Food Control 18 (2007) 503508

www.elsevier.com/locate/foodcont

Eect of light and temperature on the formation of glycoalkaloids

in potato tubers
Rita M.D. Machado *, Maria Ceclia F. Toledo, Lucila C. Garcia
Department of Food Science, Faculty of Food Engineering, State University of Campinas, UNICAMP, Caixa Postal 6121,
CEP 13083-862, Campinas, SP, Brazil
Received 28 December 2004; received in revised form 16 December 2005; accepted 16 December 2005

Abstract
This study was designed to determine the inuence of two light sources and temperature on the total glycoalkaloid (TGA) content of
potato tubers cultivated in Brazil. Tubers of cv Monaliza were exposed during 14 days to the following conditions: (1) indirect sunlight,
(2) uorescent light, (3) storage in darkness under refrigeration, and (4) storage in darkness under room temperature. The glycoalkaloids
a-solanine and a-chaconine were determined using a reversed phase C18 HPLC column with a photodiode array detector. Exposure of
potato tubers to uorescent light resulted in the highest TGA levels. Smaller tubers presented the highest TGA concentrations, irrespective of the light source and temperature. Although the TGA levels at the end of the experiments were higher than the initial content, a
steady increase of TGA concentration was observed only in tubers exposed to uorescent light. The levels of TGA found in the analysed
potato samples were below 200 mg kg 1, value that has been considered to be safe for human consumption.
 2006 Elsevier Ltd. All rights reserved.
Keywords: Glycoalkaloids; a-Solanine; a-Chaconine

1. Introduction
Potato is included among the main horticultural crops in
Brazil. Due to its broad availability and nutritional characteristics, it has been considered one of the most important
components of the human diet. Nevertheless, potato tubers
(Solanum tuberosum L.) contain two naturally occurring
toxins, a-solanine and a-chaconine, which comprise over
95% of the total glycoalkaloids (TGA) present in the
potato plant (Bushway & Ponnampalam, 1981). According
to Maga (1980), the glycoalkaloids can be found in all parts
of a potato plant. Among the tissues that contain glycoalkaloids, the skins and sprouts present the highest concentrations (Morris & Lee, 1984; Smith, Roddick, & Jones,
1996). Symptoms of glycoalkaloid poisoning include colic
pain in the abdomen and stomach, gastroenteritis, diarrhea, vomiting, fever, rapid pulse, low blood pressure,
*

Corresponding author. Tel.: +55 19 3788 2168; fax: +55 19 3788 2153.
E-mail address: rdonato@fea.unicamp.br (R.M.D. Machado).

0956-7135/$ - see front matter  2006 Elsevier Ltd. All rights reserved.
doi:10.1016/j.foodcont.2005.12.008

and neurological disorders (Morris & Lee, 1984; Slanina,

1990). Toxicity of glycoalkaloids results from anticholinesterase activity on the central nervous system and membrane disruption adversely aecting the digestive system
and general body metabolism (Jadhav, Sharma, & Salunkhe, 1981; Maga, 1980; Roddick, 1989). In vitro experiments
showed that a-solanine and particularly a-chaconine are
potent cytotoxins, acting rapidly to induce cell lysis
(Phillips et al., 1996). The cytotoxic potency of potato glycoalkaloids is considerable, particularly in the case of
a-solanine: a-chaconine mixtures, which present dierentiated synergistic action in disrupting membranes (Keukens
et al., 1996; Rayburn, Friedman, & Bantle, 1995; Roddick,
Rijnenberg, & Osman, 1988). Moreover, these synergisms
have important implications not only for the defensive role
of glycoalkaloids in potato plant but also for the safety and
acceptability of potatoes and potato products (Smith, Roddick, & Jones, 2001).
The TGA content in potato varieties commercially
available is usually below 200 mg kg 1 of fresh weight

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R.M.D. Machado et al. / Food Control 18 (2007) 503508

(FW), the recommended safety level of TGA in unprocessed potato tubers for human consumption (FAO/
WHO, 1999; Slanina, 1990). In the study conducted in Brazil by Machado and Toledo (2004), 82% of the samples of
whole potato tubers of dierent commercial varieties presented levels of TGA below 100 mg kg 1 FW. This result
is consistent with those ndings from other countries
(Friedman, Roitman, & Kozukue, 2003; Peksa, Golubowska, Rytel, Lisinska, & Aniolowski, 2002) and indicates that the commercial potatoes are usually safe for
human consumption. In addition, potato glycoalkaloids
appear to be largely unaected by home processing conditions such as baking, boiling, frying and microwaving
(Bushway & Ponnampalam, 1981; Maga, 1980).
However, the content of the glycoalkaloids can vary
greatly in dierent potato cultivars and the biosynthesis
of glycoalkaloid can be rapidly stimulated by environmental factors such as light, mechanical injury, and storage
temperature (Friedman & Dao, 1992; Griths, Bain, &
Dale, 1997; Jadhav & Salunkhe, 1975; Percival, 1999). Glycoalkaloid synthesis due to exposure to light may occur
either in the eld, at harvest or during storage (Jadhav &
Salunkhe, 1975). Several authors have demonstrated that
the TGA concentration of potato tubers exposed to light
can increase twice or three times. This means that individuals consuming large quantities of light-exposed tubers
could theoretically present adverse eects (Dale, Griths,
Bain, & Todd, 1993; Griths, Bain, & Dale, 1998; Jadhav
& Salunkhe, 1975).
It has been shown that the rate of glycoalkaloid accumulation in potato tubers can also be signicantly inuenced
by the spectral composition of the light source. In a study
conducted with the cultivar Pentland Hawk, the levels of
glycoalkaloids were four to six times higher in tubers
exposed to uorescent or sodium light sources than in
tubers exposed to mercury light sources (Percival, Dixon,
& Sword, 1994).
The aim of this work was to study the eect of light
exposure and temperature on the total glycoalkaloid
(TGA) content of potato tubers of cv Monaliza cultivated
in Brazil.
2. Materials and methods
2.1. Plant materials
Tubers of cv Monaliza, harvested three days before,
were acquired at the Central de Abastecimento de Campinas (CEASA) in April/2001 (50 kg).
Tubers were washed under running water and allowed
to dry at room temperature (25 C). After this, each
tuber was weighted and classied as small or medium,
corresponding to average weights of 68.8 4.1 and
117.6 5.9 g, respectively. Batches of 30 tubers each were
selected by size (small or medium). Each batch was placed
under the following conditions: (a) indirect sunlight exposure; (b) uorescent light exposure (lamps of 40 W, at a

Table 1
Parameters monitored during the period of 2 weeks
Condition

T (temperature)a
(C)

Light intensityb
(Lux)

Indirect sunlight exposure

Fluorescent light exposure
Storage in darkness under
refrigeration
Storage in darkness under
room temperature

2329
2430
78

131868
458960

1926

a
b

Measured during the experiment.

Measured using the Luximeter (testo 545-Lux, fc).

distance of 2 m); (c) storage in darkness under refrigeration

(78 C); (d) storage in darkness under room temperature,
with no ventilation (1926 C).
With exception of the tubers exposed to indirect sunlight, which were maintained in this condition for an average period of 10 h (daytime), all other tubers were
continuously exposed to the specied condition. Tubers
exposed to indirect sunlight and uorescent light were
rotated at 24 h intervals to ensure complete exposure to
light. Measurements during the sunlight exposure were
made in days with dierent conditions of temperature
and luminosity (sunny and cloudy days). Table 1 summarises the range of variation of the parameters monitored
during the experiments.
2.2. Sample preparation
From each batch, samples of four tubers were withdrawn at 0, 3, 7, and 14 day intervals for glycoalkaloid
analysis. The whole tubers were sliced, combined, frozen,
and freeze-dried (Freeze-drier LabConco Lyph Lock 18).
Samples were minced and stored at 15 C until analysis.
2.3. Extraction of the glycoalkaloids
The extraction procedure was based on Percival and
Dixon (1996). The freeze-dried samples (0.5 g) were placed
in a centrifuge tube and a volume of 10 ml of extraction
solution (1 l water, 20 ml acetic acid, 5 g sodium bisulphite)
was added. The samples were shaken during 15 min and
the solution claried by centrifugation (Centrifuge Hitachi
himac CR 21) at 4000 rpm for 20 min. The analysis of each
sample was made in duplicate.
2.4. Clean-up
This procedure was accomplished according to Hellenas
and Branzell (1997). A SepPak Plus C18 cartridge (Waters
Assoc., Milford, MA, USA) was conditioned with 5 ml of
acetonitrile, followed by 5 ml of extraction solution. Five
millilitres of the claried tuber extract was passed through
the column followed by washing with 4 ml water/acetonitrile, 85:15 (v/v). The glycoalkaloids were then eluted from
the cartridge with 4 ml of acetonitrile/0.022 M potassium

R.M.D. Machado et al. / Food Control 18 (2007) 503508

phosphate buer, pH 7.6, 60:40 (v/v).

nally adjusted to 5 ml with the same
injection into the liquid chromatograph,
ltered through a Millex HV 0.45 lm
Co.).

The eluate was

solution. Before
all samples were
lter (Millipore

2.5. HPLC analysis

The analysis was carried out in a HPLC apparatus
equipped with a Waters 600 Controller pump, a Waters
in-line degasser, a Waters 717 plus autosampler, and a
Waters 996 photodiode array detector. The analytical column was an ODS-Hypersil 5 lm (25 cm 4.6 mm), and the
mobile phase consisted of 65% water, 35% acetonitrile and
0.05% ethanolamine at pH 4.54.6, adjusted with orthophosphoric acid (10 vol.%). The injection volume was
20 ll, the ow rate 1.0 ml min 1 and the euent monitored
at 200 nm. HPLC grade acetonitrile was purchased from
Mallinckrodt and the water was previously puried in a
Table 2
Alfa-solanine and a-chaconine content (mg kg

505

MilliQ-Plus purication system. All other chemicals were

of analytical grade.
2.6. Glycoalkaloid quantication
Quantication was performed by comparison of the
peak areas with those from pure a-solanine and a-chaconine (Sigma Chemical Co., St Louis, USA) dissolved in
0.05 M phosphate buer (KH2PO4). The external standard
plot method was used. The concentration range of the standard curves was 0.761 lg ml 1 for a-solanine, and 0.8
64 lg ml 1 for a-chaconine. Recovery of the glycoalkaloids
added to freeze-dried samples averaged 102%.
3. Results and discussion
Table 2 shows the levels of a-solanine and a-chaconine
determined in the analysed samples, expressed on a fresh
weigh (FW) basis.

FW) of potato tubers of cv Monaliza exposed to various conditions

Condition

Sizea

Exposure time (days)

0

14

Indirect sunlight exposure

Small

a-Solanine
a-Chaconine
TGAd
C:Se
a-Solanine
a-Chaconine
TGA
C:S

14.3b 0.2c
37.1 0.4
51.4 0.6
72:28
14.3 0.2
37.1 0.4
51.4 0.6
72:28

28.5 0.2
68.4 0.7
96.9 0.5
71:29
11.7 0.8
31.0 0.2
42.7 1.0
73:27

21.8 0.9
51.5 1.4
73.3 2.4
70:30
21.2 0.4
43.4 4.3
64.6 4.8
67:33

29.2 0.6
63.3 0.7
92.5 0.1
68:32
17.6 0.6
41.3 1.6
58.9 2.3
70:30

a-Solanine
a-Chaconine
TGA
C:S
a-Solanine
a-Chaconine
TGA
C:S

14.3 0.2
37.1 0.4
51.4 0.6
72:28
14.3 0.2
37.1 0.4
51.4 0.6
72:28

17.0 0.0
42.9 0.7
59.9 0.7
72:28
14.2 1.4
35.6 3.2
49.8 4.6
71:29

36.8 2.4
66.8 0.9
103.6 1.4
64:36
22.3 0.2
45.6 1.8
67.9 1.6
67:33

42.2 3.9
65.7 7.7
107.9 11.6
61:39
34.6 1.2
68.6 7.2
103.2 8.4
67:33

a-Solanine
a-Chaconine
TGA
C:S
a-Solanine
a-Chaconine
TGA
C:S

14.3 0.2
37.1 0.4
51.4 0.6
72:28
14.3 0.2
37.1 0.4
51.4 0.6
72:28

20.6 0.3
54.4 1.4
75.0 1.7
73:27
16.2 1.2
42.5 2.2
58.7 3.5
72:28

25.0 1.3
63.4 3.2
88.4 4.3
72:28
16.3 6.3
34.5 1.5
50.8 4.8
68:32

25.2 1.2
56.6 7.5
81.8 8.7
69:31
19.3 0.5
45.7 0.7
65.0 1.2
70:30

a-Solanine
a-Chaconine
TGA
C:S
a-Solanine
a-Chaconine
TGA
C:S

14.3 0.2
37.1 0.4
51.4 0.6
72:28
14.3 0.2
37.1 0.4
51.4 0.6
72:28

21.3 2.3
55.2 2.2
76.5 4.5
72:28
19.3 0.2
52.9 2.7
72.2 3.0
73:27

21.2 0.1
55.7 2.5
76.9 2.6
72:28
17.5 1.6
42.1 5.8
59.6 7.4
71:29

16.6 1.7
44.2 1.2
60.8 3.0
73:27
16.1 0.3
40.9 1.1
57.0 1.4
72:28

Medium

Fluorescent light exposure (lamps of 40 W)

Small

Medium

Storage in darkness under refrigeration temperature (78 C)

Small

Medium

Storage in darkness under room temperature (1926 C)

Small

Medium

a
b
c
d
e

Small: tubers with average weight of 68.8 4.1 g, medium: tubers with average weight of 117.6 5.9 g.
Mean value of duplicates for each sample.
Standard deviation of the duplicates for each sample.
Total glycoalkaloid concentration.
Ratio a-chaconine:a-solanine.

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R.M.D. Machado et al. / Food Control 18 (2007) 503508

Regardless of the tuber size and exposure conditions,

TGA concentrations were higher at the end of the experiments when compared to the initial levels (Fig. 1). Smaller
tubers, irrespective of the treatment, presented the highest
concentrations of total glycoalkaloids at 14 days exposure.
This behaviour was also observed in studies accomplished
with cultivars Pentland Dell and Estima where the glycoalkaloid concentration was highly aected by the genotype
and an inverse relationship between concentration of total
glycoalkaloids and weight of individual tubers occurred
(Papathanasiou, Mitchell, & Harvey, 1999).
As shown in Fig. 1, the concentrations of glycoalkaloid
during most of the experiments uctuated; only the tubers
(small and medium sizes) exposed to uorescent light presented a continuous increase of TGA levels. In addition,
uorescent light exposure was the condition that most
induced glycoalkaloid formation, independent of the tuber
size. This treatment doubled the initial concentration of
total glycoalkaloids at the end of the experiment (maximum of 107.9 mg kg 1 at 14 days).
The total glycoalkaloid concentration of smaller tubers
exposed to indirect sunlight increased from 51.4 to
96.9 mg kg 1 (FW) in the rst three days, then decreased
to 73.3 mg kg 1 (FW) at 7 days and, nally, increased to
92.5 mg kg 1 (FW) at 14 days. A similar behaviour was
observed by Percival, Dixon, and Sword (1996) for the cul-

tivars Kerrs Pink, Pentland Hawk and Desiree. Although

an increase of the TGA level was veried after exposure
of the tubers to natural light for 21 days, the authors
reported that the TGA contents uctuated during the
experiment, with no continuous accumulation of these
compounds by the potato varieties studied.
The increase of glycoalkaloids levels in potato tubers of
lower size submitted to indirect sunlight and uorescent
light was approximately 46 times higher than that of
potato tubers stored in darkness under room temperature.
Similar results were found by Salunkhe, Wu, and Jadhav
(1972) who reported that potato exposure to sunlight or
articial light can increase the glycoalkaloid synthesis to
levels 3 or 4 times higher than those found when the potatoes are maintained in darkness.
The potatoes stored in darkness under refrigeration (7
8 C) synthesised glycoalkaloids in higher levels than those
found in tubers stored in darkness under room temperature
(1926 C). Griths et al. (1998) also observed that glycoalkaloid accumulation can occur with potato tubers
stored under low temperatures (410 C), and that the
intensity of this eect on the glycoalkaloid biosynthesis
depends of the potato variety.
According to literature, growth residual activity of
potato may occur in darkness, allowing synthesis and degradation reactions such as the glycoalkaloid biosynthesis

Indirect sunlight
Fluorescent light
Darkness under refrigeration temperature

Total glycoalkaloids (mg.kg-1)

Darkness under room temperature

110
100
90
80
70
60
50
40
30

61:39
68:32
69:31

73:27

3.5

10.5

14

Time (days)
Indirect sunlight
Fluorescent light
Darkness under refrigeration temperature

Total glycoalkaloids (mg.kg-1)

Darkness under room temperature

110
100
90
80
70
60
50
40
30

67:33

70:30
70:30
72:28

3.5

10.5

14

Time (days)

Fig. 1. TGA concentration in potato tubers of cv Monaliza exposed to the following conditions: indirect sunlight, uorescent light, storage in darkness
under refrigeration (78 C), and storage in darkness under room temperature (1926 C). (a) Small tubers: average weight of 68.8 4.1 g; (b) medium
tubers: average weight of 117.6 5.9 g. The initial ratio chaconine:solanine was 72:28 and the nal values are represented in the gure.

R.M.D. Machado et al. / Food Control 18 (2007) 503508

(Percival et al., 1996; Spoladore, Zullo, Teixeira, Coelho, &

Miranda, 1985). Although some authors (Haddadin,
Humeid, Qaroot, & Robinson, 2001; Maine, Bain, & Joyce,
1988; Percival, 1999) have used storage in darkness as a
control treatment when studying the glycoalkaloid formation in potato tubers exposed to light, our results suggest
that this procedure may not be appropriate for all potato
varieties.
Exposure to uorescent light decreased the ratio of achaconine:a-solanine from 72:28 to 61:39 for smaller tubers
and from 72:28 to 67:33 for the larger size ones. These
results indicate enhancement of a-solanine synthesis to a
greater degree than a-chaconine. This observation was
reported earlier by others for the cultivars Pentland Hawk,
Desiree, and King Edward (Percival, 1999; Percival et al.,
1994). As the toxicological potency of a-chaconine is
around 310-fold higher than that of a-solanine, a-chaconine may be more prominent in cases of potato poisoning
(Friedman, Rayburn, & Bantle, 1991). Furthermore, it is
believed that the relative proportion of glycoalkaloids
may inuence toxicity more than the absolute total glycoalkaloid concentrations with consequential implications for
the overall recommendation of 200 mg kg 1 FW for food
safety (Friedman et al., 2003).
The present investigation indicated that in the case of cv
Monaliza, irrespective of the exposure condition there is
always an accumulation of TGA by the potato tubers.
Moreover, as during marketing and sale potatoes are usually displayed under uorescent light, a substantial increase
of glycoalkaloids may occur, compromising of the tuber
quality. Although the levels of TGA found in the tubers of
cv Monaliza exposed to the dierent conditions were below
200 mg kg 1, this level has been questioned in relation to its
safety for long term exposures. Therefore, it is recommended that cultivars for human consumption possess slow
rates of glycoalkaloid accumulation in response to light and
be selected based on a low initial concentration of glycoalkaloid. Further investigation is needed to determine the TGA
levels in other cultivars commercialised in Brazil.
Acknowledgement
The authors would like to acknowledge FAPESP for
supporting this project (proc. 99/08476-5).
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