TRENDS in Plant Science

Vol.11 No.10

Accumulation and bioavailability of dietary carotenoids in vegetable crops
Dean A. Kopsell1 and David E. Kopsell2
1 2

Plant Sciences Department, The University of Tennessee, Knoxville, TN 37996, USA School of Agriculture, The University of Wisconsin – Platteville, Platteville, WI 53818, USA

Carotenoids are lipid-soluble pigments found in many vegetable crops that are reported to have the health benefits of cancer and eye disease reduction when consumed in the diet. Research shows that environmental and genetic factors can significantly influence carotenoid concentrations in vegetable crops, and that changing cultural management strategies could be advantageous, resulting in increased vegetable carotenoid concentrations. Improvements in vegetable carotenoid levels have been achieved using traditional breeding methods and molecular transformations to stimulate biosynthetic pathways. Postharvest and processing activities can alter carotenoid chemistry, and ultimately affect bioavailability. Bioavailability data emphasize the importance of carotenoid enhancement in vegetable crops and the need to characterize potential changes in carotenoid composition during cultivation, storage and processing before consumer purchase. Vegetable crops are important sources of dietary carotenoids Consumption of vegetable crops provides the human diet with many essential vitamins and minerals important for health maintenance. Vegetables also contain other food components, or phytochemicals, that provide benefits beyond normal health maintenance and nutrition. One important class of phytochemicals is the carotenoids. Carotenoids are lipid-soluble pigments found in all photosynthetic organisms. Among the naturally occurring plant pigments, carotenoids are widely distributed, with a high degree of structural diversity and large variations in biological functions [1,2]. There are >600 carotenoids found in nature, with 40 dietary carotenoids regularly consumed in the human diet [3]. Pro-vitamin A activity is the classical biological function of b-carotene, a-carotene and cryptoxanthins in mammalian systems. Health benefits attributed to carotenoids include prevention of certain cancers [4–6], cardiovascular diseases [7] and eye diseases [8,9], as well as enhanced immune system functions [10,11]. Current US Department of Agriculture (USDA) dietary guidelines recommend eating 7–9 servings of fruit and vegetables per day. However, the average consumption by adults in the USA is only 4.4 servings per day, with an estimated 42% of Americans eating less than
Corresponding author: Kopsell, D.A. ( Available online 1 September 2006.

two daily servings of fruit and vegetables (see http:// Research into carotenoid enhancement of vegetable crops to benefit human health has paralleled efforts to increase consumption of fruit and vegetables in the diet. Current methods to assess bioavailability (see Glossary) include serum measurements and in vitro digestion models. Release of carotenoid from the membranes of plant tissues facilitates intestinal absorption; however, changes in carotenoid chemistry by biotic (e.g. oxidation) and abiotic (e.g. temperature) factors can influence bioavailability. Programs to improve carotenoid levels in vegetable crops must successfully link plant physiology with accurate bioavailability assessments in human subjects. In this review, we discuss our current knowledge of carotenoid chemistry, carotenoid contributions to human health, bioavailability assessments, how pre- and postharvest practices can alter carotenoid levels and influence potential bioavailability, and recent vegetable carotenoid enhancement efforts. Carotenoid chemistry and plant biosynthetic pathway Carotenoids are C40 isoprenoid polyene compounds that form lipid soluble yellow, orange and red pigments [12,13]. Carotenoids are secondary plant compounds and can be divided into two groups: the oxygenated xanthophylls such as lutein (3R,30 R,60 R b,e-carotene-3,30 diol), zeaxanthin (3,30 R-b,b-carotene-3,30 diol) and violaxanthin (3S,5R,6S,30 S,50 R,60 S-5,5,50 ,60 -depoxy-5,6,50 ,60 -tetrahydrob, b-carotene-3,30 diol), and the hydrocarbon carotenes such as b-carotene (b, b-carotene), a-carotene (60 R, b,e-carotene) and lycopene (c, c-carotene) [12]. In higher plants, carotenoid compounds are synthesized and localized in cellular plastids where they are associated with light-harvesting complexes (LHCs) in the thylakoid membranes, or present as semicrystalline structures derived from the plastids [14,15]. Terpenes are a diverse group of plant secondary metabolites formed via acetyl-CoA (mevalonic acid pathway) or from glycolytic intermediates (methylerythritol phosphate pathway). The basic structural element of all terpenes is the 5-carbon branched isopentane. Terpenes are classified by the number of isoprene units present. Carotenoids are composed of 40 carbons and are thus classified as tetraterpenes. The carotenoid biosynthetic pathway was elucidated in the mid-1960s [15]. Currently, genes and cDNAs for the major enzymes functioning in carotenoid biosynthesis have been cloned from plant, algal and microbial sources [16]. Carotenoids are produced in

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g.22. Carotenoid pigments protect photosynthetic structures by quenching 3Chl to dissipate excess energy [21] and binding 1O2 to inhibit oxidative damage [20. along with other chloroplast carotenoids. Chylomicrons: the largest of the blood lipids consisting mostly of triglycerides and small amounts of cholesterol. phytoene desaturase and j-carotene desaturase. which arise from differences in the shape of the molecule or the overall dipole moment. and protect eye tissues [25]. Cardiovascular disease reductions were mostly associated with increased intakes of a-carotene. Reactive species include singlet oxygen (1O2). Some vegetables have a modified carotenoid biosynthetic capacity and produce compounds exclusively associated with their respective genus. however. and modulate the expression of gene determinants in the prevention of certain types of cancers. There is a great deal of research on the anticarcinogenic effects of individual carotenoids on specific types of cancers.27]. Cohort studies involving a food frequency questionnaire and serum carotenoid data have established the preventative effects of dietary carotenoids on cardiovascular diseases in separate studies of populations in Italy [28]. a-carotene and lutein) [16.20]. The reversible epoxidation–de-epoxidation reaction converting violaxanthin back to zeaxanthin via the intermediate antheraxanthin is collectively referred to as the violaxanthin cycle and is vital for energy dissipation from incoming solar radiation [15. violaxanthin and neoxanthin) or carotenoids with one b-ring and one e-ring (e. Higher carotenoid intakes can also lower risk of certain lymphomas [34]. mostly in the blue–green wavelength range. Europe [30] and Costa Rica [31]. Lipophilic: having a strong affinity for lipids. They are classified as to their density and are mainly involved in lipid transport in the aqueous blood plasma. protein thiol groups. Carotenoids function to help harvest light energy. Reactive oxygen species (ROS): chemical species that participate in oxidative reactions with polyunsaturated membrane phospholipids. hydroxyl radical (OH).18].17] (Figure 1). Two structurally similar enzymes. which becomes the substrate for the C20 geranylgeranyl diphosphate (GGPP) [17]. and visual functions [24]. Carotenoids can reduce the risk of cardiovascular disease through reductions in low-density lipoprotein oxidation and oxidative stress at locations of plaque formations. The enzyme GGPP synthase catalyzes the formation of GGPP from IPP and DMAPP [16]. Xanthophylls might also be involved in structural stabilization of LHCs and reduction of lipid peroxidation [21]. Zeinoxanthin is found in yellow corn (Zea mays) grains. b-carotene and b-cryptoxanthin. Lettuce (Lactuca species) contains lactucaxanthin. must rely on dietary sources to provide sufficient levels.11 No.10 Glossary Bioavailability: the extent to which a nutrient in a food constituent can be absorbed and used by the body after ingestion. normal reproductive performance. Chromophore: any light-absorbing molecular group that produces color in a compound. with men benefiting more from increased carotenoid intakes [33]. make the conversions of phytoene to lycopene [2.sciencedirect. and change the colorless phytoene into colored lycopene. Isomerization: biochemical or chemical reaction involving an intra-molecular shift resulting in a change in orientation of functional groups within a molecule. cohort and epidemiological study data reveal that increased levels of dietary carotenoids have no the plastids and are derived from isopentenyl diphosphate (IPP).and b-carotenes into the subgroup referred to as the xanthophylls. which is transferred to the photosynthetic reaction centers (RCs). Carotenoids can inhibit cell proliferations. The protective effects of carotenoids against lung cancers differ between the sexes. zeaxanthin. usually carbon (C).23]. www.500 Review TRENDS in Plant Science Vol. The carotenoid pathway branches at the cyclization reactions of lycopene to produce carotenoids with either two b-rings (e. IPP is isomerized to dimethylallyl diphosphate (DMAPP). The first step unique to carotenoid biosynthesis is the condensation of two molecules of GGPP to form the first C40 carotenoid. via phytoene synthase [1]. Humans cannot synthesize carotenoids . non-protein thiol groups and DNA with the potential to alter cellular functions. Recent studies have established a link between increased intake of fruits and vegetables that are high in carotenoids with reduced incidences of mortality related to cardiovascular disease. a-carotene and cryptoxanthins) and non-pro-vitamin A carotenoids (lutein. phospholipids. or even species. Data from cohort studies reveal protective effects from b-carotene intakes against the risk of tobacco-related cancers in non-smokers. By contrast. enhance the immune response. High-density and low-density lipoproteins: lipid–protein combinations with the solubility characteristics of a protein. zeaxanthin and lycopene) function as free-radical scavengers. Carotenoids and human health One of the most important physiological functions of carotenoids in human nutrition is to act as vitamin A precursors. Hydrophobic: having little or no affinity for water. b-carotene is highly concentrated close to the reaction center. Fruits and vegetables are primary sources of carotenoids in the human diet and their consumption has been associated with numerous health benefits [26. therefore.16. Epoxide: an organic molecule containing a chemical group consisting of an oxygen molecule bound to previously connected atoms. and lutein is present in several lightharvesting antennae components [19. Japan [29]. In the first step in biosynthesis. b-carotene can increase cancer risks among smokers [32]. b-carotene. capsorubin and cryptocapsin.17]. Within the thylakoid membranes of chloroplast organelles. nitric oxides (NO) and peroxyl (HOO). suppress cancer development. Further steps in xanthophyll synthesis include epoxydation reactions. which convert the hydrocarbon a. Peppers (Capsicum species) contain capsanthin. Both pro-vitamin A carotenoids (b-carotene. Pro-vitamin A carotenoid compounds support the maintenance of healthy epithelial cell differentiation. excess excitation energy can result in the formation of triplet excited chlorophyll (3Chl) and reactive singlet oxygen (1O2). The pathway advances with the additions of oxygen moieties. superoxide (O2À). They are synthesized in the intestinal mucosal cells and serve to transport lipids to tissues. phytoene. antheraxanthin.g. proteins and fat-soluble vitamins. cell transformations. carotenoids are found bound to specific protein complexes of photosystem I and photosystem II (PSII). These desaturase enzymes create the chromophore present in the carotenoid pigments. which are responsible for the vibrant fruit coloration within the genus [1]. cis (Z) isomers and trans (E) isomers often have different physical properties. In the PSII complex. When the absorption of light radiation exceeds the capacity of photosynthesis.

11 No. geranylgeranyl pyrophosphate. The yellow pigments are . Lutein and zeaxanthin are two of seven major carotenoids found in human blood serum. Precursor compounds and major carotenes and xanthophylls in the carotenoid biosynthetic pathway in plants.10 501 Figure 1.sciencedirect. isopentenyl pyrophosphate. In the retina. Abbreviations: DMAPP. lutein and zeaxanthin are chiefly responsible for the yellow pigmentation collectively referred to as macular pigment (MP) [38]. IPP. dimethylallyl pyrophosphate. they are the only carotenoids present in the retina and lens [37]. preventative effects on incidences of bladder cancers [35]. but recent results from screening trials show that lycopene levels were not associated with prostate cancer risk [36].Review TRENDS in Plant Science Vol. GGPP. The preventative effects of lycopene on prostate cancer are unclear: positive results have been demonstrated in vivo. however.

Based on their chemistry. Many of these same factors are also associated with a decreased risk of developing age-related macular degeneration. with an oxygen bound to the 5. high levels of dietary carotenoid intake and the resultant elevated serum carotenoid concentration. derive potent antioxidant activity from their high number of conjugated dienes [46]. such as lycopene. carotenoids are one of the most potent biological quenchers of singlet oxygen [45]. providing a mixture of carotenoids. In vivo antioxidant activity is determined by carotenoid structure and concentration.11 No. is more strongly associated with reduced eye disease risk than intake of individual carotenoid supplements [9]. All carotenoids exhibit cis–trans isomerization. the variability associated with the carotenoid content of vegetables. Epidemiological evidence has been inconsistent regarding the potential relationships between diet.42].40].60 double bonds in their cyclic end groups to undergo epoxidation with 1O2 (Figure 2) [26]. the complexity of measuring in vivo antioxidant behavior. are present in vegetable crop tissues. cardiovascular disease.6 and 50 . particularly the conjugated carbon–carbon double bond system. the predominant isomer of lycopene is the all-trans geometric form. Low doses of lycopene from tomato (Lycopersicon esculentum) products significantly increased serum lycopene levels and reduced lipid peroxidation in vivo [48]. botrytis).46]. The localization of carotenoid molecules in biological tissues will also influence their ability to encounter and scavenge free radicals. The physical properties of carotenoid molecules.43]. Compounds such as ascorbate.5 in cabbage (Brassica oleracea var.52–54].0:1. hence. The all-trans carotenoids in the LHCs provide efficient singlet-energy transfer to chlorophyll molecules and. This harmful oxidative damage is considered the major cause of aging and degenerative diseases such as cancer. participate mainly in lightharvesting. violaxanthin. postulated to participate in photoprotection. Acyclic carotenoids. permit the quenching of 1O2. and both isomeric groups can be found in vegetable crops (Table 1) [47. Phytoene exists predominately as the 15-cis isomer. Epoxide forms of carotenoids. a-tocopherol and carotenoids are examples of antioxidants that have the ability to quench reactive oxygen species [44]. The 15-cis carotenoids show a preference for isomerization towards the all-trans configurations upon excitation and. oxidants produced during normal metabolism and immune defense against infectious and chemical agents are responsible for damage to DNA. Carotenoid epoxidation and isomerization In the human body. capitata).10 Figure 2. which suggests that there might be a causal relationship [41. lutein and 5. Isomerase activity .46]. 1:6 in spinach (Spinacia oleracea) and 1:23 in kale (Brassica oleracea var. Conjugated double bonds at 5. acephala) [47]. The conjugated double-bond systems of the carotenoids create the light-absorbing chromophores that result in the distinctive colors associated with these plant pigments [16]. as well as the nature and concentration of the reactive oxygen species. proteins and cellular tissues [26.sciencedirect. high levels of fruit and vegetable consumption. Factors that increase MP include increased antioxidant ingestion. normal body mass index.502 Review TRENDS in Plant Science Vol. and a history of no tobacco use. genetics and environment and development of age-related macular degeneration.60 positions are highly effective at quenching singlet oxygen [45. and the RCs contain the 15-cis configurations [49–51]. Studies indicate that a high intake of a variety of vegetables. although strong associative relationships are reported. and some might even have pro-oxidant activity.60 position. The antioxidant activity of cyclic carotenoids comes from the susceptibility of 5. 1:2 in broccoli (Brassica oleracea var. The ratios of lutein epoxide:all-trans-lutein was 1. However. www. the all-trans configurations of carotenoids are the major component of the LHCs.6 or 50 . Indeed. A direct correlation between MP levels and development of macular disease has not been established [39. immune-system decline and cataracts. epoxide isomers are predicted to lack antioxidant activity because they are unable to bind 1O2. and the nutritional status of subjects used in human dietary intervention studies all affect the interpretation of the results [14. antheraxanthin. Carotenoid carbon positions and structure of zeaxanthin.44]. In the pigment–protein complexes of photosynthetic organisms.6-epoxylutein. are better suited for photoprotective functions in the RCs [49].6 and 50 . however. and diminished MP might be related to retinal damage [39.

sativa). all-trans b-carotene. Changes in the growing air temperature [75]. neolutein. 9-cis zeaxanthin. all-trans neoxanthin.83. All-trans b-carotene. potato (Solanum tuberosum subsp.79] www. 90 -cis lutein.81. 13-cis lutein. 9-cis lutein.63. tuberosum) [71]. 13-cis lutein. all-trans zeaxanthin. However. thermal degradations in carotenoid chemistry might adversely affect bioavailability in some food crops. 13-cis zeaxanthin. all-trans neoxanthin. thermal. violaxanthin.57]. lutein and zeaxanthin. and cis–trans isomers differ in their intestinal absorption in humans (Figure 3). 90 -cis neoxanthin. Food processing activities. green a Refs [54. all-trans g-carotene. b-carotene. Carotenes are entirely lipophilic molecules located in the hydrophobic cores of plant membranes. indicating a preference for cis-lycopene in intestinal absorption [58. All-trans b-carotene. all-trans zeaxanthin. irradiance level [76]. All-trans a-carotene. all-trans lutein. 9-cis zeaxanthin and 13-cis zeaxanthin. Significant differences in carotenoid accumulation among different vegetable crop species have been reported [8. composition and release of carotenoids from the food matrix. degradation or isomerization will ultimately have an influence on carotenoid biochemistry and bioavailability. greater excretions of cis-b-carotene and lower excretions of trans-b-carotene were measured in human subjects after ingestion of both raw and processed carrots (Daucus carota var.70.sciencedirect. Absorption of carotenoids in humans is passive and follows digestive pathways similar to those of lipids. all-trans zeaxanthin. 90 -cis lutein. Commodity Beans. 90 -cis lutein. xanthophylls are largely hydrophobic molecules with their polar groups at opposite ends of a non-polar carbon skeleton [84]. b-carotene. all-trans neoxanthin. 13-cis lutein. 13-cis lutein. all-trans and Spinach cis lutein epoxide.98] [67. 13-cis lutein. biotic or abiotic activities that expose carotenoid molecules to potential oxidation. 9-cis zeaxanthin and 13-cis zeaxanthin. violaxanthin. b-cryptoxanthin. Human blood plasma contains mostly all-trans carotenoids but levels of the cis form of some carotenoids are as high as 50% [55]. Kale/collards All-trans b-carotene. is present in higher plants to mediate cis to trans conversions [15]. neolutein. 90 -cis neoxanthin. The accumulation of carotenoid in plant foods is also influenced by environmental growing conditions.68] [54.and chemical-isomerization [49]. kale [61.98] [55. Processing activities usually increase bioavailability through increased release of bound carotenoids from the food matrix. All-trans b-carotene. violaxanthin. Pepper (bell) a-Carotene.82. 90 -cis lutein. indicating an absorption preference for alltrans b-carotene [60]. Factors that influence carotenoid accumulation and bioavailability Carotenoid accumulation in plant tissue appears to be shaped by the physiological.82] but decrease during senescence [1]. b-cryptoxanthin. freezing or low-temperature storage generally preserves carotenoid concentrations by reducing potential enzymatic oxidation [85. all-trans lutein. neolutein. as well as by environmental growth factors such as light. 9-cis lutein. all-trans and Lettuce cis lutein epoxide. 9-cis lutein. lutein and lycopene Carrot a-Carotene.86]. phytofluene and lycopene-5. phytoene. Carotenoid concentrations increase in leaf tissues with maturity [81. Carotenoid accumulations have been shown to increase and decrease in response to environmental manipulations. lactucaxanthin. 9-cis lutein.Review TRENDS in Plant Science Vol. such as thermal processing. Because of their lipophilic nature. 9-cis lutein.60. genetic and biochemical attributes of a plant species. neolutein. neolutein. Protein. 90 -cis neoxanthin. 13-cis lutein. all-trans and Broccoli cis lutein all affect plant carotenoid accumulations.6 diol. lettuce [70]. Tomato (raw) All-trans b-carotene. the majority of lycopene found in fresh and processed tomatoes exists in the all-trans form [56.85. 90 -cis neoxanthin. 9-cis zeaxanthin and 13-cis zeaxanthin.83].86].or membrane-bound carotenoids must first be released from tissues and dissolved in a hydrophobic .63.99. all-trans lutein.65.69]. The genetic variation for carotenoid concentrations within species might be advantageous to vegetable improvement programs. However. corn [68].73] and soybean (Glycine max) [74]. all-trans violaxanthin. 9-cis lutein. all-trans lutein. all-trans and cis lutein epoxide. irradiance photoperiod [77] and nutritional fertility [78.66].100] Carotenoids in bold text identify those reported in highest concentrations. Manipulation of cultural growing conditions and time of harvest would therefore affect the carotenoid concentrations of fruit and vegetable crops. transportation within the lipoprotein fractions. pepper [72. As mentioned previously. all-trans d-carotene.14. all-trans lutein.59]. all-trans lycopene. biochemical conversions and tissue-specific depositions. all-trans zeaxanthin. 90 -cis neoxanthin.98] [54. All-trans carotenoids in plants are susceptible to photo-. probably through isomerization or oxidation reactions [53. The increased coloration in vegetable and fruit tissues associated with maturity is often indicative of increases in carotenoid concentrations [72. By contrast. 90 -cis lutein. all-trans neoxanthin. Corn all-trans zeaxanthin and 9-cis zeaxanthin.85] [54. temperature and fertility [61–64].73.98] [54. with different results for different plant species. all-trans lutein. Similarly. Carotenoids identified and quantified in major vegetable crops Carotenoids identifieda All-trans b-carotene. as well as by the nutritional status of the ingesting host [12. all-trans lutein. all-trans zeaxanthin. j-carotene. however. amount consumed and absorption in the intestinal track.80]. lycopene in human and animal tissues exits predominately as cis-isomers. The first step in carotenoid bioavailability is release from the food matrix. mincing or liquefying result in changes to carotenoid chemistry. The bioavailability of carotenoids from plant foods is highly variable and is influenced by the species and structures of carotenoids present in the food. 90 -cis lutein. 9-cis zeaxanthin and 13-cis zeaxanthin. violaxanthin.11 No. neurosporene.98] [73] [54. all-trans lutein epoxide. capsanthin. and significant genetic variation within species has been found for carrot [67].10 503 Table 1.

48.91]. b-carotene. Carotenoid compounds can remain in the liver. a-carotene. Carotenoid absorption requires the presence of dietary fat in the small intestine.88. to the liver. Current results demonstrate that in vitro Caco-2 cells can be used to predict carotenoid bioavailability from supplements and whole foods. domain (oils. Some caveats to interpreting serum carotenoid bioavailability include: (i) serum www. After release from the food matrix. which are eventually delivered to the blood stream and. Carotenoid bioavailability is usually assessed in blood serum after ingestion in dietary trials or cohort studies. (ii) carotenoids measured in serum signify an equilibrium between intestinal absorption.11 No.sciencedirect. Recent studies have shown that absorption of carotenoids increases when they are ingested with dietary lipids [87. carotenoids are incorporated into chylomicrons. Recent studies have also used in vitro Caco-2 human intestinal cell lines to assess carotenoid bioavailability [89–91]. carotenoids in the mostly aqueous environment in plant foods must be transferred to bulk lipids or intestinal micelles in the digesta [14].87. or be transferred to low-density or high-density lipoproteins before eventual tissue-specific deposition [12].com responses to single oral doses of carotenoids are highly variable. which stimulates the release of emulsifying bile acids by the gall-bladder [12]. and (iii) high concentrations of endogenous carotenoids (i. Absorption potential is measured using standard HPLC carotenoid analysis. The occurrence and properties of carotenoid isomers in food crops can influence intestinal absorption. pure carotenoid compounds and whole food samples are brought through an in vitro digestion and reacted with Caco-2 human intestinal cells. lycopene and lutein) are already present in serum [25]. In these studies. Our own research has revealed differences in serum carotenoids and responses to macular pigment optical density (MPOD) evaluations in human subjects administered different doses of lutein from mono-molecular supplements and whole food sources.88].83. The relatively simple analysis quantifies carotenoid changes in serum at various time intervals following ingestion of whole foods or supplements. carotenoids are assimilated and oriented into lipid micelles before uptake by intestinal mucosal cells.504 Review TRENDS in Plant Science Vol. Serum carotenoid levels were . Increases in serum carotenoid concentrations typically result after ingestion of carotenoids from whole food or mono-molecular supplements [25. tissue uptake and tissue release. fats or bulk lipid emulsions) [14]. ultimately.e.60. breakdowns. After release to the enterocyte. Structures of some common all-trans and cis carotenoid isomers found in fruit and vegetable crops. Because of their hydrophobic nature.10 Figure 3.

D. and Bramley.M. Clin. et al. Nutr. 531S–534S 13 Sandmann. However. eds). A two. and Extension Service. et al. (2003) Retinoid. O. In The Photochemistry of Carotenoids (Frank. Fluxes of metabolites away from GA resulted in stunted tomato plants with abnormal pigmentation. Carl Sams. J. 4–12 14 Faulks. (2000) Relation among serum and tissue concentrations of lutein and zeaxanthin and macular pigment density. and Southon. (2002) Factors that influence the bioavailability of xanthophylls. and Erdman. F. 287–290 5 Seifried. Lipid Res.) 95.L. xanthophyll and carotene production in rice (Oryza sativa) endosperm (‘Golden Rice’) has been increased using exogenous cyclase and desaturase enzymes [97]. Biochim.D. G. et al.W. Evan Ford. Successful approaches have centered on modification of the biosynthetic pathway to change the flux and end products. 95–100 15 Fraser. 180–187 11 Hughes. US Department of Agriculture. Arch. Laura Dukach. Immunology 110. 90. 713–718 12 Zaripheh. (2005) Challenges to understanding and measuring carotenoid bioavailability. Am. Cancer Res. J. Nutr. the degree of isomerization before. Joanne Curran-Celentano. whether combating widespread vitamin A deficiency or fighting aging eye diseases. 907–910 9 Johnson. but the question still remains: will the general public accept genetic modification of raw or unprocessed foods? Programs designed to increase carotenoid bioavailability in food crops must begin with a solid understanding of carotenoid chemistry and biosynthesis. There are obvious advantages to genetic engineering techniques. glucosinolates. as well as other isoprenoid pathways and metabolic regulations. (2005) Proposed criteria for assessing the efficacy of cancer reduction by plant foods enriched in carotenoids.A.E. Joe Sheehan and Adam Wenzel for their collaborative participation in our projects.11 No. In vitro and in vivo studies show that carotenoid bioavailability is influenced by source (whole food versus supplement). (2004) The biosynthesis and nutritional uses of carotenoids. et al. Identification and cloning of genes and cDNAs in the carotenoid pathway have facilitated genetic manipulations. P.sciencedirect. Acknowledgements We thank Pam Bishop. 4295–4298 6 Finley. R. 487–502 8 be accompanied by a clear understanding of how these manipulations will ultimately affect plant physiology. Acta 1740. Education. transit time in the intestine. interactions with other carotenoid compounds. Soc.W. genetic manipulations to the carotenoid biosynthetic pathway have resulted in abnormal growth and dwarfing in tomato [93]. Advances in carotenoid biosynthesis have been facilitated using molecular approaches. A. et al. during and after absorption. (1999) Effects of carotenoids on human immune function. Bot. under Agreement No.15. References 1 Gross. Funding for our carotenoid research came in part from a grant received by the Cooperative State Research. Mark Lefsrud. 132. Catherine Gerwick. Br. care must be taken to outline specific research impacts. both stemming from the common precursor GGPP. The tomato examples show the need to understand how genetic engineering influences carotenoid precursor pools. 228–265 . J. Nutr.92]. success has also been achieved overexpressing phytoene desaturase enzymes [15]. Prog. S. Nutr. 1075–1096 7 Granado. and engineering carotenogenic behavior in tissues completely devoid of carotenoid activity [13]. (1998) Fruits and vegetables that are sources for lutein and zeaxanthin: the macular pigment in human eyes. Before selecting candidate crops for enhancement efforts. et al. (2001) Carotenoid biosynthesis and biotechnological applications. improvements in carotenoid concentrations using traditional breeding efforts are limited because of the high costs associated with analytical laboratory measurements. pp. S. Scott McElroy. A. H. (2003) Nutritional and clinical relevance of lutein in human health. Recent reviews have chronicled molecular advances in carotenoid pathway manipulation to improve biosynthesis and partitioning [13.M. pp. and the nutritional status of the human subjects [14]. Lastly. J. New York Academy of Sciences 4 Tang. 385.10 505 higher than the baseline level after ingestion of 10 mg or 30 mg lutein supplements or spinach fortified with 8 mg or 12 mg lutein per 100 g fresh weight. 63.Review TRENDS in Plant Science Vol. enzyme activities and locations of expression. Most notably. AVI/Van Nostrand Reinhold 2 DellaPenna. L. Genetic strategies have also increased b-carotene production in canola (Brassica napus) using a seed-specific promoter [95] and increased xanthophyll carotenoid production in potato using a tuber-specific promoter (epoxidation reactions were suppressed) [96]. Ophthalmol. Ann. eds). Understanding how carotenoids in the food matrix respond to food processing activities and any subsequent changes to carotenoid chemistry will be of primary importance. J. Biophys. (1999) Carotenoid synthesis and function in plants: insights from mutant studies in Arabidopsis thaliana. but they must www. (2005) Lycopene inhibits the growth of human androgen-independent prostate cancer cells in vitro and in BALB/c nude mice. John McLean. Assessment of genetic variation in several crop species can be assumed to be the beginning of improvement programs using traditional breeding methods. (1993) Biological functions of carotenoids. Kluwer 3 Bendich. 2001–52102–11254. Nutr. J.and carotenoid-enriched diets influence the ontogenesis of the immune system in mice. (Lond. (2003) The antioxidant conundrum in cancer.. 58. P. et al.A. L. threefold increase in tomato fruit-specific carotenoid accumulation has been achieved using a bacterial phytoene synthase (crtB) and a tissue-specific promoter [94].J. 135. In Carotenoids in Human Health (Canfield. Kopsell. These results were attributed to metabolic precursor competition between carotenoid and gibberellin (GA). increasing pre-existing carotenoids.M. however. Biochem. Proc. unpublished). However. 82. Most studies have demonstrated that phytoene synthase has the greatest control over fluxes in the carotenoid pathway. degree of processing. J. Biophys. (1991) Pigments in Vegetables: Chlorophylls and Carotenoids. 61–67. E. Increases in MPOD from the baseline to the end of a 12 week intervention occurred in subjects administered 30 mg lutein supplements and spinach with 12 mg lutein per 100 g fresh weight (D. 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