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Potatoes and Human Health

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Critical Reviews in Food Science and Nutrition, 49:823–840 (2009)
Copyright
C Taylor and Francis Group, LLC
ISSN: 1040-8398
DOI: 10.1080/10408390903041996

Potatoes and Human Health

MARY ELLEN CAMIRE,1 STAN KUBOW,2 and DANIELLE J. DONNELLY3

1
Department of Food Science & Human Nutrition, University of Maine, USA
2
School of Dietetics & Human Nutrition, McGill University, QC, Canada
3
Plant Science Department, McGill University, QC, Canada

The potato (Solanum tuberosum L.) tuber follows only rice and wheat in world importance as a food crop for human
consumption. Cultivated potatoes have spread from the Andes of South America where they originated to 160 countries
around the world. Consumption of fresh potatoes has declined while processed products have increased in popularity. As the
potato becomes a staple in the diets of an increasing number of humans, small differences in potato nutritional composition
will have major impacts on population health. The potato is a carbohydrate-rich, energy-providing food with little fat. Potato
protein content is fairly low but has an excellent biological value of 90–100. Potatoes are particularly high in vitamin C
and are a good source of several B vitamins and potassium. The skins provide substantial dietary fiber. Many compounds in
potatoes contribute to antioxidant activity and interest in cultivars with pigmented flesh is growing. This review will examine
the nutrient and bioactive compounds in potatoes and their impact on human health.

Keywords Cardiovascular disease, glycemic index, chlorogenic acid, obesity

This review does not intend to replace the numerous reviews were first spread to European countries, including Spain and
on potato as nutrient or food item but only to summarize the England, in the late 1500s (CDC, 1999b). The potato may have
contribution of potato to long-term human health. This humble been instrumental in preventing scurvy among early consumers,
tuber has sustained many generations and new research has including sailors, due to its relatively high vitamin C content
revealed possible directions for breeding programs to enhance (Buckenhüskes, 2005). Potatoes became so widely distributed
the nutritional composition of potatoes to meet the needs of and important, especially in certain parts of Europe, that they
specific groups of consumers. are often referred to as “European” or “Irish” potatoes. Potatoes
are now grown in 160 countries (AAFC, 2007) with over 4,000
cultivars (Hils and Pieterse, 2007). Potatoes are eaten fresh or
ORIGIN OF POTATO AS HUMAN FOOD following storage; prepared in a multiplicity of different ways
at home or commercially processed into many different foods.
The cultivated potato (Solanum tuberosum L.) originated in Despite its apparent diversity, the cultivated potato contains
South America where it has been used for food for over 10,000 only a fraction of the potential biodiversity that is present in
years (CDC, 1999a) and domesticated during pre-Columbian South American cultivars and cross-compatible wild species
times over 8,000 years ago (CIP, 2008a). Pre-Columbian food (reviewed by Bradshaw, 2007). In a comparison of 205 cul-
preparation involved the production of chuño, which was used tivars and 1220 genotypes of wild and cultivated species for
in soups and stews, much as it is still produced in the Andes dry matter, starch, resistant starch, starch granule diameter, and
today (Woolfe, 1987). Harvested potato tubers are piled on the protein content, Jansen et al. (2001) found more variability in
ground and repeatedly frozen over several nights. Skin removal wild than cultivated potatoes for all characters. For this reason,
(through trampling) is followed by soaking or leaching in run- and because the potato genome is known, there exists enor-
ning river water. This serves to remove the bitter flavors caused mous capacity through breeding to improve potato towards an
by glycoalkaloids. The final step involves sun-drying which pre- increasingly more healthy food item. The need for new cultivars
serves the chuño for up to several years. Another product, papa is primarily driven by goals of stable or improved yields un-
seca, is produced by boiling, peeling, slicing, then sun-drying der more sustainable growing conditions (primarily lower fuel
and grinding and is also used in stews and soups. Potatoes and fertilizer inputs), environmental stresses, and disease and
pest pressures. Where daily per capita availability of nutritious
Maine Agricultural & Forest Experiment Station external publication 3052. food is below recommended phytochemical intake, diet diver-
Address correspondence to Mary Ellen Camire, Department of Food Science
and Human Nutrition, University of Maine, 5735 Hitchner Hall, Orono, ME,
sification, and improved preparation and processing to increase
USA, 04469–5735. E-mail: [email protected] micronutrient bioavailability is needed (Hanson et al., 2004),
823
824 M. E. CAMIRE ET AL.

and improved potato cultivars may have a significant role here. compared with major gains in the minor food items (Bamberg
Where food security is not an overwhelming issue, consumer and del Rio, 2005).
demand is for more convenience foods, improved nutritional and Undermining the overall gains in Asian and African prosper-
health properties, including organically-grown produce, better ity over the past decade are recent soaring food prices resulting
flavor, and novel foods. from diversion of grain from feed to biofuels and associated
speculation in the commodities markets (Fresh Plaza, 2008a).
Fuel and fertilizer shortages are also impacting agricultural pro-
POTATO PRODUCTION – CURRENT AND FUTURE
duction. Potato increasingly contributes to world food secu-
TRENDS
rity and has a critical role to play in developing nations facing
The potato has achieved worldwide prominence as a food hunger. Potatoes supplement or replace grain-based diets where
item, in part, due to its tremendous yield per unit area compared rice, wheat, or corn availability has lessened or price has be-
with many other food crops. Average potato yield in metric tons come unaffordable. The potato now ranks third, behind rice and
per hectare (mtha−1 ) in North America (40.6) is far in excess wheat, for human food as the use of corn for biofuels and animal
of those achieved in Europe (17.3), Latin America (16.6), Asia feed has lessened its human food applications.
(15.7), or Africa (10.8) (2005 figures, CIP 2008b). There would Predicted global climate change for 2010–2039, with asso-
seem to be much potential for increased yield in many potato ciated increased temperatures, is expected to cause upheaval
growing areas. Higher production in parts of North America in most agricultural, including potato, growing areas (Hijmans,
is favored by many factors, including cool climate with ample 2003). Major cultural adaptations involving planting time and
rainfall, mechanization and efficiencies of scale, relatively high cultivar choice (particularly more heat-tolerant cultivars) will
inputs (fertilizer, pesticides), long growing season which favors need to be implemented to mediate impact of increased tempera-
the higher yielding long-season cultivars, and production sys- tures. However, effects will vary with different geographic areas
tems involving rotation with cereals and forages that improve where potato is grown. Areas expected to be least affected are
soil structure and discourage disease. at high latitudes (Canada, China, Russia, Scandinavia) or high
World potato growing areas are in a state of dynamic change. altitudes in the tropics (Peruvian/Bolivian Altiplano) where pro-
Global production statistics collected by FAO were tabularized duction area may increase. Areas expected to be most affected,
and mapped by CIP (2008b). The production levels in the devel- with predicted large yield declines, comprise a zone that includes
oped nations of Europe, North America, and the former Soviet southeastern Europe, Russia, and Kazakhstan. Subtropical areas
Union have declined by 30 million metric tons (mmt) within such as India and Bangladesh where potatoes are already grown
the past 16 years (183 and 156 mmt in 1991 and 2007, respec- during the coolest season will be greatly affected without much
tively). During this same interval, production has doubled in scope to mediate yield.
countries of the developing world including Asia, Africa, and
Latin America (85 and 165 mmt in 1991 and 2007, respec- POTATO VARIETIES AND CLASSIFICATION FOR
tively). The top three world leaders in potato production in 2007 FOOD USES
included China (72.0 mmt), the Russian Federation (35.7 mmt),
and India (26.3 mmt). Production in China is forecast to reach Potato varieties can be classified using a broad range of cri-
81 mmt in 2010 (Wang and Zhang, 2004). Asian production, teria. One common classification is based on the number of
overall, is increasing at about 6% annually (Bamberg and del days to maturity following planting of whole or cut seed tu-
Rio, 2005). Demand is fueled by increasing numbers of potato- bers. For example, potatoes are classified as very early (65–70),
consumers in both production and non-production areas, interest early (70–90), mid-season (90–100), late (110–130), or very
in processed potato products for a developing food service in- late (>130) (CFIA, 2008). Cultivars with longer maturity times
dustry, use as animal feed for a growing livestock industry, and generally out-yield the shorter maturity cultivars. “New” pota-
anticipated export potential to neighboring countries including toes are often short-season cultivars that are harvested early as
Japan and South Korea. It is expected that China will increas- small, tender potatoes and usually boiled for table use.
ingly export fresh potatoes and import processed potato product. Varieties may be classified based on tuber quality traits suited
The situation is similar in India, where both internal and export to specific cooking or processing activities (boiling, baking,
markets continue to increase (CommodityOnline, 2008). dehydrating, frying, etc.). These may be divided further. For
Almost half of the global potato supply is now consumed example, cultivars preferred for frying are generally separated
in Asia. The consumption per capita in Asia is on the increase into chipping (round) or French fry (elongate) types. Further
as of 2005, but is still relatively low (26 kg) compared with to this, cultivars can be classified based on storage properties.
Europe (96 kg) or North America (58 kg) (CIP, 2008b). This Some cultivars must be eaten or processed following harvest,
suggests that demand for potato in Asia could double or triple and others maintain their starch properties longer in storage.
over the next few years. As prosperity increases, consumption During storage, or prolonged storage, starch may be converted
may both increase and shift to include processed food products to reducing sugars that caramelize during the frying process.
(CIP, 2008c). As consumption increases, relatively small im- Cultivars that can be stored for longer are advantageous to the
provements in nutrition impact increasingly on consumer health food industry.
 POTATOES AND HUMAN HEALTH 825

Consumer preference may relate to tuber periderm (skin) 21.9 to 42.7 (mean of 31.2) in the cultivars (Jansen et al., 2001).
and/or flesh color. The most common colors include brown, red, This indicates some potential for increased amylose through
white, or yellow skin with white or yellow flesh. Skins that are selection among cultivated potato, and potential improvement
russet (brownish or reddish-brown with a raised “fish net”) can through breeding. As the crystalline structure of native potato
be clearly distinguished from those that are not. The “specialty” starch is generally impervious to the action of amylolytic di-
or “novelty” potatoes reflect the variation seen among South gestive enzymes, there is a substantial resistance of raw potato
American wild species that possess a broader range of pigmen- starch to digestion and so it acts physiologically as a “resistant
tation. For example, phenotypes occur with colors such as purple starch.” The resistance to amylase digestion is greatly dimin-
and blue skin and/or flesh color. Potato pigments include antho- ished when potato starch is gelatinized following cooking as
cyanins, carotenoids, and precursor flavonoids and phenolics gelatinization causes loss of crystallinity leading to the solubi-
(van Eck, 2007). These pigments are secondary metabolites that lization of the starch polymers. Starch quality traits of cooked
are important in plant defense (Hahlbrock and Scheel, 1989). potato that impact on health include the amylose:amylopectin
These substances and others contribute as antioxidants, with ratio and the degree of phosphorylation. The branched structure
important health properties that are discussed in a later section. of amylopectin allows for greater digestibility than the linear
chain structure of amylose, which leads to a higher glycemic
response. Nutritionally, a greater proportion of resistant starch
POTATO NUTRITIONAL VALUE (or more slowly digested starch) is considered advantageous as
it provides similar health benefits to fermentable fiber. Resistant
The potato has been widely accepted throughout the world as starch refers to the summation of starch and starch degradation
a staple food and is available in many forms yet many consumers products that are not absorbed in the small intestine. Upon cool-
are unaware of the healthful attributes of the tubers. The potato ing following cooking, higher-amylose starches have greater
has greater dry matter and protein per unit growing area com- retrogradation following processing compared with those with
pared with the cereals (Bamberg and del Rio, 2005). Despite this, more amylopectin. Retrogradation causes the starch to become
consumers tend to believe that potatoes are high in calories and more crystalline leading to resistance to digestive enzymes.
in fat compared with other carbohydrate sources such as rice or Higher amylose starches also reduce oil penetration, so are fa-
pasta; an incorrect assumption since potato has negligible fat and vored for use in snack foods to decrease consumer fat intake
a low energy density similar to legumes (Priestley, 2006). Edu- (Tarn et al., 2006). Covalently-bound phosphorus is present
cational and research efforts are underway to convert consumers in potato starch at greater levels (0.08%) than in other plant
to the merits of potatoes or potato products as a replacement for starches (0.02% in corn) (Li et al., 2006) and is important to the
cereals or cereal products in cooked and processed food items. physicochemical properties of starch, affecting its gelatiniza-
The reader is directed to recent reviews on nutritional aspects tion and pasting properties (Tarn et al., 2006). Additionally, the
of potato, including Buckenhüskes (2005) and the Symposium: phosphorylated glucosyl residue is not susceptible to cleavage
Enhancing the Nutritional Value of Potato Tubers (Suttle, 2008 by amylolytic enzymes, leading to the release of phosphory-
and others). Potatoes are usually eaten cooked, and most often lated oligosaccharides and reduced digestibility of gelatinized
eaten boiled and unpeeled in many regions of the world. The and raw starch (Kamasaka et al., 1995). Sucrose is the major
various nutrient components of a 100-gram serving of baked, disaccharide of potatoes while glucose and fructose are the ma-
boiled, and French-fried potatoes are presented in Table 1. jor monosaccharides. The equilibrium between free sugars and
starch changes during tuber storage and impacts processing. Too
Carbohydrates much reducing sugar leads to browning when potatoes are fried.

Cooked potatoes are a good dietary source of carbohydrates,

which make up about 75% of the total dry matter of the tuber. Protein
Starch is the predominant carbohydrate in potatoes and serves as
an energy reserve for the plant. While cultivated potato averaged Potato protein ranged from 1–1.5% of tuber fresh weight in
11.0–30.4% starch on a fresh weight basis (mean of 18.8%), the 20 cultivars tested (Ortiz-Medina, 2007). Compared with
wild species ranged from 3.8 to 39.6 with a mean of 18.1% other raw vegetable sources, potatoes are not typically consid-
(Jansen et al., 2001). However, these data were not grouped ered to be a good dietary protein source due to their low overall
based on maturity type. Maturity type is far more important than protein content. However, potato protein has excellent biolog-
remaining genetic variation for tuber yield and starch content ical value (BV), with a BV of 90–100 compared with whole
(van Eck, 2007). The late-maturing cultivars tend to produce egg (100), soybean (84), and beans (73) (Kasper, 2004; cited
much greater tuber and starch yield compared with the early- in Buckenhüskes, 2005). Patatin is the major storage protein
maturing cultivars. and an allergen for some people. This allergenicity is signifi-
Starch is packed in granules that typically contain amylose cantly reduced by heating (Koppelman et al., 2002). The sec-
and amylopectin in a ratio of 1:3. Amylose content in the wild ond major potato tuber protein after patatin is a diverse group
species was 23–37% of the total starch (mean of 29.7%) and of low molecular weight proteins that inhibit proteases of the
826 M. E. CAMIRE ET AL.

Table 1 Nutrient content of 100 g (FW) of potato prepared in common stylesa

Boiled in skin, French fries, frozen Daily Reference Value or

Nutrient Units Baked, flesh + skinb flesh onlyc then oven-heated Reference Daily Intake

Proximates
Water g 74.89 76.98 61.51
Energy kcal 93 87 172
Energy kJ 390 365 721
Protein g 2.50 1.87 2.66 50
Total lipid (fat) g 0.13 0.10 5.22
Ash g 1.33 0.92 1.90
Carbohydrate, by difference g 21.15 20.13 28.71 300
Total dietary fiber g 2.2 1.8 2.6 25
Total sugars g 1.18 0.87 0.28
Sucrose g 0.40 0.19 0.18
Glucose (dextrose) g 0.44 0.37 0.11
Fructose g 0.34 0.30 0.00
Starch g 17.27 20.13
Minerals
Calcium, Ca mg 15 5 12 1000
Iron, Fe mg 1.08 0.31 0.74 18
Magnesium, Mg mg 28 22 26 400
Phosphorus, P mg 70 44 97 1000
Potassium, K mg 535 379 451 3500
Sodium, Na mg 10 4 32 2500
Zinc, Zn mg 0.36 0.30 0.38 15
Copper, Cu mg 0.118 0.188 0.135 2
Manganese, Mn mg 0.219 0.138 0.210 2
Selenium, Se mcg 0.4 0.3 0.2 70
Vitamins
Vitamin C, total ascorbic acid mg 9.6 13.0 13.3 60
Thiamin mg 0.064 0.106 0.128 1.5
Riboflavin mg 0.048 0.020 0.031 1.7
Niacin mg 1.410 1.439 2.218 20
Pantothenic acid mg 0.376 0.520 0.522 10
Vitamin B-6 mg 0.311 0.299 0.184 2.0
Folate, total mcg 28 10 28 400
Vitamin A, RAE mcg RAE 1 0
Carotene, beta mcg 6 2
Vitamin A, IU IU 10 3 5000
Lutein + zeaxanthin mcg 30 9
Vitamin E (alpha-tocopherol) mg 0.04 0.01 0.11
Vitamin K (phylloquinone) mcg 2.0 2.1 2.5
Lipids
Fatty acids, total saturated g 0.035 0.026 1.029 20
Fatty acids, total monounsaturated g 0.003 0.002 3.237
Fatty acids, total polyunsaturated g 0.058 0.043 0.321
Cholesterol mg 0 0 0 300
a Data obtained from the USDA National Nutrient Database for Standard Reference, Release 21 (2008), accessed November, 2008.
bA medium baked potato 2.25–3.25 inches in diameter weighs 173 g. The standard U.S. labeling portion is 148 g.
c A 2.5 inch diameter boiled potato weighs 136 g without the skin.

Kunitz-type and other enzymes (Pots et al., 1999). Lysine is Lipids and Dietary Fiber
at greater levels compared with cereal proteins and the sulfur
containing amino acids (methionine and cystine) are at lower Lipids are only a tiny fraction of potato weight, amounting
levels. Chakraborty et al. (2000) described genetically modified to approx. 0.15 g/150 g fresh weight (FW); less than cooked
(GM) potato transformed with the amaranth gene AmA1. These rice (1.95 g) or pasta (0.5 g) (Priestley, 2006). Dietary fiber is
GM potato produced 35–45% more protein than control with supplied by cell walls, especially the thickened cell walls of the
2.5 to 4-fold increased lysine, methionine, cysteine, and tyro- periderm (peel) which makes up 1–2% of the tuber. These non-
sine. These potatoes are being grown in India and are reaching lignified fibers may have a role to play in reducing cholesterol
the final testing stages (Gilani and Nasim, 2007). levels (Lazarov and Werman, 1996).
 POTATOES AND HUMAN HEALTH 827

Minerals Table 2 Antioxidant compounds in potatoes

Category Individual compounds

The minerals present in greatest concentrations in raw potato
Proteins patatin
include (mg/g FW): potassium (564), phosphorus (30–60), cal-
Vitamins ascorbic acid
cium (6–18) (Burton, 1989; Buckenhüskes, 2005). The % RDA dehydroascorbic acid
for these minerals is 22, 6, and 6, respectively (White and folic acid
Broadley, 2005). Skin-on potatoes are considered a good dietary Phenolic acids caffeic acid
source of potassium. Rats fed a diet high in sodium (2% sodium chlorogenic acid
p-coumaric acid
chloride) were better able to retain calcium and magnesium and
ferulic acid
increase urinary pH when fed cooked potato instead of wheat gallic acid
starch due to the high potassium citrate content of the potato protocatechuic acid
(Narcy et al., 2006). Potatoes contain relatively little phospho- vanillic acid
rus in the form of phytate. Since phytate forms unabsorbable Flavonoids (Flavan-3-ols, catechin
Flavonols and Anthocyanins)a cyanidin
complexes with key minerals such as zinc and iron, potatoes
delphinidin
have better bioavailability for those minerals relative to many malvidin
other plant foods with substantially higher phytic acid content. malvidin-3-(p-coumaroyl rutinoside)
Biofortification is a relatively recent initiative to improve health pelargonidin
in impoverished populations through breeding of micronutrient- peonidin
petunidin
rich staple food crops (White and Broadley, 2005). It may be
rutin
possible to increase the mineral fraction of potato tubers through Carotenoids (Xanthophylls)b antheraxanthin
improved selection among cultivars. For example, zinc is criti- B-carotene (trace)
cally important to cognitive skills; its range among cultivars was lutein
found to be much wider than previously known, (12.5–20 µg/g neoxanthin
violaxanthin
DW = 160% spread) (Brown, 2008). Similarly, iron deficiency
zeaxanthin
is common among impoverished populations and not uncom-
a Inpotatoes with red, blue or purple color.
mon in the developed world. Brown (2008) determined that iron b Withthe exception of lutein, B-carotene, and violaxanthin found in potatoes,
levels varied significantly among cultivars (18–65 µg/g DW
with yellow- orange flesh.
= 373% spread); well beyond previous estimates. Curiously, a
link was determined between greater iron levels and red-skinned genated xanthophyll that protects against macular degeneration,
cultivars. Improved breeding and mineral fertilization are al- the leading cause of visual impairment and blindness in older
ternative means to increase potato mineral status (White and North American adults (Seddon et al., 1994). Potatoes with yel-
Broadley, 2005; Brown, 2008). low flesh contain primarily lutein, with a trace of β-carotene
and various other pigments including violaxanthin, zeaxanthin,
Vitamins and others (Brown, 2008) (Table 2). Carotenoid content ranges
from 57–750 µg/150 g FW; there are more carotenoids in yel-
The predominant vitamin in potatoes is vitamin C (ascorbic low than white fleshed varieties (Buckenhüskes, 2005). The
acid), which ranges in content between 84 to 145 mg per 100 g orange-fleshed potatoes contain zeaxanthin in addition to lutein
dry weight depending on cultivar, planting site, and storage (Brown et al., 2003). Breeding may increase carotenoid content,
conditions (Augustin, 1975). Vitamin C is important to iron as wild species may contain these pigments in greater concen-
availability, a mineral that tends to be limiting in the human trations (Brown, 2008). In lettuce, both β-carotene and lutein
diet (Brown, 2008). Potatoes constitute an important dietary were highly correlated, so selection for higher levels of one
source of vitamin C in many areas of the world. Also present could lead to increase in both (Mou, 2005). This relationship
are several B vitamins (folic acid, niacin, pyridoxine, riboflavin, has not been examined for potato. It may also be possible to
and thiamin) and potatoes can be described as a good dietary transform potato to increase β-carotenoid content. To this end,
source of pyridoxine (vitamin B6 ). Ducreux et al. (2005) reported the transformation of two cv.,
Carotenoids and their derivative xanthophylls are diverse Desiree and Mayan Gold, with a phytoene synthase gene from
lipid-soluble pigments. In potato, xanthophylls are the most Erwinia using a patatin promoter. Subsequently, Van Eck et al.,
abundant carotenoids (Brown, 2008). Two of these pigments, (2007) enhanced β-carotene content by RNAi-mediated silenc-
present in low concentration in cultivated potato (β-carotene ing of the β-carotene hydroxylase gene that converts β-carotene
and lutein), have an important role to play in eye health. Vita- to the less useful zeaxanthin. However, the β-carotene to retinol
min A deficiency is widespread; more than 124 million children conversion efficiency (21 µg of β-carotene per 1 µg retinol)
around the world are deficient in vitamin A (Humphrey et al., proposed for developing countries suggests that a combination
1992) leading to various ailments, including blindness, and re- of strategies will be necessary to sufficiently improve the pro-
sulting in premature death. The most potent dietary source of vitamin A content of potato for populations at risk of vitamin A
vitamin A (pro-vitamin A) is β-carotene. Lutein is an oxy- deficiency (Van Eck et al., 2007).
828 M. E. CAMIRE ET AL.

Phytochemicals Table 3 Potential health effects of glycoalkaloids, primarily α–solanine and

α–chaconine

Phytochemicals are secondary products of plant metabolism, Beneficial effects Harmful effects
many of which are implicated in human health as antioxidants. decreased serum glucose cholinesterase inhibition
These vary in amount and composition among potato cultivars antibiotic teratogenicity
(Al-Saikhan et al., 1995; Brown et al., 2003; 2005; 2008). Chu cancer cell growth inhibition intestinal inflammation
et al. (2002) estimated 124.5 mg vitamin C equivalent/150 g FW reduced serum cholesterol levels nausea
anti-inflammatory
potato; 13% from vitamin C and the balance from carotenoid
anti-pyretic
and phenolic substances.
Anthocyanins are red, blue, and purple pigments that can be
found in the skin and flesh of certain cultivars. These are im- mary potato lectin. Wounding of the tubers produced additional
portant in plant and human health, contributing important anti- chitin-binding compounds (Millar et al., 1992). Recently, a
oxidant properties (Brown et al., 2003; Brown, 2005; 2008). mechanism of lectin interaction with immunoglobulin E was
Total anthocyanin concentrations in Andean potatoes ranged proposed to explain the non-allergic hypersensitivity of some
from 14–16,330 µg/g DW (Andre et al., 2007). Anthocyanins consumers to potatoes (Pramod et al., 2007).
are water-soluble and usually occur in their glycosylated and Allergies to potatoes appear to be relatively uncommon.
acetylated forms. Reyes and Cisneros-Zevallos (2007) evalu- Patatin (Sol t 1) is the primary storage protein (Shewry, 2003)
ated extracts from red and purple potatoes and concluded that and the major allergen in potatoes. Patatin may induce allergic
they were a potential source of natural color. The genetic sys- symptoms during potato peeling (Seppälä et al., 1999). Four
tems influencing anthocyanin production in potatoes have been other proteins (Sol t 2, Sol t 3.0101, Sol t 3.0102, and Sol t
summarized by De Jong (1991) and Brown (2005). Usually, cul- 4) are related to soybean trypsin inhibitors and may cause re-
tivars high in anthocyanins are low in carotenoids and vice versa, actions in atopic children (Seppälä et al., 2001). Patatin may
although breeding efforts were able to overcome this negative be cross-reactive for persons with allergy to latex, and children
correlation (Brown, 2008). with atopic dermatitis appear to have increased sensitivity to this
Phenolic acids and polyphenols play important roles in plant potato protein (Schmidt et al., 2002). Twelve infants with atopic
health, cooking properties, and human health (Friedman, 1997). dermatitis and suspected potato allergy were found to have pos-
Chlorogenic acid is the predominant phenolic compound in itive responses to both raw and cooked potato (Majamaa et al.,
potatoes, as in other Solanaceous species such as eggplant 2001). De Swert and colleagues (2007) reported that most in-
(Prohens et al., 2007). Chlorogenic acid, catechin, caffeic acid, fants with potato allergy develop tolerance to cooked potato at
ferulic acid, gallic acid, and malvidin were the major phenolics 4 years of age and that potato allergy could be a risk factor
isolated from methanol/water extracts (Reddivari et al., 2007; for future pollen allergy. An investigation of extracts from wild
Leo et al., 2008). Total antioxidant activity was correlated with and GM potatoes revealed an incidence of positive skin prick
total phenolic content. In addition to the free-form phenolic responses in 5.7% of persons with diagnosed allergy disorders
compounds such as chlorogenic acid and caffeic acid, bound- (Lee et al., 2006). No differences were observed between the
form phenolics such as ferulic acid that is ester-linked to cell wild and GM potato extracts. Boiling and exposure to simu-
wall polysaccharides is also present in potatoes and contributes lated gastric fluid greatly reduced protein and IgE-binding. This
to the radical scavenging activity of potato (Nara et al., 2006). agreed with previous reports of allergic responses to consump-
tion of raw potatoes. Potatoes were recommended as a “safe”
food for persons with an allergy to lipid transfer protein, a highly
Natural Toxicants and Allergens cross-reactive plant allergen (Asero et al., 2007).

Glycoalkaloids are nitrogenous compounds contained by

potatoes and other members of the Solanaceae family. The POTATOES AS FOOD
primary glycoalkaloids in potatoes are α-chaconine and α-
solanine. The ratio of these compounds varies and they are Potatoes are prepared by consumers in an overwhelming va-
localized in the skin. New cultivars are screened for their total riety of means. Baking, boiling, dehydrating, and frying are
glycoalkaloid content (TGA) due to concerns about the effects of employed world-wide. Commercial processes are addressed in
these compounds on human health. Friedman (2006) reviewed many texts including Woolfe (1987), Lisińska and Leszczyński
these compounds and noted that their effects in humans can be (1989), and Gopal and Khurana (2006). Examples of commer-
both deleterious and beneficial (Table 3). Grunenfelder et al. cial potato products are listed in Table 4.
(2006) dispelled the myth that green potatoes contain higher
levels of glycoalkaloids. Chlorophyll and glycoalkaloids both
accumulate in potato skins in the presence of light but they do Processing Effects on Nutrient Composition
so independently.
Lectins are glycoproteins that bind chitin and therefore serve Cooking or processing of potatoes greatly improves the di-
as plant protectants. S. tuberosum agglutinin (STA) is the pri- gestibility of potato starch, which has very low digestibility in
 POTATOES AND HUMAN HEALTH 829

Table 4 Processed potato products leaching alone does not result in significant losses of potas-
Category Products sium (Bethke and Jansky, 2008). Boiling cubed potatoes reduced
potassium levels by an average of 50%; shredding potatoes to
Fresh/refrigerated hash browns increase surface area before cooking resulted in 75% loss of the
gnocchi
steamed or baked whole, halved, cubed
mineral. Conversely, persons with normal kidney function who
steamed and mashed wish to maximize potassium should cook whole potatoes and
pancakes and latkes minimize boiling time to reduce leaching. Due to their naturally
cake high ratio of potassium to sodium content, potatoes are useful
knishes in sodium-restricted diets.
Frozen cakes
French fries, home fries
Fresh-cut, minimally-processed potatoes are available in
hashed browns many forms, including diced, sliced, and shredded. Polyphenol
pierogies oxidase may react with substrates (tyrosine, chlorogenic acid) to
Dried cubes discolor the cut pieces. Enzymatic browning could lead to sub-
flakes (instant mashed) stantial losses of amino acids, which could be minimized by im-
flour
granules
mediate cooking of the cut potatoes or their immersion in water.
starch Cutting potatoes increased the activity of phenylalanine ammo-
slices nia lyase (PAL) with subsequent increase in antioxidants (Reyes
Shelf-stable bread et al., 2007). Quercetin glycosides, total flavonols, and caffeic
chips/crisps acid derivatives increased in fresh-cut potatoes stored at 4◦ C
extruded snacks
sticks
for 6 days (Tudela et al., 2002). Light inhibited wound-induced
Beverages vodka phenolic biosynthesis, suggesting that opaque packaging may
wine be desirable for fresh-cut potatoes to maximize antioxidants in
products destined to become home fries and mashed potatoes.
the raw state since potato starch granules have a β-crystalline Wounded cv. All-Blue retained more anthocyanins when stored
structure that is resistant to amylase digestion (Englyst et al., in the dark than in the presence of light (Reyes and Cisneros-
1992). In general, unpeeled potatoes that undergo cooking have Zevallos, 2003). New potatoes produced fewer phenolic com-
better nutrient retention than do peeled potatoes and size reduc- pounds than did potatoes from long-term storage. Cutting pota-
tion brings about further losses. Boiling cut or peeled potatoes toes leads to loss of ascorbic acid in the presence of oxygen.
leads to loss of water-soluble vitamins and minerals due to their Excessive enzymatic browning coupled with the loss of vitamin
leaching out into the cooking water. Losses of minerals occur C makes the normal atmosphere impractical for commercial use
only via boiling with the greatest losses occurring for potassium (Tudela et al., 2003). Vacuum packaging of fresh-cut potatoes
and copper (Finglas and Faulks, 1984). Baking, roasting, and retained up to 89% of the original vitamin C content and light
frying generally result in lower losses of vitamins than boiling. color was maintained.
Conventional frying or chipping of potatoes in oil such as for Compared with uncooked potatoes, all methods of cook-
French fries and chips typically increases fat content within a ing produced significant losses of total flavonoids (Tudela
short period. The fiber content of potatoes, which is nutritionally et al., 2002). Boiling, steaming, and frying led to retention
significant, is reduced greatly by peeling. Ascorbic acid concen- of 4–16 mg of quercetin derivatives and 6–30 mg of caffeic
tration is reduced as cooking time and temperature increase (Han acid derivatives per serving; microwaving resulted in fewer
et al., 2004). Microwave cooking produced the smallest losses flavonoids than did boiling and steaming. Boiling potatoes in
of ascorbic acid in three Korean cultivars; boiling caused losses varying volumes of water did not seem to affect the magni-
of 77.2–88.4%. tude of chlorogenic acid that leached into the cooking water
Persons who undergo kidney dialysis must limit their potas- (Andlauer et al., 2003). Despite losses, cooked potatoes con-
sium consumption. Patients have been advised to soak pota- tribute substantial amounts of flavonols to the diet. Brown (2005)
toes in water to help leach some of this mineral out. However, reviewed storage and cooking factors impacting potato antioxi-
dants. After-cooking darkening (ACD) is a defect for some cul-
tivars that are exposed to air after processing (boiling, baking,
frying, and dehydrating). This color change involves oxidation
Table 5 Strategies for controlling acrylamide formation in potato productsa
of ferrous-chlorogenic acid to a bluish-grey ferri-dichlorogenic
Intervention compound that depends on the ratio of chlorogenic to citric acid
Selection of cultivars low in asparagine and glucose (Wang-Pruski and Nowak, 2004). Cultivar and storage duration
Removal of precursors prior to processing were more important than management practices in affecting
Optimization of food processing to prevent acrylamide formation ADC (Wang-Pruski et al., 2007).
Addition of ingredients that inhibit acrylamide formation Glycoalkaloid content is affected by certain types of process-
Removal of acrylamide post-processing
ing. Friedman and McDonald (1999) reviewed the effect of pro-
a Adapted from Friedman and Levin (2008). cessing methods on glycoalkaloids in potato products. Peeling
830 M. E. CAMIRE ET AL.

and slicing contributed most to reducing total glycoalkaloid are screened to have safe glycoalkaloid levels before consumers
(TGA) content during French fry production (Pêska et al., 2006). evaluate them. High glycoalkaloid content results in bitter taste
However, washing and blanching significantly reduced nitrates. and even burning sensations (Friedman, 2006). Phenolic acids
Early-season cultivars tended to have lower TGA levels, and a contribute to bitter flavors in some potatoes. Taste thresholds
doubling of nitrogen fertilizer increased TGA by 10% (Tajner- of chlorogenic and ferulic acid in dehydrated mashed potatoes
Czopek et al., 2008). Peeling and cooking resulted in greater were 82 and 62 mg/L, respectively (Work and Camire, 1996).
losses of the more toxic solanine compared with chaconine. Addition of margarine to the mashed potatoes increased the taste
threshold for chlorogenic acid only. Glutamate, aspartate, and
nucleotides contributed to off-flavor in boiled potatoes (Morris
Use of Processed Peels as a Food Item et al., 2008).

Potatoes used for chipping usually have thin skins that are
relatively easily removed through abrasion. In the French fry in- Food Safety Concerns
dustry, potato peels are removed through steam-peeling. Since
dried peels are 50% dietary fiber (Camire et al., 1993) and a Reported food safety issues have been relatively rare for
good source of phenolic acids and other antioxidants (Kanatt potatoes and their products. Quality assurance procedures are in
et al., 2005), several researchers have investigated their use as place in many potato-producing nations to address food safety
food additives to improve nutritional quality or to retard lipid concerns for products such as potato. For example, the Can-
oxidation. Potato peel aqueous extract was demonstrated to be Trace initiative in Canada enables identification and response
equally effective to BHA in preventing oxidation of sunflower to food safety issues and concerns from the farm-level through
oil, which was attributed to the high content of polypheno- to the retail and food service industries (AAFC, 2007). Prior to
lic compounds. For example, freeze-dried aqueous potato ex- harvest, various chemical fertilizers are incorporated into the soil
tract contained phenolic acid content of 3.43% that consisted to stimulate plant growth. Pesticides are applied to the leaves
of chlorogenic, caffeic, gallic, and protocatechuic acids at con- of plants to deter predation from Colorado potato beetle and
centrations of 50.3%, 41.7%, 7.8%, and 0.21%, respectively, of other insects, and inhibit fungal growth, including late blight.
the total phenolic content (Rodriguez de Sotillo et al., 1994). Post-harvest, sprout inhibitors may be applied. Isopropyl N -(3-
Extruded peels were more effective in minimizing peroxide val- chlorophenyl) carbamate (also called chlorpropham or CIPC) is
ues in oatmeal cookies than dried peels at the same levels of the primary sprout inhibitory agent used (Kleinkopf et al., 2003).
addition (10%) (Arora and Camire, 1994). Potato peel extracts At the time of harvest, the peel may have residual nitrates and
were used to limit lipid oxidation in irradiated lamb meat (Kanat chemical sprout inhibitors (Lang, 1992). Washing, blanching,
et al., 2005). and cooking all helped to reduce pesticide levels in Egyptian
Peels processed by extrusion cooking after drying bound bile potatoes (Soliman, 2001).
acids (Camire et al., 1993) and the carcinogen benzo[a]pyrene
in vitro (Camire et al., 1995). An extract from potato peels
ameliorated oxidative damage to rat erythrocytes and human Contaminants Following Cooking
erythrocyte membranes (Singh and Rajini, 2008). Inclusion of
10% potato peels in a diet fed to diabetic rats helped normalize A review on the microbiology of potato products noted that
some aspects of oxidative stress (Singh et al., 2005). many pathogenic bacteria and fungi have been isolated from
While potato flesh contains free phenolics, peels contain potato (Doan and Davidson, 2000). The surface of raw tubers
bound:free phenolic acids in a ratio of about 1:3 (Nara et al., may be contaminated with pathogens from the soil in which
2006). The bound forms are present as β-glycosides that are the potatoes were grown or during handling following harvest.
resistant to upper gastrointestinal digestion but can be absorbed Microbial problems in cooked potatoes are not common, al-
following colonic microbial metabolism (Chu et al., 2000). Peels though they can be problematic in some processed products.
may not contain the only antioxidant by-products of potato pro- Bacillus cereus was found in 10–40% of dehydrated potatoes,
cessing. Hydrolyzed potato protein, a by-product of the potato and poses a problem for reconstituted mashed potatoes (King
starch industry, retarded oxidation in beef patties by >40% et al., 2007). Salmonella strains were identified in potatoes pre-
(Wang and Xiong, 2005). pared for home dehydration (DiPersio et al., 2005). Blanching
prior to dehydration greatly reduced microbial counts.
Similarly, microbial problems in prepared foods containing
Sensory Qualities potato are unusual, unless these products are improperly han-
dled. For example, Clostridium botulinum growth in gnocchi
Potato sensory properties and methods of analysis have been made with potato flakes was controlled by 8◦ C storage and
reviewed by Arvanitoyannis et al. (2008). Much of the literature inclusion of sorbate at 0.09% (w/w) (Del Torre et al., 2004).
on sensory properties of potatoes has been conducted as part of A heat-resistant type of Staphylococcus aureus was associated
established potato breeding programs. Typically, new cultivars with the illness of more than 100 persons in India who consumed
 POTATOES AND HUMAN HEALTH 831

fried potato balls; the food handler apparently spread the bacte- other hand, it is lauded for its contribution to preventing malnu-
ria to the food (Nema et al., 2007). Several recalls of refrigerated trition and promoted as a healthy food item. This is primarily
potato products such as mashed potatoes and home fries due to due to the presence of vitamins and minerals, and the content of
contamination with Listeria monocytogenes have occurred in antioxidants, with their free-radical scavenging characteristics.
the U.S. in the past few years. Ozone treatment of potatoes These antioxidants may slow the onset of age-related chronic
offers promise as a means to control pathogens in processed diseases including certain cancers, cardiovascular disease, and
potatoes (Selma et al., 2006). diabetes.

Acrylamide Potato and Energy Metabolism

Swedish researchers first reported that acrylamide, a common The glycemic index (GI) is a research tool developed by
industrial chemical, can be found in many heated carbohydrate- Jenkins et al. (1981) to assess the impact on blood sugar lev-
rich foods (Tareke et al., 2002). French fries and potato chips els from a serving of any food containing 50 grams of car-
had some of the highest acrylamide levels reported, spurring bohydrate. Procedures for GI and related measures have been
numerous studies on the mechanisms of acrylamide formation reviewed (Kendall et al., 2006; Monro and Shaw, 2008). GI
in foods as well as its mitigation (Lineback et al., 2006). There is values for potato vary greatly due to compositional differences
generally consensus that asparagine as a free amino acid reacts among cultivars and methods of food preparation. For example,
with reducing sugars in the presence of heat to form acrylamide GI values ranged from 56 to 94 for eight British cultivars (Henry
through a complex pathway involving Maillard and related re- et al., 2005). Lower GI values were reported for waxy than for
actions (Stadler et al., 2002; Zyzak et al., 2003). Although firm or floury types. New potatoes, which are smaller than most
potatoes generally contain both free asparagine and reducing potatoes, had lower GI compared with other cultivars (Sohl
sugars, there exists a wide variability in these acrylamide pre- and Brand-Miller, 1999). When boiled red potatoes were served
cursors among potato cultivars. Kennebec and White potatoes hot to volunteers, a GI of 89.4 was found based on glucose =
have very low levels of both asparagine and reducing sugars 100 scale (Fernandes et al., 2005). When cooking is followed by
(Vivanti et al., 2006). Friedman and Levin (2008) have sum- cooling, amylose retrogrades to produce resistant starch. The GI
marized intervention strategies to control acrylamide levels in response was only 56.2 when cooking was followed by refrig-
potatoes and other foods (Table 5). While it appears prudent eration of 12–24 hours. This phenomenon slows postprandial
to limit acrylamide in heat processed potato products, there is glucose release from cooked potatoes. Low GI diets have been
still controversy surrounding the role of dietary acrylamide in associated with decreased risk of obesity, cardiovascular dis-
carcinogenesis. Epidemiology has revealed no association with ease, and type-2 diabetes. In addition, intervention trials have
dietary acrylamide and risks for breast (Hogervorst et al., 2007; demonstrated improvements in certain metabolic risk factors
Olesen et al., 2008), bladder and prostate (Hogervorst et al., (Aston, 2006). However, despite commercial efforts to exploit
2008a), or gastrointestinal cancers (Hogervorst et al., 2008b). GI, the value of GI to prevent or treat obesity and diseases re-
Some increased risk for endometrial and ovarian (Hogervorst mains controversial. The need for improved methodology was
et al., 2007), and renal cell cancers (Hogervorst et al., 2008a), cited by van Bakel et al. (2009) to obtain better estimates of GI
have been identified. Mucci and Wilson (2008) noted that none and glycemic load (GL) for epidemiology research.
of these studies found a risk due to consumption of a spe- Potatoes may have a role in controlling appetite and therefore
cific group of foods, including potato products. Physiologically- weight gain, by contributing to satiety. Satiety is the feeling of
based pharmacokinetic/pharmacodynamic modeling could not fullness and the loss of hunger that occur after eating. Many
support a link between dietary acrylamide exposure and human factors influence satiety, including the rate of gastric emptying,
neurotoxicity; lifetime excess risks ranged 1.0-3.9 × 10−4 av- and the proportion of macronutrients in the food. Foods that
erage human tumor incidence (Doerge et al., 2008). The U.S. increase satiety are thought to promote weight control by de-
Food and Drug Administration researchers concluded that con- laying subsequent meals and total calories consumed. A study
sumers should continue to follow the advice to eat a balanced with 13 volunteers compared the effects of isocaloric test meals
and varied diet that is low in trans fat and saturated fat and rich containing potatoes prepared by several methods: oven-baked
in whole grains, fruits, and vegetables. French-fries, boiled and mashed with varying quantities of wa-
ter (Leeman et al., 2008). Postprandial satiety after 3 hours was
least for the French fries and greatest for the boiled potatoes on
CONTRIBUTION OF POTATOES TO HUMAN HEALTH an equivalent-energy basis, but no differences were found when
various types of potato were fed as carbohydrate-equivalent
Beyond the nutritional properties of a quality staple food, meals. Men fed several test meals consumed less energy during
potato may have a role to play in human health over a lifetime the boiled potato arm of the study than when pasta or white rice
of consumption. Potato has been implicated in contributing to was eaten as the side dish (Erdmann et al., 2007). The potato
diabetes and obesity because of its high glycemic index. On the meal resulted in lower post-prandial serum insulin and higher
832 M. E. CAMIRE ET AL.

ghrelin levels. This latter study contrasts with short-term in- many countries. The importance of potato in contributing vita-
tervention studies. These have generally indicated that high GI min C is partly because they can be stored, allowing potatoes to
meals decrease satiety, and increase the return of hunger and be a regular item in the diet. It is estimated that potatoes provide,
energy intake at a later meal, in comparison with low glycemic on average, over 50% of the daily ascorbic acid requirement in
index meals containing potatoes (Roberts, 2000). Conversely, the USA and about 20% of the dietary intake in Europe (Love
longer-term studies on the effects of low and high GI diets, and Pavek, 2008). In developing countries, seasonal variations
matched for fiber and macronutrient composition, among free- in plasma ascorbate have been related to deterioration in ascor-
living subjects have not shown effects on body weight. For bic acid content of potatoes in locations where refrigeration
example, a randomized crossover intervention study that in- is relatively unavailable (FAO, 2002). In Glasgow, Scotland,
cluded two consecutive 12–week periods showed that lesser or the WHO MONICA project compared plasma vitamin C levels
greater ad libitum-fed glycemic index diets made equivalent in with items from a food frequency questionnaire (Wrieden et al.,
macronutrient composition, fiber content, or energy density, had 2000). Citrus products and other vegetables were major sources
no impact on body weight, fat mass, satiety, or energy intake of vitamin C for persons with optimal levels of the vitamin, but
in overweight/obese women (Aston et al., 2008). A systemic potatoes and fried potatoes (chips) contributed about 17–18 mg
review of 31 short-term and 20 long-term clinical intervention per day of ascorbic acid in the diets of adults with low and
trials demonstrated no consistent impact of low versus high GI marginal vitamin plasma C. The researchers concluded that di-
diets on either satiety or body weight control (Raben, 2002). etary advice to avoid eating fried potatoes should also contain
In a short-term study, obese children consumed significantly recommendation of other foods to replace the vitamin C pro-
more energy after a test lunch containing rapidly-digested car- vided by the chips. Cooked, peeled, or unpeeled potatoes, offer
bohydrates (mashed potato, meat, nectar) than after one with 11–12 mg vitamin C per 100 g product (Biesalski, 2005; cited
more slowly digested carbohydrates (spaghetti, meat, orange) in Buckenhüskes, 2005). Potatoes are a component of many
(Alviña and Araya, 2004). It seems that appetite was stimulated traditional Indian recipes. Vegetable dishes containing potatoes
and increased caloric intake promoted by the more rapidly di- contribute about 34 mg of ascorbic acid per 100 g in the Punjab
gested meal. However, more research is required to determine region (Gupta and Bains, 2006). Loss of vitamin is affected by
whether high GI meals can generate similar findings on energy cooking method, so opportunity may exist to increase nutrient
intake on a longer-term basis. retention via education about different cooking procedures.
Potato chips and French fries have been implicated by several Potatoes contain significant amounts of folic acid (folate;
nutritionists as major contributors to obesity as these products vitamin B9 ). Potato consumption was significantly associated
contain a high fat and caloric content. In a Canadian study of with adequate serum folate status in Cretan adults (Hatzis et al.,
elementary school-aged children in a First Nations community, 2006). Consumption of at least 122 g of potatoes produced the
obese children ate 50% more French fries on a weight basis same odds ratio (0.41) for low serum folate as did consumption
(p ≤ 0.05, analysis of covariance) than the normal-weight chil- of at least 190 g of other vegetables, suggesting that potato
dren (Receveur et al., 2008). Children’s preference for French should be promoted more heavily for its folate contribution.
fries may be related to food neophobia and avoidance of veg- Human dietary intake of vitamin A and the essential miner-
etables (Dovey et al., 2008). als iron and zinc is often insufficient. These deficiencies or low
In summary, potato is a healthy component of a varied diet. intakes usually occur in populations where diets are primarily
As a starch food item, it should be consumed in moderation and plant-based (Nicolle et al., 2004). Increased level of micronu-
without excess lipid additions. A meal containing potato con- trients in vegetables such as potato would most benefit these
tributes to satiety. Potato servings do not in themselves promote populations. Iron deficiency is a global health concern for chil-
obesity; excess starchy food and especially food laden with high- dren and premenopausal women. Although potato is not typi-
calorie lipid additions is the culprit. Excluding potatoes from the cally considered a good source of dietary iron, iron uptake is
diets of persons attempting weight loss is not warranted based enhanced by ascorbic acid. So, increasing ascorbic acid levels in
on their carbohydrate content. Samaha et al. (2007) concluded vegetables such as potato may contribute to alleviating human
that carbohydrate restriction would be of greatest benefit for iron deficiencies (Nicolle et al., 2004; Brown, 2008). Alterna-
persons with insulin resistance syndromes. tively, several native Andean varieties contained significant lev-
els of iron and zinc (Burgos et al., 2007). Genotype had a greater
influence on mineral content than environment, suggesting that
Prevention of Nutrient Deficiencies new varieties could be developed with enhanced iron and/or
zinc content. Cooking did not generally reduce concentrations
Vitamin C is often lacking in the diet of individuals with- of either mineral. The researchers calculated that consumption
out access to fresh produce. Potato had a role in prevention of of genotypes with the highest iron level could provide 6.88 and
scurvy from its first contact with Europeans. Despite destruc- 1.72 mg per day for women and children, respectively. Provision
tion of ascorbic acid during cooking and a moderate content of 38% of the Daily Value for iron would qualify this potato for
compared with some other fruits and vegetables, potato plays the following U.S. iron content claims: excellent source of iron,
a critical nutritional role as the primary source of vitamin C in rich in iron, and high in iron.
 POTATOES AND HUMAN HEALTH 833

Zinc deficiency impairs development. Andean potato vari- Consumption of foods rich in antioxidants is expected to
eties vary in their zinc content, but some are relatively high in increase antioxidant levels in vivo. Malondialdehyde (MDA)
this mineral (Burgos et al., 2007). Based on a daily consump- levels as an index of in vivo lipid peroxidation were higher in
tion of 200 g by Peruvian children aged 1–3 years, and 800 g the plasma of elderly Spanish citizens who ate 29.8 grams of
by women, the variety with the greatest zinc content could theo- potatoes or more daily (Lasheras et al., 2003). Glycoalkaloids
retically provide 29% and 87%, respectively, of the correspond- or other toxins in potatoes and differences in cooking methods
ing recommended nutrient intake (RNI) for those demographic were proposed as sources of the oxidative damage. A review of
groups. MDA as a biomarker for oxidative stress concluded that day-to-
Potato has an important role to play in preventing malnutri- day and within-subject variation limited the applicability of the
tion, especially in impoverished areas of the world. Although assay for assessment of individuals (Nielsen et al., 1997). While
potato has a relatively low energy density, it has negligible fat, the assay may be suitable for estimating oxidative stress in a
contains quality protein, fiber, and vitamins, especially vitamin population, it is not clear whether the 162 subjects in the Spanish
C and the B-group vitamins, minerals, especially potassium, study were an adequate sample of the population. However,
and important phytochemicals, many of which have antioxidant residents of the Canary Islands consume an average of 143.2
properties. Potato is important to global nutrition and health. grams of potatoes per person per day, which is higher than
that for the general Spanish population (del Mar Verde Méndez
et al., 2004). Potatoes contribute an estimated 19% and 14%
Antioxidants and their Relation to Health of the recommended amount of flavonoids for men and women
in the Canary Islands, based upon the levels of (+)-catechin
Reactive oxygen species are indicated to be key modulat- measured.
ing factors responsible for the development of many important
age-related and inflammatory disease conditions such as arthri-
tis, atherosclerosis, cancers, cardiovascular diseases, diabetes, Potato and Prevention of Cancer
gastrointestinal disorders and neurodegenerative dysfunctions
(Packer, 1995). Potatoes contain a diverse mixture of antiox- Consumption of produce is associated with reduced risks for
idants (Brown, 2005; 2008) (Table 3), which exhibit multiple prostate cancer, which is a leading cause of cancer deaths among
antioxidant activities including superoxide scavenging capabil- men in the United States. Extracts from four specialty pota-
ity, ferrous ion chelating effects, and strong reducing capacity toes and the anthocyanin fraction from genotype CO112F2-2
(Singh and Rajini, 2004). Polyphenolic compounds provide a reduced cell growth and induced apoptosis in both androgen-
large portion of the antioxidant action of potatoes and their ex- dependent (LNCaP) and androgen-independent (PC-3) prostate
tracts by scavenging and neutralizing free radicals, decompos- cancer cell lines (Reddivari et al., 2007). The anthocyanins ap-
ing lipid peroxides, and quenching singlet oxygen (Cao et al., pear to cause mitochondrial release of the proteins Endo G
1997). Although phenolic compounds contributed greatly to the and AIF that promote apoptosis. Likewise, intake of antho-
total antioxidant activity in four Italian early potato cultivars, cyanins from purple and red potatoes might play a protective
carotenoids and ascorbic acid were also major contributors (Leo role against stomach cancer. Intake of steamed purple and red
et al., 2008). Differences in antioxidant content were found due potatoes repressed the growth of mouse stomach cancer induced
to cultivar and growing site. by benzo(a)pyrene (Hayashi et al., 2006). Isolated anthocyanins
Potato cultivars and wild Solanum species with flesh that induced apoptosis in human stomach cancer cell lines as well
is purple or red and solidly-pigmented are particularly high as suppressing benzo(a)pyrene-induced mouse stomach cancer
in anthocyanin content (Brown, 2005). Antioxidant capacity proliferation indicating that anthocyanins were the bioactive
has been directly related to anthocyanin content in potatoes anti-tumor components. No induction of apoptosis was observed
(Brown et al., 2003). Drum-dried potato flakes from both a upon exposure to normal lymphocytes prepared from healthy
light purple (KM) and darker purple (H92) potato cultivar had volunteers. The exact anthocyanins involved in the anti-tumor
significant in vitro antioxidant activity, but only the antioxidant properties remain to be delineated as there are approximately
activity of sera from rats fed the darker flakes were higher than ten and eight different types of pigment present in the purple
those of rats fed a control diet (Han et al., 2006b). Liver lipid and red potato anthocyanin fractions, respectively.
oxidation was also lower in the H92-fed animals. In a related The role of vegetable consumption in prevention of can-
study, rats fed a purple potato (cv. Shadow-Queen) diet had cer has not reached consensus in the scientific community.
lower oxidation levels in both sera and liver, but white potatoes Methanol-water extracts from four Italian short-season potato
(cv. Toyoshiro) also reduced serum urate levels (Han et al., cultivars inhibited the proliferation of MCF-7 breast cancer cells
2007a). Extracts from a red potato cultivar (Hokkai no. 91) (Kallio et al., 2008). Extracts from cv. Nicola were most effec-
ameliorated galactosamine-induced liver damage in rats (Han tive, but concentrations above 1 × 10−3 µg gallic acid equiv-
et al., 2006a). Potato flakes from the red cv. Northern-Ruby alents µL−1 stimulated cell growth. Potato glycoalkaloids in-
reduced serum lipid oxidation and increased hepatic superoxide hibited the growth of human colon (HT29) and liver (HepG2)
dismutase (SOD) mRNA (Han et al., 2007b). cancer cells (Lee et al., 2004), and human cervical, liver
834 M. E. CAMIRE ET AL.

lymphoma, and stomach cancer cells (Friedman et al., 2005). sumption. Drawbacks to this study included the differences in
Alpha-chaconine reduced lung cancer metastasis in vitro by rat lipid metabolism compared with that of humans, and the ex-
suppressing metabolic pathways (Shih et al., 2007) and induced tremely high levels of potato in the test diet. The lowered plasma
apoptosis of human colon cancer cells via activation of caspase- cholesterol concentrations in potato-fed rats could be partly re-
3 and inhibition of signaling compound ERK 1/2 (Yang et al., lated to lipid lowering properties of potato protein (Morita et al.,
2006.) 1997). Potato protein was demonstrated to exert potent fecal bile
Although cooking inactivates lectins, these compounds may acid and neutral steroid excretion properties relative to other
also help to reduce cancer risks. Lectins induce apoptosis and proteins including soy and casein, which was related to the
limit the synthesis of proteins, DNA, and RNA in cancer cells relatively lower methionine content and methionine-to-glycine
(De Mejı́a and Prisecaru, 2005). Potato lectin reduced the vi- ratios in potato protein.
ability of human hepatoma, human choriocarcinoma, mouse Another rat feeding study compared three diets: potato-,
melanoma, and rat osteosarcoma cells, but was not as effective starch-, and sucrose-based (Robert et al., 2008). Rats on the
as other lectins with different carbohydrate-binding characteris- potato diet for 3 weeks had lower cholesterol and triglycerides
tics (Wang et al., 2000). While in vitro assays have limitations, than those fed the control or sucrose diets. Antioxidant status
these findings suggest that glycoalkaloids and lectins in potatoes and short chain fatty acids were greater in the potato-fed
may not be as menacing as once thought. animals. Potato starch phosphorylation may also play a role in
Fried potato products may contain acrylamide. For example, controlling serum lipids. Kanazawa and colleagues (2008) fed
Korean potato chips were found to contain as much as 4.0 µg/kg rats on diets containing different types of starch. Serum-free
acrylamide (Lee and Shim, 2007). The controversy surrounding fatty acids, triglycerides, and liver triglycerides were less in the
the risks of consumption of dietary acrylamide were discussed animals fed a Hokkaikogane potato starch with a phosphorus
previously. content of 8,136 mg/L. Triglyceride HDL was also less in that
In summary, relatively little has been published regarding the treatment group. Potato starch increased fecal bile acid excre-
long-term cancer–related health effects of potato in diets of con- tion but starch type had no effect on cecal short chain fatty acid
sumers around the world. Scattered, relative short-term studies synthesis or pH. So, it appears that potato starch that contains a
have implicated potato anthocyanins, glycoalkaloids, and lectins high level of phosphorus exhibits lipid-lowering properties but
as anti-tumor agents while concerns have been expressed regard- not the cecal fermentation-promoting effects of resistant starch.
ing trace quantities of acrylamide in cooked starches. The plasma lipid-lowering effects could be attributable to
slower digestion of gelatinized high-phosphorus potato starch.
The phosphate residue on the carbon-3 hydroxyl group imparts
Potato and Prevention of Cardiovascular Disease relative resistance to hydrolysis by α-amylase (Takeda et al.,
1983).
Although many factors affect the status of the heart and cir- Potato processing by-products can be rich sources of dietary
culatory system, maintenance of normal serum lipids and blood fiber. Potato peels from a French fry processing plant were
pressure are essential for health. The many adverse effects evaluated for their ability to bind bile acids in vitro (Camire
of dietary fat on the cardiovascular system were reviewed by et al., 1993). Although dried peels bound some bile acids, ex-
Damjanovi and Barton (2008). Potato products with high lipid trusion cooking enhanced peel binding of cholic, deoxycholic,
content should be minimized in the diet. Ideally, for cardiovas- and glycocholic acids; binding of deoxycholic acid was highly
cular benefit, potato should be prepared to contain low fat and correlated with total dietary fiber and insoluble dietary fiber
sodium. Relatively high levels of potassium are needed to coun- content. All peels bound a smaller percentage of bile acids than
teract the effect of sodium and protect against hypertension. did the drug cholestyramine. Residue from potato starch pro-
Like leguminous seeds and various root vegetables (Nicolle duction in Japan had different effects on lipid metabolism of
et al., 2004), potato is an excellent potassium source. Retention rats fed the pulps at a 15% level for 4 weeks (Hashimoto et al.,
of potassium during cooking was best in whole boiled potatoes 2006). Residue from cv. Benimaru potatoes increased fecal bile
compared with cut boiled product (McGinnis, 2008). acid production while cv. Hokkaikogane pulp inhibited fatty
Rats fed potato peels for 4 weeks showed less plasma choles- acid synthesis.
terol (40% decrease) and less hepatic fat cholesterol (30% de- Inflammation is another risk factor for cardiovascular dis-
crease) compared with cellulose-fed rats (Lazarov and Werman, ease. Although many biomarkers for inflammation exist, high-
1996). Rats fed a diet with 78% potato for 3 weeks to evalu- sensitivity C-reactive protein (CRP) has received the most at-
ate the vegetable’s role in lipid metabolism had lower plasma tention as a predictor for cardiovascular disease. When BMI
cholesterol and triglycerides and reduced liver cholesterol than was controlled for in a study of Australian adults, potato intake
control rats (Robert et al., 2006). The test animals excreted was associated with lower CRP levels (Hickling et al., 2008).
more neutral sterol, particularly coprostanol, indicating a pos- Potato antioxidants may be responsible for lowering inflamma-
sible mechanism for cholesterol reduction. Plasma vitamin E tion. Rats fed isolated potato peptides showed greater serum
and FRAP antioxidant levels were greater in the potato-fed rats, HDL-cholesterol and fecal steroid output and less non-HDL
demonstrating additional cardiovascular benefit to potato con- cholesterol (Liyanage et al., 2008). The results were attributed
 POTATOES AND HUMAN HEALTH 835

to inhibition of cholesterol absorption, possibly via suppression potato intake and incidence of type-2 diabetes (Williams et al.,
of micellar solubility of cholesterol. 1999; Hodge et al., 2004; Liu et al., 2004).
Peptides isolated from potato tubers exhibited angiotensin- The diets of adults in the Attica region of Greece were ex-
converting enzyme (ACE) inhibition in vitro (Pihlanto et al., amined for patterns associated with measures of metabolic syn-
2008). Inhibition was lowest in isolates from immature (new) drome (Panagiotakos et al., 2007). Principal component anal-
potatoes, and increased in isolates from mature tubers, sprouted ysis grouped fried, baked, and boiled potatoes together with
potatoes, and commercial potato processing by-products. ACE beef, pork, and poultry for a component (numbered 2) that
inhibition was 44–94%, with IC50 values of 0.018–0.086 explained 11.7% of the variance in the model. Multivariate
mg/mL. regression found a positively significant association between
Little has been published on the role of potato in prevention of component 2 and waist circumference and a negatively signifi-
cardiovascular disease. For maximum health, potato should be cant one for high-density lipoprotein cholesterol. Associations
prepared with conservative lipid additions and the peels should between component 2 and systolic blood pressure, blood glu-
be eaten for their fiber content. There is some evidence that cose, and triglycerides were not significant. Since potatoes are
potato protein, resistant starch, and phosphorylated starch also often consumed with meats, these trends are not unexpected.
contribute to cholesterol lowering properties. Phytochemicals, Consumption of potatoes does not necessarily lead to increased
especially antioxidants, were implicated in reducing inflamma- girth; persons who eat a “meat and potatoes” diet may have
tion, a risk for cardiovascular disease. other life patterns that affect health.
An examination of the diets of adolescents with the sugges-
tive subtitle of “Trading candy for potato chips?” reported that
Potato and Prevention of Diabetes teens with type-1 diabetes consumed more fat and less carbohy-
drate compared with their non-diabetic peers (Helgeson et al.,
The incidence of diabetes (diabetes mellitus) is increasing 2006). Twenty-four hour diet recalls were used to estimate the
world-wide both in the developed and developing economies. nutrient intake of the adolescents, a problematic assessment
Type-2 diabetes involves 90% of diabetics, and is character- technique, particularly when dealing with this age group. Con-
ized by insulin resistance and often associated with obesity and sumption of potato chips or any other specific food was not
dyslipidemia. The development of diabetes and its progressive reported in the publication.
complications come about through unregulated elevated blood Potato peel extracts, which contain a rich content of polyphe-
sugar levels (hyperglycemia). Management of diabetes involves nolic antioxidants reduced hyperglycemia, oxidative stress, and
oral administration of synthetic hypoglycemic agents. How- overall food consumption in diabetic rodents when fed at 10%
ever, these are not always effective, may have side-effects, and of the diet (Singh et al., 2005a). Plasma glucose levels in dia-
may not prevent long-term vascular complications (nephropa- betic rats fed 5% and 10% potato peel powder were 16% and
thy, neuropathy, retinopathy, etc.) (Spiller and Sawyer, 2006). 33% lower than the control diabetic animals. Other significant
Hyperglycemia induces oxidative stress; circulating markers of improvements in the peel-fed animals included reduced urine
free-radical-induced damage are increased and antioxidant de- output and serum alanine amino transferase (ALT). The com-
fenses are reduced. bined dietary fiber and polyphenol content of peels merits further
The combination of available carbohydrates and anti-diabetic investigation as a therapeutic aid for diabetes. Two antioxidants,
factors such as antioxidants complicates evaluation of the role caffeic and chlorogenic acids, both found at high concentrations
of potatoes in prevention and management of diabetes. Potato, in potato extracts, were implicated in prevention of type-2 dia-
and especially French fry consumption, was modestly associ- betes (Paynter et al., 2006) and cardiovascular disease (Morton
ated with an increased risk for type-2 diabetes in the Nurses’ et al., 2000). Chlorogenic acid appears to slow gut glucose ab-
Health Study (Halton et al., 2006). For potatoes in general, sorption because 3 weeks of intravenous infusion of chlorogenic
but not French fries, the highest quintile of potato consump- acid to obese, hyperlipidemic, insulin resistant (fa/fa) Zucker
tion was associated with greater relative risks (1.22) in obese rats lowered the postprandial peak response to a glucose chal-
women (BMI ≥ 30), but not (0.95) for women who were not lenge (Rodriguez de Sotillo and Hadley, 2002). Administration
obese (BMI < 30 kg/m2 ). Risks associated with French fry con- of chlorogenic acid appears to increase insulin sensitivity rather

R
sumption were significant for all women. In contrast, despite an than affecting insulin release. Svetol is a trade-marked item
association of GI and glycemic load with risk for developing dia- containing chlorogenic acid, now approved in Norway and the
betes, potato intake was associated with a lowered risk of type-2 UK for addition to coffee, gum, and mints, to promote weight
diabetes in the Shanghai Women’s Health Study, which was loss (Svetol, 2008). Chlorogenic acid selectively inhibits hep-
a population-based prospective cohort study of 74,942 women atic glucose-6-phosphatase (Arion et al., 1997), a rate-limiting
aged 40 to 70 years (Villegas et al., 2007). The contrasting find- enzyme in gluconeogenesis. A potato protein, proteinase in-
ings were explained based on a relatively low intake of potato in hibitor 2 (PI2) is incorporated into a weight loss supplement,


R
the Chinese population and a differing pattern of potato prepara- Slendesta , as this protein acts as an appetite suppressant by
tion involving less fat and frying compared with Western dietary stimulating the release of the peptide cholecystokinin, that in-
patterns. Other studies have reported no association between creases satiety (Hill et al., 1990).
836 M. E. CAMIRE ET AL.

The incidence of obesity and diabetes are on the increase cholesterol-lowering properties. Phytochemicals, especially an-
world-wide. More studies are needed to confirm the link between tioxidants, were implicated in reducing inflammation, a risk for
potato dietary fiber and polyphenolic content in prevention or cancer, cardiovascular disease, and diabetes. The incidence of
therapy for diabetes. diabetes is on the increase world-wide. More studies are needed
to confirm the link between potato dietary fiber and polyphenolic
content in prevention or therapy for diabetes.
Other Health Effects Increased knowledge of the composition of phenolic com-
ponents and antioxidant activity of various potato cultivars will
An analysis of dietary factors associated with the onset of lead to greater awareness by the food industry and consumers re-
overactive bladder in men revealed a hitherto unknown rela- garding potatoes as “functional foods” and possibly change food
tionship between urinary incontinence and potato consumption industry practices and consumer habits regarding utilization of
(Dallosso et al., 2004). The odds ratio increased from 1.00 for specific “high antioxidant” potato cultivars. Emerging research
0–5 servings per week to 1.48 for 8 or more servings per week perspectives suggest that the polyphenolic components are key
(p = 0.05). The researchers could offer no firm explanations for food constituents involved in the prevention of chronic diseases.
the finding, but the effects of high potato consumption on uri- This research could drive the development of informative con-
nary control, and possible prostate involvement, merits further sumer labeling regarding polyphenolic content. Such labeling
investigation. will lead to consumer trends towards the purchase of potato cul-
tivars with higher polyphenolic content and lead processors to
attempt to maintain or enrich these components within potato
SUMMARY products. The optimal phytochemical panel still needs to be
evaluated since a variety of phenolic compounds coexist within
The potato stem tuber is one of the world’s most popular potato cultivars. However, only limited knowledge is available
food items, now grown in more than 160 countries around the about their synergistic or antagonistic interactions, including
world. Potatoes can be prepared in a plethora of different ways. with ascorbic acid. More information is required regarding nu-
As a staple in the diets of an increasing number of humans, trient interactions within the complex mixtures found in various
small differences in potato nutritional composition impacts on potato cultivars and co-consumed foods since the effects of these
population health. The potato is a nutrient dense food that sup- compounds cannot be considered in isolation from one other.
plies significant nutrients without too many calories. It has a More information on the bioavailability of phenolic compounds
role in preventing malnutrition in impoverished areas, and con- in potatoes is also needed.
tributes to health where food is ample. In populations where
potato consumption is high, potatoes can make an important di-
etary contribution of phenolic compounds, ascorbic acid, potas- REFERENCES
sium, as well as moderate contributions of dietary fiber (when
skins are eaten), magnesium, phosphorus, and B-vitamins. Se- AAFC (Agriculture and Agri-Food Canada) (2007). Canadian Potato Situation
lection for cultivars with greater iron and zinc content is likely and Trends 2006–2007. Horticulture and Special Crops Division, Food Value
to significantly improve the coonsumption of these important Chain Bureau, Markets and Trade Team. 36 pp.
minerals especially in view of the relatively low phytate con- Al-Saikhan, M. S., Howard, L. R., and Miller, J. C. (1995). Antioxidant activity
and total phenolics in different genotypes of potato (Solanum tuberosum L.)
tent of potatoes—of particular importance where food choice J. Food Sci., 60:341–343.
is limited. These nutritional contributions make the potato an Alviña, M. and Araya, H. (2004). Rapid carbohydrate digestion rate produced
important and generally underestimated foodstuff in the diet of lesser short-term satiety in obese preschool children. Eur. J. Clin. Nutr.,
many populations worldwide and underline its potential role in 58:637–642.
Andlauer, W., Stumpf, C., Hubert, M., Rings, A., and Fürst, P. (2003). Influence
fighting malnutrition in impoverished areas. A meal containing
of cooking process on phenolic marker compounds of vegetables. Int. J.
potatoes contributes to satiety. Potato servings do not in them- Vitam. Nutr. Res., 73:152–159.
selves contribute to obesity, which is a complex problem with Andre, C. M., Oufir, M., Guignard, C., Hoffmann, L., Hausman, J. F., Evers,
numerous contributing factors. D., and Larondelle, Y. (2007). Antioxidant profiling of native Andean potato
Relatively little has been published regarding the long-term tubers (Solanum tuberosum L.) reveals cultivars with high levels of beta-
carotene, alpha-tocopherol, chlorogenic acid, and petanin. J. Agric. Food
health effects of potato in diets of consumers around the world.
Chem., 55:1039–1049.
Scattered, relatively short-term studies have implicated potato Arion, W. J., Canfield, W. K., Ramos, F. C., Schindler, P. W., Burger, H. J.,
anthocyanins, glycoalkaloids, and lectins as anti-tumor agents Hemmerle, H., Schubert, G., Below, P., and Herling, A. W. (1997). Chloro-
in cancer cell lines while concerns have been expressed re- genic acid and hydroxynitrobenzaldehyde: new inhibitors of hepatic glucose
garding trace quantities of acrylamide in cooked starches. For 6-phosphatase. Arch. Biochem. Biophys., 339:315–322.
Arora, A. and Camire, M. E. (1994). Performance of potato peels in muffins
maximum cardiovascular health, potato should be prepared with
and cookies. Food Res. Int., 27:15–22.
conservative lipid additions and the peels should be eaten for Arvanitoyannis, I. S., Vaitsi, O., and Mavromatis, A. (2008). Potato: a compara-
their fiber content. There is some evidence that potato protein, tive study of the effect of cultivars and cultivation conditions and genetic mod-
resistant starches, and phosphorylated starches also contribute ification on the physico-chemical properties of potato tubers in conjunction
 POTATOES AND HUMAN HEALTH 837

with multivariate analysis towards authenticity. Crit. Rev. Food Sci. Nutr., CommodityOnline (2008). Potato boom in Gujarat. http://www.
48:799–823. commodityonline.com/news/topstory/newsdetails.php?id=5661. Visited
Asero, R., Mistrello, G., Roncarolo, D., and Amato, S. (2007). Detection of June, 2008.
some safe plant-derived foods for LTP-allergic patients. Int. Arch. Allergy Dallosso, H., Matthews, R. J., McGrother, C. W., Donaldson, M. M. K., Shaw,
Immunol., 144:57–63. C., and Leicester, MRC Incontinence Study Group. (2004). The association
Aston, L. M. (2006). Glycaemic index and metabolic disease risk. Proc. Nutr. of diet and other lifestyle factors with the onset of overactive bladder: a
Soc., 65:125–134. longitudinal study in men. Pub. Health Nutr., 7:885–891.
Aston, L. M, Stokes, C. S., and Jebb, S. A. (2008). No effect of a diet with Damjanovi, M., and Barton, M. (2008). Fat intake and cardiovascular response.
a reduced glycaemic index on satiety, energy intake and body weight in Curr. Hypertens. Rep., 10:25–31.
overweight and obese women. Int. J .Obes. (Lond)., 32:160–165. De Jong, H. (1991). Inheritance of anthocyanin pigmentation in the cultivated
Augustin, J. (1975). Variations in the nutritional composition of fresh potatoes. potato: a critical review. Am. Potato J., 68:585–593.
J. Food Sci., 40:1295–1299. De Mejı́a, E. G. and Prisecaru, V. I. (2005). Lectins as bioactive plant proteins:
Bamberg, J. and del Rio, A. (2005). Conservation of Potato Genetic Resources. a potential in cancer treatment. Crit. Rev. Food Sci. Nutr., 45:425–445.
In: Genetic Improvement of Solanaceous Crops Vol. 1: Potato. Chap. 1, pp. Del Mar Verde Méndez, C., Rodrı́guez Delgado, M. A., Rodrı́guez Rodrı́guez,
1–38. Razdan, J.K. and Mattoo, A.K., Eds. Science Pub. Inc. E. M., and Dı́az Romero, C. (2004). Content of free phenolic compounds in
Bethke, P. C. and Jansky, S. H. (2008). The effects of boiling and leaching cultivars of potatoes harvested in Tenerife (Canary Islands). J. Agric. Food
on the content of potassium and other minerals in potatoes. J. Food Sci., Chem., 52:1323–1327.
75:H80–H85. Del Torre, M., Stecchini, M. L., Braconnier, A., and Peck, M. W. (2004).
Bradshaw, J. E. (2007). Potato breeding strategy. In: Potato Biology and Biotech- Prevalence of Clostridium species and behaviour of Clostridium botulinum
nology. Advances and Perspectives. Chap. 8 pp. 157–177. Vreugdenhil, D. in gnocchi, a REPFED of Italian origin. Int. J. Food Micro., 96:115–131.
Ed. Elsevier, London. De Swert, L. F., Cadot, P., and Ceuppens, J. L. (2007). Diagnosis and natural
Brown, C. R. (2008). Breeding for phytonutrient enhancement of potato. Am. J. course of allergy to cooked potatoes in children. Allergy, 62:750–757.
Potato Res., 85:298–307. DiPersio, P. A., Kendall, P. A., Yoon, Y., and Sofos, J. N. (2005). Influence
Brown, C. R. (2005). Antioxidants in potato. Am. J. Potato Res., 82:163–172. of blanching treatments on Salmonella during home-type dehydration and
Brown, C. R., Culley, D., Yang, C.-P., Durst, R. and Wrolstad, R. (2005). storage of potato slices. J. Food Prot., 68:2587–2593.
Variation of anthocyanin and carotenoid contents and associated antioxidant Doan, C.H. and Davidson, P.M. (2000). Microbiology of potatoes and potato
values in potato breeding lines. J. Am. Soc. Hort. Sci., 130:174–180. products: a review. J. Food Prot., 63:668–683.
Brown, C. R., Wrolstad, R., Durst, R., Yang, C.-P, and Clevidence, B., (2003). Doerge, D. R., Young, J. F., Chen, J. J., DiNovi, M. J., and Henry, S.
Breeding studies in potatoes containing high concentrations of anthocyanins. H. (2008). Using dietary exposure and physiologically based pharmacoki-
Am. J. Potato. Res., 80:241–250. netic/pharmacodynamic modeling in human risk extrapolations for acry-
Buckenhüskes, H. J. (2005). Nutritionally relevant aspects of potatoes and potato lamide toxicity. J. Agric. Food Chem., 56:6031–6038.
constituents. In: Potato in Progress: Science Meets Practice. Ch. 1, pp. 17–45. Dovey, T. M., Staples, P. A., Gibson, E. L., and Halford, J. C. G. (2008).
Haverkort, A.J. and Struik, P.C., Eds. Wageningen Academic Pub. Food neophobia and ‘picky/fussy’ eating in children: a review. Appetite,
Burgos, G., Amoros, W., Morote, M., Stangoulis, J., and Bonierbale, M. (2007). 50:181–193.
Iron and zinc concentration of native Andean potato cultivars from a human Ducreux, L. J. M., Morris, W. L., Hedley, P. E., Shepherd, T. Davies, H. V., Mil-
nutrition perspective. J. Sci. Food Agric., 87:668–675. lam, S. and Taylor, M. A. (2005). Metabolic engineering of high-carotenoid
Burton, W. G. (1989). The Potato. 3rd ed. Longman Scientific & Technical, potato tubers containing enhanced levels of ß-carotene and lutein. J. Exp.
Harlow, UK. Bot., 56:81–89.
Camire, M. E., Zhao, J., Dougherty, M. P. and Bushway, R. J. (1995). In vitro Englyst, H. N., Kingman, J. H. and Cummings, J. H. (1992). Classification and
binding of benzo[α]pyrene by extruded potato peel. J. Agric. Food Chem., measurement of nutritionally important starch fractions. Eur. J. Clin. Nutr.,
43:970–973. 46: S33–S50.
Camire, M. E., Zhao, J., and Violette, D. A. (1993). In vitro binding of bile acids Erdmann, J., Hebeisen, Y., Lippl, F., Wagenpfeil, S., and Schusdziarra, V.
by extruded potato peels. J. Agic. Food Chem., 41:2391–2394. (2007). Food intake and plasma ghrelin response during potato-, rice- and
Cao, G., Sofic, E., and Prior, R. L. (1997). Antioxidant and prooxidant behav- pasta-rich test meals. Eur. J. Nutr., 46:196–203.
ior of flavonoids: structure-activity relationships. Free Radical Biol. Med., FAO. (2002). Vitamin C. In: Human Nutrition and Mineral Requirements. Chap.
22:749–760. 6, Report of a joint FAO/WHO expert consultation, Bangkok, Thailand.
CDC (1999a). The Potato, Then and Now. Early Beginnings. Canada’s Dig- Fernandes, G., Velangi, A., and Wolever, T. M. S. (2005). Glycemic index
ital Collections archived by Library and Archives Canada. http://epe.lac- of potatoes commonly consumed in North America. J. Am. Diet. Assoc.,
bac.gc.ca/100/205/301/ic/cdc/potato/history/beginnings.asp 105:557–562.
CDC (1999b). The Potato, Then and Now. Potato migration to Europe. Canada’s Finglas, P. M., and Faulks, R. M. (1984). Nutritional composition of UK retail
Digital Collections archived by Library and Archives Canada. http://epe.lac- potatoes, both raw and cooked. J. Sci Food Agric., 35:1347–1356.
bac.gc.ca/100/205/301/ic/cdc/potato/history/migration.asp. Visited June, Fresh-Plaza (2008a). Let them eat spuds: potatoes – the world’s new staple?
2008. http://www.freshplaza.com/news detail.asp?id=20631. Visited June, 2008.
CFIA (2008). Maturity. http://www.inspection.gc.ca/english/plaveg/potpom/ Fresh-Plaza (2008b). Bangladesh: Potato rots in Chandpur cold storages as
var/chacare.shtml#mature. Visited June, 2008. overloading raises temperature. http://www.freshplaza.com/news detail.asp?
Chakraborty, S., Chakraborty, N., and Datta, A. (2000). Increased nutritive value id=20833. Visited June, 2008.
of transgenic potato by expressing a nonallergenic seed albumin gene from Friedman, M. (1997). Chemistry, biochemistry, and dietary role of potato
Amaranthus hypochondriacus. Proc. Natl. Acad. Sci., 97:3724–3729. polyphenols. A review. J. Agric. Food Chem., 45:1523–1540.
Chu, Y.-F., Sun, J., Wu, X., and Liu, R. H. (2002) Antioxidant and antiprolifer- Friedman, M. (2006). Potato glycoalkaloids and metabolites: roles in the plant
ative activities of common vegetables. J. Agric. Food Chem., 50:6910–6916. and in the diet. J. Agric. Food Chem., 54:8655–8681.
CIP, 2008a. International Year of the Potato, Origins. http://www.potato2008. Friedman, M., Lee, K. R., Kim, H. J., Lee, I. S., and Kozukue, N. (2005).
org/en/potato/origins.html. Visited June, 2008. Anticarcinogenic effects of glycoalkaloids from potatoes against human
CIP (2008b). International Year of the Potato, Potato World. http://www. cervical, liver, lymphoma, and stomach cancer cells. J. Agric. Food Chem.,
potato2008.org/en/world/index.html. Visited June, 2008. 53:6162–6169. Erratum in: J. Agric. Food Chem., 53(21):8420.
CIP (2008c). International Year of the Potato, Uses of Potato http://www. Friedman, M. and Levin, C. E. (2008). Review of methods for the reduction of di-
potato2008.org/en/potato/utilization.html. Visited June, 2008. etary content and toxicity of acrylamide. J. Agric. Food Chem., 56:6113–6140.
838 M. E. CAMIRE ET AL.

Friedman, M. and McDonald, G. M. (1999). Postharvest changes in glycoalka- Hodge, A. M., English, D. R., O’Dea, K., Giles, G. G. (2004). Glycemic
loid content of potatoes. Adv. Exp. Med. Biol., 459:121–143. index and dietary fiber and the risk of type 2 diabetes. Diabetes Care.,
Gilani, G. S. and Nasim, A. (2007). Impact of foods nutritionally enhanced 27:2701–2706.
through biotechnology in alleviating malnutrition in developing countries. J. Hogervost, J. G. F., Schouten, L. J., Konings, E. J. M., Goldbohm, R. A., and
AOAC Int., 90:1440–1441. van den Brandt, P. A. (2007). A prospective study of dietary acrylamide
Gopal, J. and Khurana, S. M. P. (Eds.). (2006). Handbook of Potato Production, intake and the risk of endometrial, ovarian, and breast cancer. Cancer
Improvement, and Postharvest Management. Haworth Press, NY, 605 pp. Epidemiol Biomarkers Prev., 16:2304–2311.
Grunenfelder, L. A., Knowles, L. O., Hiller, L. K., and Knowles, N. R. Hogervost, J. G. F., Schouten, L. J., Konings, E. J. M., Goldbohm, R. A., and
(2006). Glycoalkaloid development during greening of fresh market potatoes van den Brandt, P. A. (2008a). Dietary acrylamide intake and the risk of
(Solanum tuberosum L.). J. Agric. Food Chem., 54:5847–5854. renal cell, bladder and prostate cancer. Am. J. Clin. Nutr., 87:1428–1438.
Gupta, S. and Bains, K. (2006). Traditional cooked vegetable dishes as Hogervost, J. G. F., Schouten, L. J., Konings, E. J. M., Goldbohm, R. A., and
important sources of ascorbic acid and ß-carotene in the diets of Indian urban van den Brandt, P. A. (2008b). Dietary acrylamide intake is not associated
and rural families. Food Nutr. Bull., 27:306–310. with gastrointestinal cancer risk. J. Nutr., 138:2229–2236.
Hahlbrock, K. and Scheel, D. (1989). Physiology and molecular biology of Humphrey, J. H., West, Jr. K. P., and Sommer, A. (1992). Vitamin A deficiency
phenylpropanoid metabolism. Annu. Rev. Plant Physiol. Plant Mol. Biol., and attributable mortality among under 5 year olds. World Health Org. Bul.,
40:347–369. 70:225–232.
Halton, T. L., Willett, W. C., Liu, S., Manson, J. E., Stamfer, M. J., and Hu, F. Jansen, G., Flame, W., Schv̈ler, K., and Vandrey, M. (2001). Tuber and starch
B. (2006). Potato and French fry consumption and risk of type 2 diabetes in quality of wild and cultivated potato species and cultivars. Potato Res.,
women. Am. J. Clin. Nutr., 83:284–290. 44:137–146.
Han, J. S., Kozukue, N., Young, K. S., Lee, K. R., and Friedman, M. (2004). Jenkins, D. J., Wolever, T. M., Taylor, R. H., Barker, H., Fielden, H., Baldwin,
Distribution of ascorbic acid in potato tubers and in home-processed and J. M., Bowling, A. C., Newman, H. C., Jenkins, A. L., and Goff, D. V. (1981).
commercial potato foods. J. Agric. Food Chem., 52:6516–6521. Glycemic index of foods: a physiological basis for carbohydrate exchange.
Han, K.-H., Hashimoto, N., Hashimoto, M., Noda, T., Shimada, K.-I., Lee, Am. J. Clin. Nutr., 34:362–366.
C.-H., Sekikawa, M., and Fukushima, M. (2006a). Red potato extract pro- Kallio, P., Kolehmainen, M., Laaksonen, D. E., Pulkkinen, L., Atalay, M.,
tects from D-galactosamine-induced liver injury in rats. Biosci. Biotechnol. Mykkänen, H., Uusitupa, M., Poutanen, K., and Niskanen, L. (2008).
Biochem., 70:2285–2288. Inflammation markers are modulated by responses to diets differing in
Han, K.-H., Sekikawa, M., Shimada, K.-I., Hashimoto, M., Hashimoto, N., postprandial insulin responses in individuals with the metabolic syndrome.
Noda, T., Tanaka, H., and Fukushima, M. (2006b). Anthocyanin-rich purple Am. J. Clin. Nutr., 87:1497–1503.
potato flake extract has antioxidant capacity and improves antioxidant Kamasaka, H., Uchida, M., Kusaka, K., Yoshikawa, K., Yamamoto, K.,
potential in rats. Brit. J. Nutr., 96:1125–1133. Okada, S., and Ichikawa, T. (1995). Inhibitory effect of phosphorylated
Han, K.-H., Matsumoto, A., Shimada, K.-I., Sekikawa, M., and Fukushima, oligosaccharides prepared from potato starch on the formation of calcium
M. (2007a). Effects of anthocyanin-rich purple potato flakes on antioxidant phosphate. Biosci. Biotech. Biochem., 59:1412–1416.
status in F344 rats fed a cholesterol-rich diet. Brit. J. Nutr., 98:914–921. Kanatt, S. R., Chander, R., Radhakrishna, P., and Sharma, A. (2005). Potato
Han, K.-H., Shimada, K.-I., Sekikawa, M., and Fukushima, M. (2007b). peel extract- a natural antioxidant for retarding lipid peroxidation in radiation
Anthocyanin-rich red potato flakes affect serum lipid peroxidation and hep- processed lamb meat. J. Agric. Food Chem., 53:1499–1504.
atic SOD mRNA level in rats. Biosci. Biotechnol. Biochem., 71:1356–1359. Kanazawa, T., Atsumi, M., Mineo, H., Fukushima, M., Nishimura, N., Noda,
Hanson, P. M., Yang, R., Wu, J., Chen, J., Ledesma, D., and Tsou, S. C. S. T., and Chiji, H. (2008). Ingestion of gelatinized potato starch containing
(2004). Variation for antioxidant activity and antioxidants in tomato. J. Am. a high level of phosphorus decreases serum and liver lipids in rats. J. Oleo
Soc. Hort. Sci., 129:704–711. Sci., 57:335–343.
Hashimoto, N., Ito, Y., Han, K. H., Shimada, K., Sekikawa, M., Topping, D. Kendall, C. W., Augustin, L. S., Emam, A., Josse, A. R., Saxena, N., and
L., Bird, A. R., Noda, T., Chiji, H., and Fukushima, M. (2006). Potato pulps Jenkins, D. J. (2006). The glycemic index: methodology and use. Nestle
lowered the serum cholesterol and triglyceride levels in rats. J. Nutr. Sci. Nutr. Workshop Ser. Clin. Perform. Program., 11:43–56.
Vitaminol. (Tokyo)., 52:445–450. King, N. J., Whyte, R., and Hudson J. A. (2007). Presence and significance of
Hatzis, C. M., Bertsias, G. K., Linardakis, M., Scott, J. M., and Kafatos, A. G. Bacillus cereus in dehydrated potato products. Food Prot., 70:514–520.
(2006). Dietary and other lifestyle correlates of serum folate concentrations Kleinkopf, G. E., Oberg, N. A. and Olsen, N. L. (2003). Sprout inhibition
in a healthy adult population in Crete, Greece: a cross-sectional study. Nutr. in storage: Current status, new chemistries and natural compounds. Am. J.
J., 5:5. Potato Res., 80:317–327.
Hayashi, K., Hibasami, H., Murakami, T., Terahara, N., Mori, M., Tsukui, A. Koppelman, S. J., van Koningsveld, G. A., Knulst, A. C., Gruppen, H., Pigmans,
(2006). Induction of apoptosis in cultured human stomach cancer cells by I. G. A. J., and de Jongh, H. H. J. (2002). Effect of heat-induced aggregation
potato anthocyanins and its inhibitory effects on growth of stomach cancer on the IgE binding of patatin (Sol t 1) is dominated by other potato proteins.
in mice. FSTR., 12:22–26. J. Agric. Food Chem., 50:1562–1568.
Helgeson, V. S., Viccaro, L., Becker, D., Escobar, O., and Siminerio, L. (2006). Lang, S. L. (1992). Potato skins found to have chemical residues. Cornell
Diet of adolescents with and without diabetes: Trading candy for potato Focus, 1:30.
chips? Diabetes Care, 29:982–987. Lasheras, C., Gonzalez, S., Huerta, J., Lombardia, C., Ibañez, R., Patterson, A.
Henry, C. J. K., Lightowler, H. J., Strik, C. M., and Storey, M. (2005). and Fernandez, S. (2003). Food habits are associated with lipid peroxidation
Glycaemic index values for commercially available potatoes in Great Britain. in an elderly population. J. Am. Diet. Assoc., 103:1480–1487.
Brit. J. Nutr., 94:917–921. Lazarov, K. and Werman, M. J. (1996). Hypocholesterolaemic effect of potato
Hickling, S., Hung, J., Knuiman, M., Divitini, M., and Beilby, J. (2008). Are peel as a dietary fibre source. Med. Sci. Res., 24:581–582.
the associations between diet and C-reactive protein independent of obesity? Lee, B. M. and Shim, G. A. (2007). Dietary exposure estimation of
Prev. Med, 47:71–76. benzo[a]pyrene and cancer risk assessment. J. Toxicol. Environ. Health A.,
Hijmans, R. J. (2003). The effect of climate change on global potato production. 70:1391–1394.
Am. J. Potato Res., 80:271–280. Lee, K.-R., Kozukue, N., Han, J.-S., Park, J.-H., Chang, E., Baek, E.-J., Chang,
Hill, A. J., Peikin, S. R., Ryan, C. A., and Blundell, J. E. (1990). Oral J.-S., and Friedman, M. (2004). Glycoalkaloids and metabolites inhibit the
administration of proteinase inhibitor II from potatoes reduces energy intake growth of human colon (HT29) and liver (HepG2) cancer cells. J. Agric.
in man. Physiol. Behav., 48:241–246. Food Chem., 52:2832–2839.
Hils, U. and Pieterse, L., Eds. (2007). World Catalogue of Potato Varieties. Lee, S. K., Ye, Y. M., Yoon, S. H., Lee, B. O., Kim, S. H., and Park, H. S.
Agrimedia Press, Agrimedia, GmbH, Clenze, Germany. (2006). Evaluation of the sensitization rates and identification of IgE-binding
 POTATOES AND HUMAN HEALTH 839

components in wild and genetically modified potatoes in patients with content in white, yellow, purple, orange, and dark-orange carrot cultivars. J.
allergic disorders. Clin. Mol. Allergy, 4:10. Am. Soc. Hort. Sci., 129:523–529.
Leeman, M., Ostman, E., and Björck, I. (2008). Glycaemic and satiating Nielsen, F., Mikkelsen, B. B., Nielsen, J. B., Andersen, H. R., and Grandjean,
properties of potato products. Eur. J. Clin. Nutr., 62:87–95. P. (1997). Plasma malondialdehyde as biomarker for oxidative stress:
Leo, L., Leone, A., Longo, C., Lombardi, D. A., Raimo, F., Zacheo, G. (2008). reference interval and effects of life-style factors. Clin. Chem., 43:1209–
Antioxidant compounds and antioxidant activity in "early potatoes." J. Agric. 1214.
Food Chem., 56:4154–4163. Olesen, P. T., Olsen, A., Frandsen, H., Frederiksen, K., Overvad, K., and
Li, X.-Q., Scanlon, M. G., Liu, Q., Coleman, W. K., (2006). Processing and Tjønneland, A. (2008). Acrylamide exposure and incidence of breast cancer
Value Addition. In: Handbook of Potato Production, Improvement, and among postmenopausal women in the Danish Diet, Cancer and Health Study.
Postharvest Management. Chap. 14, pp. 523–555. Gopal, J. and Khurana, Int. J. Cancer., 122:2094–2100.
S.M.P. Eds. Food Products Press, Binghamton, NY. Ortiz-Medina, E. (2007). Potato tuber protein and its manipulation by chimeral
Lineback, D., Pariza, M. W., Coughlin, J., Davies, C., Kettlitz, B., Robin, L. disassembly using specific tissue explantation for somatic embryogenesis.
P., Schmidt, D., Scimeca, J., and Stadler, R. (2006). Acrylamide in Food. Ph.D. Diss., McGill University, Montreal, 152 pp.
Issue Paper No. 32, 16 pp. Council for Agricultural Science and Technology, Packer, L. (1995). Oxidative stress, antioxidants, aging and disease. pp 1–14.
Ames, IA. In: Molecular and Cell Biology Updates (Oxidative Stress and Aging).
Lisińska, G. and Leszczyński, W. (1989). Potato Science and Technology. Cutler, R.G., Packer, L., Bertram, J., and Mori, A., Eds. Birkhauser Verlag,
Elsevier Applied Science, London, 391 pp. Basel, Switzerland.
Liu, S., Serdula, M., Janket, S.-J., Cook, N. R., Sesso, H. D., Willett, W. C., Panagiotakos, D. B., Pitsavos, C., Skoumas, Y., and Stefanadis, C. (2007). The
Manson, J. E., and Buring, J. E. (2004). A prospective study of fruit and association between food patterns and the metabolic syndrome using principal
vegetable intake and the risk of type 2 diabetes in women. Diabetes Care., component analysis: the ATTICA study. J. Am. Diet. Assoc., 107:979–987.
27:2993–2996. Paynter, N. P., Yeh, H. C., Voutilainen, S., Schmidt, M. I., Heiss, G., Folsom, A.
Liyanage, R., Han, K. H., Watanabe, S., Shimada, K., Sekikawa, M., Ohba, R., Brancati, F. L., and Kao, W. H. (2006). Coffee and sweetened beverage
K., Tokuji, Y., Ohnishi, M., Shibayama, S., Nakamori, T., and Fukushima, consumption and the risk of type 2 diabetes mellitus: the atherosclerosis risk
M. (2008). Potato and soy peptide diets modulate lipid metabolism in rats. in communities study. Am. J. Epidemiol., 164:1075–1084.
Biosci. Biotechnol. Biochem., 72:943–50). Pȩska, A., Gołubowska, G., Aniołowski, K., Lisińka, G., and Rytel, E. (2006).
Love, S. L. and Pavek, J. J. (2008). Positioning the potato as a primary food Changes of glycoalkaloids and nitrate contents in potatoes during chip
source of vitamin C. Am. J. Potato Res., 85:277–285. processing. Food Chem., 97:151–156.
Majamaa, H., Seppälä, U., Palosuo, T. Turjanmaa, K., Kalkkinen, N., and Pihlanto, A., Akkanen, S., and Korhonen, H. J. (2008). ACE-inhibitory
Reunala, T. 2001. Positive skin and oral challenge responses to potato and and antioxidant properties of potato (Solanum tuberosum). Food Chem.,
occurrence of immunoglobulin E antibodies to patatin (Sol t 1) in infants 109:104–112.
with atopic dermatitis. Pediatr. Allergy Immunol., 12:283–288. Pots, A. M., Gruppen, H., van Diepenbeek, R., van der Lee, J. J., van Boekel,
McGinnis, L. (2008). Potassium and potato preparation. USDA Research M. A. J. S., Wijngaards, G., and Voragen, A. G. J. (1999) The effect of
Service. http://www.ars.usda.gov/is/pr/2008/080624.htm. Visited June, 2008. storage of whole potatoes of three cultivars on the patatin and protease
Millar, D. J., Allen, A. K., Smith, C. G., Sidebottom, C., Slabas, A. R., and inhibitor content; a study using capillary electrophoresis and MALDI-TOF
Bolwell, G. P. (1992). Chitin-binding proteins in potato (Solanum tuberosum mass spectrometry. J. Sci. Food Agric., 79:1557–1564.
L.) tuber. Characterization, immunolocalization and effects of wounding. Pramod, S. N., Venkatesh, Y. P., and Mahesh, P. A. (2007). Potato lectin
Biochem. J., 283:813–821. activates basophils and mast cells of atopic subjects by its interaction with
Morris W. L., Ross H. A., Ducreux, L. J., Bradshaw J. E., Bryan G. J., and core chitobiose of cell-bound non-specific immunoglobulin E. Clin. Exp.
Taylor M. A. (2008). Umami compounds are a determinant of the flavor of Immunol., 148:391–401.106.
potato (Solanum tuberosum L.). J. Agric. Food Chem., 55:9627–9633. Priestley, H. (2006). How to think like consumers. . . and win! In: Potato
Morita, T., Oh-hashi, A., Takei, K., Ikai, M., Kasaoka, S., and Kiriyama, S. developments in a changing Europe. Chap. 20 pp. 189–198. Haase, N.U. and
(1997). Cholesterol-lowering effects of soybean, potato and rice proteins Haverkort, A.J. Eds. Wageningen Academic Pub.
depend on their low methionine contents in rats fed a cholesterol-free Prohens, J., Rodrı́guez-Burruezo, A., Raigón, M. D. and Nuez, F. (2007). Total
purified diet. J. Nutr., 127:470–477. phenolic concentration and browning in a collection of different varietal
Morton, L. W., Caccetta, R. A.-A., Puddey, I. B., and Croft, K. D. (2000). types and hybrids of eggplant: implications for breeding for higher nutritional
Chemistry and biological effects of dietary phenolic compounds: relevance quality and reduced browning. J. Am. Soc. Hort. Sci., 132:638–646.
to cardiovascular disease. Clin. Exp. Pharmacol. Physiol., 27:152–159. Raben, A. (2002) Should obese patients be counselled to follow a low-glycaemic
Monro, J. A. and Shaw, M. (2008). Glycemic impact, glycemic glucose index diet? Obesity Rev. 3:245–256.
equivalents, glycemic index, and glycemic load: definitions, distinctions, Receveur, O., Morou, K., Gray-Donald, K., and Macaulay, A. C. (2008).
and implications. Am. J. Clin. Nutr., 87:237S-243S. Consumption of key food items is associated with excess weight among
Mou, B. (2005). Genetic variation of beta-carotene and lutein contents in elementary-school-aged children in a Canadian first Nations community. J.
lettuce. J. Am. Soc. Hort. Sci., 130:870–876. Am. Diet. Assoc., 108:362–366.
Mucci, L. A. and Wilson, K. M. (2008). Acrylamide intake through diet and Reddivari, L., Hale, H. A., and Creighton-Miler, J. (2007). Determination of
human cancer risk. J. Agric. Food Chem., 56:6013–6019. phenolic content, composition and their contribution to antioxidant activity
Nara, K., Miyoshi, T., Honma, T., and Koga, H. (2006). Antioxidative activity in specialty potato selections. Am. J. Potato Res., 84:275–282.
of bound-form phenolics in potato peel. Biosci. Biotechnol. Biochem., Reddivari, L., Vanamala, J., Chintharlapalli, S., Safe, S. H., and
70:1489–1491. Miller, J. C. Jr. (2007). Anthocyanin fraction from potato extracts
Narcy, A., Robert, L., Mazur, A., Demigné, C, and Rémésy, C. (2006). Effect is cytotoxic to prostate cancer cells through activation of caspase-
of potato on acid-base and mineral homeostasis in rats fed a high-sodium dependent and caspase-independent pathways. Carcinogenesis, 28:2227–
chloride diet. Br. J. Nutr., 95:925–932. 2235.
Nema, V., Agrawal, R., Kasmboj, D. V., Goel, A. K., and Singh, L. (2007). Iso- Reyes, L. F. and Cisneros-Zevallos, L. (2003). Wounding stress increases the
lation and characterization of heat resistant enterotoxigenic Staphylococcus phenolic content and antioxidant capacity of purple-flesh potatoes (Solanum
aureus from a food poisoning outbreak in Indian subcontinent. Intl. J. Food tuberosum L.). J. Agric. Food Chem., 51:5296–5300.
Microbiol., 117:29–35. Reyes, L. F., and Cisneros-Zevallos, L. (2007). Degradation kinetics and
Nicolle, C., Simon, G., Rock, E., Amouroux, P., and Rémésy, C. (2004). colour of anthocyanins in aqueous extracts of purple- and red-flesh potatoes
Genetic variability influences carotenoid, vitamin, phenolic, and mineral (Solanum tuberosum L.). Food Chem., 100:885–894.
840 M. E. CAMIRE ET AL.

Reyes, L. F., Villareal, J. E., and Cisneros-Zevallos, L. (2007). The increase in

R
Svetol, 2008. Svetol http://www.svetol.com/visited June 30, 2008.
antioxidant capacity after wounding depends on the type of fruit or vegetable Takeda, Y., Hizukuri, S., Ozono, Y., Suetake, M. (1983). Actions of porcine
tissue. Food Chem., 101:254–1262. pancreatic and Bacillus subtilis alpha amylases and Aspergillus niger
Robert, L., Narcy, A., Rayssiguier, Y., Mazur, A., and Rémésy, C. (2008). Lipid glucoamylase on phosphorylated (1–4)-alpha-D-glucan. Biochem. Biophys.
metabolism and antioxidant status in sucrose vs. potato-fed rats. J. Am. Coll. Acta, 749:302–311.
Nutr., 27:109–116. Tareke, E., Rydberg, P., KArlsson, P., Eriksson S., and Törnqvist, M. (2002).
Robert, L., Narcy, A., Demigne, C., Mazur A., and Rémésy, C. (2006). Entire Analysis of acrylamide, a carcinogen formed in heated foodstuffs. J. Agric.
potato consumption improves lipid metabolism and antioxidant status in Food Chem., 50:4998–5006.
cholesterol-fed rat. Eur. J. Nutr., 45:267–274. Tarn, T. R., Tai, G. C. C., and Liu, Q. (2006). Quality improvement. In: Hand-
Robert, L., Narcy, A., Rayssiguier, Y., Mazur A., and Rémésy, C. (2008). Lipid book of Potato Production, Improvement, and Postharvest Management. Ch.
metabolism and antioxidant status in sucrose vs. potato-fed rats. J. Am. Coll. 5, pp. 147–178. Gopal, J. and Khurana, S.M. P., Eds. Haworth Press Inc., NY.
Nutr., 27:109–116. Tajner-Czopek, A., Jarych-Szyszka, M., and Lisińska, G. (2008). Changes in
Roberts, S. B. (2000). High-glycemic index foods, hunger, and obesity: is there glycoalkaloids content of potatoes destined for consumption. Food Chem.,
a connection? Nutr. Rev., 58:163–169. 106:706–711.
Rodriguez de Sotillo, D., and Hadley, M., (2002). Chlorogenic acid modifies Tudela, J. A., Cantos, E., Espı́n, J. C., Tomás-Barberán, F. A., and Gil, M. I.
plasma and liver concentrations of: cholesterol, triacylglycerol, and minerals (2002). Induction of antioxidant flavonol biosynthesis in fresh-cut potatoes.
in (fa/fa) Zucker rats. J. Nutr. Biochem., 13:717–726. Effect of domestic cooking. J. Agric. Food Chem., 50:5925–5931.
Rodriguez de Sotillo, D., Hadley, M., and Holm, E. T., (1994). Phenolics in Tudela, J. A., Hernández, J. A., Gil, M. I., and Espı́n, J. C. (2003). L-galactono-
aqueous potato peel extract: extraction, identification and degradation. J. gamma-lactone dehydrogenase and vitamin C content in fresh-cut potatoes
Food Sci., 59:649–651. stored under controlled atmospheres. J. Agric. Food Chem., 51:4296–4302.
Schmidt, M. H., Raulf-Heimsoth, M., and Posch, A. (2002). Evaluation of Van Eck, H. J. (2007). Genetics of morphological and tuber traits. In: Potato
patatin as a major cross-reactive allergen in latex-induced potato allergy. Biology and Biotechnology: Advances and Perspectives. Chap. 6, pp.
Ann. Allergy Asthma Immunol., 89:613–618. 91–115. Vreugdenhil, D. Ed., Elsevier, London.
Seddon, J. M., Ajani, U. A., Sperduto, R. D., Hiller, R., Blair, N., Burton. T. C., Van Eck, J., Conlin, B., Garvin, D. F., Mason, H., Navarre, D. A., and Brown,
Farber, M. D., Gragoudas, E. S., Haller, J. and Miller, D. T. (1994). Dietary C. R. (2007). Enhancing beta-carotene content in potato by RNAi-mediated
carotenoids, vitamins A, C, and E, and advanced age-related macular degen- silencing of the beta-carotene hydroxylase gene. Am. J. Potato Res.,
eration. Eye Disease Case-Control Study Group. JAMA, 272:1413–1420. 84:331–342.
Selma, M. V., Beltrán, D., Chacón-Vera, E., and Gil, M. I. (2006). Effect of Villegas, R., Liu, S., Gao, Y. T., Yang, G., Li, H., Zheng, W., and Shu, X.
ozone on the inactivation of Yersinia enterocolitica and the reduction of O. (2007). Prospective study of dietary carbohydrates, glycemic index,
natural flora on potatoes. J. Food Prot., 69:2357–2363. glycemic load, and incidence of type 2 diabetes mellitus in middle-aged
Seppälä, U., Alenius, H., Turjanmaa, K., Reunala, T., Palosuo, T., and Kalkki- Chinese women. Arch. Intern. Med., 167:2310–2316.
nen, N. (1999). Identification of patatin as a novel allergen for children with Vivanti, V., Finotti, E., and Friedman M. (2006) Level of acrylamide precursors
positive skin prick test responses to raw potato. J. Allergy Clin. Immunol., asparagine, fructose, glucose, and sucrose in potatoes sold at retail in Italy
103:165–171. and in the United States. J. Food Sc., 71:C81-C85.
Seppälä, U., Majamaa, H., Turjanmaa, K., Helin, J., Reunala, T., Kalkkinen, Wang, Q. and Zhang, W. (2004). China’s potato industry and potential impacts
N., and Palosuo, T. (2001). Identification of four novel potato (Solanum on the global market. Am. J. Potato Res., 81:101–110.
tuberosum) allergens belonging to the family of soybean trypsin inhibitors. Wang, H., Ng, T. B., Ooi, V. E. C., and Liu, W. K. (2000). Effects of lectins
Allergy., 56:619–626. with different carbohydrate-binding specificities on hepatoma, choriocarci-
Shewry, P. R. (2003). Tuber storage proteins. Ann. Bot. (Lond)., 91:755–769. noma, melanoma and osteosarcoma cell lines. Inl. J. Biochem. Cell Biol.,
Shih, Y.-W., Chen, P.-S., Wu, C.-H., Jeng, Y.-F., and Wang, C.-J. (2007). 32:365–372.
α-Chaconine-reduced metastasis involves a PI3K/Akt signaling pathway Wang, L. L., and Xiong, Y. L. (2005). Inhibition of lipid oxidation in cooked
with downregulation of NF-κB in human lung adenocarcinoma A549 cells. beef patties by hydrolyzed potato protein is related to its reducing and radical
J. Agric. Food Chem., 55:11035–11043. scavenging ability. J. Agric. Food Chem., 53:9186–9192.
Singh, N., Kamath, V., and Rajini, P. S. (2005a). Attenuation of hyperglycemia Wang-Pruski, G. and Nowak, J. (2004). Potato after-cooking darkening. Am. J.
and associated biochemical parameters in STZ-induced diabetic rats by di- Potato Res., 81:7–16.
etary supplementation of potato peel powder. Clin. Chim. Acta, 353:165–175. Wang-Pruski, G., Zebarth, B. J., Leclerc, Y., Arsenault, W. J., Botha, E. J.,
Singh, N., Kamath, V., and Rajini, P. S. (2005b). Protective effect of potato Moorehead, S., and Ronis, D. (2007). Effect of soil type and nutrient man-
peel powder in ameliorating oxidative stress in streptozotocin diabetic rats. agement on potato after cooking darkening. Am. J. Potato Res., 84:291–299.
Plant Foods Hum. Nutr., 60:49–54. White, P. J. and Broadley, M. R. (2005). Biofortifying crops with essential
Singh, N. and Rajini, P. S. (2008). Antioxidant-mediated protective effects of mineral elements. Trends Plant Sci., 10:586–593.
potato peel extract in erythrocytes against oxidative damage. Chemico-Biol. Williams, D. E., Wareham, N. J., Cox, B. D., Byrne, C. D., Hales, C. N., and
Interact., 173:97–104. Day, N. E. (1999). Frequent salad vegetable consumption is associated with
Singh, N. and Rajini, P. S. (2004). Free radical scavenging activity of an a reduction in the risk of diabetes mellitus. J. Clin. Epidemiol., 52:329–335.
aqueous extract of potato peel. Food Chem., 85:611–616. Woolfe, J. A. (1987). The Potato in the Human Diet. Cambridge University
Samaha, F. F., Foster, G. D., and Makris, A. P. (2007). Low-carbohydrate Press. p. 143–149.
diets, obesity, and metabolic risk factors for cardiovascular disease. Curr. Work, T. M. and Camire, M. E. (1996). Phenolic acid detection thresholds in
Atheroscler. Rep., 9:441–447. processed potatoes. Food Qual. Pref., 7:271–274.
Sohl, N. L. and Brand-Miller, J. (1999). The glycaemic index of potatoes: the ef- Wrieden, W. L., Hannah, M. K., Bolton-Smith, C., Tavendale, R., Morrison,
fect of variety, cooking methods and maturity. Eur. J. Clin. Nutr., 53:249–254. C., and Tunstall-Pedoe, H. (2000). Plasma vitamin C and food choice in the
Soliman, K. M. (2001). Changes in concentration of pesticide residues in pota- third Glasgow MONICA population survey. J. Epidemiol. Comm. Health,
toes during washing and home preparation. Food Chem. Toxicol., 39:887–891. 54:355–360.
Spiller, H. A. and Sawyer, T. S. (2006). Toxicology of oral antidiabetic Yang, A.-A., Paek, S.-H., Kozukue, N., Lee, K.-R., and Kim, J.-A. (2006).
medications. Am. J. Health Syst. Pharm., 63:929–938. α-Chaconine, a potato glycoalkaloid, induces apoptosis of HT-29 human
Suttle, J. (2008). Symposium introduction: Enhancing the nutritional value of cancer colon cancer cells through caspase-3 activation and inhibition of ERK
potato tubers. Am. J. Potato Res., 85:266. 1/2 phosphorylation. Food Chem. Toxicol., 44:839–846.

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