Healthy Aspects of Potatoes As Part of The Human

Potato Research (2008) 51:239258
DOI 10.1007/s11540-008-9111-4
Healthy Aspects of Potatoes as Part of the

Human Diet
N. U. Haase
Received: 16 July 2008 / Accepted: 19 August 2008 /

Published online: 4 November 2008
# EAPR 2008
Abstract Potatoes are constituents of many diets. Nutritionalists identify several

positive aspects but also discuss some adverse reactions. Therefore, healthiness of potato
food has to be established taking into account new knowledge about natural constituents
and food-borne substances. This paper presents data of three main areas: carbohydrates,
toxins and antioxidants. The glycaemic behaviour of prepared potatoes has discredited
the general understanding of potatoes as a healthy foodstuff. Boiled or steamed potatoes
contain a large amount of rapidly available starch, but alongside genotype driven
variability some preparation steps may also have an influence. The glycaemic load as the
most relevant criterion for healthy subjects is relatively low. Potatoes may contain
toxins, either natural (e.g. glycoalkaloids) or food-borne toxins (e.g. acrylamide).
Minimization strategies have been developed for several potato dishes to reduce the
intake substantially. Consumer handling particularly determines the specific level with
those toxins. Antioxidants are a potent source of health promoting reactions in humans.
They are present in potatoes, but specific concentrations are related to several aspects,
e.g. plant growth, time interval after lifting, genotype, and kind of preparation. Again,
the way in which consumers handle the potatoes is relevant. In summary, potatoes are
very well suited for our modern diet, but consumers need advice to ensure that they are
stored and prepared in the most appropriate way.
Keywords Acrylamide . Antioxidants . Glycaemic index . Glycoalkaloids . Potato .
Vitamin C
Introduction
The human diet has reached the focus of popular debate. Malnutrition exists in many
regions of the world but, particularly in developed countries, we observe an
N. U. Haase (*)
Max Rubner-Institute, Schuetzenberg 12, 32756 Detmold, Germany
e-mail: potato.detmold@mri.bund.de
240
increasing rate of super nutrition, so that more and more people are suffering from
obesity. As a result, several food categories have come under discussion, even though
a food itself cannot be healthy or unhealthy. In many cases our eating behaviour is not
rational and officially available guidelines are not followed. Potatoes are also included
in this behaviour. Therefore, a comparison of healthy and unhealthy aspects in this
paper will provide arguments to avoid any misinterpretation.
Potatoes are available all around the world, however, specific acceptance is
different between countries and population subgroups (e.g. social classes and age
groups). Next to freshly boiled or steamed potatoes, consumers request increasing
amounts of potato products.
Updated evaluations of the different food categories by nutritionists have resulted
in a downgrading of potatoes and potato products (Willett 2001). Nevertheless,
potatoes have a nutritionally favourable composition with respect to several
nutrients.
An analysis of the composition clearly identifies that boiled potatoes are a good
source of protein and carbohydrates, but because of the high water content, the
nutrient density is low. Minerals and vitamins are valuable constituents. For instance
a typical meal with 200 g of boiled potatoes (at least three medium sized tubers) will
contribute to our daily diet
&
&
&
&
&
&
&
&
6% of energy (Guideline Daily Amount (GDA): 9,629 kJ),

0.3% of fat (GDA: 70 g),
11% of carbohydrates (GDA: 270 g),
9% of protein (GDA: 50 g),
11% of dietary fibres (GDA: 25 g),
28% of potassium (GDA: 3 g),
47% of vitamin C (Recommended Dietary Allowance (RDA): 60 mg),
8% of vitamin B2 (RDA: 1.2 mg).
Furthermore, several other constituents, e.g. secondary plant metabolites, are

relevant.
Processed potato foods like French fries are also accepted worldwide, but
composition tends to be less healthy, as for several snacks such as potato crisps. This
snack food offers a lifestyle feeling, especially by its special taste, but of course it is
not a basic food.
Recently, the newly obtained knowledge of food-borne toxicants and food
contaminants has extended the discussion about healthy nutrition. Last but not least,
rational aspects are always accompanied by social and mental aspects, which make it
more complicated to arrive at the right choice of a so called healthy diet. The
consumers will even enjoy their food!
Raw potatoes contain protease inhibitors, which are active against trypsin. These
inhibitors interfere with the digestion of proteins. Using raw potatoes or a potato
protein concentrate as food or feed will lead to a decreased protein efficiency.
Trypsin inhibitor (TI) concentration can be up to 174 g TI kg1 protein (Baker et al.
1982), but heating destroys the TI. Therefore, this topic will not be discussed here.
Potatoes also contain multiple heat-labile proteins which can induce immediate
hypersensitivity reactions (Jeannet-Peter et al. 1999). Peeling of raw potatoes may
cause allergic symptoms in adults, such as contact dermatitis, sneezing and
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wheezing. For children, potatoes as food may cause various allergic reactions.
Patatin, the main storage protein in potatoes with a high nutritional value, induces
such a reaction in sensitive children (Seppl et al. 1999), but further studies are
needed to confirm the clinical importance of patatin as a food allergen.
Material and Methods

General
Dry matter (DM) content of boiled and lyophilised samples and of potato products
was determined by water loss in an oven at 105 C until constant weight (American
Association of Cereal Chemists (AACC) 1993a). Dry matter content of raw potatoes
was determined by drying on quartz sand (American Association of Cereal Chemists
(AACC) 1993b).
The Glycaemic Approach
Potato samples of the 2005 harvest with a cooking behaviour ranking from firm and
fairly firm to floury (German classification system) were either bought from a local
retailer (Cilena, Ditta, Princess) or collected from a field trial for cultivar testing
(Agria, Albatros, Bintje, Carmona, Saturna).
Unpeeled or peeled samples of five to six tubers each were boiled in a typical
cooking pot with sufficient water. Some samples were chilled overnight in a
refrigerator at +4 C to enhance starch retrogradation. Microwave heating of
unpeeled tubers from the same samples was carried out in a domestic microwave
oven (Miele, Gtersloh, Germany).
Potato products were processed on semi-technical processing lines with suitable
cultivars (three cultivars each) toward dehydrated potatoes, potato crisps and parfried French fries, which had to be finished before analysis.
Rapidly and slowly available glucose (RAG and SAG, respectively), free glucose
and sucrose released glucose of the prepared samples were analysed in triplicate
according to Englyst et al. (2000) with minor modifications.
Glucose concentration in all samples was determined enzymatically with a
glucose-oxidase-peroxidase-reagent according to Karkalas (1985) (incubation for
75 min at room temperature in the dark, and measuring at 510 nm in a
spectrophotometer (Specord 200, Analytik Jena, Germany). Calibration with pure
glucose was in the range between 3 and 30 mg kg1.
Total glucose content was determined separately. Polysaccharides were split to
glucose by heat, lye, and enzymatic degradation.
Resistant starch (RS) content was calculated according to Englyst et al. (2000)
(RS=0.9(TG(RAG+SAG)) and is defined as non-digested starch in the human
small intestine.
Glycaemic index (GI) was calculated according to a statistical regression formula
created from data of Englyst et al. (1996a). Glycaemic load (GL) was calculated for
a serving size of 150 g potatoes or potato products (French fries, rehydrated
potatoes). The crisps serving size was 50 g.
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Toxins
Glycoalkaloids were analyzed by an HPLC procedure. Lyophilized and ground
samples were homogenized with a mixture of water/acetic acid/sodium bisulphite,
95:5:0.5 (v/v/w) (Hellens et al. 1995b). Extraction was with SPE cartridges (SepPak
C18 (Waters Associates, Milford, MA, USA)) after conditioning with acetonitrile.
Glycoalkaloids were eluted with the HPLC mobile phase (acetonitrile/0.022 mol m3
potassium phosphate buffer (pH=7.6), 55:45 v/v). The HPLC separation was carried
out by a ODS Hypersil 5 m phase (C18) in a 2504.6 mm column. Signals were
detected at 202 nm (Kontron Instruments, UV VIS) by a flow rate of 25109 m3 s1
(Kontron Instruments, HPLC pump model 420).
Acrylamide concentration of potato crisps was determined by a GCMS
method in the EI mode. To extract acrylamide from the food material, 50
106 m3 of distilled water were added to 4.00 g of ground sample by ultrasonic at
60 C. After centrifugation (44,145 m s1, 600 s) the supernatant solution was
defatted by petroleum ether and clarified by 5106 m3 Carrez I and II solution,
each. Acrylamide was salted out with sodium chloride and extracted from the
aqueous phase by threefold extraction with 30106 m3 ethyl acetate, each. These
organic phases were filtrated by water-repellent filters (MN 616 WA 1/4
(Macherey-Nagel, Germany)) to separate water and then evaporated to about
0.5106 m3 before using for GCMS. GCMS analysis was carried out using the
electron ionization mode (EI, 70 eV) on a Hewlett Packard Instrument Model 5890
Series II/5989 A. For the determination of acrylamide and D3-acrylamide the SIMtechnique (selected ion monitoring) was used and the identification was achieved
by the ions with masses (m/z) 74, 71, 58 and 55. The quantification was carried out
by ions with masses 74 and 71. The separation was achieved with a DB-23
capillary column (J&W Scientific Products, Germany) (300.25103 m i.d.,
0.25106 m film thickness). The carrier gas was helium at a flow rate of 16.7
109 m3 s1.
The column temperature was initially kept at 80 C for 120 s and then increased
from 80 C to 220 C at 0.17 C s1. The final temperature was held for 60 s. Other
operating conditions were a split/splitless injector (splitless, temperature 240 C), an
interface temperature of 250 C, and an ion source temperature of 200 C. For
quantification deuterated acrylamide was used as internal standard.
Ascorbic Acid
Ascorbic acid (L-AA and DHA) was analyzed by means of a colorimetric method
according to Roche Diagnostics, Mannheim, Germany: L-AA reduces the
tetrazolium salt MTT (3-(4,5-dimethylthiazolyl-2)-2,5-diphenyltetrazolium bromide)
in the presence of the electron carrier PMS (5-methylphenazinium methosulfate) at
pH 3.5 to a formazan (MTT-formazan), which is determined by means of its light
absorbance at 578 nm (Specord 200, Analytik Jena, Germany). After reduction of
DHA to L-AA in the presence of dithiothreite (DTT), the difference between totalAA and L-AA content is DHA.
Because of different dry matter concentrations of individual samples, all values
are expressed on a dry matter basis.
243
Antioxidant Capacity
The antioxidant capacity was estimated with the FRAP-procedure according to
Benzie and Strain (1996). After a lengthened extraction procedure (7,200 s at 37 C)
with respect to phenolic compounds of the potato (Pulido et al. 2000), samples were
filtered (folded paper filter), and an aliquot was stained: 90109 m3 filtrate and
270109 m3 aqua bidest. were combined with 2.7106 m3 of the FRAP reagent
(50106 m3 of 100 mol m3 Na-acetate-buffer+5106 m3 10 mmol TPTZ (2,4,6tripyridyl-s-triazine)+5106 m3 20 mmol FeCl3 6H2O) and measured at 593 nm
(Specord 200, Analytik Jena, Germany). Trolox was used for calibration.
Results and Discussion

Carbohydrates
Available potato carbohydrates are present as starch or as free sugars, whereas nonavailable carbohydrates are considered as dietary fibres. Starch is the main source
with a typical concentration range between 8 and 29% in fresh weight (Lisinska and
Leszczynski 1989). Genotype and environment may have a strong influence upon
specific levels. Potatoes with a low to medium content of starch are often preferred
for fresh consumption (firm to fairly firm cooking type), whereas higher starch levels
are requested by the food industry. Raw potato starch has a high amount of nondigestible units, on the other hand gelatinized starch is easily available. Resistant
starch (RS) is not available for human digestion, but fermentation towards short
chain fatty acids (SCFA) takes place. The RS level in potato food is generally low,
but may increase throughout a longer stay after preparation or by storage of precooked potatoes (e.g., potato salad, par-fried French fries (Goni et al. 1997)). Raw
potato starch has a very high level of RSbut only few people are going to eat raw
potatoes. Some sugars in the potato may be involved in heat-induced reactions (e.g.,
Maillard reaction) resulting in adverse effects (see below). Furthermore, the debate
on low-carb food has enveloped potatoes, too. Principally, the absolute
carbohydrate concentration in potatoes is low, but a range is still given. If consumers
want to select low-carb potatoes, they may choose firm cooking tubers.
Nutritionists have downgraded the relevance of potatoes within our diets
especially in view of the glycaemic behaviour, because all the available
carbohydrates are relatively rapidly available and can be absorbed as monosaccharides and metabolized by the body (Ludwig 2000; Willett 2001). As a consequence
the postprandial blood glucose response (Englyst et al. 1996b) is elicited. The
resulting glycaemic index (GI) of boiled potatoes, calculated as the relationship
between food-borne blood sugar and reference index (glucose or white bread), ranks
between 20 and 100, indicating potatoes as a food with medium to high GI. The
overall insulin mediated glucose uptake is based upon the glucose level in
consequence of the carbohydrate concentration in a food. Therefore, the term
glycaemic load (GL) combines GI and the concentration of carbohydrates. The
glycaemic load of potatoes is low or medium. Several nutrition guidelines were
developed within the last few years reflecting the GI or GL of a single food (e.g. a
244
low glycaemic index pyramidLOGI (Ludwig 2000) and a healthy eating pyramid
(Willett 2001)). Potatoes appear near the top of those pyramids and the guidelines
suggest their limited consumption, but the reason for that positioning is based upon
U.S. preferences of convenience food consumption (esp. French fries, baked
potatoes, potato mash) (Buyken and Kroke 2006). The GI measurement itself
implicates several points of criticisms (Brouns et al. 2005). For instance, the
availability of GI-data does not reflect the high variance of carbohydrates in
potatoes. GI is not a stable feature of a food, but is considerably modified by its
processing, composition, preparation, the composition of a mixed food and many
other factors which renders it difficult to precisely predict the actual blood glucose
increase in an individual situation (Hauner 2006). GI of potatoes may be changed
dramatically by parallel consumption of proteins and fats (Gulliford et al. 1989).
Also a high intra-individual variability exists in response to a carbohydrate load
(Jenkins 2007). Finally, a relationship between GI, GL and cancer risk could not be
validated until now (Michaud et al. 2005; WHO/FAO 2003).
Soh and Brand-Miller (1999) have described the influence of cultivar and
preparation type to the glycaemic index of potatoes (Table 1). GI in this overview
was between 65 and 101, but only the difference between lowest and highest value
reached a statistical significance. Another study by Fernandes et al. (2005) has
expanded the experiment toward the time interval between preparation and
consumption (Table 2), pointing out a GI between 56 (boiled cold potatoes) and
89 (boiled hot potatoes). This principal range of GI data from potatoes reflects a
collection of international data (Foster-Powell et al. 2002).
To overcome the time and money consuming in vivo measurements of the GI, an
in vitro technique was developed. Englyst et al. (1996b) have suggested measuring
the rapidly and slowly available glucose (RAG and SAG, respectively). BrandMiller (2007) states, that the GI of a food cannot be guessed, it must be tested
according to the standardized in vivo method. Nevertheless, Englyst et al. (2003)
have described, that GI values of cereal products can be explained by the RAG and
SAG contents. A high SAG content identifies low-GI foods which are rich in slowly
released carbohydrates for which health benefits have been proposed.
Table 1 GI values of test foods (n=9 or 10)
Food
Cultivar
Dsire, peeled and boiled
Pontiac, peeled and boiled
Sebago, peeled and boiled
Cooking method
Pontiac, peeled and boiled
Pontiac, peeled, boiled and mashed
Pontiac, peeled and microwaved
Pontiac, peeled and baked
Maturity
New, unpeeled and boiled
New, canned and microwave heated
Modified from Soh and Brand-Miller (1999)
GIs.d.
10115
889
877
889
919
799
9311
7812
659
245
Table 2 GI values for 50 g available carbohydrate portions of seven potatoes tested by a cohort of 12
subjects
Food
GIs.d.
Baked Russet potato

Instant mashed potato
Roasted California white potato
Baked Prince Edward Island white potato
Boiled red potato (hot)
Boiled red potato (cold)
French fried potatoes
76.58.7xyz
87.78.0xy
72.38.2xyz
72.84.5xyz
89.47.2x
56.25.3z
63.65.5yz
Modified from Fernandes et al. (2005).

significantly (P<0.05)
xyz
means not sharing the same letter superscript differ
The fate of dietary carbohydrates in eight potato cultivars was determined in vitro
according to the protocol of Englyst et al. (2000) by its rapidly available (RAG),
slowly available (SAG), and total glucose (TG) content. Different preparation and
processing techniques were included for three cultivars each. The relationship
between RAG and TG varied within 55 and 78% irrespective of the individual
cooking behaviour. The preparation techniques indicated a cultivar-specific
behaviour. Peeling enhanced the relative RAG in two of three cultivars. Chilling
after boiling reduced the glucose release in one cultivar only. Microwave heating had
a positive effect upon SAG, but RAG was also enhanced in two of three cultivars.
Dehydrated potatoes (DP), French fries (FF), and potato crisps (PC) were processed
from three suitable cultivars each. Differences between cultivars were negligible.
RAG and SAG were related to TG for a better comparability between the products.
A concurrent comparison of boiled and processed tubers from identical cultivars of
the same lot identified specific changes during processing (Fig. 1).
Processing of dehydrated potatoes and of crisps resulted in a faster RAGavailability compared with the alternately boiled samples. RAG of French fries was
definitely reduced, indicating a shift in physical behaviour.
Fig. 1 Rapidly available glucose (RAG) in boiled and processed tubers of three cultivars; relative data:
means, and standard deviation (DP dehydrated potatoes, FF French fries, PC potato crisps, *_B boiled
samples)
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Total glucose content (TG) was positively correlated with the glycaemic index
(GI, calculated) (R2 =0.58) in processed samples. All firm and fairly firm cooking
cultivars had a high GI, whereas one of the floury cooking cultivars had the lowest
GI (Saturna) and the other one the highest GI of all cultivars (Albatros) (Table 3).
Different preparation types indicated cultivar dependent reactions. Cilena had a
lower GI after boiling and chilling (application C), whereas Bintje had a lower GI
after microwave heating.
Dehydration and crisping showed stable or increased GI compared with boiling
(variant A), while French fries had a lower GI.
The range of GL was absolutely between 17 and 27 with one single exception (35
for Albatros) (Table 3). GL data of the products were not comparable with the
corresponding GL of boiled samples. Both, higher and lower values were found for
French fries and potato crisps, while dehydrated potatoes showed a trend for slightly
increased values.
Dehydration and crisps production resulted in an increase of RAG, whereas in
French fries a significant decrease of RAG and, simultaneously, a significant
increase of SAG were present. RS was different between cultivars, but not between
preparation and processing techniques. Calculated data of the glycaemic index and
glycaemic load reflected comparable trends shown with temporal glucose release.
The investigated set of potato cultivars clearly indicates the need of specific
considerations to prevent inaccuracies in rating of potato.
Toxins (Natural vs. Food-borne)
Glycoalkaloids
Potato tubers contain several glycoalkaloids, especially -solanine and -chaconine.
They belong to the class of solanidine tri-glycosides and account for more than 95%
of total glycoalkaloids (TGA) in potatoes. They are haemolitically active like
saponin and highly toxic to humans (Jadhav 1983). Symptoms involve gastrointes-
Table 3 Glycaemic index (GI) and glycaemic load (GL) of eight potato cultivars, calculated according to
Englyst et al. (2003)
Cultivar
Marabel
Cilena
Ditta
Agria
Bintje
Carmona
Saturna
Albatros
GI (preparation/processing)
GL (preparation/processing)
74
78
77
79
78
70
65
84
86
88
71
81
69
74
76
80
65
87
79
83
64
61
63
89
83
76
17
23
23
20
18
21
24
35
20
20
20
18
17
21
18
20
21
27
23
21
23
26
26
22
22
21
Cells without values: no analysis. Serving size: 150 g, dehydrated potatoes after rehydration; crisps: 50 g
(applications: A boiled with skin; B boiled without skin; C boiled without skin and chilled over night at 4
C; D microwave heating with skin; processing E dehydration; F French fries; G potato crisps)
247
tinal disturbances by membrane damages (Roddick 1987) and neurological disorders

by inhibition of the enzyme acetylcholinesterase (Bushway 1987). An oral dose of
12 mg kg1 body weight initiates a poisoning while a dose of 36 mg kg1 body
weight may be lethal (Morris and Lee 1984). Serious glycoalkaloid poisoning of
humans is rare, but it is suspected that mild poisoning is more prevalent than
supposed. It seems to be rarely diagnosed or treated, because the symptoms are
similar to those of other gastrointestinal disorders (Smith et al. 1996). Otherwise
anti-carcinogenic effects of glycoalkaloids against human cervical, liver, lymphoma,
and stomach cancer cells will get pharmacological interest (Friedman et al. 2005).
TGA play an important role by natural defence strategies of plants (Roddick 1988).
The level within potato plants is not equal. High levels are reported in leaves (up to
1,000 mg kg1 Fresh Weight (FW)) and sprouts (up to 7,300 mg kg1 FW). In whole
tubers TGA concentrations between 20 and 350 mg kg1 FW are reported (cited by
Maga 1980). Levels above 200 mg kg1 FW may be a potential health risk, which
was proposed as early as 1924 (Bmer and Mattis 1924). Mechanical impacts and
weather are relevant factors for increased values (Cepl and Zrust 1996). In Sweden,
the cultivar Magnum Bonum had an extreme TGA enrichment during a cold and
rainy period before lifting (Hellens et al. 1995a). Maturity stage at lifting, tuber size
and storage conditions have an influence, too (Cieslik and Praznik 1998). High TGA
concentrations in the potato flesh will lead to a bitter taste, a natural barrier against
non-controllable potato consumption. With respect to the uneven distribution of
TGA even within one single tuber, the level within the peel fraction easily reaches a
toxic level, whereas the peeled tuber is still usable (Table 4).
Green or partly green tubers may have increased TGA concentrations because of
a de novo synthesis of TGA, but increase may be different between cultivars (Fig. 2)
(Haase 1999).
With respect to different tuber sizes, a correction formula between size and mass
was proposed by Engel et al. (1998): ln TGA=ln a+(bln m).
Dale et al. (1998) have found an increase of TGA-values between 27 and 130% in
case of blackspot bruise. Also mechanical damage and light stress increase TGAlevel (Olsson 1996).
TGA may affect the flavour of boiled potatoes at higher concentrations. Sinden et
al. (1976) have described a lower critical level of 140 mg TGA kg1 FW following
an astringent and bitter taste.
With respect to several unsolved questions in combination with factors of TGAbiochemistry and toxicity, it was proposed to reduce the acceptable level in fresh
tubers down toward 6070 mg kg1 FW (Parnell et al. 1984).
Table 4 Glycoalkaloid distribution within potato tubers (Haase 1999)

Tuber section
-Solanine
(mg kg1 FW)
-Chaconine
(mg kg1 FW)
Total glycoalkaloids
(mg kg1 FM)
Relationship Sol.:
Chac.
Skin
Peel
Kernel
1,285
795
15
1,764
1,378
12
3,049
2,173
27
1:1.4
1:1.7
1:0.8
248
Fig. 2 Glycoalkaloid content of potato lots from seven cultivars, separated into faultless (f) and green
tubers (g)
Heat Induced Toxicants

In the last few years, acrylamide formation has been intensively investigated with
regard of findings in several food categories, including fried and baked potato
products. It is mainly formed in food by the reaction of asparagine with reducing
sugars as part of the Maillard reaction (Stadler et al. 2004; Mottram et al. 2002;
Weisshaar and Gutsche 2002; Becalski et al. 2003). In the early Maillard pathway, the
glycoconjugate of asparagine and sugar undergoes a Strecker-type degradation leading
to azomethine ylides, that affords acrylamide after a -elimination reaction of the
decarboxylated Amadori compound (Yaylayan et al. 2003). Furthermore, a -proton
in the formation of the corresponding vinylogous compounds is important (Stadler et
al. 2004). Other precursors have also been proposed, such as 3-aminopropionamide (3APA) (Zyzak et al. 2003). More detailed studies demonstrated that 3-APA is a key
transient intermediate, i.e. furnishes acrylamide in a thermally-driven deamination
reaction in very high yield (Granvogl and Schieberle 2007). It was also shown that 3APA can be generated in an enzyme-catalysed reaction directly through decarboxylation of asparagine (Granvogl et al. 2004).
Efficiency of the acrylamide reaction is low (Stadler et al. 2004) and
concentration inside a food represents a snapshot of parallel synthesis and
degradation (Biedermann et al. 2002a). Trials with different food matrices have
shown that further specific factors are involved, like recipe components, pH-value of
food, fermentation products, and pre-treatments (washing, blanching, enzymatic
degradation) (Amrein et al. 2003; Becalski et al. 2003; Biedermann et al. 2002a, b).
Acrylamide is under suspicion to be mutagenic via its metabolic glycidamid. Also
neurotoxic and reproductive genotoxic properties are under discussion (Dybing
2007). The lifetime risk of cancer based on acrylamide was estimated by animal
experiments with 1.3103 to 5.2103 (average and high consumers, respectively)
(corresponds to 13 to 52 cancer diseases per 10,000 individuals) (Dybing and Sanner
2003). Furthermore, a positive association was seen between acrylamide-hemoglobin
249
levels and estrogen receptor positive to breast cancer (Olesen et al. 2008), but other
studies did not show this (Mucci et al. 2003; Pelucchi et al. 2006). Acrylamide
uptake is between 0.5 and 1 g acrylamide kg1 body weight and day1 for average
consumers, but high consumers may take up till 48 g kg1 body weight and day1
(Dybing 2007). The margin of exposure (MOE) as a ratio between benchmark dose
lower confidence limit (BMDL) and estimated intake in humans was calculated
between 300 and 75 (average and high consumers, resp.), using a BMDL of 0.3 mg
kg1 body weight and day1 for induction of mammary tumours in rats (FAO/WHO
2005). The FAO/WHO committee of experts on food additives (JECFA) considered
these MOEs to be low for a compound that is genotoxic and carcinogenic and that
they may indicate a human health concern. Therefore, despite of contradictory data
from epidemiological studies (U.S. Environmental Protection Agency 1999; Bader et
al. 2005; Calleman 1996; Pelucchi et al. 2006), the ALARA principle (as low as
reasonably achievable) is worthwhile being applied.
Acrylamide containing animal feeding-stuff may enhance the total exposure, but
first experiments with chicken have shown a low carry-over rate (Halle et al. 2006).
Smokers have a high acrylamide intake from tobacco, which may over-ride any
additional effects from different eating habits (Bader et al. 2005).
Different tools have been identified to lower the acrylamide formation in heated
potato food, including selection of potato cultivar, potato storage regime, process
control (thermal input, pre-processing), final preparation, and colour control (Haase
et al. 2004). Most of them have been collected in a toolbox system (Confederation of
the food and drink industries of the EU (CIAA) 2006; European Commission DG
Health and Consumer Protection 2007). A strong correlation was found between
acrylamide formation and the concentration of reducing sugars in the potatoes. It
was further improved by including the concentration of asparagine (Amrein et al.
2003). In fact, asparagine concentrations in fresh potatoes are typically between
2,000 and 4,000 mg kg1. Concentrations of reducing sugars range between 100 and
3,000 mg kg1 after harvesting and may reach 20,000 mg kg1 after currently used
storage at low temperature (Amrein et al. 2003; Haase et al. 2004; Haase 2006).
Traditionally, potato processors advise plant breeders to develop cultivars with low
sugar content, and they have been encouraged to check potatoes either by fry test or
sugar determination before accepting deliveries.
To minimise losses from spoilage and shrinkage, the potatoes are stored at low
temperatures. However, low temperatures tend to increase the sugar levels,
particularly when stored below 6 C, through a process known as cold induced
sweetening (CIS) (Sowokinos and Preston 1988), resulting in dark flavoured and
bitter tasting fried products. For potatoes to be held in storage for the longest period
prior to processing, the minimum storage temperature used by producers of French
fries or potato crisps is generally 6 C.
Storage at higher temperature may result in breaking the dormancy of the tubers.
Sprouting can be controlled through the use of suppressants, such as chloropropham
(CIPC). For organically grown potatoes, compatible growth inhibitors are used.
Reconditioning by warming the tubers to some higher temperatures (e.g. 12 C)
for a few weeks reduces the sugar levels, but does not restore initial levels.
Slicing or cutting potatoes to a specific surface to volume ratio is particularly
important, because acrylamide formation typically occurs on the surface layer of the
250
products. Coarse-cut French fries strips (1414 mm) with a surface to volume ratio of
3.3 cm1 resulted in significant lower (P<0.05) amounts of acrylamide than fine-cut
strips (88 mm) (Matthus et al. 2004). The same is valid for sliced products (crisps),
available from 1.2 mm to 1.75 mm. It must, however, be pointed out that the
organoleptic properties may change, since larger pieces are cooked for a longer period.
The removal of fine cuts from the outer sphere of the tubers contributes to
acrylamide reduction, because they tend to be overheated and the concentration of
reducing sugars in the peripheral region of the tubers is higher.
Soaking or blanching potato slices or sticks prior to frying can reduce acrylamide
formation in the final product by removing the precursors (Grob et al. 2003). For
producers of French fries, the blanching process is one of the major steps in the
production. It is important to remove reducing sugars from the surface of the potato
sticks and to inactivate enzymes, particularly amylase. During the final stage of
blanching, the pH of the blanching water is lowered by the addition of sodium
pyrophosphate, which can also contribute to the reduction of acrylamide formation.
In the case of potato crisps, blanching (1) impacts the texture and flavour of the
fried product (Confederation of the food and drink industries of the EU (CIAA)
2006), (2) increases the oil content (Pedreschi and Moyano 2005), and (3) leaches
out vitamin C.
Using lower thermal input into the frying or roasting process by decreasing the
cooking temperatures and/or cooking time usually reduces acrylamide levels (Grob
et al. 2003).
A slight increase of the remaining moisture of potato crisps may reduce
acrylamide without loss of crispiness, but a trial under standardized conditions
showed no correlation with the acrylamide concentration (Haase 2007). Acrylamide
formation can, furthermore, be minimized by careful selection of temperature, in
particular by a decrease towards the end of the frying process.
It is impossible to produce potato crisps without some degree of colour variation,
because natural differences in the sugar levels of individual tubers or the flow
characteristics in the fryer can result in the production of darker crisps with higher
levels of acrylamide.
Frying under reduced pressure (7.9103 kg m1 s2) in a vacuum fryer enables
lower frying temperatures to be used (approx. 130 C). Crisps with 97% lower
acrylamide content were obtained (Granda et al. 2004), but also a modified final
frying step may reduce the acrylamide content for about 90% (Table 5) (Haase and
Lindhauer 2005).
Table 5 Effect of vacuum frying upon potato crisps quality
Atmospheric
Vacuum
Product
Temp.
[C]
Res. moisture
[%]
Pressure
[kg m1 s2]
Temp.
[C]
Res. moisture
[%]
Fat
[%]
Colour
[L-value]
Acrylamide
[ kg1]
160
160
160
160
160
2.5
9
4.5
4.5
9
2104
1104
1104 5104
1104
100
90
90
100
2.5
2.7
3.2
4.9
3.3
36
43.8
44.9
47.1
41.7
67
64
66
63
66
2,140
610
486
468
279
251
The acrylamide concentration in par-fried French fries and other potato products
produced for final cooking at home or in a restaurant, are typically very low, because
acrylamide is formed primarily during final frying with a non-linear rate of
formation with increased temperature (Grob et al. 2003). At lower frying temperatures and longer frying times, acrylamide concentrations were reduced, but there is
a lower temperature limit below which quality is affected, such as by higher fat
uptake and poorer texture (Taeymans et al. 2004).
Latest data have shown that bound 3-monochloropropane-1,2-diol (3-MCPD
esters) is present in some fried potato products. The potato itself is not the problem
rather the refined frying oil. At the moment the amount of release during human
digestion is still unclear.
Acrylamide has become a signal effect, but the highly variable reaction pathways
during heating of food should be considered. Therefore, we can expect to identify a
number of other heat induced chemicals with potentially adverse health effects.
Antioxidants (Vitamin C)
Vitamins of the potato are well known to contribute substantial amounts of the
recommended daily intake. Vitamin C, which is present as L-ascorbic acid (L-AA)
and as mono-dehydro ascorbic acid (MDHA), particularly has been studied
intensively (Davey et al. 2000).
The biological function of L-AA is focused on the antioxidant properties. In humans,
both forms, L-AA as well as MDHA, are active. MDHA can be reduced to L-AA. The
plasma level of L-AA in large sections of the population is sub-optimal. Therefore, the LAA level of plant food should be improved or losses may be minimized.
Several growing factors have an influence upon the final vitamin C level. Next to
the right choice of a cultivar, fertilization, and weather conditions have effects, too
(Kolbe 1997; Weber and Putz 1998; Rogozinska et al. 2005).
Following the post-harvest period, the sum of L-AA and MDHA declines,
because some MDHA molecules convert to dehydro ascorbic acid (DHA), which
undergoes decomposition toward 2,3-diketo gulonic acid, which is no longer active.
An experiment with three different cultivars stored at two different temperature
regimes (4 and 8 C) confirmed the results. The first storage section especially
indicated a fast decline, whereas a more stable level was reached after three months.
Slightly warmer stored tubers (8 C) had a slower decline of vitamin C than the 4 C
samples (Fig. 3).
Preparation is best when boiling potatoes with skin on. However, high losses can
occur during potato preparation, because vitamin C is heat sensitive. The warm
keeping of boiled potatoes may reduce the initial level for about 40%. Leaching is
also a reason for a reduced concentration, especially in processing lines. While
potatoes for French fries production had a relatively low reduction, tubers for crisps
or flakes production had a loss of 60% (Fig. 4).
Antioxidant Capacity
Several secondary constituents of the potato have an antioxidant activity, which
contributes to the physiological defence against oxidative and free-radical-mediated
252
Fig. 3 Vitamin C levels in tubers of three cultivars over the storage period, following two strategies (+4
and +8 C storage)
reactions. After vitamin C, several secondary constituents, e.g. flavonoids,

carotenoids, and other phenolic compounds, are involved in these reactions (Chu
et al. 2000; Chaillou and Nazareno 2006). Wang et al. (1996) have shown a limited
contribution (less than 15%) of vitamin C to the total antioxidant activity in several
fruits. Another study has estimated that vitamin C extracted from an unknown potato
sample contributes 13% to the total antioxidant activity (Chu et al. 2002).
Nevertheless, our knowledge about potato antioxidants is still low. At least a diet
with antioxidant rich food will result in a high antioxidant level in the blood (Cao et
al. 1998; Welch et al. 2005), which will reduce the incidence of degenerative
diseases like cancer and atherosclerotic heart disease.
Fig. 4 Remaining vitamin C level (relative values) in a number of potato preparations ((Weber and Putz
1998), modified, and (Haase and Weber 2003)) (FF: French fries)
253
Data from antioxidant potato compounds are also rare. Most of them are available
from raw materials. Flesh colour (e.g. dark yellow, blue or red) in particular has been
reported to induce different levels of antioxidants (Reyes et al. 2005; Gromes and
Herrmann 2008). Coloured cultivars contain higher amounts of anthocyanins and
flavonoids compared to white or yellow flesh tubers (Lewis et al. 1998), whereby the
antioxidant level increases considerably (Lachman and Hamouz 2005). However,
white flesh cultivars may also contain several colourless compounds that are
probably either flavonoids or phenolic acids with a high antioxidant potential (Hale
2003). Phenotype of coloured cultivars does not seem to be stable all the time. Reyes
and Cisneros-Zevallos (2003) have found an influence of the growing location upon
anthocyanin concentration. These authors have also described a significant increase
in case of physiological stress during storage. The concentration of phenolic acids
accounted for relatively little of the total antioxidant activity (R2 =0.18). In a ranking
of different phenolic acids for antioxidant capacity, caffeic acid and chlorogenic acid
were the lowest (Hale 2003). Carotenoids include primarily lutein, zeaxanthin, and
violaxanthin (Lu et al. 2001; Nesterenko and Sink 2003), whereas the amounts of or -carotene are negligible (Gross 1991). Antioxidative behaviour of single
carotenoids is still a matter of research. In comparison with lycopene, both lutein
and zeaxanthin were about as half as effective (Miller et al. 1996; Bhm et al. 2002).
Generally, the preparation of potatoes will lower the antioxidants, because of heat
sensitivity of most substances involved. Long heating (e.g. boiling) reduces more
than short preparations (e.g. microwave heating, frying), but an average remaining
antioxidant capacity of 75% is reported (Blessington et al. 2005). Experiments with
bread crust have shown a de novo synthesis within the Maillard reaction
(Lindenmeier and Hofmann 2004). Consequently, fried potato products may have
enhanced values in relation to boiled tubers, but valid data are still under research. In
comparison with legumes and fruits, potatoes offer a low but substantial contribution
to the daily requirement for antioxidants (Halvorsen et al. 2002). Methodical studies
Fig. 5 Antioxidant capacity in stored potato tubers from three cultivars, stored at +4 C (left) or +8 C
(right)
254
have shown that potato antioxidative capacity is often undervalued with respect to
the slow release of phenolic substances (Pulido et al. 2000).
In contrast to a vitamin C decline (see Fig. 3), the total antioxidant capacity
increased over the storage season, as was found in an experiment (Fig. 5). Lewis et
al. (1999) have described the same phenomenon in coloured cultivars. They
explained it with a de novo synthesis of anthocyanins, which happened especially in
relation to cold sweetening, because sugars are an anthocyanin precursor.
Conclusion
Potato tubers offer several nutritional benefits. Potatoes provide a substantial
contribution to the daily supply of minerals, vitamins, and secondary compounds as
well as carbohydrates and protein. Potatoes have the highest satiety index of all plant
foods. Alongside these positive aspects only few adverse factors are identified,
including the existence of glycoalkaloids and acrylamide, but the right treatment can
reduce these levels substantially. Unfortunately, many consumers have a limited
knowledge about all these facts. Education campaigns could help to promote the
nutritionally positive methods of preparation and to increase the daily potato intake.
References
American Association of Cereal Chemists (AACC) (1993a) Approved methods of the AACC. Method 4415a (Moistureair oven method). AACC, St Paul, MN, The Association, St. Paul, MN
American Association of Cereal Chemists (AACC) (1993b) Approved methods of the AACC: Method
4460 (Moisturedrying on quartz sand). The Association, St. Paul, MN, USA
Amrein TM, Bachmann S, Noti A, Biedermann M, Barbosa MF, Biedermann-Brem S, Grob K, Keiser A,
Realini P, Escher F, Amado R (2003) Potential of acrylamide formation, sugars, and free asparagine in
potatoes: a comparison of cultivars and farming systems. J Agric Food Chem 51:55565560
Bader M, Hecker H, Wrbitzky R (2005) Querschnittsstudie zur ernhrungs- und tabakrauchbedingten
Belastung mit Acrylamid. Dtsch Arztebl 102:A2640A2643
Baker EC, Rackis JJ, Mustakas GC, Strolle EO (1982) Development of a pilot-plant process for the
preparation of a trypsin inhibitor-rich fraction from potatoes. Ind Eng Chem Prod Res Dev 21:8082
Becalski A, Lau BPY, Lewis D, Seaman SW (2003) Acrylamide in foods: occurrence, sources, and
modelling. J Agric Food Chem 51:802808
Benzie IFF, Strain JJ (1996) The ferric reducing ability of plasma (FRAP) as a measure of antioxidant
power: The FRAP assay. Anal Biochem 239:7076
Biedermann M, Biedermann-Brem S, Noti A, Grob K (2002a) Methods for determining the potential of
acrylamide formation and its elimination in raw materials for food preparation, such as potatoes. Mitt
Geb Lebensmittelunters Hyg 93:653667
Biedermann M, Noti A, Biedermann-Brem S, Mozzetti V, Grob K (2002b) Experiments on acrylamide
formation and possibilities to decrease the potential of acrylamide formation in potatoes. Mitt Geb
Lebensmittelunters Hyg 93:668687
Blessington T, Hale AL, Scheuring DC, Miller JC (2005) Effects of cooking, storage and gamma
irradiation on antioxidant activity in potato (Solanum tuberosum L.). Am J Potato Res 82:58
Bhm V, Puspitasari-Nienaber NL, Ferruzzi MG, Schwartz SJ (2002) Trolox equivalent antioxidant
capacity of different geometrical isomers of alpha-carotene, beta-carotene, lycopene, and zeaxanthin. J
Agric Food Chem 50:221226
Bmer A, Mattis H (1924) Der Solaningehalt de Kartoffeln. Z Unters Nahr Genussm Gebrauchsgegenstnde 47:97127
Brand-Miller J (2007) The glycemic index as a measure of health and nutritional quality: an Australian
perspective. Cereal Foods World 52:4144
255
Brouns F, Bjorck I, Frayn KN, Gibbs AL, Lang V, Slama G, Wolever TMS (2005) Glycaemic index
methodology. Nutrition Res Rev 18:145171
Bushway RJ (1987) Inhibition of acetyl cholinesterase by solanaceaeus glycoalkaloids and alkaloids. Am
Potato J 64:409413
Buyken AE, Kroke A (2006) Glykmischer Index von KartoffelnMythos und Wirklichkeit. Ernhrung
im Focus 6:254257
Calleman CJ (1996) The metabolism and pharmacokinetics of acrylamide: implications for mechanisms of
toxicity and human risk estimation. Drug Met Rev 28:527590
Cao G, Booth SL, Sadowski JS, Prior RL (1998) Increase in human plasma antioxidant capacity after
consumption of controlled diets high in fruit and vegetables. Am J Clin Nutr 68:10811087
Cepl J, Zrust J (1996) Content of glycoalkaloids as affected by variety, nitrogen fertilization and
mechanical damage of tubers. Abstracts of Conference Papers, Posters and Demonstrations of the
13th Triennial Conference of the EAPR, Veldhoven, 1996, pp. 532533
Chaillou LL, Nazareno MA (2006) New method to determine antioxidant activity of polyphenols. J Agric
Food Chem 54:83978402
Chu Y, Chang C, Hsu H (2000) Flavonoid content of several vegetables and their antioxidant activity. J
Sci Food Agric 80:561566
Chu YF, Sun J, Wu XZ, Liu RH (2002) Antioxidant and anti proliferative activities of common
vegetables. J Agric Food Chem 50:69106916
Cieslik E, Praznik W (1998) Changes of glycoalkaloid content in potato tubers of selected varieties during
vegetation and storage. Polish J Food Nutr Sci 48:417422
Confederation of the food and drink industries of the EU (CIAA) (2006) CIAA acrylamide toolbox,
URL: http://www.ciaa.be/documents/brochures/CIAA_Acrylamide_Toolbox_Oct2006.pdf (access 2503-07), Brussels
Dale MFB, Griffiths DW, Bain H (1998) Effect of bruising on the total glycoalkaloid and chlorogenic acid
content of potato tubers of five cultivars. J Sci Food Agric 77:499505
Davey MW, van Montagu M, Inze D, Sanmartin M, Kanellis A, Smirnoff N, Benzi IJJ (2000) Review:
Plant L-ascorbic acid: chemistry, function, metabolism, bioavailability and effects of processing. J Sci
Food Agric 80:825860
Dybing E (2007) Risk assessment of acrylamide. In: Eisenbrand G, Engel K-H, Grunow W, Hartwig A,
Knorr D, Knudsen I, Schlatter J, Schreier P, Steinberg P, Vieths S (eds) Thermal processing of food:
potential health benefits and risks. Wiley, Weinheim, pp 7589
Dybing E, Sanner T (2003) Risk assessment of acrylamide in foods. Toxicol Sci 75:715
Engel KH, Gerstner G, Ross A (1998) Investigation of glycoalkaloids in potatoes as example for the
principle of substantial equivalence. In: Novel foods regulation in the European Union, BGVV,
Berlin, pp 197209
Englyst HN, Kingman SM, Hudson GJ, Cummings JH (1996a) Measurement of resistant starch in vitro
and in vivo. Brit J Nutr 75:749755
Englyst HN, Veenstra J, Hudson GJ (1996b) Measurement of rapidly available glucose (RAG) in plant
foods: a potential in vitro predictor of the glycaemic response. Brit J Nutr 75:327337
Englyst KN, Hudson GJ, Englyst HN (2000) Starch analysis in food. In: Meyers RA (ed) Encyclopedia of
analytical chemistry. Wiley, Chichester, pp 42464262
Englyst KN, Vinoy S, Englyst HN, Lang V (2003) Glycaemic index of cereal products explained by their
content of rapidly and slowly available glucose. Brit J Nutr 89:329339
European Commission DG Health and Consumer Protection (2007) Food contaminantsacrylamide.
http://europa.eu.int/comm/food/food/chemicalsafety/contaminants/acrylamide_en.htm
FAO/WHO (2005) Joint FAO/WHO expert committee on food additives. 64th meeting. Report 817 Feb.,
Rome
Fernandes G, Velangi A, Wolever TMS (2005) Glycemic index of potatoes commonly consumed in North
America. J Am Diet Assoc 105:557562
Foster-Powell K, Holt SHA, Brand-Miller JC (2002) International table of glycemic index and glycemic
load values: 2002. Am J Clin Nutr 76:556
Friedman M, Kap-Rang L, Hyun-Jeong K, In-Seon L, Kozukue N (2005) Anticarcinogenic effects of
glycoalkaloids from potatoes against human cervical, liver, lymphoma, and stomach cancer cells. J
Goni I, Bravo L, Larrauri JA, Saura Calixto F (1997) Resistant starch in potatoes deep-fried in olive oils.
Food Chem 59:269272
Granda C, Moreira RG, Tichy SE (2004) Reduction of acrylamide formation in potato chips by lowtemperature vacuum frying. J Food Sci 69:E405E411
256
Granvogl M, Jezussek M, Khler P, Schieberle P (2004) Quantitation of 3-aminopropionamide in potatoes

a minor but potent precursor in acrylamide formation. J Agric Food Chem 52:47514757
Granvogl M, Schieberle P (2007) Quantification of 3-aminopropionamide in cocoa, coffee and cereal
products. Eur Food Res Tech 225:857863
Grob K, Biedermann M, Biedermann-Brem S, Noti A, Imhof D, Amrein T, Pfefferle A (2003) French fries
with less than 100 g/kg acrylamide. A collaboration between cooks and analysts. Eur Food Res Tech
217:185194
Gromes R, Herrmann M-E (2008) Antioxidative Kapazitt von Kartoffeln. Ernaehrungs-Umschau, pp 7882
Gross J (1991) Pigments in vegetables: chlorophylls and carotenoids. Van Nostrand Reinhold, New York
Gulliford MC, Bicknell EJ, Scarpello JH (1989) Differential effect of protein and fat ingestion on bloodglucose responses to high-glycemic-index and low-glycemic-index carbohydrates in noninsulindependent diabetic subjects. Am J Clin Nutr 50:773777
Haase NU (1999) Glycoalkaloide in Kartoffelknollen. Kartoffelbau 50:4447
Haase NU (2006) The formation of acrylamide in potato products. In: Skog K, Alexander J (eds)
Acrylamide and other hazardous compounds in heat-treated foods. Woodhead, Cambridge, pp 4159
Haase NU (2007) Minimization strategies: acrylamide. In: Eisenbrand G, Engel K-H, Grunow W, Hartwig
A, Knorr D, Knudsen I, Schlatter J, Schreier P, Steinberg P, Vieths S (eds) Thermal processing of
food: potential health benefits and risks. Wiley, Weinheim, pp 155174
Haase NU, Lindhauer MG (2005) Minimization strategies in potato food. In: Bund fr Lebensmittelrecht
und Lebensmittelkunde e.V.(BLL), Forschungskreis der Ernhrungsindustrie e.V. (FEI) (ed)
Development of new technologies to minimize acrylamide in food., URL: http://www.bll.de/
download/themen/kontaminanten/acrylamid/acrylamid.pdf, Bonn, pp 4556
Haase NU, Weber L (2003) Ascorbic acid losses during processing of French fries and potato chips. J
Food Eng 56:207209
Haase NU, Matthus B, Vosmann K (2004) Aspects of acrylamide formation in potato crisps. J Appl Bot
Food Qual 78:144147
Hale AL (2003) Screening potato genotypes for antioxidant activity, identification of the responsible
compounds, and differentiating Russet Norkotah strains using AFLP and microsatellite marker
analysis. PhD dissertation, Texas A&M University, College Station, TX. Cited in: Brown CR (eds)
Antioxidants in Potato; Am J Potato Res 82 (2005):163172
Halle I, Ihling M, Lahrsson-Wiederholt M, Klaffke H, Flachowsky G (2006) Carry-over of acrylamide
from feed (heated potato product) to eggs and body tissues of laying hens. J Verbraucherschutz und
Lebensmittelsicherheit 1:290293
Halvorsen BL, Holte K, Myhrstad MCW, Barikmo I, Hvattum E, Fagertun Remberg S, Wold AB, Haffner
K, Baugerod H, Frost Andersen L, Moskaug JO, Jacobs DJ, Blomhoff R (2002) A systematic
screening of total antioxidants in dietary plants. J Nutr 132:461471
Hauner H (2006) Glykmischer Index und glykmische Last: relevant fr die Gesundheit. Aktuel Ernaehr
Med 31:4248
Hellens KE, Branzell C, Johnsson H, Slanina P (1995a) High-levels of glycoalkaloids in the established
Swedish potato variety Magnum-Bonum. J Sci Food Agric 68:249255
Hellens KE, Branzell C, Johnsson H, Slanina P (1995b) Glycoalkaloid content of early potato varieties. J
Jadhav SJ (1983) Naturally occurring toxic alkaloids in foods. CRC Crit Rev Toxicol 9:21103
Jeannet-Peter N, Piletta-Zanin P-A, Hauser C (1999) Facial dermatitis, contact urticaria, rhinoconjunctivitis, and asthma induced by potato. Am J Contact Dermat 10:4042
Jenkins AL (2007) The glycemic index: looking back 25 years. Cereal Foods World 52:5053
Karkalas J (1985) An improved enzymatic method for the determination of native and modified starch. J
Kolbe H (1997) Einflufaktoren auf die Inhaltsstoffe der Kartoffel. VII Vitamine. Kartoffelbau 48:3439
Lachman K, Hamouz K (2005) Red and purple coloured potatoes as a significant antioxidant source in
human nutritiona review. Plant, Soil Environ 51:477482
Lewis CE, Walker JRL, Lancaster JE, Sutton KH (1998) Determination of anthocyanins, flavonoids and
phenolic acids in potatoes. I: Coloured cultivars of Solanum tuberosum L. J Sci Food Agric 77:4557
Lewis CE, Walker JRL, Lancaster JE (1999) Changes in anthocyanin, flavonoid and phenolic acid
concentrations during development and storage of coloured potato (Solanum tuberosum L.) tubers. J
Lindenmeier M, Hofmann T (2004) Influence of baking conditions and precursor supplementation on the
amounts of the antioxidant pronyl-L-lysine in bakery products. J Agric Food Chem 52:350354
Lisinska G, Leszczynski W (1989) Potato science and technology. Elsevier, London
257
Lu WH, Haynes K, Wiley E, Clevidence B (2001) Carotenoid content and color in diploid potatoes. J Am
Soc Hort Sci 126:722726
Ludwig DS (2000) Dietary glycemic index and obesity. J Nutr 130:280S283S
Maga JA (1980) Potato glycoalkaloids. CRC Crit Rev Food Sci Nutr 12:371405
Matthus B, Haase NU, Vosmann K (2004) Factors effecting the concentration of acrylamide during deepfat frying of potatoes. Eur J Lipid Sci Technol 106:793801
Michaud DS, Fuchs CS, Liu SM, Willett WC, Colditz GA, Giovannucci E (2005) Dietary glycemic load,
carbohydrate, sugar, and colorectal cancer risk in men and women. Cancer Epidemiol Biomarkers
Prev 14:138143
Miller NJ, Sampson J, Candeias LP, Bramley PM, RiceEvans CA (1996) Antioxidant activities of
carotenes and xanthophylls. Febs Lett 384:240242
Morris SC, Lee TH (1984) The toxicity and teratogenicity of Solanaceae glycoalkaloids, particularly those
of the potato (Solanum tuberosum): a review. Food Technol Aust 36:118124
Mottram DS, Wedzicha BL, Dodson AT (2002) Acrylamide is formed in the Maillard reaction. Nature
419:448449
Mucci LA, Dickman PW, Steineck G, Adami HO, Augustsson K (2003) Dietary acrylamide and cancer of
the large bowel, kidney, and bladder: absence of an association in a population-based study in
Sweden. Br J Cancer 88:8489
Nesterenko S, Sink KC (2003) Carotenoid profiles of potato breeding lines and selected cultivars.
HortScience 38:11731177
Olesen PT, Olsen A, Frandsen H, Frederiksen K, Overvad K, Tjonneland A (2008) Acrylamide exposure
and incidence of breast cancer among postmenopausal women in the Danish Diet, Cancer and Health
study. Int J Cancer 122:20942100
Olsson K (1996) Attempts to unveil potato clones disposed to stress-induced accumulation of
glycoalkaloids in the field. EA. In: 13th Triennial Conference of the European Association for Potato
Research (eds) Abstracts of Conference Papers, Posters and Demonstrations, European Association
for Potato Research, Veldhoven, NL, pp 538539
Parnell A, Bhuva VS, Bintcliffe EJB (1984) The glycoalkaloid content of potato varieties. J Natl Inst
Agric Bot 16:535542
Pedreschi F, Moyano P (2005) Oil uptake and texture development in fried potato slices. J Food Eng
70:557563
Pelucchi C, Galeone C, Levi F, Negri E, Franceschi S, Talamini R, Bosetti CN, Giacosa A, La Vecchia C
(2006) Dietary acrylamide and human cancer. Int J Cancer 118:467471
Pulido R, Bravo L, Saura-Calixto F (2000) Antioxidant activity of dietary polyphenols as determined by a
modified ferric reducing/antioxidant power assay. J Agric Food Chem 48:33963402
Reyes LF, Cisneros-Zevallos L (2003) Wounding stress increases the phenolic content and antioxidant
capacity of purple-flesh potatoes (Solanum tuberosum L.). J Agr Food Chem 51:52965300
Reyes LF, Miller JJ, Cisneros-Zevallos L (2005) Antioxidant capacity, anthocyanins and total phenolics
in purple- and red-fleshed potato (Solanum tuberosum L.) genotypes. Am J Potato Res 82:271
277
Roddick JG (1987) Antifungal activity of plant steroids. In: Fuller G, Nes WD (eds) Ecology and
metabolism of plant lipids (ACS Symposium Series 325). American Chemical Society, Washington,
DC, pp 286303
Roddick JG (1988) Synergistic interaction between potato glycoalkaloids solanine and chaconine in
relation to destabilisation of cell membranes: ecological implications. J Chem Ecol 14:889902
Rogozinska I, Wszelaczynska E, Wichrowska D (2005) Effect of bioelements (Mg, N, K) and herbicides
on vitamin C content in potato tubers. Part 1: Vitamin C content in potato tubers immediately after
harvest. J Elementol 10:9991008
Seppl U, Alenius H, Turjanmaa K, Reunala T, Palosuo T, Kalkkinen N (1999) Identification of patatin
as a novel allergen for children with positive skin prick test responses to raw potato. J Allergy Clin
Immunol 103:165171
Sinden SL, Deahl KL, Aulenbach BB (1976) Effect of glycoalkaloids and phenolics on potato flavor. J
Food Sci 41:520523
Smith DB, Roddick JG, Jones JL (1996) Potato glycoalkaloids: some unanswered questions. Trends Food
Sci Technol 7:126131
Soh NL, Brand-Miller J (1999) The glycaemic index of potatoes: the effect of variety, cooking method and
maturity. Eur J Clin Nutr 53:249254
Sowokinos JR, Preston DA (1988) Maintenance of potato processing quality by chemical maturity
monitoring (CMM). Station Bulletin 586, University of Minnesota
258
Stadler RH, Robert F, Riediker S, Varga N, Davidek T, Devaud S, Goldmann T, Ha J (2004) In-depth
mechanistic study on the formation of acrylamide and other vinylogous compounds by the Maillard
reaction. J Agric Food Chem 52:55505558
Taeymans D, Wood J, Ashby P, Blank I, Studer A, Stadler RH, Gonde P, van Eijck P, Lalljie S, Lingnert
H, Lindblom M, Matissek R, Mueller D, Tallmadge D, O'Brien J, Thompson S, Silvani D, Whitmore
T (2004) A review of acrylamide: an industry perspective on research, analysis, formation, and
control. CRC Crit Rev Food Sci Nutr 44:323347
U.S. Environmental Protection Agency (1999) Integrated risk information system (IRIS) on acrylamide.
National Center for Environmental Assessment, Office of Research and Development, Washington
DC
Wang H, Cao G, Prior RL (1996) Total antioxidant capacity of fruits. J Agric Food Chem 44:701705
Weber L, Putz B (1998) Vitamin-C in Kartoffeln. Kartoffelbau 49:278281
Weisshaar R, Gutsche B (2002) Formation of acrylamide in heated potato products - model experiments
pointing to asparagine as precursor. Dtsch Lebensm-Rundsch 98:397400
Welch RW, Price RK, Lee AM, Strain JJ (2005) Uptake and antioxidant activity of oat and wheat
phenolics in humans. In: ICC (eds) Cerealsthe future challengebook of abstracts, 1515
WHO/FAO (2003) Nutrition, diet, and prevention of chronic diseases. WHO Technical Report Series 916.
World Health Organization, Geneva
Willett WC (2001) Eat, drink and be healthy: the Harvard Medical School guide to healthy eating. Simon
& Schuster, New York
Yaylayan VA, Wnorowski A, Locas CP (2003) Why asparagine needs carbohydrates to generate
acrylamide. J Agric Food Chem 51:17531757
Zyzak DV, Sanders RA, Stojanovic M, Tallmadge DH, Eberhart BL, Ewald DK, Gruber DC, Morsch TR,
Strothers MA, Rizzi GP, Villagran MD (2003) Acrylamide formation mechanism in heated foods. J

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Potato Research (2008) 51:239258