Bioavailability of Nutrients
HC Schönfeldt, B Pretorius, and N Hall, University of Pretoria, Pretoria, South Africa
ã 2016 Elsevier Ltd. All rights reserved.
Background
Correct assessment of the adequacy of nutrient intakes from
the diet requires not only knowledge about nutrient content of
ingested foods but also the extent to which the nutrient is
available for absorption and utilization in the human body.
Bioavailability is the technical term used to convey the fact
that not all of the nutrients ingested will be absorbed,
irrespective of whether consumed in the form of food or sup-
plements. Bioavailability aims to describe the effects of a
sequence of metabolic events, including digestion, solubili-
zation, absorption, uptake and release, enzymatic transforma-
tion, secretion, and excretion, on nutrient utilization. Thus, the
supply of nutrients – and increasingly nonnutrient bioactive
compounds – to the human body depends not only on the
amount in a food but also on its bioavailability. Understand-
ing bioavailability helps to optimize diets and set appropriate
recommendations.
The bioavailability of macronutrients, that is, carbohy-
drates, proteins, and fats, is usually high with more than 90%
of the amount ingested being absorbed and utilized in the
body. On the other hand, for micronutrients, such as vitamins
and minerals, and bioactive phytochemicals such as flavonoids
and carotenoids, absorption and utilization can vary widely
after ingestion.
Defining Bioavailability
Until a nutrient passes from the digestive system into the
bloodstream, it has little or no value. Bioavailability can be
explained as the amount of a nutrient absorbed from the gut
that becomes available for normal physiological functions or
storage.
The Variability of Nutrient Bioavailability
The bioavailability of nutrients is highly variable and can be
influenced by numerous factors, including physiochemical
properties, such as chemical binding form; the matrix in
which the nutrient is incorporated; the presence or absence of
other food components that enhance or inhibit absorption;
metabolization after absorption; host-related factors (includ-
ing state of health, genetic factors, age, and lifestyle); and other
individual factors.
Enhancers and Inhibitors
Nutrients can interact with one another and with other dietary
components at the site of absorption, affecting a change in
bioavailability or – if enhancers and inhibitors cancel each
other out – a nil effect. Enhancers can act in different ways,
Encyclopedia of Food and Health http://dx.doi.org/10.1016/B978-0-12-384947-2.00068-4
such as keeping a nutrient soluble or protecting it from inter-
action with inhibitors. For example, because carotenoids are
fat-soluble, adding small quantities of fat or oil to a meal
(3–5 g) improves their bioavailability. Similarly, meat, fish,
and poultry, while containing highly bioavailable iron, are
also known to enhance the absorption of iron from other
foods ingested at the same time. Although this meat factor has
yet to be identified, it has been suggested that muscle protein
may exert an influence.
Inhibitors, on the other hand, may reduce bioavailability
by binding the nutrient in a form that is not recognized by
uptake systems on the surface of intestinal cells, rendering the
nutrient insoluble and, thus, unavailable for absorption, or by
competing for the same uptake system. For example, phytic
acid is abundant in certain plant foods (e.g., pulses, whole-
grain cereals, seeds, and nuts) and strongly binds minerals,
such as calcium, iron, and zinc in soluble or insoluble com-
plexes, which are unavailable for absorption. Ways to reduce
phytic acid content of foods include fermentation (e.g., exten-
sive leavening of whole-wheat bread dough) or the soaking
and germination of pulses.
The inhibitory effect of food constituents can also be used
advantageously, as in the case of phytosterols. These com-
pounds can be extracted from certain plant foods and added
in higher doses (about 2 g per portion) to a variety of other
foods (e.g., spreads and fermented milk drinks) to reduce the
absorption of cholesterol, be it from dietary sources or pro-
duced in the human body.
Bioavailability of Specific Nutrients
Protein–energy malnutrition, vitamin A, and iron deficiencies
are some of the most common forms of malnutrition experi-
enced globally and often coexist. For effective intervention and
dietary guidelines, the bioavailability of these nutrients, as
supplied from different food sources, is particularly important.
The bioavailability of these nutrients is distinct and influenced
by different factors, providing good examples of the complex-
ity of nutrient bioavailability.
Protein and Amino Acids
Although protein is a macronutrient that is easily absorbed by
the human body, its bioavailability is directly linked to digest-
ibility. To be most bioavailable, a meal needs to supply all the
required essential amino acids in the correct proportions.
Amino acids are the central units in protein metabolism.
They are incorporated into various proteins and converted to
metabolically essential compounds, such as nucleic acids,
creatine, and porphyrins. Of the 20 amino acids in human
proteins, 12 can be made by the body and are known as non-
essential amino acids. The remaining eight (8) (isoleucine,
401
402 Bioavailability of Nutrients
leucine, lysine, methionine, phenylalanine, threonine, trypto-
phan, and valine) must be obtained from the diet and, thus,
are termed essential or indispensable amino acids. Sufficient
intakes of essential amino acids and adequate amounts of
nitrogen for the body to produce the nonessential amino
acids are important for protein metabolism.
Protein quality
The nutritional quality of food proteins varies, depending on
the essential amino acid composition. Foods that contain
essential amino acids at levels that facilitate tissue growth and
repair are known as complete protein foods, supplying high-
quality proteins. Amino acids containing sulfur (including
methionine and cysteine) most commonly limit the nutri-
tional value (quality) of proteins in the human diet. Concen-
trations of these sulfur-containing amino acids are, generally,
considered lower in legumes and fruits than in food of animal
origin. The roles of these amino acids in the human body are
crucial as methionine is the amino acid initiating the synthesis
of almost all eukaryotic proteins and cysteine (due to its ability
to form sulfur bonds) has an important role in protein struc-
ture. Other essential amino acids, lysine and tryptophan, are
also consistently found at lower concentrations in plant-based
rather than animal-based foods; for example, tryptophan and
lysine are limiting in corn; lysine in wheat, sorghum, and other
cereals; and methionine in soybeans and other legumes. For
further reading on the global protein quality debate, refer to
the 2011 report of the FAO Expert Consultation on dietary
protein quality evaluation in human nutrition (FAO, 2011).
In addition to protein quality, digestibility (absorption),
chemical integrity, and inhibitors are three key properties of
foods that can influence the bioavailability of amino acids.
Amino acid digestibility
Amino acid digestibility explains the proportion of amino
acids consumed that is absorbed. It is not a fixed value, but
reflects the interaction between food and the host and, there-
fore, may be subject to individual variation. Although the
digestibility (absorption) of macronutrients, including pro-
tein, is relatively high, protein utilization is influenced by
total dietary energy intake and the quality of protein in terms
of meeting metabolic demand.
Chemical integrity
Chemical integrity describes the proportion of amino acids
absorbed in a form that can be utilized. Some amino acids
present in foods may be structurally unavailable (i.e., absorbed
in a form that cannot be utilized). This is most likely to be
encountered in foods that are heat-treated, oxidized, or sub-
jected to other processes that limit amino acid bioavailability.
Heat treatment leads to the formation of Maillard compounds
and reduced lysine availability. Oxidization of sulfur-
containing amino acids reduced the bioavailability of
tryptophan and threonine. High pH induces racemization of
L-amino acid residues to D-isomers and formation of cross-
linked amino acids, such as lysinoalanine, which also reduces
bioavailability.
Inhibitors
Many foods contain bioactive (protein or nonprotein) sub-
stances that may inhibit amino acid bioavailability by affecting
either digestibility or utilization postabsorption. These inhibi-
tors may be naturally occurring (e.g., tannins, phytates, trypsin
inhibitors, glucosinolates, and isothiocyanates), formed dur-
ing processing (e.g., D-amino acids and lysinoalanine), or
formed during genetic modification of crops (e.g., lectins in
lentils) (lectins suppress growth at low levels and are toxic at
high levels).
Vitamin A
Vitamin A is a generic term used for a group of structurally
related chemical compounds known as retinoids, which may
be naturally occurring or synthetic compounds with or without
vitamin A biological activity. Figure 1 shows the chemical
structures of some retinoids.
Vitamin A is often used as a general term for all compounds
that exhibit the biological activity of retinol. Vitamin A activity
in the diet derives from two sources: preformed vitamin A, such
as retinyl esters or retinoids, and provitamin A carotenoids,
such as b-carotene, a-carotene, and b-cryptoxanthin. Although
an essential nutrient, vitamin A is needed only in small
amounts and is necessary for normal functioning of the visual
system, growth and development, and maintenance of epithe-
lial cellular integrity, immune function, and reproduction.
Retinol (preformed vitamin A)
Vitamin A in vivo is, generally, found in the free alcohol form
(retinol) or esterified with a fatty acid (retinyl ester). Available
in a pure form by chemical synthesis or as vitamin A palmitate
or acetate, it is a pale yellow solid, which dissolves freely in oils
and fats, but is insoluble in water. When vitamin A intake is
adequate, more than 90% of the total body vitamin A is located
in the liver, which releases the nutrient into the circulation as
needed. The major dietary forms of vitamin A are long-chain
fatty acid esters of retinol, commonly found in foods of animal
origin, such as glandular meats, liver and fish liver oils (espe-
cially), egg yolk, and whole milk and dairy products.
Preformed vitamin A is absorbed in the small intestine.
Generally, the bioavailability of retinol is high, ranging from
70% to 90%. Factors such as dietary fat and intestinal infec-
tions can affect the absorption of vitamin A. Products of fat
digestion (e.g., fatty acids, monoglycerides, cholesterol, and
phospholipids) and secretions in bile (e.g., bile salts and
hydrolytic enzymes) are essential for the efficient solubili-
zation of retinol. The absorption of retinol appears to be
reduced in individuals with diarrhea and intestinal infections
or infestations.
Carotenoids
Carotenoids are lipid-soluble plant pigments found in photo-
synthetic plants and animal tissues. About 600 carotenoids
have been isolated and characterized in nature, and about
10% of these can be metabolized to vitamin A in a variety of
animal species, including humans. Both provitamin A carot-
enoids, such as a- and b-carotenes and cryptoxanthins, and
non-provitamin A carotenoids, such as lutein, zeaxanthin, and

H3C CH3 CH3 CH3
CH3
all-trans retinol
H 3C CH3 CH3 CH3
CH3
retinyl ester
H3C CH3 CH3
CH3 H3C
11-cis retinal
Figure 1 Chemical structures of different retinoids. All-trans retinol is by definition vitamin A, and 1 mg of all-trans retinol is equal to 1 retinol
equivalent (RE). When a fatty acyl group is esterified to the hydroxyl terminus of all-trans retinol, retinyl ester is formed for storage. The most abundant
retinyl esters are those of palmitic, oleic, stearic, and linoleic acids. Retinyl acetate and palmitate are often used as dietary supplements, but do not
occur naturally. Retinol can be reversibly oxidized to retinal, the 11-cis isomer, which is essential for the visual cycle. Reproduced from Packer, L.,
Kraemer, K., Obermüller-Jevic, U. and Sies, H. (eds.) (2005). Carotenoids and Retinoids: Molecular aspects and health issues, Illinois: AOCS Press.
lycopene, are present in the human blood and tissues and have
a variety of functions. The structures of these carotenoids are
shown in Figure 2. Provitamin A carotenoids are an important
source of dietary vitamin A that are found primarily in dark-
green leafy vegetables, such as spinach, and in orange and
yellow vegetables and fruit, such as carrots, mango, and
papaya, although their bioavailability is significantly more
variable than that of preformed vitamin A (retinol).
The bioavailability of carotenoids is affected by various
factors. Different carotenoids have different levels of vitamin
A activity depending upon the efficiency of their absorption and
the rate of their conversion to vitamin A. Recent research has
shown that the bioavailability of traditional dietary sources of
b-carotene is considerably lower (around half to a quarter) than
was previously assumed. Conversion factors for estimating vita-
min A obtained from plant foods were revised from 6:1 to 12:1
(mg b-carotene–retinol activity equivalent (RAE)) and 24:1 for
other provitamin A carotenoids in a mixed diet. There is also a
wide variation in vitamin A equivalency ratios, which can be
affected by food- and diet-related factors and health, nutri-
tional, and genetic characteristics among human populations.
Various diet- and host-related factors affect the bioavailabil-
ity of carotenoids. These factors have been evaluated and
extensively reported by Castenmiller and West (1998), De
Pee et al. (1998), Van Het Hof et al. (2000), and Yeum and
Russell (2002).
Bioavailability of Nutrients 403
OH
O
O
R CH3
O
Acetate
R
O
R CH2(CH2)13 CH3
Palmitate
N
CH3
Bioavailability of carotenoids
The main diet-related factors influencing carotenoid bioavail-
ability are the food matrix, amounts ingested, and habitual diet
type. The nutritional status, health, and genetic characteristics
of human populations can also affect the absorption and bio-
availability of carotenoids.
Release of the carotenoids from the food matrix is important in
the absorption process. The rupture of the plant cell walls by
processing (e.g., heating or pureeing) promotes the release of
b-carotene from the cells before and during digestion and, there-
fore, facilitates solubilization and absorption. Generally, the
bioavailability of b-carotene from fruits is higher than for vegeta-
bles, as the cell wall structure in fruits is usually weaker than in
most vegetables and leaves. Furthermore, inhibitors of carotenoid
absorption present in fruit are also less than in leafy vegetables.
The composition of the diet (due to nutrient-to-nutrient
interactions) affects, to a large extent, the absorption of carot-
enoids. The second step in absorption, which may affect bio-
availability, involves the incorporation of carotenoids into
mixed micelles. Among other factors, the formation of these
micelles is dependent on the presence of fat in the intestine.
Therefore, ingestion of fat, along with carotenoids, is thought
to be crucial although only a small amount of fat is necessary to
enhance carotenoid absorption. As expected, fat-soluble com-
pounds that cannot be absorbed, such as sucrose polyester
(a fat replacer), reduce carotenoid absorption. Also, as dietary
404 Bioavailability of Nutrients

H3C
H3C CH3 CH3 CH3

CH3 CH3 H3C CH3

CH3
a-Carotene

CH3
CH3 CH3 CH3
H3C

CH3
CH3 CH3 CH3

CH3
b-Carotene

H3C
H3C CH3 CH3 H3C

CH3 H3C H3C CH3

HO CH3
b-Cryptoxanthin

H3C OH
H3C CH3 CH3 CH3

CH3 CH3 H 3C CH3

HO CH3
Zeaxanthin

H3C OH
H3C CH3 CH3 CH3

CH3 CH3 H3C CH3

HO CH3
Lutein

H3C OH
H3C CH3 CH3 CH3

CH3 CH3 H 3C CH3

HO CH3

Lycopene
Figure 2 Chemical structures of major provitamin A carotenoids (a-carotene, b-carotene, and b-cryptoxanthin) and non-provitamin carotenoids
(lutein, zeaxanthin, and lycopene) found in food.

fiber content increases, the absorption of b-carotene decreases.
Dietary fiber reacts with bile acids and, thereby, decreases the
absorption of fat and fat-soluble nutrients. The presence of
dietary fiber in vegetables and fruits may explain in part the
reduced bioavailability of carotenoids from plant foods.
Simultaneous ingestion of carotenoids may also reduce
absorption of any of the carotenoids due to interactions at
the intestinal level. Studies on simultaneous ingestion of carot-
enoids indicate that lutein may interfere with the absorption of
b-carotene resulting in reduced bioavailability. It has also been
found that with pharmaceutical doses of b-carotene, conver-
sion of b-carotene to vitamin A decreases as the oral dose of b-
carotene increases. Further research is required to identify the
mechanisms behind these interactions.
Furthermore, the absorption of carotenoids is highly likely
to be dependent on vitamin A status of the host. Consuming
b-carotene-rich foods leads to an increase in serum retinol
levels only when these are initially low. The serum response

to b-carotene is higher in women than in men; however, this
could in part be attributed to differences in body weight and
composition. Intestinal helminthic infections are associated
with malnutrition, which may be mediated through impaired
fat absorption and reduced vitamin absorption, particularly of
vitamin A.
Iron
Minerals (and other nutrients) exist in different chemical forms
in food, which can influence bioavailability. Iron is a classic
example: there are two primary forms of dietary iron, namely,
heme and nonheme. The former is only found in animal
products, such as red meat, fish, and poultry. The heme iron
content of animal sourced foods is estimated at 40% of the
total iron, but data suggest that a portion of red meat provides
considerably more heme iron than a portion of white fish, for
example. Heme iron is a component from hemoglobin and
myoglobin (see Figure 3), which explains why it is only found
in animal tissue. Nonheme iron is found mostly in plant-based
foods and makes up the remaining estimated 60% of iron
found in animal products.
The types of iron (heme or nonheme) notably influence
bioavailability. Approximately 90% of dietary iron is con-
sumed in the nonheme form. However, due to low bioavail-
ability, it constitutes only approximately half the iron absorbed
by the body. The absorption of nonheme iron is usually much
lower than heme iron. In general, the rate of nonheme iron
absorption is related to its solubility in the upper part of the
small intestine. The presence of soluble enhancers, such as
ascorbic acid, and inhibitors, such as phytates, polyphenols,
and calcium, consumed in the same meal will have a notable
effect on the amount of nonheme iron absorbed. Heme iron is
much less affected by other dietary factors and contributes
more significantly to absorbable iron.
Inhibitors and enhancers of iron bioavailability
Nonheme iron that enters the common iron pool in the diges-
tive tract is absorbed to the same extent, depending on the
balance between inhibitors and enhancers and the iron status
of the individuals.
OH
O inverse relationship between iron status and iron absorption.
CH3
O
N CH3
HO
N- Fe2+ N-
CH2
H3C N cidin is a regulatory hormone secreted by the liver, which
CH3
CH2
Figure 3 Chemical structure of heme iron as found in food from animal
sources.
Bioavailability of Nutrients 405
Phytate and polyphenols in plant-based diets are the main
inhibitors of nonheme iron absorption. The negative effect of
phytate on iron bioavailability is dose-dependent, and any
food processing and preparation methods, such as milling,
heating, soaking, germination, and fermentation, which
degrade phytate to varying extents, will have a positive effect
on iron absorption. Controversies exist on the inhibitory effect
of oxalic acid in spinach and cabbage and nondigestible car-
bohydrates in beans on iron absorption, as these foods are also
good sources of ascorbic acid, which enhances iron absorption.
Calcium and dairy products have also been shown to have a
negative effect on nonheme iron absorption, but what separates
these from other inhibitors is the ability to also inhibit heme
iron absorption. Single-meal studies show a negative effect of
calcium on iron absorption, but multimeal studies, with a vari-
ety of other inhibitors and enhancers, indicate calcium has only
limited impact on iron absorption. In a recent study, the two
major milk protein fraction (casein and whey) and egg protein
(albumin) were reported to have a negative effect on iron
absorption in humans. Although phytate has been shown to
be the major inhibitor in soy, even after complete phytate
digestion in soy protein isolates, significant inhibition of iron
absorption can be observed. Thus, in soy, it was concluded that
both phytate and a protein fraction inhibit iron absorption.
Ascorbic acid has been shown convincingly to enhance iron
bioavailability in a dose-dependent manner. This effect is
largely due to its ability to reduce ferric to ferrous iron. Ascorbic
acid has also been shown, at least partially, to counteract the
inhibitory effects of both phytate and polyphenols on non-
heme iron absorption.
Small amounts of meat are recognized to enhance the
absorption of nonheme iron from plant foods, although the
mechanism(s) is unknown. Studies also support the enhancing
effect of cysteine-containing peptides following the proteolysis
of meat muscle.
Vitamin A and b-carotene can enhance nonheme iron
absorption and increase hemoglobin levels, although several
studies suggest this is only observed in iron-deficient individ-
uals. Host-related factors that influence the absorption of heme
and nonheme iron include mainly iron status, other nutritional
deficiencies, infection, genetic disorders, and physiological state.
As with inhibitors and enhancers, the iron status of an
individual mainly influences the absorption of nonheme
iron, while heme iron absorption is less affected. There is an
Protein–energy malnutrition, riboflavin, and vitamin A defi-
ciencies have also been shown to impair iron metabolism and
absorption; correction of these nutritional deficiencies will
improve iron absorption.
Iron deficiency often coexists in a burden of disease in
seemingly well-fed, overweight populations, due in part to
iron absorption being reduced by the peptide, hepcidin. Hep-
inhibits iron absorption. It secretion is increased in chronic
inflammation and obesity.
Achlorhydria might also be a substantial cause of iron
deficiency, mainly in elderly people, among whom atrophic
gastritis is common and gastric acid secretion is low. Gastric
acid is needed to maintain ferric iron in solution and make it
bioavailable. However, heme iron does not appear to be
406 Bioavailability of Nutrients
affected by the lack of acid and is normally absorbed in indi-
viduals with atrophic gastritis.
Other common causes of reduced iron absorption and/or
iron deficiency are mucosal atrophy in celiac disease and,
possibly, Helicobacter pylori infection, although no consensus
has been reached. For further reading on iron bioavailability,
refer to Heath and Fairweather-Tait (2002), Zimmermann and
Hurrell (2007), and Hurrell and Egli (2010).
Conclusions
Nutrients react differently once ingested and absorption can be
influenced by a variety of factors, including quality of the food
source and the matrix in which it is consumed; the composi-
tion of the whole meal, inhibitors, and enhancers; and the
status of the host. Although bioavailability is only a partial
measure of the benefit from a nutrient, this factor quantifies
the amount entering the bloodstream. Once in the blood-
stream, the nutrient must cross cell membranes before it can
nourish cells. In addition to nutrient content of foods, nutrient
bioavailability should also be taken into consideration when
nutrition-sensitive policies, nutrition interventions, and die-
tary guidelines are developed. However, it should be noted
that bioavailability is not a constant value and needs to be
considered with caution in the context of multiple factors,
both intrinsic and extrinsic, which can affect the bioavailability
from food- and nonfood sources of nutrients.
See also: Amino Acids: Metabolism; Ascorbic Acid: Physiology and
Health Effects; Carotenoids: Occurrence, Properties and Determination;
Carotenoids: Physiology; Iron: Biosynthesis and Significance of Heme;
Protein: Digestion, Absorption and Metabolism; Retinol: Physiology;
Retinol: Properties and Determination.
Further Reading
Castenmiller JJM and West CE (1998) Bioavailability and bioconversion of carotenoids.
Annual Review of Nutrition 18: 19–38.
Castenmiller JJM, West CE, Linssen JPH, van het Hof KH, and Voragen AGJ (1999) The
food matrix of spinach is a limiting factor in determining the bioavailability of ß-carotene
and to a lesser extent of lutein in humans. Journal of Nutrition 129: 349–355.
Dary O and Mora JO (2002) Food fortification to reduce vitamin a deficiency:
international Vitamin A consultative group recommendations. Journal of Nutrition
132: 2927S–2933S.
De Pee S, West CE, Permaesih D, Martuti S, Muhilal, and Hautvast JGAJ (1998) Orange
fruit is more effective than are dark-green leafy vegetables in increasing serum
concentrations of retinol and ß-carotene in schoolchildren in Indonesia. American
Journal of Clinical Nutrition 68: 1058–1067.
FAO (2011) Dietary protein quality evaluation in human nutrition. Report of an FAO
Expert Consultation. 31 March – 2 April, Auckland, New Zealand. FAO Food and
Nutrition Paper 92’, Food and Agricultural Organisation, Rome.
Friedman M (1996) Nutritional value of proteins from different food sources. Journal of
Agricultural and Food Chemistry 44: 6–29.
Harrison EH (2012) Mechanisms involved in the intestinal absorption of dietary
vitamin A and provitamin A carotenoids. Biochimica et Biophysica Acta
1821: 70–77.
Haskell MJ (2012) The challenge to reach nutritional adequacy for vitamin A: ß-carotene
bioavailability and conversion – evidence in humans. American Journal of Clinical
Nutrition 96(Suppl): 1193S–1203S.
Heath A-L and Fairweather-Tait SJ (2002) Clinical implications of changes in the
modern diet: iron intake, absorption and status. Best Practice & Research. Clinical
Haematology 15(2): 225–241.
Hurrell R and Egli I (2010) Iron bioavailability and dietary reference values. American
Journal of Clinical Nutrition 91(Suppl): 1461S–1467S.
Millward DJ, Layman DK, Tomé D, and Schaafsma G (2008) Protein quality
assessment: impact of understanding of protein and amino acid needs for
optimal health. American Journal of Clinical Nutrition 87(Suppl):
1576S–1581S.
Otten JJ, Hellwig JP, and Meyers LD (2006) Dietary reference intakes: the essential
guide to nutrient requirements. Washington: Institute of Medicine, ISBN 0-309-
10091-7.
Packer L, Kraemer K, Obermüller-Jevic U, and Sies H (eds.) (2005) Carotenoids and
retinoids: molecular aspects and health issues Illinois: AOCS Press.
van het Hof KH, West CE, Weststrate JA, and Hauvast JGAJ (2000) Dietary factors
that affect the bioavailability of carotenoids. Journal of Nutrition 130(3):
503–506.
Yeum KJ and Russell RM (2002) Carotenoids bioavailability and bioconversion. Annual
Review of Nutrition 22: 483–504.
Zimmermann MB and Hurrell RF (2007) Nutritional iron deficiency. Lancet
370: 511–520.
Relevant Websites
http://www.eufic.org/article/en/artid/Nutrient-bioavailability-food/ – European Food
Information Council.
http://www.fao.org – Food and Agricultural Organization of the United Nations .
http://www.harvestplus.org – HarvestPlus.
http://www.ifpri.org – International Food Policy Research Institute.
http://www.usda.gov – United States Department of Agriculture.
http://www.who.org – World Health Organization.