9
Critical Review Nutrients, bioactive non-nutrients and anti-nutrients in potatoes Barbara Burlingame *, Beatrice Mouille ´ , Ruth Charrondie ` re Food and Agriculture Organization of the United Nations, viale delle Terme di Caracalla, 00153 Rome, Italy 1. Introduction Potato biodiversity is vast: approximately 5000 varieties of potatoes are known. Most cultivated varieties belong to the species Solanum tuberosum. However, about 10 other Solanum species are cultivated, and 200 wild species have been recorded. Conventional agriculture and climate change may threaten the survival of these wild relatives: it is forecast that as many as 12% will become extinct as their growing conditions deteriorate. If climate changes drastically, the area where wild potatoes grow naturally could be reduced by as much as 70% (FAO, 2008). According to the latest FAOSTAT statistics (FAO, 2009a), potatoes account for only about 2% of the world’s dietary energy supply. As a staple food, potatoes are less important globally than rice, wheat and maize, which account for 20%, 18% and 5%, respectively. During the past 50 years, potato production and consumption has been rather steady. FAOSTAT reports continue to increase around the world. Potatoes have a more dominant place in the diets of people in developed countries than those in developing countries. Averaged across the world, potatoes account for only 170 kJ (41 kcal) per person per day for the developing world, yet the number is 540 kJ (130 kcal) for developed world. Countries in the Commonwealth of Independent States (CIS) rely most heavily on potatoes for their energy intakes, with Belarus topping the list at 1250 kJ (300 kcal) per person per day. Seven other CIS (Common- wealth of Independent States) countries and Portugal follow closely behind, each averaging more than 840 kJ (200 kcal) per person per day from potatoes. Potatoes are the subject of many popular culture conceptions and misconceptions. At various times in recent history, they have had the reputation of being nutritionally both ‘‘bad’’ and ‘‘good’’. During its phase of having a nutritionally bad reputation, potatoes were marked as being ‘‘fattening’’ and contributing to diabetes, obesity and its co-morbidities. Many more campaigns have promoted potatoes as good, including the famous, ‘‘Potatoes are not fattening’’ campaign of the United Kingdom, ‘‘Goodness Unearthed’’ and ‘‘Healthy MR. POTATO HEAD’’ campaign of the US potato industry (US Potato Board, 2009a, 2009b), the ‘‘Spudtacular Show’’ and ‘‘Baked Potato Campaign’’ of New Zealand (HEHA, 2009). The food components most frequently highlighted in nutrient content claims are energy, e.g. ‘‘one medium-sized potato has 110 calories’’; fat, e.g. ‘‘100% fat free; saturated fat, trans fatty acids, and cholesterol, e.g. ‘‘fat free’’’, ‘‘0 grams trans fat’’, no cholesterol; sodium, e.g. ‘‘salt free’’. Other nutrients frequently declared are potassium, fibre, Vitamin C, B6 and various antioxidants (US Potato Board, 2009c). Journal of Food Composition and Analysis 22 (2009) 494–502 ARTICLE INFO Article history: Received 29 September 2009 Accepted 29 September 2009 Keywords: Solanum spp. Potato Variety Wild species Biodiversity Cultivar differences Nutrient composition Plant genetic diversity Micronutrients Bioactive non-nutrients Anti-nutrients Antioxidant composition Food composition Food analysis ABSTRACT Globally, potatoes account for only about 2% of the food energy supply, yet they are the predominant staple for many countries. In developed countries, potatoes account for 540 kJ (130 kcal) per person per day, while in developing countries, it is only 170 kJ (42 kcal) per person per day. In addition to energy, which is derived almost entirely from their carbohydrate content, many varieties of potatoes contribute nutritionally important amounts of dietary fibre (up to 3.3%), ascorbic acid (up to 42 mg/100 g), potassium (up to 693.8 mg/100 g), total carotenoids (up to 2700 mcg/100 g), and antioxidant phenols such as chlorogenic acid (up to 1570 mcg/100 g) and its polymers, and anti-nutrients such as a-solanine (0.001–47.2 mg/100 g); and lesser amounts of protein (0.85–4.2%), amino acids, other minerals and vitamins, and other beneficial and harmful bioactive components. Nutrient content depends on a number of factors, with variety being among the most important. Potato biodiversity is vast, with more than 4000 known varieties. Most belong to the species Solanum tuberosum, but another 10 species are cultivated and 200 wild species have been identified. Modern agricultural practices and climate change are contributing to the loss of potato biodiversity, and thus the loss of the genes coding for nutrient biosynthetic pathways. Knowledge of differences in nutrient composition of potatoes related to their genetic diversity will help guide strategies that may contribute to reducing biodiversity loss and improving food and nutrition security. ß 2009 Published by Elsevier Inc. * Corresponding author. Tel.: +39 06 570 53728. E-mail address: [email protected] (B. Burlingame). Contents lists available at ScienceDirect Journal of Food Composition and Analysis journal homepage: www.elsevier.com/locate/jfca 0889-1575/$ – see front matter ß 2009 Published by Elsevier Inc. doi:10.1016/j.jfca.2009.09.001

Nutrients, bioactive non-nutrients and anti-nutrients in potatoes

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Journal of Food Composition and Analysis 22 (2009) 494–502

Critical Review

Nutrients, bioactive non-nutrients and anti-nutrients in potatoes

Barbara Burlingame *, Beatrice Mouille, Ruth Charrondiere

Food and Agriculture Organization of the United Nations, viale delle Terme di Caracalla, 00153 Rome, Italy

A R T I C L E I N F O

Article history:

Received 29 September 2009

Accepted 29 September 2009

Keywords:

Solanum spp.

Potato

Variety

Wild species

Biodiversity

Cultivar differences

Nutrient composition

Plant genetic diversity

Micronutrients

Bioactive non-nutrients

Anti-nutrients

Antioxidant composition

Food composition

Food analysis

A B S T R A C T

Globally, potatoes account for only about 2% of the food energy supply, yet they are the predominant

staple for many countries. In developed countries, potatoes account for 540 kJ (130 kcal) per person per

day, while in developing countries, it is only 170 kJ (42 kcal) per person per day. In addition to energy,

which is derived almost entirely from their carbohydrate content, many varieties of potatoes contribute

nutritionally important amounts of dietary fibre (up to 3.3%), ascorbic acid (up to 42 mg/100 g),

potassium (up to 693.8 mg/100 g), total carotenoids (up to 2700 mcg/100 g), and antioxidant phenols

such as chlorogenic acid (up to 1570 mcg/100 g) and its polymers, and anti-nutrients such as a-solanine

(0.001–47.2 mg/100 g); and lesser amounts of protein (0.85–4.2%), amino acids, other minerals and

vitamins, and other beneficial and harmful bioactive components. Nutrient content depends on a

number of factors, with variety being among the most important. Potato biodiversity is vast, with more

than 4000 known varieties. Most belong to the species Solanum tuberosum, but another 10 species are

cultivated and 200 wild species have been identified. Modern agricultural practices and climate change

are contributing to the loss of potato biodiversity, and thus the loss of the genes coding for nutrient

biosynthetic pathways. Knowledge of differences in nutrient composition of potatoes related to their

genetic diversity will help guide strategies that may contribute to reducing biodiversity loss and

improving food and nutrition security.

� 2009 Published by Elsevier Inc.

Contents lists available at ScienceDirect

Journal of Food Composition and Analysis

journa l homepage: www.e lsev ier .com/ locate / j fca

1. Introduction

Potato biodiversity is vast: approximately 5000 varieties ofpotatoes are known. Most cultivated varieties belong to the speciesSolanum tuberosum. However, about 10 other Solanum species arecultivated, and 200 wild species have been recorded. Conventionalagriculture and climate change may threaten the survival of thesewild relatives: it is forecast that as many as 12% will becomeextinct as their growing conditions deteriorate. If climate changesdrastically, the area where wild potatoes grow naturally could bereduced by as much as 70% (FAO, 2008).

According to the latest FAOSTAT statistics (FAO, 2009a),potatoes account for only about 2% of the world’s dietary energysupply. As a staple food, potatoes are less important globally thanrice, wheat and maize, which account for 20%, 18% and 5%,respectively. During the past 50 years, potato production andconsumption has been rather steady. FAOSTAT reports continue toincrease around the world. Potatoes have a more dominant place inthe diets of people in developed countries than those in developingcountries. Averaged across the world, potatoes account for only170 kJ (41 kcal) per person per day for the developing world, yet

* Corresponding author. Tel.: +39 06 570 53728.

E-mail address: [email protected] (B. Burlingame).

0889-1575/$ – see front matter � 2009 Published by Elsevier Inc.

doi:10.1016/j.jfca.2009.09.001

the number is 540 kJ (130 kcal) for developed world. Countries inthe Commonwealth of Independent States (CIS) rely most heavilyon potatoes for their energy intakes, with Belarus topping the list at1250 kJ (300 kcal) per person per day. Seven other CIS (Common-wealth of Independent States) countries and Portugal followclosely behind, each averaging more than 840 kJ (200 kcal) perperson per day from potatoes.

Potatoes are the subject of many popular culture conceptions andmisconceptions. At various times in recent history, they have had thereputation of being nutritionally both ‘‘bad’’ and ‘‘good’’. During itsphase of having a nutritionally bad reputation, potatoes weremarked as being ‘‘fattening’’ and contributing to diabetes, obesityand its co-morbidities. Many more campaigns have promotedpotatoes as good, including the famous, ‘‘Potatoes are not fattening’’campaign of the United Kingdom, ‘‘Goodness Unearthed’’ and‘‘Healthy MR. POTATO HEAD’’ campaign of the US potato industry(US Potato Board, 2009a, 2009b), the ‘‘Spudtacular Show’’ and‘‘Baked Potato Campaign’’ of New Zealand (HEHA, 2009).

The food components most frequently highlighted in nutrientcontent claims are energy, e.g. ‘‘one medium-sized potato has 110calories’’; fat, e.g. ‘‘100% fat free; saturated fat, trans fatty acids, andcholesterol, e.g. ‘‘fat free’’’, ‘‘0 grams trans fat’’, no cholesterol;sodium, e.g. ‘‘salt free’’. Other nutrients frequently declared arepotassium, fibre, Vitamin C, B6 and various antioxidants (US PotatoBoard, 2009c).

B. Burlingame et al. / Journal of Food Composition and Analysis 22 (2009) 494–502 495

Rarely are varieties differentiated in nutrient declarations orcontent claims, yet there are well-known content differences.Under the Codex Alimentarius food labelling guidelines for‘‘source’’ and ‘‘good source’’ of nutrients (Codex Alimentarius,2007), some varieties would qualify for the claims, while otherswould not. The purpose of this review is to examine the literatureon potato composition at the level of the genetic resource andcompile the data illustrating that the vast existing biodiversity ofpotatoes also reflects vast differences in nutrient content.

2. Methods

2.1. Compilation of nutrient composition data of potatoes by variety

A thorough literature search was performed to gather existinginformation on nutrient composition of potatoes by variety. Whilemany environmental, agricultural and post-harvest factors such asclimate, fertilizer use, maturity at harvest, storage, processing andpreparation influence the nutrient content, this review focusesmainly on the measured differences among varieties. However, theinfluence of variety cannot be assured for many of the data.National Food Composition tables were searched, as were nutritionand agriculture journals. In most food composition data resources,part (flesh, flesh + peel) and state (e.g. cooked or raw) are generallyreported, but varietal name is often missing.

Differences in water content will likely account for many of thedifferences in nutrient content, because values are present on a per100 g basis, fresh weight, unless otherwise noted.

2.2. Nutrient identification—methods and expressions

Among the different data sources, significant differenceswere encountered in the data expressions and conversionfactors. Therefore it was necessary to standardize these datafor making comparisons. For example, nitrogen to proteinconversion factors are mostly standardized as 6.25 for potatoes,some researchers have suggested numbers as low as 5.7 and ashigh as 7.5 (Vigue and Li, 1975; Desborough and Weiser, 1974).The variation in units used to calculate micronutrient composi-tion was extensive. Some of the units encountered were mg/g,g and mg/100 g, mg/kg, mg/100 kg, % dmb, ppm and %. All nutri-ents were standardized to g, mg, or mcg per 100 g fresh weight,unless otherwise noted. Differences in the data reported in thispaper and those found in the source materials result from theserecalculations.

3. Results and discussion

Tables 1–4 provide summary data on the varietal differencesfound in potatoes. The full data set is to be published as an on-linedatabase (FAO, 2009b).

In addition to energy, which is derived almost entirely fromtheir carbohydrate content, many varieties of potatoes contributenutritionally important amounts of other nutrients and bioactivenon-nutrients. Although nutrient content depends on a number offactors, variety is thought to be among the most significant factors(Toledo and Burlingame, 2006). Important nutrients in differentvarieties of potatoes include dietary fibre, up to 3.7% found in theskin of Red Pontiac variety (Mullin et al., 1993), the highest valuefound in flesh is for the variety Runa, grown in Argentina, whichhas a value of 3.3 g/100 g (Jimenez et al., 2009), ascorbic acid, up to42 mg/100 g in the Korean grown Chaju (Han et al., 2004),potassium, with up to 693.8 mg/100 g found in Azucena of Spain(Casanas Rivero et al., 2003b), total carotenoids, with high-endvalues exceeding 2000 mcg/100 g for several varieties includingPhureja group 1 of Peru (Bonierbale et al., 2009).

3.1. Water

The water content of fresh weight potatoes varies for a numberof reasons, variety being one. Values reported in the literature formoisture content of fresh weight potatoes, raw, range from 63 to87%. The lowest values was reported for the variety Chandramukhi,grown in India (Chatterjee et al., 2006); the highest for the PucaQuitish variety grown in Peru (Ritter et al., 2008). It is likely thatsome of the differences in nutrient composition are related to thedifferences in water content.

3.2. Carbohydrates

More than 95% of the energy in uncooked/unprocessed potatocomes from its carbohydrate content. Data from studies on 53different varieties of potato show starch contents as low as 9.1% foran Indian cultivar Kufri Bahar (Negi and Nath, 2002), and as high as22.6% for another Argentine variety, Imilla Negra S. tuberosum ssp.Andigena (Jimenez et al., 2009).

3.3. Protein

Protein content showed a wide range of values, from a low ofless than 1% for the Argentine cultivar Revolucion (Jimenez et al.,2009), to a high of 4.2% for the Spanish Roja Rinon (Ritter et al.,2008).

4. Special issue of JFCA on potato

This special issue covers a range of topics, from new data onnutrition and biodiversity issues to topics relevant to food safety.The importance of the potato as one of the world’s major staplecrops is reflected in the geographic origin of the researchcontributions which include Algeria, Argentina, Brazil, Denmark,France, Holland, India, Italy, New Zealand, Philippines, Poland,Turkey and U.S.A.

Several articles quantify the nutrients and antioxidants.Burgos et al. (2009b) determined the total and individualcarotenoid profiles in raw tubers of a sample of 23 accessionsof Solanum phureja cultivated potatoes in two papers in this issue,first by spectrophotometry and HPLC. The authors identify twoaccessions which contains the highest concentration of zea-xanthin (1290 mg 100 g�1 FW) reported thus far for potatoes,including genetically modified potatoes; see Fig. 1 (p. 500). Thecompanion paper by Bonierbale et al. (2009) presents a newmethod to estimate total and individual carotenoid concentra-tions by near-infrared reflectance spectroscopy (NIRS), and showsthat this method is useful for characterizing a large germplasmcollection at low cost and in reduced time compared to HPLC.Burgos et al. (2009a) look at how the ascorbic acid concentrationof native Andean potato varieties is affected by environment,cooking and storage; in another paper by a slightly different teamof researchers, Burgos et al. (2009c) study the protein, iron, zincand calcium concentrations of potatoes following traditionalprocessing as ‘‘chuno’’, and they conclude that the addition ofwater used in the preparation of this traditionally freeze–driedpotato in the Peruvian highlands contributes to higher calciumlevels.

In another paper from the International Potato Center in Peru,Andre et al. (2009) present their data on the influence ofenvironment and genotype on polyphenol compounds and in

vitro antioxidant capacity of native Andean potatoes (S. tuberosum

L.). They conclude that the hydrophilic antioxidant capacityobserved in potato extracts is the result of various and complexinteractions that could be attributed to phenolic compounds, toother non-identified molecules, and to synergistic as well as

Table 1Varietal differences in macronutrient composition. Data in this table have combined values from skin, flesh, or the whole potato. Cf. the full set of food composition tables on potato nutrients (FAO, 2009b).

Range Average Variety with highest value Taxonomic

groupa

Variety with lowest value Taxonomic

groupa

Notes Refb

(highest)

Refb (lowest)

Energy (n = 9) 240–423 kJ/100 g 340.00 Imilla Negra (Argentina) Adg Spunta (Argentina) Tbr 1 1

Moisture (n = 51) 62.68–87.0 g/100 g 78.50 Puca Quitish (Peru) Adg Chandramukhi (India) Tbr 3 4

Lipid (n = 15) 0.05–0.51 g/100 g 0.20 Imilla Colorada (Argentina) Adg Spunta (Argentina) Tbr 1 1

Protein (n = 41) 0.85–4.2 g/100 g 2.12 Roja Rinon (Spain) Tbr/T Revolucion (Argentina) Adg 3 1

Total fiber (n = 31) 0.3–3.67 g/100 g 2.04 Red Pontiac* – skin

(Eastern Canada)

Tbr Katahdin *– flesh (Maine, USA) Tbr In flesh, Runa (Argentina)

Adg has highest value of

3.3 g/100 g (ref 1)

5 7

Soluble fiber (n = 13) 0.87–1.22 g/100 g 1.04 Yukon Gold*

(Eastern Canada)

Tbr Red Pontiac* (Eastern Canada) Tbr 5 5

Insoluble fiber (n = 1) 0.41–2.53 g/100 g 1.31 Red Pontiac*-skin

(Eastern Canada)

Tbr Kennebec*-flesh Eastern Canada Tbr In flesh, Russet Burbank

(Canada) Tbr has highest

value of 0.7 g/100 g (ref 5)

5 5

Starch (n = 53) 9.1–22.6 g/100 g 16.13 Imilla Negra (Argentina) Adg Kufri Bahar (India) Tbr 1 9

Glucose (n = 14) 0.015–0.34 g/100 g 0.12 Arinda (Bologna, Italy) Tbr FL1815 (Oregon, USA) Tbr Jelli (Bologna, Italy) has

next lowest value

0.02 g/100 g (ref 10)

10 11

Fructose (n = 15) 0.00–0.18 g/100 g 0.07 Frinka (Bologna, Italy) Tbr Sponta (Bologna, Italy) Tbr 10 10

Sucrose (n = 19) 0.08–1.39 g/100 g 0.40 Marabel (Bologna, Italy) Tbr Ptarmigan (Oregon, USA) Tbr 10 11

Amylose (n = 5) 3.5–4.3 g/100 g 3.86 Bonita (Canary Islands, Spain) Adg Cara (Canary Islands, Spain) Tbr/T 13 13

The number of varieties used to calculate the values is indicated by n. (The number of studies is often greater than this value, as where data for the same variety were available from different studies, the mean value calculated was

used.) All data were taken from the references and were expressed in fresh weight for raw potatoes; an asterisk indicates that the value was calculated from dry weight. The place of origin is indicated in brackets.

References: (1) Jimenez et al. (2009), (2) Tudela et al. (2002), (3) Ritter et al. (2008), (4) Chatterjee et al. (2006), (5) Mullin et al. (1993), (6) Augustin et al. (1978), (7) Toma et al. (1979), (8) Bacigalupo et al. (2004), (9) Negi and Nath

(2002), (10) Finotti et al. (2006), (11) Rodriguez-Saona and Wrolstad (1997), (12) Casanas Rivero et al. (2003a), (13) Casanas et al. (2002), (14) del Mar Verde Mendez et al. (2004), (15) Casanas Rivero et al. (2003b), (16) Andre et al.

(2007a), (17) Andre et al. (2007b), (18) Burgos et al. (2009a), (19) Burgos et al. (2007), (20) Zhang et al. (1997), (21) Im et al. (2008), (22) Cambie and Ferguson (2003), (23) Delaplace et al. (2008), (24) Lu et al. (2001), (25) Sikora and

Cieslik (1999), (26) Cieslik and Sikora (1998), (27) Chun et al. (2006), (28) Han et al. (2004), (29) Hajslova et al. (2005), (30) Brown et al. (2008), (31) Campos et al. (2006), (32) Bonierbale et al. (2009), (33) Burgos et al. (2009b), (34)

Hale (2003), (35) Breithaupt and Bamedi (2002), (36) Andersen et al. (2002), (37) Donnelly et al. (2001), (38) Kalogeropoulos et al. (2007) (39) Linnemann et al. (1985), (40) Friedman et al. (2003), (41) Thybo et al. (2000), (42) Hale

et al. (2008), (43) Brown (2008), (44) Lewis et al. (1998), (45) Konschuh et al. (2007), (46) Brown et al. (2003), (47) Konschuh et al. (2005), (48) Brown (2006), (49) Phillippy et al. (2004), (50) Sarquis et al. (2000), (51) Sengul et al.

(2004), (52) Groza et al. (2005).a Solanum species used: S. tuberosum, S. brevidens, S. acaule, and S. commersonii. S. tuberosum L.: Tbr; S. tuberosum subsp. tuberosum: Tbr/T; S. tuberosum subsp. andigena: Adg; S. chaucha: Cha; S. phureja subsp. phureja: Phu; S.

stenotomum ssp. stenotomum: Stn/S; S. stenotomum ssp. goniocalyx: Gon/G.b Ref: reference.

B.

Bu

rling

am

eet

al./Jo

urn

al

of

Foo

dC

om

po

sition

an

dA

na

lysis

22

(20

09

)4

94

–5

02

49

6

Table 2Varietal differences in selected micronutrient composition.

Range Average Variety with highest

nutrient content

Taxonomic

groupa

Variety with lowest

nutrient content

Taxonomic

groupa

Refb

(highest)

Refb

(lowest)

Thiamin (B1) (n = 5) 0.08–0.11 mg/100 g 0.09 Katahdin*

(Maine, USA)

Tbr Norchip*

(North Dakota, USA)

Tbr 6 6

Riboflavin (B2) (n = 5) 0.022–0.36 mg/100 g 0.07 Russet Burbank*

(Idaho USA)

Tbr Katahdin* (USA) Tbr 6 6

Niacin (B3) (n = 5) 0.62–2.07 mg/100 g 1.26 Russet Burbank*

(USA)

Tbr White rose (Davis,

California, USA, USA)

Tbr 6 19

Vitamin B6 (n = 4) 0.15–0.3 mg/100 g 0.24 Norchip*

(North Dakota, USA)

Tbr Pontiac* (North

Dakota, USA)

Tbr 6 6

Folic acid (n = 4) 10.3–15.3 mcg/100 g 12.42 Russet Burbank*

(USA)

Tbr Katahdin* (Maine, USA) Tbr 6 6

Calcium (n = 127) 1.35–27.8 mg/100 g 10.59 Jancko Anckanchi*

(Huancayo, Peru)

Ajanhuiri Marta (Poland) Tbr 17 26

Phosphorus (n = 26) 33–126 mg/100 g 71.43 Unknown CIP703315

(Peru)

Gon/G Kufri Bahar (India) Tbr 3 9

Sodium (n = 31) 0.1–7.7 mg/100 g 2.87 Peluca (Spain) Tbr/T Ibicenda (Spain) Tbr 15 3

Potassium (n = 53) 239–693.8 mg/100 g 443.30 Azucena (Spain) Adg Monalisa (Spain) Tbr 15 3

Magnesium (n = 47) 10.8–37.6 mg/100 g 20.21 Puca Huaryo

(Lima, Peru)

Cha Monalisa (Spain) Tbr 3 3

Manganese (n = 28) 0.05–0.39 mg/100 g 0.13 Bryza (Poland) Tbr Fala (Poland) Tbr 25 25

Iron (n = 177) 0.14–10.4 mg/100 g 1.04 Peluca (Spain) Tbr/T Sipancachi (Bolilvia) Adg 15 3

Zinc (n = 156) 0.22–0.76 mg/100 g 0.41 Roscanela* (Peru) Adg Cara (Spain) Tbr/T 16 15

Copper (n = 27) 0.05–0.15 mg/100 g 0.11 Bonita (Spain) Adg Cara (Spain Tbr/T 15 15

The number of varieties used to calculate the values is indicated by n. (The number of studies is often greater than this value, as where data for the same variety were available

from different studies, the mean value calculated was used.) All data were taken from the references and were expressed in fresh weight for raw potatoes; an asterisk indicates

that the value was calculated from dry weight. The place of origin is indicated in brackets.

References: (1) Jimenez et al. (2009), (2) Tudela et al. (2002), (3) Ritter et al. (2008), (4) Chatterjee et al. (2006), (5) Mullin et al. (1993), (6) Augustin et al. (1978), (7) Toma et al.

(1979), (8) Bacigalupo et al. (2004), (9) Negi and Nath (2002), (10) Finotti et al. (2006), (11) Rodriguez-Saona and Wrolstad (1997), (12) Casanas Rivero et al. (2003a), (13)

Casanas et al. (2002), (14) del Mar Verde Mendez et al. (2004), (15) Casanas Rivero et al. (2003b), (16) Andre et al. (2007a), (17) Andre et al. (2007b), (18) Burgos et al. (2009a),

(19) Burgos et al. (2007), (20) Zhang et al. (1997), (21) Im et al. (2008), (22) Cambie and Ferguson (2003), (23) Delaplace et al. (2008), (24) Lu et al. (2001), (25) Sikora and

Cieslik (1999), (26) Cieslik and Sikora (1998), (27) Chun et al. (2006), (28) Han et al. (2004), (29) Hajslova et al. (2005), (30) Brown et al. (2008), (31) Campos et al. (2006), (32)

Bonierbale et al. (2009), (33) Burgos et al. (2009b), (34) Hale (2003), (35) Breithaupt and Bamedi (2002), (36) Andersen et al. (2002), (37) Donnelly et al. (2001), (38)

Kalogeropoulos et al. (2007) (39) Linnemann et al. (1985), (40) Friedman et al. (2003), (41) Thybo et al. (2000), (42) Hale et al. (2008), (43) Brown (2008), (44) Lewis et al.

(1998), (45) Konschuh et al. (2007), (46) Brown et al. (2003), (47) Konschuh et al. (2005), (48) Brown (2006), (49) Phillippy et al. (2004), (50) Sarquis et al. (2000), (51) Sengul

et al. (2004), (52) Groza et al. (2005).a Solanum species used: S. tuberosum, S. brevidens, S. acaule, and S. commersonii. S. tuberosum L.: Tbr; S. tuberosum subsp. tuberosum: Tbr/T; S. tuberosum subsp. andigena:

Adg; S. chaucha: Cha; S. phureja subsp. phureja: Phu; S. stenotomum ssp. stenotomum: Stn/S; S. stenotomum ssp. goniocalyx: Gon/G.b Ref: reference.

B. Burlingame et al. / Journal of Food Composition and Analysis 22 (2009) 494–502 497

antagonistic effects between all these components. They alsoreport that the high stability of the ranking of cultivars acrossenvironment in terms of phenolic content and antioxidant capacityindicates that the native Andean cultivars could be confidentlyused within breeding programs aiming at improving the health-promoting value of potato.

The phenolic content and antioxidant capacity of Philippinepotato (S. tuberosum) tubers is presented by Rumbaoa et al. (2009)who analyzed four potato varieties for total phenolic content andantioxidant activities to provide baseline data for Philippine potatovarieties. The phenotypic, agronomic and nutritional character-istics of seven varieties of Andean potatoes are presented byJimenez et al. (2009).

How nutrients in potato are affected by various cooking andprocessing techniques is also addressed. Nisha et al. (2009) look atthe kinetics of niacin degradation in potato (S. tuberosum L.) over atemperature range of 50–120 8C. The degradation kinetics of niacinin normal open pan cooking, pressure-cooking and a newlydeveloped and patented fuel-efficient ‘‘EcoCooker’’ are presented,with the results indicating that the niacin degradation is of asimilar magnitude irrespective of the cooking method.

A somewhat different (although still related to human health)characteristic of potatoes is addressed by Thompson et al. (2009),who look at the phytochemical composition of potatoes, and theinhibition of induced breast cancer in rats fed on freeze–driedpotato powder. They report that rats fed on the Mountain Rosecultivar had a 23% reduction in cancer incidence (p = 0.009) and a49% reduction in cancer multiplicity (p = 0.004) with evidence of adose dependent effect on cancer multiplicity.

The effect of agro-techniques on nutrients is discussed byBuono et al. (2009), who look at the tuber quality and nutritionalcomponents of ‘‘early’’ potato subjected to chemical haulmdesiccation, a technique which is being increasingly used as ameans of simplifying harvesting and helping to manipulate tubersize distribution and tuber skin-set. Their results indicate thatchemical desiccation does not affect the levels of Vitamin C and ofother molecules with antioxidant properties as well as those oftotal soluble carbohydrates, although it does induce a significantdecrease in starch storage and an increase in the percentage ofresistant starch.

Toxicological issues are dealt with in several articles in thisissue. Food processing induced acrylamide formation in Frenchfries and potato crisps are looked at in two articles by Romani et al.(2009) and Serpen and Gokmen (2009), respectively. However, thelatter article also looks at the positive aspects of frying, by studyingthe antioxidants formed during this process. A fast, simple andaccurate method to evaluate potato chip quality using infraredspectroscopy is developed by Shiroma and Rodriguez-Saona(2009). Majcher and Jelen (2009) present a comparison ofsuitability of SPME, SAFE and SDE methods for isolation of flavourcompounds from extruded potato snacks. Another food safetyrelated issue is addressed in the article by Knuthsen et al. (2009),who make a survey of the distribution and contents of totalglycoalkaloids in potatoes on the Danish market. Their studyincludes many different and relevant potato varieties studiedduring a 6-year period.

Although potatoes are not commonly classified as a high-nitratecontaining vegetable, because of the large amounts consumed,

Table 3Varietal differences in antioxidant composition.

Range Average Variety with

highest

Cultivar (Cv),

Andean cultivar,

wild Solanum

species etc.

Taxonomic

groupa

Variety with lowest

nutrient

Cultivar (Cv),

Andean cultivar,

wild Solanum

species, etc.

Taxonomic

groupa

Notes Refb

(highest)

Refb

(lowest)

Vitamin E

(a-tocopherol)

(n = 24)

0.07–0.52 mg/100 g 0.22 Pulu* (Peru) Andean Cv Adg Wila Huaka Lajra*

(Huancayo, Peru)

Andean Cv Adg Unknown (USA)

Tbr also has

0.07 mg/100 g

(ref 27)

17 17

Ascorbic acid

(n = 157)

2.8–42 mg/100 g 17.1 Chaju (Korea) Cv Tbr Liseta (Spain) Cv Tbr 28 2

Vitamin C (n = 78) 6.5–34 mg/100 g 16.7 Maria Cruz

(Peruvian Andes:

Inyaya Alto, La

Victoria and

Aymara)

native Andean

Cv

Tbr Gon/G Krasa (Czech Republic) Cv Tbr 18 29

Total carotenoid

(n = 541)

ND–2690 mcg/100 g 328 Unknown (USA) Cross between

91E22 and

Yema de

Huevo

Tbr Gogon Magash (Peru) Native

Andean Cv

Adg Phureja group 1

(Comas, Huancayo;

Peru) has next

highest value of

2017 mcg/100 g

(ref 32)

30 31

Violaxanthin

(n = 72)

Tr–960 mcg/100 g 117 Amarilla del

Centro* (Peru)

Andean Cv S. stenotomum Unknown CIP 703566

(Peru)

Andean

landraces

Phu Bintje (Germany)

Tbr has next lowest

value of 3.3 mcg/

100 g (ref 35)

17 33

Antheraxanthin

(n = 48)

N.D.–376 mcg/100 g 105 Chaucha Amarilla

yellow fleshed

(Peru)

Andean

landraces

Phu Pichea Papa* (Peru) Andean Cv Adg 33 17

Lutein (n = 447) 8.2–875 mcg/100 g 228 S. pinnatisectum

275231 TAX 31

(Sturgeon Bay,

Wisconsin, USA)

Wild S. pinnatisectum S. pinnatisectum 186553

HERB 3.1–.7 (Sturgeon

Bay, Wisconsin, USA)

Wild S. pinnatisectum 34 34

Zeaxanthin

(n = 97)

Tr–1290 mcg/100 g 111 Unknown

CIP703566 (Peru)

Andean

landraces

Phu Chaucha tomate (Peru) Andean

landraces

Phu Phureja group 1

(Comas, Huancayo;

Peru) has next

highest value of

1037 mcg/100 g

(ref 32)

33 33

b-Carotene

(n = 408)

0.0–111 mcg/100 g 13.5 S. gourlayi 500049

FFAO 24 (Sturgeon

Bay, Wisconsin,

USA)

Wild S. gourlayi Phureja group 1 (Peru) _ Phu 34 32

Total phenolic

acids (n = 297)

0.2–580 mg/100 g 74.1 MN17922

(Minnesota, USA)

– Tbr Unknown Greek Tbr

(Greece)

_ Tbr Dark Red Norland

(Minnesota, USA) has

next highest value of

460.0 mg/100 g (ref 36).

Atlantic (Youngyang,

Korea) Korean Cv has

next lowest value of

0.49 mg/100 g (ref 21).

36 38

Caffeic acid

(n = 156)

10–157000 mcg/

100 g

67.5 S. pinnatisectum

184774 FFAO 52

(Sturgeon Bay,

Wisconsin, USA)

Wild S. pinnatisectum Atlantic (Springlake,

Texas, USA)

Korean Cv Tbr 34 21

Chlorogenic acid

(n = 156)

0.0–144600 mcg/

100 g

269 Purple Peruvian*

(Berkeley Bowl,

California)

Tbr S. brachistotrichum

(Weslaco and Miami,

USA)

Wild S. brachistotrichum 21 42

B.

Bu

rling

am

eet

al./Jo

urn

al

of

Foo

dC

om

po

sition

an

dA

na

lysis

22

(20

09

)4

94

–5

02

49

8

Total anthocyanins

(n = 115)

0.0–508 mg/100 g 41.3 Urenika (Lincoln,

New Zealand)

Cv Tbr Whitu (Lincoln,

New Zealand)

Cv Tbr 44 44

Antioxidant activity

(hydrophilic)

(n = 488)

43–892 mcg Trolox

equivalent/g

386.6 S. pinnatisectum

PNT bulked

(Sturgeon Bay,

Wisconsin, USA)

Wild S. pinnatisectum S. brachistrotichum

255528TAX42

Wild S. brachistrotichum 34 34

The number of varieties used to calculate the values is indicated by n. (The number of studies is often greater than this value, as where data for the same variety were available from different studies, the mean value calculated was

used.) All data were taken from the references and were expressed in fresh weight for raw potatoes; an asterisk indicates that the value was calculated from dry weight. The place of origin is indicated in brackets. This table shows

ascorbic acid and Vitamin C separately because some studies reported Vitamin C values only, some had the value for Vitamin C taken as the sum of values for ascorbic acid and dehydroascorbic acid (DHA).

References: (1) Jimenez et al. (2009), (2) Tudela et al. (2002), (3) Ritter et al. (2008), (4) Chatterjee et al. (2006), (5) Mullin et al. (1993), (6) Augustin et al. (1978), (7) Toma et al. (1979), (8) Bacigalupo et al. (2004), (9) Negi and Nath

(2002), (10) Finotti et al. (2006), (11) Rodriguez-Saona and Wrolstad (1997), (12) Casanas Rivero et al. (2003a), (13) Casanas et al. (2002), (14) del Mar Verde Mendez et al. (2004), (15) Casanas Rivero et al. (2003b), (16) Andre et al.

(2007a), (17) Andre et al. (2007b), (18) Burgos et al. (2009a), (19) Burgos et al. (2007), (20) Zhang et al. (1997), (21) Im et al. (2008), (22) Cambie and Ferguson (2003), (23) Delaplace et al. (2008), (24) Lu et al. (2001), (25) Sikora and

Cieslik (1999), (26) Cieslik and Sikora (1998), (27) Chun et al. (2006), (28) Han et al. (2004), (29) Hajslova et al. (2005), (30) Brown et al. (2008), (31) Campos et al. (2006), (32) Bonierbale et al. (2009), (33) Burgos et al. (2009b), (34)

Hale (2003), (35) Breithaupt and Bamedi (2002), (36) Andersen et al. (2002), (37) Donnelly et al. (2001), (38) Kalogeropoulos et al. (2007) (39) Linnemann et al. (1985), (40) Friedman et al. (2003), (41) Thybo et al. (2000), (42) Hale

et al. (2008), (43) Brown (2008), (44) Lewis et al. (1998), (45) Konschuh et al. (2007), (46) Brown et al. (2003), (47) Konschuh et al. (2005), (48) Brown (2006), (49) Phillippy et al. (2004), (50) Sarquis et al. (2000), (51) Sengul et al.

(2004), (52) Groza et al. (2005).a Solanum species used: S. tuberosum, S. brevidens, S. acaule, and S. commersonii. S. tuberosum L.: Tbr; S. tuberosum subsp. tuberosum: Tbr/T; S. tuberosum subsp. andigena: Adg; S. chaucha: Cha; S. phureja subsp. phureja: Phu; S.

stenotomum ssp. stenotomum: Stn/S; S. stenotomum ssp. goniocalyx: Gon/G.b Ref: reference.

Table 4Varietal differences in organic acids, phytates, asparagine and anti-nutrient composition.

Range Average Variety with highest value Taxonomic

groupa

Variety with lowest value Taxonomic

groupa

Notes Refb

(highest)

Refb

(lowest)

Oxalic acid (n = 5) 20.4–38.3 mg/100 g 27.8 Colorada Adg Negra (South America) Tbr 13 13

Malic acid (n = 14) 34.3–161 mg/100 g 97.1 Marabel Tbr Kerr’s Pink Tbr/T 10 13

Citric acid (n = 14) 116–656 mg/100 g 389 Sponta Tbr Bonita Adg 10 13

Fumaric acid (n = 5) 2.23–3.89 mg/100 g 3.28 Negra (South America) Tbr Cara Tbr/T 13 13

Asparagine (n = 15) 15.6–458 mg/100 g 245 Frinka Tbr Agata Tbr 10 10

Phytate (n = 8) 0.02–0.047 g/100 g 0.037 Russet Norkota (Aberdeen, Idaho USA) Tbr Yellow Finn (Idaho, USA) Tbr 49 49

a-Solanine (n = 64) 0.001–47.2 mg/100 g 5.63 Wild-unknown (Mexico) S. iopetalum Marfona (Ataturk, Turkey) Tbr Next highest value for a known

variety is for Nortena* (Mexico),

26.92 mg/100 g (ref 49)

50 51

a-Chaconine (n = 66) 0.07–64.7 mg/100 g 8.51 Nortena* (Mexico) Tbr Marfona (Ataturk, Turkey) Tbr 49 51

Total Glycoalkaloid

(n = 75)

0.071–175 mg/100 g 20 Desiree (Aberdeen,

Idaho USA, skin)

Tbr Marfona (Ataturk, Turkey, Flesh) Tbr Highest value in the flesh: 32.5,

for May Queen (Tbr, Japan) (ref 40)

8 51

The number of varieties used to calculate the values is indicated by n. (The number of studies is often greater than this value, as where data for the same variety were available from different studies, the mean value calculated was

used.) All data were taken from the references and were expressed in fresh weight for raw potatoes; an asterisk indicates that the value was calculated from dry weight. The place of origin is indicated in brackets.

References: (1) Jimenez et al. (2009), (2) Tudela et al. (2002), (3) Ritter et al. (2008), (4) Chatterjee et al. (2006), (5) Mullin et al. (1993), (6) Augustin et al. (1978), (7) Toma et al. (1979), (8) Bacigalupo et al. (2004), (9) Negi and Nath

(2002), (10) Finotti et al. (2006), (11) Rodriguez-Saona and Wrolstad (1997), (12) Casanas Rivero et al. (2003a), (13) Casanas et al. (2002), (14) del Mar Verde Mendez et al. (2004), (15) Casanas Rivero et al. (2003b), (16) Andre et al.

(2007a), (17) Andre et al. (2007b), (18) Burgos et al. (2009a), (19) Burgos et al. (2007), (20) Zhang et al. (1997), (21) Im et al. (2008), (22) Cambie and Ferguson (2003), (23) Delaplace et al. (2008), (24) Lu et al. (2001), (25) Sikora and

Cieslik (1999), (26) Cieslik and Sikora (1998), (27) Chun et al. (2006), (28) Han et al. (2004), (29) Hajslova et al. (2005), (30) Brown et al. (2008), (31) Campos et al. (2006), (32) Bonierbale et al. (2009), (33) Burgos et al. (2009b), (34)

Hale (2003), (35) Breithaupt and Bamedi (2002), (36) Andersen et al. (2002), (37) Donnelly et al. (2001), (38) Kalogeropoulos et al. (2007) (39) Linnemann et al. (1985), (40) Friedman et al. (2003), (41) Thybo et al. (2000), (42) Hale

et al. (2008), (43) Brown (2008), (44) Lewis et al. (1998), (45) Konschuh et al. (2007), (46) Brown et al. (2003), (47) Konschuh et al. (2005), (48) Brown (2006), (49) Phillippy et al. (2004), (50) Sarquis et al. (2000), (51) Sengul et al.

(2004), (52) Groza et al. (2005).a Solanum species used: S. tuberosum, S. brevidens, S. acaule, and S. commersonii. S. tuberosum L.: Tbr; S. tuberosum subsp. tuberosum: Tbr/T; S. tuberosum subsp. andigena: Adg; S. chaucha: Cha; S. phureja subsp. phureja: Phu; S.

stenotomum ssp. stenotomum: Stn/S; S. stenotomum ssp. goniocalyx: Gon/G.b Ref: reference.

B.

Bu

rling

am

eet

al./Jo

urn

al

of

Foo

dC

om

po

sition

an

dA

na

lysis

22

(20

09

)4

94

–5

02

49

9

Fig. 1. Genetic resources and farmers’ varieties of Solanum phureja spp. potatoes. Photo by kind courtesy of Walter Amoros and the International Potato Center, Lima, Peru.

B. Burlingame et al. / Journal of Food Composition and Analysis 22 (2009) 494–502500

they can make a significant contribution to the daily intake ofnitrate in the diet. For this reason, Ierna (2009) investigates theinfluence of potato variety and harvest date on tuber nitratecontent for off-season production in three varieties of potato,Arinda (mid-early), Spunta (mid-late) and Mondial (late).

While potatoes are an important source of carbohydrate, waysof reducing the glycaemic impact of potatoes is an importantresearch area. Monro et al. (2009) look at potato genotypedifferences in nutritionally distinct starch fractions after cooking,and cooking plus storing cool. Their results suggest that theglycaemic impact of some potatoes may be substantially reducedby cool-storing after cooking. Djabali et al. (2009) look at therelationship between potato starch isolation methods and kineticparameters of hydrolysis by free and immobilised a-amylase.

With agricultural research becoming increasingly targetedtowards improving human nutrition, understanding the foodconsumption habits of populations is essential for orientingresearch. This subject is addressed by Rose et al. (2009), whocombine food composition with household expenditure data inorder to understand the role of potatoes in the Peruvian diet.

5. Conclusion

The compositional data for varieties of potatoes shown in thisreview and in the new papers presented in this Special Issue of theJournal of Food Composition and Analysis, illustrate the impor-tance of the potato from a genetic resource perspective. As withmany species of foods, varietal differences are often significant,

B. Burlingame et al. / Journal of Food Composition and Analysis 22 (2009) 494–502 501

making a compelling case for food composition databases andother sources of nutrient data, to be conscientious in developingsampling strategies, in documenting the food sample, and inreporting compositional data at the level of the variety. Thisapproach was adopted by the International Rice Commission (IRC),as follows (FAO, 2002):

� Existing biodiversity of rice varieties and their nutritionalcomposition need to be explored before engaging in transgenics.� Nutrient content needs to be among the criteria in cultivar

promotion.� Cultivar-specific nutrient analysis and data dissemination should

be systematically undertaken.

These endorsements by the IRC represent a useful model that can beused for all food crops, including potatoes. When nutritionallysignificant differences within the species are recognized, then it isclear that the potato is actually many different hidden treasures.

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