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Bioavailability of dietary minerals

Article  in  Biochemical Society Transactions · September 1996

DOI: 10.1042/bst0240775 · Source: PubMed

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6 Blunden, G., Roch, 0. G., Rogers, D. J., Coker,
R. D., Bradburn, N. and John, A. E. (1991) Med.
1,ab. Sci. 48, 271-282
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Poisonous Plants in Britain, HMSO, London
9 Schlimrne, E. and Meisel, H. (1995) Nahrung 39,
1-20
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Allergy Immunol. 6,95-97
11 Littlewood, J. T., Gibb, C., Glover, V., Sandler, M.,
Davies, P. T. and Rose, F. C. (1988) Lancet i,
558-559
12 Peatfield, R. C. (1995) Headache 35,355-357
13 British Nutrition Foundation (1995) Iron Nutri-
tional and Physiological Significance, Chapman &
Hall, London
14 Fraser, D. R. (1995) Lancet 345, 104-107
15 Wasserman, R. H., Henion, J. D., Haussler, M. R.
and McCain, T. A. (1976) Science 194, 853-855
16 Gomez-Sanchez, C. E., Gomez-Sanchez, E. P. and
Yamakita, N. (1995) Semin. Nephrol. 15, 106-115
17 Anonymous (1995) MRC Institute for Environment
and Health IEH Assessment on Environmental
Oestrogens: Consequences to Human Health and
Wildlife, Assessment Al, Page Bros., Norwich
Bioavailability of dietary minerals
S. J. Fairweather-Tait
Institute of Food Research, Norwich Laboratory, Norwich Research Park, Colney, Norwich NR4 7UA, U.K.
Introduction
Many inorganic nutrients, including iron, zinc
and calcium, are incompletely and variably ab-
sorbed from the diet, and these differences in
behaviour can be characterized in terms of bio-
availability. T h e tems ‘bioavailability’ or ‘avail-
ability’ are sometimes used loosely to describe a
number of ill-defined measurements relating to
the absorption, retention or utilization of
minerals, and even, misguidedly, to data gener-
ated from in vitro studies. T h e concept of bio-
availability was originally derived from
pharmacokinetic studies where measurements of
the uptake of drugs into the circulation are
made. However, with inorganic nutrients present
in the diet, the generally accepted definition of
bioavailability is the proportion of the total that is
utilized for normal body functions. Mineral bio-
availability is determined by dietary factors
Bioactive Components of Food
18 Brown, C. A., Bolton-Smith, C., Woodward, M. and
Tunstall-Pedoe, H. (1993) J. Epidemiol. Commu-
nity Health 47, 171-175
19 Heckers, H., Gobel, U. and Kleppel, U. (1994)
J. Intern. Med. 235, 192-193 775
20 Sanders, T. A. (1993) Proc. Nutr. SOC.52, 457-472
21 Sanders, T. A. (1991) Proc. Nutr. Soc. 50, 513-517
22 Doll, R. and Peto, R. (1981) J. Natl. Cancer Inst.
66, 1191-1308
23 Thorogood, M., Mann, J., Appleby, P. and
McPherson, K. (1994) Br. Med. J. 308, 1667-1670
24 Block, G., Patterson, B. and Subar, A. (1992) Nutr.
Cancer 18, 1-29
25 Rowe, P. M. (1996) Lancet 347,69
26 Haapakoski, J., Rautalaahti, M., Hartman, A. M.
and Palmgren, J. (1995) Am. J. Clin. Nutr. 62,
1427s-1430s
27 Key, T. J. (1994) Proc. Nutr. SOC.53,605-614
28 Forman, D. (1991) in Relevance to Human Cancer
of N-nitroso Compounds, Tobacco Smoke and
Mycotoxins (O’Nell, I. K., Chen, J. and Bartsch,
H., eds.), pp. 22-32, International Agency for
Research on Cancer, Lyon
29 Hague, A., Elder, D. J., Hicks, D. J. and Paraskeva,
C. (1995) Int. J. Cancer 60,400-406
Received 18 March 1996
known to enhance or inhibit uptake into the
body, as modified by host-related variables which
regulate absorption and subsequent metabolism
of the element. T h e complexity of these inter-
actions necessitates careful planning of experi-
mental protocols that are clearly focused on
specific questions.
Methodological approaches
Iron was the first mineral to be selected for
studies on bioavailability. Bioassays for iron were
first published in 1926, shortly followed by
chemical tests for ‘available iron’. A wide range
of techniques have subsequently been developed
to investigate iron bioavailability [ 13, as summar-
ized in Table 1. Iron is the only inorganic
nutrient for which it is possible to obtain a direct
measure of bioavailability; this is accomplished
using radio- or stable isotopes of iron to label the
I996
 Biochemical Society Transactions
Table I
Techniques used to assess mineral bioavailability
Faecal monitoring following oral isotopes (luminal
776
disappearance)
Facecal and urinary monitoring following oral isotopes
(retention)
True absorption measured by a double-isotope technique
(calcium)
Intestinal perfusion using a triple lumen tube (zinc)
Plasma tolerance curves following oral isotopes
Rate of repletion following depletion (e g haemoglobin
repletion with iron)
Haemoglobin incorporation of iron isotopes (iron)
In vitro methods, e g dialysable iron intestinal vesicle
preparations, cell lines (e g CaCo-2)
food iron and follow its utilization in the body.
Since most newly absorbed iron is incorporated
into the haemoglobin of reticulocytes within
10- 14 days, haemoglobin incorporation of iso-
topically labelled iron can be used to quantify
bioavailability. T h e method depends on estimates
of blood volume and percentage incorporation of
absorbed iron into red blood cells, both of which
can be accurately measured using radioisotopes
~ ~ 3 1 .
T h e bioavailability of other inorganic
nutrients is more difficult to study, and assess-
ments of bioavailability are generally based upon
measurements of absorption using isotopic
labels. Faecal monitoring can be used to measure
the luminal disappearance of isotopes, and
apparent absorption is calculated by subtracting
the quantity of isotope appearing in the faeces
from the oral dose [eqn. (l)]:
FA = (I-F ) / I
where FA is fractional absorption, I is intake of
isotope and F is faecal excretion of isotope.
T h e completeness of the faecal collection
associated with the unabsorbed isotope must be
established using non-absorbable markers, such
as radio-opaque pellets [4].Rare earth elements
have recently been introduced as faecal markers
for mineral absorption studies [S], and results
suggest that their excretion exactly parallels that
of zinc, magnesium [6] and iron [7] isotopes.
Further validation work for other inorganic ele-
ments is in progress in our laboratory.
Whole-body retention of an element can be
determined from whole-body counting if a
Volume 24
y-emitting radioisotope is used to label the food
mineral. Alternatively, for other isotopes, reten-
tion ( R )can be calculated from measurements of
urinary excretion [eqn. (Z)]:
R =I - (F + U ) (2)
where U is urinary excretion of isotope.
T h e calculation of true absorption requires
the measurement of the quantity of absorbed iso-
tope excreted via the intestine or in the urine to
be made. Endogenous secretions can make a sig-
nificant contribution to total excretion for some
of the inorganic elements, such as calcium,
copper and zinc. This problem has been solved
for calcium by using a double stable isotope
technique [8] derived from a radioisotope
method [9]. A known quantity of a stable isotope
of calcium (e.g. 42Ca) is given by intravenous (iv)
injection just after the calcium test meal contain-
ing a different stable isotope (e.g. "Ca) has been
consumed. True absorption (TA) is calculated by
measuring the isotope enrichment of the urine
24 h later [eqn. (3)]. T h e validity of applying this
method to measure true absorption of other
minerals (e.g. zinc and selenium) is under
investigation.
TA = (44Ca,la/42Ca,la)
(42Ca1,./44Ca,,ral)
x (A%XS"Ca/A%XS"Ca) (3)
where na is the natural abundance of the isotope
(atom%), iv and oral refers to the mode of
administration, and A%XS is the measured
enrichment.
Dietary factors
Inorganic nutrients must be present in a suitable
form to be taken up into the absorptive cells of
the intestine. First, the element must be solubi-
lized from the food matrix; this generally takes
(1) place in the acid environment of the stomach and
continues as the chyme moves down the intest-
inal tract. Chemical reactions that occur in the
gastrointestinal lumen play an important role in
modifying the availability of an element for
absorption into the mucosal cells. T h e strength
of binding of minerals to luminal constituents
will determine whether or not they can be trans-
ported into the mucosal cells.
One of the most potent modifiers of mineral
bioavailability is myo-inositol 1,2,3,4,5,6-hexakis-
dihydrogen phosphate (phytate, IP(,). It binds
cations, making them unavailable for absorption;
1 mol of phytate can complex with up to 6 mol of
Cu2+, up to 5 rnol of CaZf, 4 mol of Fe3+ and

3.5-4 mol of Zn2+ [10,11]. T h e metal-IP4-6
complexes are less soluble than metal-IPI-3 and
will precipitate above pH 3.5. T h e major site of
mineral absorption is the duodenum, with a pH
of 6-7.4. Within this pH range the metal-IP4-6
complex has minimum solubility and the mineral
will therefore be unavailable for absorption. IPS
and IP, have inhibitory effects on zinc absorption
whereas the lower IPS have little or no effect
[ 121. T h e individual binding affinities of
inorganic elements with phytate are ranked as
follows: Cuzt >ZnZt >MnZt >Fe3' >Ca2+.
Animal studies indicate that the bioavail-
ability of dietary zinc can be predicted from the
phytate/zinc molar ratios [13], and it has been
suggested that daily phytatehnc molar ratios
> 15 may be associated with an increased risk of
zinc deficiency [14]. More than one metal can
complex with phytate: for example calcium and
magnesium form a salt with phytate commonly
referred to as phytin [15]. T h e presence of cal-
cium exacerbates the antagonistic effect of phy-
tate on zinc because the di-metal complex is
more stable than the mono-metal phytate com-
plex. In solution, maximum chelation of zinc
occurs with calciudphytate molar ratios > 6 :1
and decreases with decreasing molar ratio [16].
T h e critical calciudphytate molar ratio for maxi-
mum zinc precipitation in humans is not known,
but it has been suggested that daily ([phytate]
x [Ca])/[Zn] ratios exceeding 200 mmol/l000
kcal (4.2 MJ) may adversely affect zinc balance in
man [14]. Most Western diets are below these
critical molar ratios, but some vegetarian diets
[17] and diets in developing countries [14] may
contain enough phytate to reduce significantly
dietary zinc bioavailability and hence increase the
risk of zinc deficiency in vulnerable groups.
Many investigations into mineral bioavail-
ability are concerned with improving the dietary
supply of minerals. Thus the dietary factors that
modulate absorption are of primary interest, and
the experimental design should incorporate a
means of controlling all other variables that affect
bioavailability (see below). Most dietary constit-
uents that modify the absorbability of minerals
have different effects depending on their chemi-
cal form and the properties of the mineral. A
summary of the main dietary constituents that
increase or decrease absorption, excretion or
utilization of calcium, magnesium, iron, zinc,
copper, selenium and chromium is given in
Table 2 [18].
Other mechanisms whereby dietary sub-
Bioactive Components of Food
stances affect mineral uptake are changes in
valency (e.g. Fe3+/Fe2+),competitive interactions
between minerals for transport proteins, and
effects on intestinal or mucosal function. T h e
physicochemical form of the mineral may have an 777
important effect on its availability for absorption.
For example, haem and non-haem iron are ab-
sorbed by two independent pathways. Haem is
absorbed intact into the mucosal cell, where it is
broken down by haem oxygenase and the
released iron enters the common pool in the cell.
Efficiency of absorption is fairly constant
(20-30%) and is little affected by dietary and
host-related variables. Bioavailability of non-haem
iron, on the other hand, is greatly influenced by
external factors and differs considerably between
different foods, according to the presence of
enhancers or inhibitors of absorption [19], as
illustrated in Table 3. T h e proportion of a
mineral that is available for absorption depends
upon the composition of the entire meal, and
attempts have been made to estimate the avail-
ability of dietary iron based upon the amounts of
enhancing substances [ZO]. However, this model
needs to be developed further to include the
effects of inhibitory substances, as discussed by
Siegenberg et al. [Zl].
Host-related variables
There are a number of mechanisms whereby the
body is able to maintain homoeostasis by regulat-
ing the absorption, retention, endogenous excre-
tion and utilization of inorganic nutrients.
Clearly these can only operate on minerals that
are in a form available for absorption in the
lumen of the intestine. These physiological vari-
ables play an important role in determining the
metabolism of the nutrient element and hence
its utilization, i.e. bioavailability.
Host-related factors that influence mineral
bioavailability are: age, sex, stage of growth, preg-
nancy, lactation, nutritional status and disease.
These fall into two general categories that relate
to (i) physiological requirements and (ii) meta-
bolic functions. Thus age, sex, stage of growth
(including pregnancy and lactation) and body
levels of the inorganic nutrients or other modify-
ing compounds (e.g. vitamin D status with
respect to calcium) have a profound effect on
mineral metabolism. When requirements are
high, efficiency of absorption is up-regulated and
endogenous losses are reduced in order to con-
serve body levels of the element. Conversely,
when requirements are lower, as for example
I996
 Biochemical Society Transactions

when stores of iron have accumulated in the
body, efficiency of absorption is down-regulated
and, wherever possible, losses are increased in
order to maintain homoeostasis. However, the
778 period of time for adaptive responses to attain a
steady-state is not known and may occasionally
cause problems in the design and/or interpreta-
tion of results from bioavailability studies.
Deficiency is almost always accompanied
by an apparent ‘increase’ in measured bioavail-
ability, which has nothing, of course, to do with
the physicochemical form of the element in the
food/diet. In order to control for the inter-indi-
vidual variation in efficiency of absorptionllevel of
endogenous excretion, absorption of the mineral
from test foods should be compared with a
standard reference dose, and results expressed as
a ratio of the two. A widely used method of deal-
ing with bioavailability data has been developed
for iron, whereby an allowance is made for dif-
ferent levels of efficiency of absorption, knowing
that individuals who are borderline iron-deficient
(i.e. with virtually no iron stores) will absorb
40% of iron from a 3 mg dose of iron as ferrous
ascorbate [22]. Dietary absorption is corrected to
a mean reference value of 40% in each subject by
multiplying by 4O/R, where R is the reference
dose absorption. An alternative method has been
suggested based upon serum ferritin concentra-
tion (as an index of iron stores), since serum
ferritin concentrations bear a close inverse rela-
tionship to iron absorption [23]. The mean

Table 2
Examples of dietary factors that increase (+) or decrease (-) Ca, Mg, Fe, Zn,
Cu, Se, Mn and Cr absorption (A), excretion (E) or utilization (U) [ I7

Dietary constituent Modifying influence Inorganic nutrient

Quantity of mineral in the diet Dose Ca, Mg, Fe, Zn, Cu, Se
Phytate -A Ca, Mg, Zn, Fe, Cr, Mn
+E cu
Phosphate -A Mg, Fe, Mn
Oxalate -A Ca
+A Cr
Polyphenols -A Fe
(hydrolysable tannins)
Thiols +A Selenite
Ascorbic acid +A Fe, Cr, selenite
-U cu
Meat protein +E Ca
-E cu
+A Zn, Fe, Mg
Casein -A Fe
Some amino acids +A Mg, Cr
- A (methionine) Selenomethionine
Sodium +E Ca
Simple sugars +A Cr
Fructose +A Ca, Mg, Cu, Zn, Fe, Mn
Guar gums +E Se
Other metals -A +
Zn Fe/Sn
+
Cr Zn/Fe/Va/Ca
+
Fe Ca/Mn/Zn
+
Cu Zn
+
Mn Ca/Fe
+
Se heavy metals
+E +
Cu Mo
-U +
Cu Fe/Cd
Antacids -A cu

Volume 24
 Bioactive Components of Food

Table 3

Examples of foods that enhance (+) or inhibit (-) non-haem iron absorption

Food Degree of effect Active substance

779
Meat +++ Cysteine-containing peptides
Citrus fruit +++ Ascorbic and citric acids
Cauliflower ++ Ascorbic acid
Beer ++ Ethanol, lactic acid
Sauerkraut ++ Lactic acid
Soya sauce + Fermentation products, cysteine, glutathione
Wheat bran _ _ _ Phytate
Tea _ _ _ Tannins
Nuts, legumes --- Phytate, polyphenols
Oregano ___ Polyphenols
Milk chocolate -- Phytate, calcium, polyphenols
Eggs - Phosphoprotein, albumin
Spinach - Polyphenbols, oxalic acid

serum ferritin value is obtained for the experi-
mental group and inserted in the following for-
mula:
log A, = log A, + log F , - log F ,
where A, is corrected dietary absorption, A, is
observed absorption, F , is observed serum ferri-
tin and F , is mean serum ferritin.
Disease states also influence mineral metab-
olism, either directly by raising or lowering
physiological requirements or as a secondary
effect, caused by changes in intestinal function
or metabolic pathways. Drugs also modify
mineral metabolism, e.g. penicillamine reduces
copper absorption [24]. Examples of diseases
that affect one or more minerals include infec-
tion, diarrhoea and vomiting, malabsorption syn-
dromes, protein-energy malnutrition, coeliac
disease, Crohn’s disease, chronic renal failure,
diabetes, hyper- or hypo-parathyroidism (Ca),
biliary cirrhosis, pancreatitis, Menkes’ syndrome
and Wilson’s disease (Cu).
Isotopic labelling of foods
In order to follow the metabolism of an inorganic
nutrient in a food, it is essential that it is labelled
with an isotope. This can be done by biosyn-
thetically incorporating the isotope into the
growing food (intrinsic labelling) or, more easily,
by mixing the label into the food shortly before
consumption (extrinsic labelling) when it
exchanges with the native element in the food.
T h e latter technique has been shown to be valid
in most instances for non-haem iron, but haem
iron must be labelled separately. An extrinsic
label can also be used in many instances for
other minerals, such as zinc, calcium and copper,
but it cannot be used for selenium, because
exchange does not take place between different
chemical forms. Radioisotopes with a high
specific radioactivity behave as true traces, and if
y emitters are available, these offer the simplest
method of studying mineral bioavailability. How-
ever, because of the hazards associated with
radioisotopes, methods have been developed to
employ stable isotopes as labels for bioavailability
studies [4,25].
Two important questions must be asked
with regard to the use of stable isotopes, namely
(i) what is the appropriate method (intrinsic or
extrinsic) for labelling the native element, and
(ii) in the case of extrinsic labelling, does the
addition of a significant quantity of the element
to the food change its properties and affect the
measurement of absorption/bioavailability? In
order to detect isotope enrichment in target
samples (e.g. blood, faeces, urine) it is necessary
to give a minimum dose of isotopically enriched
element. T h e best way of avoiding the need to
add high quantities of isotope to test foods is to
perform multiple dosing, but this has the dis-
advantage of prolonged collection periods for
faeces and urine. Another factor to be taken into
consideration is the high cost of stable isotopes.
For example, although iron bioavailability from
weaning foods can be measured in infants by
stable isotope haemoglobin incorporation [26],

I996
 Biochemical Society Transactions
the amount of isotope needed to enrich red blood
cells in adults is prohibitive. Great care must be
taken in the design of mineral bioavailability
studies to ensure that all the above factors are
780
taken into consideration.
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The metabolism of dietary nitrites and nitrates
R. Walker
School of Biological Sciences, University of Surrey, Guildford GU2 SXH, U.K.
Dietary exposure
Diet constitutes an important source of exposure
to nitrite and particularly nitrate. T h e major
dietary source of nitrate is vegetables. Lettuce,
spinach, celery and beetroot commonly contain
more than 1 g of NO;/kg fresh weight and,
depending on growing conditions (most notably
temperature and light intensity, and to a lesser
degree fertilizer use), may reach concentrations
of 3-4 g/kg fresh weight [l]. Nitrate concentra-
Volume 24
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13 Fordyce, E. J., Forbes, R. M., Robbins, K. R. and
Erdman, Jr., J. W. (1987) J. Food Sci. 52, 440-444
14 Ferguson, E. L., Gibson, R. S., Thompson, L. U.
and Ounpuu, S. (1989) Am. J. Clin. Nutr. 50,
1450-1456
15 Maga, J. A. (1982) J. Agric. Food Chem. 30, 1-9
16 Wise, A. (1983) Nutr. Abstr. Rev./Rev. Clin. Nutr.
53, 791-806
17 Bindra, G. S., Gibson, R. S. and Thompson, L. U.
(1986) Nutr. Res. 6, 475-483
18 Fairweather-Tait, S. and Hurrell, R. (eds.) (1996)
Nutr. Res. Rev., in the press
19 Report of the British Nutrition Foundation’s Task
Force (1995) Iron: Nutritional and Physiological
Significance, Chapman & Hall, London
20 Monsen, E. R., Hallberg, L., Layrisse, M. and
Hegsted, D. M., Cook, J. D., Mertz, W. and Finch,
C. A. (1978) Am. J. Clin. Nutr. 31, 134-141
21 Siegenberg, D., Baynes, R. D., Bothwell, T. H.,
Macfarlane, B. J., Lamparelli, R. D., Car, N. G.,
MacPhail, P., Schmidt, U., Tal, A. and Mayet, F.
(1991) Am. J. Clin. Nutr. 53, 537-541
22 Magnusson, B., Bjorn-Rasmussen, E., Hallberg, L.
and Rossander, L. (1981) Scand. J. Haematol. 27,
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(1991) Am. J. Clin. Nutr. 54, 717-722
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Sandstrom, B., eds.), pp. 15-21, Academic Press,
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26 Fairweather-Tait, S., Fox, T., Wharf, S. G. and
Eagles, J. (1995) Pediatr. Res. 37, 389-394
Received 14 February 1996
tions in other vegetables, such as potatoes and
cabbage, normally fall into the range 0.1-1 g/kg,
but the amount consumed means that these
make a substantial contribution to dietary intake.
Nitrite occurs in plants at low concentrations,
usually between 1 and 2 mg/kg fresh weight and
rarely in excess of 10 mg/kg. However, potatoes
have been reported to contain 2-60 mg/kg with a
mean concentration of 19 mg/kg [Z].
Estimates of mean nitrate intake range from