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ABSTRACT Dietary phytonutrients found in vegetables and able (17). Although potentially beneficial to human health in
1424 Am J Clin Nutr 2000;72:1424–35. Printed in USA. © 2000 American Society for Clinical Nutrition
BITTER TASTE AND PHYTONUTRIENTS 1425
that they are selectively bred out of plants and routinely removed Genetic linkage studies in humans linked the ability to taste
from processed foods (20, 26). Indeed, the low amounts of bitter PROP with a locus at 5p15 (44). The ability to taste PTC and
plant compounds in the current diet largely reflect the achieve- PROP, once thought to be transmitted as a dominant taster gene
ments of the agricultural and food industries (26). The debitter- (45), may involve more than one locus. Genetic linkage studies
ing of plant foods has long been a major sensory concern for suggested that PROP tasting may involve both a specific sensitivity
food science (27). to PROP and a more general bitter taste responsiveness (46). The
early studies segregated PTC and PROP tasters from nontasters on
the basis of their tasting of PTC or PROP crystals or the bimodal
BITTER TASTE AND FOOD REJECTION distribution of detection thresholds for PROP solutions (47). The
Abnormal bitterness tends to be equated with dietary danger, wide variability of the tasters’ responses led Bartoshuk (41) to pro-
and rightly so. Rancid fats, hydrolyzed proteins, plant-derived pose the existence of PROP “supertasters.” PROP supertasters,
alkaloids, and other toxins generally have an unpleasant bitter most of them women, were identified by the high ratio of perceived
taste (37). Microbial fermentation also results in bitter-tasting bitterness intensity of PROP to the perceived saltiness of sodium
compounds (37). Bitterness has been reported in such diverse chloride solutions. On average, supertaster women had more fungi-
TABLE 1
Some bitter phytonutrients in common plant foods1
Phytonutrient class Typical component Taste quality Food source Amount present Reference
Phenolic compounds
Flavanones Naringin Bitter Grapefruit, flavedo 2701–4319 mg/kg 55
Grapefruit, albedo 1301–15 592 mg/kg
Grapefruit, pith 13 285–17 603 mg/kg
Grapefruit, seeds 295–2677 mg/kg
Immature grapefruit 97 920–144 120 mg/kg FW 56, 57
Grapefruit juice 300–750 mg/L 56
Oroblanco juice 346–489 mg/L 58
Melogold juice 413–580 mg/L 58
Flavones Tangeretin Bitter Orange fruit 0–30 mg/kg DW 59
Orange juice 0.6 mg/L
(Continued)
BITTER TASTE AND PHYTONUTRIENTS 1427
TABLE 1 (Continued)
Phytonutrient class Typical component Taste quality Food source Amount present Reference
Triterpenes
Limonoid aglycones Limonin Bitter Lemon juice 12.2 mg/L 56
Orange juice 9.7 mg/L
Grapefruit juice 11.4 mg/L
Tangerine juice 34.7 mg/L
Grapefruit, flavedo 6.1–42 mg/kg
Grapefruit, albedo 11.6–65 mg/kg
Grapefruit, pith 103–525 mg/kg
Grapefruit, seeds 1188–1885 mg/kg
Nomilin Bitter Grapefruit juice 0.1–1.6 mg/L 58
Oroblanco juice 0.4–0.8 mg/L
Melogold juice 0.9–1.8 mg/L
nins complex with proteins, starches, and digestive enzymes and are depending on the type of the glycoside chain. Naringin, a flavanone
thought to reduce the nutritional value of foods (8). neohesperidoside, and neohesperidin are very bitter, whereas hes-
peridin is tasteless. On the other hand, neohesperidin dihydrochal-
cone is intensely sweet. Naringin concentrations are highest in young
BITTER FLAVONOIDS IN CITRUS FRUIT leaves and in the pulp (albedo) of immature fruit (59). Although typ-
Flavonoids in citrus fruit include flavanones (naringin), flavones ical concentrations of naringin in grapefruit juice are 400 mg/L ,
(nobiletin), and flavonols (quercetin). Polymethoxylated flavones problems with early-season grapefruit juice can raise naringin con-
(tangeretin and nobiletin) are concentrated in the skin of unripe fruit centrations to a less acceptable 1200 mg/L (80). Blending of juices
and are the constituents of bitter citrus oils (79). Bitter flavonoids means that bitterness affects even more of the crop.
may act through bactericidal activity or by making the plant unpalat- Limonin, a triterpene, is responsible for the so-called delayed
able (17). Some flavonoids are very bitter whereas others are not, bitterness of citrus juices. A tasteless limonin precursor,
1428 DREWNOWSKI AND GOMEZ-CARNEROS
FIGURE 4. Relation between bitterness intensity (on a scale of 0–100) and the proportion of consumers who remarked that the product was “too
bitter” or had “poor taste.” Data from reference 36.
1430 DREWNOWSKI AND GOMEZ-CARNEROS
most bitter varieties. Bitter taste had been identified as the main
reason for avoidance of vegetables and was reported as being the
least well tolerated for Brassica vegetables, spinach, squash, and
onions (30, 73, 103). The current health-oriented push toward
selective breeding of phytonutrient-rich and therefore more bit-
ter varieties runs counter to the published studies on vegetables
and consumer acceptance (30, 73, 103).
Creation of new and less bitter cultivars has also occurred
with other crops. A wholesale reduction of glucosinolates in
such oilseed crops as rapeseed and Crambe abyssinica was car-
ried out successfully worldwide (105). The development of a
transgenic citrus fruit, free of limonin, was also described in
FIGURE 5. Relation between progoitrin and sinigrin content of the literature (106).
Some scientists have argued that these modifications may
TABLE 2
Summary of industrial debittering processes to remove bitter phytonutrients from foods
Phytonutrient class Typical compound Food Debittering process Reference
Flavanones Naringin Grapefruit juice Adsorption to polymers 74, 107, 108
Naringenin Orange juice Passage through resins 56, 109, 110
Flavones Tangeretin Passage through enzyme matrix 111, 112
Nobiletin Passage through microbial mass 113–116
Sinensetin Accelerated ripening with ethylene 116
Triterpenes Limonin Use of cyclodextrin polymers 117, 118
Genetic engineering: transgenic citrus trees 106
Flavans Catechin, epicatechin Tea Fermentation 65, 82
Chocolate Removal of chocolate liquor 84
Catechin polymers Wine Precipitation with proteins: egg white, casein, 53, 54
gelatin, isinglass
chemoprotective role of phytochemicals. Chemoprotective agents which is derived from glucoraphanin. Inducer potency has been
generally belong in 1 of 3 categories: those that block metabolic linked to concentrations of bitter sinigrin, progoitrin, glucobras-
activation of carcinogens, those that prevent the formation of car- sicin, gluconapin, and glucoraphanin. As a result, several clinical
cinogens from precursors, and those that suppress neoplasia in studies attempted to feed patients large amounts of Brassica veg-
cells previously exposed to carcinogens (123, 124). Phytonutrients etables daily. In some cases, respondents were fed 300 g Brus-
and their metabolites elicit a variety of biological activities, acting sels sprouts/d for 3 wk or 500 g broccoli/d for 12 d (11). Other
as antioxidants, phytoestrogens, or enzyme inducers (125). studies used a mix of Brussels sprouts, cabbage, broccoli, and
Among the most promising compounds under study are bitter phe- cauliflower in the amount of 400 g/d (11).
nols and polyphenols, flavonoids, isoflavones, and glucosinolates. Epidemiologic outcomes of diets high in vegetables and fruit
Phenolic acids, catechins, flavonols, and polymeric antho- were summarized in a recent report (4). Research has focused on
cyanins in wine were reported to have antioxidant activity (126, vegetables rather than fruit, with special emphasis on green leafy
127). Although the bioavailability of phenols and polyphenols is a and Brassica vegetables, citrus fruit, soybeans, and red wine.
matter for further research, only small amounts of food polyphe- Population-based studies (9, 129) showed that the risk of coro-
nols estimated at ≤ 30 mg/d are absorbed (17). However, even such nary heart disease was reduced at higher estimated intakes of
low amounts may have a potent antioxidant effect in vivo (17). flavonoids (apigenin, kaempferol, luteolin, myricetin, and
Even at very dilute concentrations (10 mol/L) in vitro, phenolic quercetin). Data on diet and colon cancer (64) also show a signi-
compounds had antioxidant effects (128). ficant decline in risk with higher consumption of vegetables,
Plant flavonoids may protect against LDL oxidation through a green leafy vegetables, and cabbage. There is a general consen-
reduction of free radicals, chelation of metal ions, or protection or sus that a diet higher in plant foods than is the current norm is
regeneration of -tocopherol (9, 17, 123). Studies of their anticar- associated with improved health and reduced disease risk. How-
cinogenic action focused on the activation of enzymes involved in ever, the most potent plant products are also likely to be the most
the metabolism of xenobiotics (phase 2 enzymes). Naringin, the bitter and therefore the most aversive to the consumer.
bitter flavonoid component of grapefruit juice, or its metabolites,
are reported to inhibit the activity of cytochrome P-450 (49).
Glucosinolates and isothiocyanates are also regarded as dietary THE DILEMMA OF THE FOOD INDUSTRY
protectors against cancer (7). Isothiocyanates inhibit the activa- The competing demands of taste and health pose a dilemma
tion of carcinogens by cytochrome P-450 (phase 1) enzymes and for the food industry. The major determinant of food selection is
promote detoxification of activator carcinogens by inducing taste. Foods that are bitter, acrid, or astringent tend to be rejected
phase 2 enzymes. Phase 2 enzymes inactivate carcinogens by by the consumer—and generally for the right reasons. The
neutralizing their toxic properties and speeding their elimination instinctive rejection of bitter taste may not be modifiable because
from the body (33). Some phase 2 enzymes function as inhibitors it is a key mechanism for survival.
of cytochrome P-450 (7). The results of early studies (18, 19) and Bitter taste is the main reason for the rejection of diverse food
more recent data suggest that one of the most potent inducers is products (23, 30). Bitterness of cruciferous vegetables has been
sulforaphane, or 4-methylsulfinylbutyl isothiocyanate (33), linked repeatedly to their low acceptance (30, 48). Focus groups
1432 DREWNOWSKI AND GOMEZ-CARNEROS
conducted by the National Cancer Institute before the 5-A-Day molecular level (137). However, it now appears that the sec-
project found that dislike of certain vegetables was a major bar- ondary and tertiary functions are linked, if not at odds with each
rier to vegetable consumption (130). In other studies, bitter Bras- other. For the most part, plant-based phytochemicals are bitter
sica vegetables were thought to taste good only if sauces were and therefore aversive to the consumer. The discovery of a fam-
added, that is, after the addition of fat, sugar, or salt (131). ily of ≥ 50 different bitter-taste receptors only confirms how
Excessive bitterness lowers the acceptablity of citrus juices (81). important sensitivity to bitter taste was to evolution and sur-
The taste of soy proteins and some soy products is generally vival. Research on plant-based functional foods presents several
described as objectionable (35). Bitterness of wine is reduced by challenging opportunities for cancer biology and nutrition sci-
aging and the removal of phenols and polyphenols (53, 54). The ence. However, issues of taste and behavioral nutrition ought to
food industry has engaged in these practices for years in be considered as well.
response to consumer demand.
Not all bitterness is rejected automatically (132). In selected
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