Bioconversion of dietary provitamin A carotenoids to vitamin A in

humans1–5
Guangwen Tang

ABSTRACT alone (10) or with vitamin E (11) or vitamin A (12), does not
Recent progress in the measurement of the bioconversion of dietary decrease the risk of cancer or cardiovascular disease, and might

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provitamin A carotenoids to vitamin A is reviewed in this article. even be harmful to smokers or former asbestos workers. Thus, it
Methods to assess the bioavailability and bioconversion of provitamin may be that b-carotene and other carotenoids promote health
A carotenoids have advanced significantly in the past 10 y, specifically when taken at physiologic amounts in foods, but have adverse
through the use of stable isotope methodology, which includes the use properties when given in high doses and under highly oxidative
of labeled plant foods. The effects of the food matrix on the bioconver- conditions. Furthermore, the health benefits of food b-carotene
sion of provitamin A carotenoids to vitamin A, dietary fat effects, and consumed at a physiologic amount as an intact molecule and/or
the effect of genotype on the absorption and metabolism of b-carotene its cleavage product (vitamin A) should be investigated to de-
have been reported recently. A summary of the major human studies termine the relative importance of each potentially favorable
that determined conversion factors for dietary b-carotene to retinol property in various populations. The conversion of dietary
is presented here, and these data show that the conversion efficiency of b-carotene to vitamin A may relate to preformed vitamin A in
dietary b-carotene to retinol is in the range of 3.6–28:1 by weight. the diet (13); that is, the conversion may be less efficient when
There is a wide variation in conversion factors reported not only vitamin A has been provided from other dietary sources. The
between different studies but also between individuals in a particular issue of the efficiency of conversion of provitamin A carotenoids
study. These findings show that the vitamin A value of individual into retinol and other retinoids is therefore of interest for us to
plant foods rich in provitamin A carotenoids may vary significantly better understand the nutritional value of dietary provitamin A
and need further investigation. Am J Clin Nutr 2010;91(suppl): carotenoids.
1468S–73S.
It is well established that after an oral dose of b-carotene, both
intact b-carotene and its metabolite retinol can be found in the
INTRODUCTION circulation. In humans, conversion of b-carotene into vitamin A
takes place predominantly in the intestine and less so in other
Dietary provitamin A carotenoids are a major source of our
tissues. The ratio of the amount of b-carotene given in an oral
vitamin A needs. Vitamin A is an essential vitamin for the
dose to the amount of vitamin A derived from this b-carotene
promotion of general growth, maintenance of visual function,
dose is defined as the b-carotene-to-vitamin-A conversion factor
regulation of differentiation of epithelial tissues, and embryonic
or b-carotene equivalent to vitamin A.
development (1). Vitamin A can be obtained from food, either as
For a healthy population, the major factors that affect the
preformed vitamin A in animal products, such as eggs and dairy
bioavailability of food carotenoids and the bioconversion of food
products, or as provitamin A carotenoids, mainly b-carotene in
plant products, such as green leafy and yellow-colored vegeta-
1
bles and orange-colored fruit. From the Jean Mayer US Department of Agriculture Human Nutrition
In Western societies, the provitamin A carotenoids derived Research Center on Aging, Tufts University, Boston, MA.
2
from plants provide ,30% of daily vitamin A intake, whereas Presented at the workshop ‘‘Micronutrient Bioavailability: Priorities and
Challenges for Setting Dietary Reference Values,’’ held in Barcelona,
preformed vitamin A derived from animal products provides
Spain, 11–12 June 2009.
.70% daily vitamin A intake (2). In contrast, in developing 3
Any opinions, findings, conclusions, or recommendations expressed in
countries, provitamin A carotenoids in vegetables and fruit this publication are those of the author and do not necessarily reflect the view
provide .70% of daily vitamin A intakes (3). of the US Department of Agriculture, nor does mention of trade names,
Epidemiologic data have shown that diets rich in carotenoid- commercial products, or organizations imply endorsement by the US Gov-
containing foods are associated with decreased risk of certain ernment.
4
types of chronic diseases, such as cancer (4), cardiovascular Supported by the US Department of Agriculture under Cooperative
disease (5), age-related macular degeneration (6, 7), and cataracts Agreements 58–1950-7–707, a grant from NIH NIDDK DK620021, and
USAID.
(8, 9). The disease-preventing activity of b-carotene and other 5
Address correspondence and reprint requests to G Tang, Jean Mayer
provitamin A carotenoids could be ascribed either to their US Department of Agriculture Human Nutrition Research Center on
conversion into retinoid or to their activity as intact molecules. Aging, Tufts University, 711 Washington Street, Boston, MA 02111; E-mail:
The results of several human intervention studies, however, in- [email protected].
dicate that high-dose supplementation with b-carotene, either First published online March 3, 2010; doi: 10.3945/ajcn.2010.28674G.

1468S Am J Clin Nutr 2010;91(suppl):1468S–73S. Printed in USA. Ó 2010 American Society for Nutrition
 DIETARY PROVITAMIN A CAROTENOIDS CONVERSION TO VITAMIN A 1469S
provitamin A carotenoids to vitamin A in humans are food alence was, therefore 2:1 by weight. In these studies, all subjects
matrices, food preparation, and the fat content of a meal (14). had been made deficient in vitamin A, so it cannot be de-
Recent studies reported that the conversion efficiency of dietary termined whether a 3.8- or 2-lg equivalence of b-carotene to
b-carotene is in the range of 10 to 28:1 by weight (15–19). 1 lg retinol is applicable in vitamin-A–sufficient individuals.
These data indicated that the bioconversion of b-carotene to
vitamin A was not as efficient as expected, and, as a result, the
Food and Nutrition Board recently revised the estimated effi- CHANGES IN CONCENTRATION OF SERUM VITAMIN A
ciency factor for the conversion of dietary b-carotene to vitamin The measurement of changes in serum vitamin A concen-
A from 6:1 by weight (20) to the new value of 12:1 by weight trations after the feeding of synthetic b-carotene or food rich in
(21). However, this new conversion ratio must be regarded provitamin A carotenoids has been used in populations with
as temporary and could well change, as more data become lower vitamin A status. The conversion of b-carotene into vi-
available. tamin A cannot be estimated accurately in well-nourished hu-
On the other hand, preformed vitamin A from animal origins or mans by the assessment of changes in serum retinol after
from supplements can be absorbed and stored in the human body supplementation with unlabeled b-carotene, because of the in-

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very effectively. A recent report (22) showed that among women ability to distinguish newly formed retinol from retinol derived
who did not take supplemental vitamin A, retinol from food was from body reserves. In addition, it is well known that blood
significantly associated with fracture risk (relative risk: 1.69; retinol concentrations are homeostatically controlled in well-
95% CI: 1.05, 2.74; P for trend: 0.05, for 1000 compared with nourished individuals. Nevertheless, many investigations in
,400 lg/d). However, b-carotene did not contribute signifi- populations with normally low vitamin A intakes have reported
cantly to fracture risk (relative risk: 1.22; 95% CI: 0.90, 1.66; blood retinol responses to acute or chronic b-carotene supple-
P for trend: 0.10, for 6300 compared with ,2550 lg/d). That ments (25, 26). Changes in serum retinol concentrations were
is, long-term intake of a diet high in retinol may promote the seen (16) in vitamin A–deficient (’0.7 lmol/L) anemic school
development of osteoporotic hip fractures in women. Because children aged 7–11 y, who were fed 1 of 4 supplements: 1) 556
of such a narrow safe range of food retinol, it is important to retinol equivalents (RE)/d from retinol-rich foods (n = 48), 2)
study the vitamin A value of food provitamin A carotenoids for 509 RE/d from fruit (n = 49), 3) 684 RE/d from vegetables (n =
humans. 45), or 4) 44 RE/d from low-retinol and low-carotene foods (n =
It is a challenge to study the conversion of b-carotene to vi- 46) 6 d/wk for 9 wk. The changes in serum retinol showed that
tamin A at physiologic doses and even more of a challenge to the consumption of fruit (diet 2) or vegetables (diet 3) resulted in
study this conversion at dietary intake amounts because of the increases of 0.12 and 0.07 lmol/L, respectively, in serum reti-
inability to distinguish newly formed retinol from body reserves. nol, whereas the group who consumed foods rich in preformed
Different methods have been developed and used to determine vitamin A (diet 1) showed an increase of 0.23 lmol/L. The
these bioconversion factors in various populations with different relative mean conversion factor of vegetable b-carotene into ret-
vitamin A nutritional status. The most widely used methods and inol was calculated, by weight, as 26:1 and that of b-carotene from
the results obtained from the application of these methods are orange fruit as 12:1. Use of a similar approach (17) showed that,
described and evaluated below. for breastfeeding women, the conversion factors of b-carotene
into retinol were, by weight, 12:1 for fruit and 28:1 for green leafy
vegetables.
DEPLETION-REPLETION METHOD
In early studies, a depletion-repletion method was used, which
involved measurement of the repletion doses of b-carotene and CHANGES IN WHOLE-BODY STORES OF VITAMIN A
vitamin A that were needed to reverse vitamin A deficiency in (PAIRED DEUTERATED RETINOL DILUTION)
depleted adults. This approach is no longer acceptable. A de- Advances in isotope technology have facilitated the measure-
pletion study (23) was conducted in 16 healthy subjects between ment of changes in body stores of vitamin A after the feeding of
the ages of 19 and 34 y (7 additional subjects served as positive dietary provitamin A carotenoids; specifically, paired deuterated-
controls). After 12 mo of depletion, only 3 of the subjects were retinol-dilution (DRD) tests can be used to measure the conversion
vitamin A deficient; a blood concentration ,0.35 lmol/L efficiency in food-based intervention studies. For populations with
(10 lg/dL) and deterioration in dark adaptation were used to marginal-to-normal vitamin A status, changes in serum retinol
define "unmistakably deficient" subjects. Of the 3 subjects with concentrations may not be a sensitive indicator of vitamin A status.
vitamin A deficiency, 2 were given b-carotene and one was Instead, DRD can be used to measure changes of total body stores
given preformed vitamin A. Daily doses of 1500 lg b-carotene of vitamin A. A DRD method was used in a study of children with
or 390 lg retinol for 3 wk to 6 mo were sufficient to reverse marginal-to-normal vitamin A status, who participated in a food-
vitamin A deficiency in these subjects. Therefore, from this based intervention with either green-yellow vegetables or light-
human study, the b-carotene/vitamin A equivalence was de- colored vegetables with low carotene content (18). The serum
termined to be 3.8:1 by weight. In 1974, another vitamin A carotenoid concentrations of children fed green-yellow vegetables
depletion-repletion study in human subjects was reported (24). increased, whereas the serum concentration of vitamin A did
Eight healthy male subjects aged 31–43 y were depleted in vi- not change. In contrast, the DRD tests carried out before and after
tamin A within 359–771 d. Five subjects were then given vita- the vegetable intervention showed that the body stores of vitamin
min A and 3 subjects were given b-carotene. Daily doses of 600 A were stable in the group fed green-yellow vegetables (n = 10)
lg retinol or 1200 lg b-carotene were required to cure vitamin but decreased in the group fed light-colored vegetables (n = 8).
A deficiency. In this study, the b-carotene-to-vitamin-A equiv- Over a 10-wk period, a loss of 7 mg vitamin A from body stores
1470S TANG

was seen in the children fed light-colored vegetables that con-

tained little b-carotene, but 275 mg b-carotene from green-yellow
vegetables prevented this loss. From this paired DRD test, it
was calculated that 27 lg b-carotene from vegetables was
equivalent to 1 lg retinol. This conversion factor is similar to that
reported in another study, which measured changes in serum vi-
tamin A concentration after consumption of carotenoids from
vegetables (17).
The paired DRD technique has also been used (15) to measure
change in the vitamin A pool size after 60-d supplementation with
750 RE/d as either retinyl palmitate, b-carotene, sweet potato, or
Indian spinach, compared with a control that contained no ret-
inol or carotene (n = 14/group). Vitamin A equivalency factors
of 6:1 for b-carotene in oil, 10:1 for b-carotene in Indian spin-

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ach, and 13:1 for b-carotene in sweet potato were determined. A
recent study used a mixed-vegetable intervention and the paired FIGURE 1. Conversion of 2H8 b-carotene and 2H8 retinyl acetate into 2H4
retinol and 2H8 retinol, respectively.
DRD test to measure changes in vitamin A pool size (27). The
results showed that the conversion factors were better than 12:1
for b-carotene and 24:1 for other provitamin A carotenoids.
contains b-carotene, as long as it is labeled properly. Such an
isotope reference method (with a known amount of 2H8 retinyl
CONSUMPTION OF INTRINSICALLY LABELED acetate as the reference material) can be used to define the vi-
DIETARY PROVITAMIN A CAROTENOIDS tamin A activity of various vitamin A precursors, synthetic
To achieve an accurate assessment of carotenoid bioabsorption b-carotene (30), or provitamin A carotenoids in vegetables or
and a subsequent vitamin A value from a food source, food other plants in humans.
material is required in which the carotenoids have been en- This method has been used to determine the vitamin A value of
dogenously or intrinsically labeled with a low-abundance stable endogenously labeled spinach and carrot carotenoids (19). For
isotope. Plant carotenoids can be intrinsically labeled either these experiments on carotenoid absorbability and conversion, we
through the addition of a carbon-stable isotope presented in the recruited 7 healthy male adults who had normal vitamin A status
gas atmosphere as 13CO2 or through the addition of a hydrogen- and were nonsmokers. They were given pureed and cooked
stable isotope presented to the roots in the form of heavy water, spinach (n = 14, 7 men and 7 women) and pureed and cooked
2
H2O. For 2H2O, plants can be labeled via hydroponic growth carrot (n = 7 men) grown hydroponically in 25 atom % 2H2O.
(28) on a nutrient solution composed of a fixed 2H2O percentage With the use of 3.0 mg labeled retinyl acetate as a reference dose,
to achieve the labeling of the carotenoids. This allows pre- we observed that provitamin A carotenes in carrots have greater
sentation of the carotenoids in their normal cellular compart- vitamin A potency than those in spinach. The 300 g labeled
ments, and the isotopic label enables identification of those spinach and 100 g labeled carrots each contained ’0.11 mg (all-
serum carotenoids (or derived retinol), which come from the trans)-b-carotene, and it was assumed, as usual, that a-carotene
specific food in question. The deuteration of intrinsically labeled and (cis)-b-carotene, which were also present, had half the ac-
plant 2Hn-b-carotene has been shown to be distributed randomly tivity of (all-trans)-b-carotene (21). The retinol equivalences
throughout the carotenoid molecules (29). were determined to be 21 lg spinach b-carotene or 15 lg carrot
We have developed an isotope reference method to quanti- b-carotene to 1 lg retinol.
tatively determine the retinol equivalence of b-carotene (either With a similar approach, dried Spirulina powder was studied in
synthetic pure b-carotene or b-carotene contained in a food). In humans. Spirulina is an alga rich in b-carotene. With the use of
the isotope reference method, through the use of a known hydroponically grown and intrinsically deuterium-labeled Spir-
amount of labeled vitamin A, such as retinyl acetate-d8, as ulina, and labeled retinyl acetate as a reference dose, a study was
a reference and the comparison of the amount of retinol formed conducted in Chinese adults (n = 10 men) with normal vitamin A
in vivo from a vitamin A precursor (eg, 2H8 b-carotene), we may status (31). The volunteers (average age, 48 y) each took 5 g dried
determine quantitatively the vitamin A value of the vitamin A Spirulina powder that contained 4.3 mg b-carotene. When com-
precursor, as shown in Figure 1. Therefore, the retinol equiva- pared with a reference dose of 2.0 mg 13C10 retinyl acetate in oil
lence of the 2H8 b-carotene dose = (AUC2H4 retinol/AUC2H8 retinol) · administered in a capsule, 4.5 mg Spirulina b-carotene provided
Dose2H8 retinyl acetate, where AUC is the area under the curve of 1 mg retinol.
the circulating tracer measured compared with time. For ex- Very recently, a human study that used Golden Rice b-carotene
ample, in the serum of a volunteer, if, 21 d after a 6-mg 2H8 was reported (32): 65–98 g (130–200 g cooked weight) of hy-
b-carotene oral dose and a 3-mg 2H8 retinyl acetate oral dose, droponically grown Golden Rice that contained 0.99–1.53 mg of
the area under the 2H4 retinol curve (derived from the 2H8 b-carotene was given to 5 healthy volunteers (3 women and 2
b-carotene) is measured as 10 units and the area under the 2H8 men) in the United States. With the use of the isotope reference
retinol curve (from the 2H8 retinyl acetate) is measured as 10 method, in comparison with the 13C10 retinyl acetate reference
units, we can say that 6 mg of synthetic b-carotene is nutri- dose, Golden Rice b-carotene provided 0.24–0.94 mg retinol.
tionally equivalent to 3 mg retinyl acetate. That is, we will be Thus, the conversion factor of Golden Rice b-carotene to retinol
able to define the vitamin A activity of b-carotene or a food that is 3.6:1 with a range of 1.6 to 6.4:1 by weight. Therefore,
 DIETARY PROVITAMIN A CAROTENOIDS CONVERSION TO VITAMIN A 1471S
b-carotene derived from Golden Rice is effectively converted to spinach leaves is in the form of pigment proteins located in
vitamin A in humans. chloroplasts, whereas in carrots the b-carotene is in a crystal
form in chromoplasts (42). In addition, the efficient conversion
of Spirulina b-carotene may be due to its simple cell structure,
OTHER METHODS which is composed of protein and peptidoglycans that are di-
In the 1960s, a few studies were carried out in humans that gested easily (43). Similarly, rice has a simple and easily di-
investigated the vitamin A activity of b-carotene with the use of gestible food matrix, which allows for a high bioavailability and
radioisotope-labeled b-carotene. Studies by Goodman et al (33) bioconversion of b-carotene to vitamin A.
and Blomstrand and Werner (34) have provided most of our Intake of b-carotene at different dosage amounts will affect the
knowledge about how humans absorb and metabolize b-carotene. conversion efficiency of b-carotene when given as b-carotene oil
Such radioisotope methods can no longer be used for ethical capsules (44). In this study a subject who took 6 mg b-carotene in
reasons. oil capsules showed a conversion factor of 3.8:1, whereas the
The measurement of b-carotene and retinyl esters in post- same subject who took a dose of 126 mg b-carotene in oil showed
prandial chylomicron fractions after the feeding of food rich in a conversion factor of 55:1 by weight. It is well known that di-

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provitamin A carotenoids was developed to study a single dose etary fat is a critical factor that affects bioavailability and bio-
of b-carotene. Postprandial chylomicron response curves of conversion of b-carotene. However, the amount of fat needed in
b-carotene and retinyl esters in blood were measured after the diet to facilitate the absorption of carotenoids should be
a single dose of b-carotene supplement in oil or from vegetables studied. A recent study on the influence of amounts of dietary fat
(35–37). In these studies, triacylglycerol-rich lipoproteins on the bioavailability and bioconversion of provitamin A car-
(TRLs) with density ,1.006 g/mL were separated and analyzed otenoids in yellow and green leafy vegetable meals showed that
to evaluate the absorption efficiency of b-carotene (intact and, carotenoid-rich yellow and green leafy vegetables need a certain
after central cleavage, as retinyl palmitate). The efficiency of minimum amount of fat (2.4 g fat/meal) to ensure the absorption
absorption of b-carotene by each subject was calculated by of fat-soluble provitamin A carotenoids and to improve vitamin A
measurement of the areas under the curve (AUC, nmol
h/L) of nutritional status (27).
b-carotene and retinyl ester concentrations in postprandial TRL Our study also showed that there is a correlation of the
fractions collected hourly. These curves were compared with b-carotene-to-vitamin-A conversion factor with the BMI in in-
hypothetic AUC after an intravenous dose of the same amount of dividual subjects (30). That is, conversion factors of the subjects
b-carotene, on the assumption that the b-carotene disappearance who received synthetic b-carotene were significantly correlated
follows a first-order elimination from blood with a chylomicron with BMI. This suggests that subjects with more body fat have
remnant half-life of 11.5 min (35). To compensate for the var- a lower capability to convert b-carotene to vitamin A.
iability of TRL recovery, deuterium-labeled vitamin A was used We have observed large variations in the bioconversion of
(38) as an extrinsic standard. A subject was given raw carrots dietary b-carotene to vitamin A, which may be related to the
that contained 9.8 lmol (5 mg) b-carotene and 5.2 lmol genetic characteristics of the subjects. Because the enzyme re-
a-carotene (2.8 mg), together with 7 lmol (2 mg) [2H4]-retinyl sponsible for b-carotene conversion into retinol is b-carotene
acetate, and the concentrations of b-carotene, a-carotene, and 15,15’-monoxygenase (BCMO1), genetic polymorphisms in the
labeled and unlabeled retinyl esters in the TRL were measured BCMO1 gene may contribute to the poor converter phenotype.
at various time points up to 7 h. With the assumption that ab- Very recently, it was reported (45) that 2 common nonsyn-
sorption of labeled retinyl acetate was ’80% of the dose, it was onymous single nucleotide polymorphisms (R267S and A379V)
calculated that 0.8 lmol of the carrot b-carotene was absorbed had been identified and in vitro biochemical characterization of
intact and that 1.5 lmol of unlabeled retinyl esters were formed the recombinant 267S + 379V double mutant showed a decreased
from the carrot dose. The mass equivalency of carrot b-carotene catalytic activity of BCMO1 by 57% (P , 0.001). A further
to vitamin A was, therefore, 13:1 by weight (without consider- assessment of the responsiveness to a pharmacologic dose of
ation of the contribution from 5.2 lmol of a-carotene to vitamin b-carotene in female volunteers confirmed that carriers of the
A). If the contribution of a-carotene is considered, the ratio is 379V and 267S + 379V variant alleles showed a 69% decrease
higher (16:1), with the assumption that a-carotene has half the in their ability to convert b-carotene and a 240% increase in
activity of b-carotene. fasting plasma b-carotene concentration. Therefore, there is ge-
netic variability in b-carotene metabolism. This may provide an
explanation for the molecular basis of the poor converter phe-
FACTORS THAT AFFECT THE BIOCONVERSION OF notype within the population. Multiple single nucleotide poly-
b-CAROTENE morphisms and genes that might influence b-carotene status
Review articles that have evaluated the factors that affect the warrant further study.
conversion of b-carotene to vitamin A have been published by
Castenmiller and West in 1998 (39), van het Hof et al in 2000
(40), and Yeum and Russell in 2002 (41). For bioconversion of CONCLUSIONS
dietary b-carotene, the major factors are food matrix, food Provitamin A carotenoids from various foods have been shown
processing, and fat in the diet. As mentioned above, even though to have an almost 8-fold difference in b-carotene conversion
the spinach and carrots were both pureed and cooked, the in vivo factors (on a weight basis) that ranged from 3.6:1 to 28:1 with
study showed that spinach b-carotene had a conversion factor of Golden Rice and leafy vegetables, respectively (Table 1), and
21:1, whereas carrot b-carotene had a conversion factor of 15:1. thus have different values in terms of vitamin A nutrition. The
This was due to differences in the food matrix: b-carotene in major factor that affects the vitamin A value of plant provitamin
1472S TANG
TABLE 1
Summary of studies to determine a conversion factor for b-carotene in food sources to vitamin A1
Conversion
factor (by
Food matrix Method Dose weight) Reference

Fruit, n = 49; vegetables, n = 45; Changes of serum retinol concentration Fruit: 509 RE/d; vegetables: 684 RE/d; 12:1 16
retinol-rich foods, n = 48 (ages 7–11 y) in vitamin A–deficient (’0.7 lmol/L) vitamin A–rich foods: 556 RE/d 26:1
anemic schoolchildren
Green/yellow vegetables, n = 10; Total body stores of vitamin A before Green/yellow vegetables (206 mg 27:1 18
light-colored vegetables, n = 8 and after the vegetable intervention calculated trans-b-carotene) to prevent
in schoolchildren (aged 5.3–6.5 y) the decrease of 7.7 mg in liver stores
with normal or marginal vitamin A
status
Sweet potato, Indian spinach, b-carotene Mean changes of total body stores of Sweet potato: 750 lg RE; Indian spinach: 13:1 15
capsule, or retinyl palmitate; all, vitamin A before and after a 750 lg RE; b-carotene capsule: 750 lg 10:1

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n = 14 60-d intervention in adult men RE; retinyl palmitate: 750 lg RE 6:1
compared with the mean changes
in the retinyl palmitate group
[2H]-Labeled spinach, and vitamin A in Comparison of AUC responses to the Calculated 11 mg trans-b-carotene 21:1 19
oil capsule, n = 14 spinach and the vitamin A reference from 300 g pureed, cooked spinach, and
dose in adults 3 mg [2H8]-vitamin A
[2H]-Labeled carrot, and vitamin A in Comparison of AUC responses to the Calculated 11 mg trans-b-carotene 15:1 19
oil capsule, n = 7 carrot and the vitamin A reference from 100 g pureed, cooked carrot, and
dose in adults 3 mg [2H8]-vitamin A
Fruit, n = 69; leafy vegetables, n = 70; Changes of serum retinol concentration Fruit: 4.8 mg trans-b-carotene; vegetables: 12:1 17
retinol-rich foods, n = 70; control, in lactating women after they ate fruit 5.6 mg trans-b-carotene; retinol-rich 28:1
n = 68 or vegetables, or took preformed diet: 610 lg retinol; control: 0.6 mg
vitamin A b-carotene and 1 lg retinol
Spirulina powder, n = 10 Comparison of AUC responses to the 4.3 mg Spirulina trans-b-carotene 4.5:1 31
Spirulina and the vitamin A reference
dose in adults
Golden Rice, n = 5 Comparison of AUC responses to a Rice meal that contained 1–1.5 mg rice 3.6:1 32
Golden Rice meal and the vitamin A b-carotene
reference dose in adults
1
RE, retinol equivalents; AUC, area under the curve.

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