs) (acetate, butyrate,

and propionate) and gases like hydrogen (methane and carbon

dioxide). Most of the gases produced by colonic fermentation

are absorbed into the circulation, but about 10% is present in

the breath on exhalation. Since hydrogen breath is not specific

for only RS fermentation, but does indeed measure fermenta-

tion from any dietary fiber source, care has to be taken when

interpreting these results for RS content in a food/diet.

Direct collection of ileal effluents would be expected to give

more accurate results than hydrogen breath measurement

especially when different forms of dietary fibers and RS may

be present in a diet. Analysis of the ileal effluent allows for

quantification of undigested carbohydrate and potential iden-

tification of the different sources of dietary fiber within the

effluent. Most ileostomy models are, however, based on the

use of patients that had their colon removed due to health

issues and their digestive patterns may therefore not be repre-

sentative of a healthy population. Analysis of ileal effluent

from healthy individuals should provide the most accurate

data, but these measurements are difficult and intrusive and

can therefore not be done on large scales.

It is generally accepted that any in vitro method used to

measure RS should give values in line with those obtained

with ileostomy patients. In 1986, Berry developed a new

in vitro method by modification of a method previously devel-

oped by Englyst, Wiggins, and Cummings in 1982. This mod-

ified method was subsequently confirmed to give results in line

with healthy ileostomy subjects. In 1992, Englyst developed a

method that could measure rapidly digestible starch (RDS),

slowly digestible starch (SDS), and RS. This method employed

incubation of the samples at 37 �C with pancreatic amylase

and amyloglucosidase and measures the amount of RS by

subtracting the sum of RDS and SDS from the total starch

present in the sample. Several new methods for the measure-

ment of RS were developed during the European Research

Program EURESTA. Even though many different methods

were developed during the 1990s for the measurement of RS,

none of these methods were successfully subjected to interla-

boratory evaluation to be adopted as an official AOACmethod

of analysis. In 2002, McCleary andManaghan developed a new

RS method by modifying existing methods to obtain a method

that would yield robust, reproducible results that correlated

with results from ileostomy patients. This method underwent

interlaboratory evaluation and was later adopted as AOAC

Official Method 2002.02. An Integrated method for the mea-

surement of total dietary fiber was later developed and was

published in 2007. This method allows the accurate measure-

ment of all dietary fibers including RS. The enzymatic incuba-

tion step uses pancreatic a-amylase and more closely simulates

digestion in the human digestive tract and yields RS values in

line with those obtained with AOAC Official Method 2002.02

and with results from ileostomy patients. This method was

accepted as AOAC Method 2009.01.

Sources of RS and Intake Levels

Of all unprocessed foods, the green banana has the highest RS

content of between 47% and 57%. While flour can be prepared

from green bananas and incorporated into different foods as a

source of RS, it is important to keep in mind that food proces-

sing methods such as cooking may gelatinize the starch gran-

ules and destroy the structures responsible for the enzyme

resistance originally present and responsible for RS formation.

Raw potato starch is also high in RS, but since raw potato starch

is seldom consumed, it is not a good source of RS. However,

cooling of cooked potatoes or potato products can lead to

retrogradation of the starch and may lead to formation of

new RS (type 3) if the right conditions (heat, moisture, time,

and starch concentration) are met. The level of RS in different

legumes can vary widely and RS content in the legumes as

consumed depends on processing and legume variety.

Murphy et al. examined the RS intake in the United States

by comparing reported RS contents of different foods with

dietary recall records from participants in the 1999–2002

National Health and Nutrition Examination Survey. They

showed that the RS concentration varies greatly, even within

the same food category. This could be due to natural differ-

ences (cultivar), differences in food preparation methods, and

also differences in analytic methods used to determine the

Table 1 Classification of resistant starch (RS)

RStype Source of enzyme resistance Examples

1 Whole grains Seeds, grains2 Raw starch granule structure Green banana, high

amylose, potato3 Structures formed on

reassociation or retrogradationof starch after heat treatment

Cooked pasta, cookedpotato

4 Chemical modifications of thestarch

Modified starch

5 Starch lipid complex Amylose-containing starch

2 CARBOHYDRATES | Resistant Starch and Health

amount of RS present. The RS content of different breads varied

between 0.1% in wheat rolls and 4.5% in pumpernickel bread.

Ready-to-eat breakfast levels contained between 0% (rice

cereal) and 6.2% (puffed wheat) of RS and cookies/crackers

had between 0.2% (oatmeal) and 2.8% (crackers and crisp

bread) of RS. The category of cooked cereal and pastas had

the widest range of RS contents between 0.4% (noodles and

chow mein) and 11.3% (oats and rolled uncooked). Within

the vegetable category, the lowest RS content was shown for

sweet corn (cooked/canned) and potatoes (slow-cooked) with

0.3% while the highest content was observed for fried potatoes

with 2.8%. In the legume category, RS content ranged from

0.6% for chickpeas (cooked/canned) to 4.2% for white beans

(cooked/canned).

Comparison of the RS content of different foods and the

food intake records showed that Americans on average con-

sumed an estimated 3–8 g of RS per day. In Europe, the intake

of RS has been estimated to be between 3 and 6 g per day while

the intake in developing countries has been estimated to be

between 30 and 40 g per day.

Health Benefits of RS

Gut Health

By definition, RS escapes digestion in the human small intes-

tine. RS therefore passes into the colon where like other dietary

fibers, it may be fermented. Fermentation of carbohydrates in

the colon leads to the production of SCFAs. SCFAs are the

metabolic products of anaerobic bacterial fermentation and

consist of butyrate, propionate, and acetate.

The SCFAs produced by bacterial fermentation in the

human gut are the preferred fuel for the cells lining the colon

(colonocytes) and have been shown to lower luminal pH,

increase the excretion of bile acids, and help prevent the devel-

opment of abnormal colonic cells. A lower pH has been asso-

ciated with protection against colorectal cancer and has been

shown to inhibit the conversion of primary to secondary bile

acids. Secondary bile acids are thought to promote tumors as

they are cytotoxic to colonic cells. It is therefore reasonable to

assume that RS may confer health benefits to gut health by

production of SCFA, which in turn have a positive effect on pH

and reduced production of secondary bile acids. A low pH has

also been found to suppress the growth of potentially patho-

genic organisms and reduce the absorption of toxic com-

pounds (such as ammonia).

Individual SCFAs have also been shown to have more spe-

cific actions in the colon. Most RS is readily fermented into

SCFA and many are a good source of butyrate. Butyrate is a

preferred energy source for colonocytes and helps drive the

uptake of electrolytes and water. Intake of butyrate producing

RS as part of oral rehydration solutions has been shown to

significantly reduce the recovery time for children with watery

diarrhea. Butyrate has also been shown to help maintain bar-

rier function in the gut by enhancing mucin production. The

barrier function is important to reduce bacterial translocation,

which may be a key element in the development of diseases

such as inflammatory bowel disease (IBD). Intake of RS has

been suggested to be beneficial for the reduction of IBD.

Butyrate has been shown to induce apoptosis, decreasing

the risk of cancer development. Animal studies have shown

that an increased supply of butyrate to the large bowel

decreases the induction of large bowel tumors in rats. How-

ever, despite the fact that the data strongly suggest the link

between butyrate and reduced risk of colonic cancer, studies

examining the link between RS intake and colonic cancer have

not been able to prove such a relationship. One reason for this

may be the importance of RS intake levels. Studies examining

the butyrate production from RS intake have shown significant

increase in fecal butyrate when 22 g of RS was consumed per

day, but no changes in butyrate or pH levels were observed

when 12.5 g of RS was consumed per day. The level of butyrate

produced may also depend on the type of RS consumed.

Another aspect of gut health often linked to dietary fibers is

laxation and fecal bulk. An increase of fecal bulk is thought to

be beneficial and is often reported as a potential health benefit

of different dietary fibers. Increases in fecal mass are important

for relief of constipation and prevention of diverticulosis and

other disorders such as hemorrhoids. The effect of RS on fecal

bulk has been studied and an increase of fecal output was

reported from different studies. Phillips et al. reported an

average increase in fecal output of 42% and also showed that

the fecal output increased with increased RS intake. This

increase fecal output could be due to increased water in the

fecal material, undigested and unfermented starch and dietary

fibers, or possible bacteria from the digestive tract. Increased

fecal nitrogen was observed in diets high in RS indicating either

an increase in bacterial mass or protein excretion.

Metabolic Effects

Several studies have shown that replacement of digestible car-

bohydrate with RS leads to a reduced blood glucose response.

This has formed the basis for approval of a European Food

Safety Authority (EFSA) claim for RS (EFSA, ID 681, April

2011). While this claim was granted, there was insufficient

evidence for the effects of RS when the glycemic load (amount

of digestible carbohydrate) remains constant. In 2012,

Robertson reviewed several studies on the effect of RS addition

to the diet when the available carbohydrate load was held

constant and noted a significant inconsistency between the

studies reviewed. While some found no effect of RS on glyc-

emia and significant reduction in insulin secretion, others

found the opposite, that is, a reduction in glycemia and an

elevation in postprandial insulinemia. The difference in these

results could be due to differences in the RS tested or simply be

due to the difficult relationship between glucose absorption,

clearance, and hepatic release, which makes accurate measure-

ments and conclusions difficult. Another important factor to

consider is the sources of RS used in different studies. RS can

come from foods such as beans, high-amylose corn starch, or

potatoes, and these foods all have different physiochemical

properties that can affect the response to RS ingestion. The fat

content of the diets is also important as fat is known to have a

significant impact on the glycemic response to ameal. A careful

review of studies with appropriate fat and protein levels

showed that RS consumption does indeed lead to a small

decrease in postprandial glycemia and a larger attenuation of

postprandial insulinemia. These acute changes in plasma

CARBOHYDRATES | Resistant Starch and Health 3

glucose and insulin concentrations caused by the addition of

RS to the diet could have important metabolic implications.

Both hyperglycemia and hyperinsulinemia are affected in the

development of insulin resistance. Increases in plasma glucose

concentrations have been linked to increased concentrations of

free fatty acids. Glucose uptake in the muscles and increased

glucose uptake in the adipose tissue have been linked to

hyperinsulinemia.

The potential chronic effects of RS intake on metabolism

and associated diseases are even more interesting than the

acute effects discussed earlier. There are animal data showing

a significant relationship of RS inclusion in the diet and

improved insulin sensitivity. Decreased insulin sensitivity, or

insulin resistance, has severe metabolic consequences as it is

strongly associated with several diseases known as metabolic

syndrome. Metabolic syndrome describes diseases such as type

2 diabetes, obesity, coronary heart disease, hypertension, and

dyslipidemia.

Even if RS only leads to reduction in blood glucose

response when digestible carbohydrates are replaced, such an

effect may have significant benefits to human health. Diabetes

affects about 8% of US population and is increasing globally.

Diabetes is characterized by hyperglycemia that can subse-

quently lead to systemic tissue toxicity. Risk factors for diabetes

include increased glucose response (both fasting and postpran-

dial), decreased insulin sensitivity, and obesity. These risk

factors may be reversible with lifestyle modifications that

have been shown to be effective in delaying the onset of type

2 diabetes. One lifestyle change could be to reduce the glyce-

mic load of the diet that could be achieved by replacing ordi-

nary starch in foods with RS.

RS has been shown to have significant effects of serum

triglyceride and cholesterol concentrations. Several animal

studies have shown significantly lower plasma cholesterol and

triglyceride concentrations when RS is included in the diet in

replacement of digestible starch. A decrease in fasting choles-

terol and triglycerides was observed when RS was consumed

chronically (5–14 weeks). The mechanism responsible for

these effects seems to be increased bile acid excretion caused

by consumption of RS, but some study showing reduced

plasma cholesterol and triglyceride concentrations even com-

pared to the bile acid sequestrant cholestyramine suggests that

the effects of RS extend beyond just bile acid secretion. The

effects of RS on cholesterol and blood triglyceride levels seem

to be mediated through a combination of enhancement of bile

acid secretion, a significant decrease in cholesterol absorption,

and an increase in hepatic LDL receptor expression. Both high

blood lipid and cholesterol levels are associated with coronary

heart disease and the effects of RS consumption may therefore

have important health ramifications.

The effects of RS consumption on satiety and weight man-

agement are not as clear. One major reason for the current

global obesity epidemic is the overconsumption of energy, and

therefore, strategies that can manage the energy intake may be

successful in reducing obesity. Dietary fiber is one way to

decrease the energy value of foods if it is used to replace

digestible carbohydrates that have a higher caloric value.

Some dietary fibers have also been linked to increased satiety

and lower BMI, but not all fibers have the same effects. RS is a

type of dietary fiber and it would therefore be reasonable to

expect some effect on satiety and/or total energy intake when

RS is used to replace digestible starch in the diet. Animal

studies have shown the production of satiety hormones

GLP-1 and PYY when RS was added to the diet, which suggests

an effect of RS on satiety. However, only a few human studies

have been conducted to examine the effect of RS on satiety and

the results were mixed. Some studies have shown an increase in

satiety after RS consumption, but others have shown little or

no effect. Satiety seems to be linked to changes in plasma

glucose concentrations and studies that showed a decrease in

plasma glucose/insulin response to a high RS meal also

showed an increase in satiety while those studies that showed

no change in plasma glucose also showed no difference in

satiety. Overall, there is indication that there is at least a weak

association between RS consumption and satiety. Some animal

studies have shown reduced energy intake when RS was added

to the diet. Aziz et al. showed that body weight was reduced by

40% in obese rats that were fed with a diet high in RS com-

pared with rats fed with a low-RS diet. While this study seems

to indicate a real positive effect of RS on energy intake and

potentially obesity management, the levels of RS used for this

study (23.4%) may not be realistic in a human diet. Long-term

studies in humans with realistic RS inclusion levels are

required to get a better understanding of the potential for RS

to help in the management and reduction of the current obe-

sity epidemic.

Summary/Conclusions

RS is a dietary fiber and can be used to replace digestible starch

in foods. Many animal studies have been conduced to examine

the effect of RS consumption. RS may be fermented in the

colon and the resulting SCFA production has been linked to a

number of health benefits such as reduced pH, which has been

associated with the suppression of growth of potentially path-

ogenic organisms and reduction of the absorption of toxic

compounds. Fermentation of RS has been shown to produce

relatively high levels of the SCFA butyrate that has been linked

to reduced risk of colon cancer. RS fermentation has also been

linked to benefits in the risk reduction of IBD by strengthening

of the gut barrier via increased production of mucin. RS has

been shown to increase fecal bulk, which is important for relief

of constipation and prevention of diverticulosis and other

disorders such as hemorrhoids.

An EFSA claim was granted in 2011 for reduced blood

glucose response when RS is used to replace digestible carbo-

hydrate in the diet (EFSA, ID 681, April 2011). The effect of RS

on glycemia and insulinemia is less clear, and studies have

shown that different effects are possible due to study designs

(e.g., amount of fat and protein in diets) and amount of RS

tested. Overall, it does seem that RS consumption does indeed

lead to a small decrease in postprandial glycemia and a more

significant attenuation of postprandial insulinemia that could

have important metabolic implications. Both hyperglycemia

and hyperinsulinemia are affected in the development of

insulin resistance, which in turn has been linked to several

diseases known as metabolic syndrome, for example, type 2

diabetes, obesity, coronary heart disease, hypertension, and

dyslipidemia.

4 CARBOHYDRATES | Resistant Starch and Health

RS has been shown to have significant effects on serum

triglyceride and cholesterol concentrations. The effects of RS

on cholesterol and blood triglyceride levels seem to be medi-

ated through a combination of enhancement of bile acid secre-

tion, a significant decrease in cholesterol absorption, and an

increase in hepatic LDL receptor expression. Both high blood

lipid and cholesterol levels are associated with coronary heart

disease and the effects of RS consumption may therefore have

important health ramifications.

The effects of RS consumption on satiety and weight man-

agement are not as clear. RS is a type of dietary fiber and it

would therefore be reasonable to expect some effect on satiety

and/or total energy intake when RS is used to replace digestible

starch in the diet. Only a few human studies have been con-

ducted to examine the effect of RS on satiety and the results

were mixed. Overall, there is indication that there is at least a

weak association between RS consumption and satiety, but

long-term studies in humans with realistic RS inclusion levels

are required to get a better understanding of the potential for

RS to help in the management and reduction of the current

obesity epidemic.

Exercise for Revision

• What are the different types of resistant starch?

• What is the link between resistant starch consumption and

colon cancer?

• How does resistant starch affect insulin sensitivity and why

is this important?

Exercises for Readers to Explore the Topic Further

While many animal studies have been conducted examining

the relationship of resistant starch intake and various health

benefits, a more limited number of human clinical studies

have been conducted. The results of these clinical studies are

sometimes contradictory, and strong conclusions on the health

benefits of resistant starch are therefore difficult in some areas.

More human studies with careful study designs are needed in

the future.

See also: The Basics: The Grains that Feed the World; Grainsaround the World: Grain Production and Consumption, Overview;Food Grains and the Consumer: Grains and Health; Food Grainsand Well-being: Functional Foods, Overview; Food Grains andthe Consumer: Consumer Trends in Consumption; Grains and Health– Misinformation and Misconceptions; ; Carbohydrates:Carbohydrate Metabolism; Beta glucans and health.

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