Journal of the Science of Food and Agriculture J Sci Food Agric 79 :32–36 (1999)

Variation in glycoalkaloid concentration of

potato tubers harvested from mature plants
Fokion Papathanas iou,1,s Samuel H Mitchell2 and Barbara M R Harvey1,*
1 Department of Applied Plant Science , The Queen’s Univers ity of Belfas t , Newforge Lane , Belfas t BT9 5PX , UK
2 Food Science , Department of Agriculture for Northern Ireland , Newforge Lane , Belfas t BT9 5PX , UK

Abstract : Individual tubers from mature plants of cultivars Pentland Dell and Estima were harvested
from experimental plots in two successive years and analysed for a-solanine, a-chaconine and total
glycoalkaloid concentration (a-solanine and a-chaconine combined) using high-performance liquid
chromatography. Mean tuber glycoalkaloid concentration per plant was strongly aþ ected by genotype
and was much higher in plants of Pentland Dell (10.0 and 16.3 mg per 100 g fresh weight in 1994 and
1995, respectively) than Estima (4.0 and 4.3 mg per 100 g fresh weight in 1994 and 1995, respectively).
The variation between plants in mean tuber glycoalkaloid concentration, expressed as coefficient of
variation, was similar for the two cultivars in both years. Variation within plants was higher for
cultivar Pentland Dell than Estima for both years but it was statistically signiücant only in 1995.
Tuber position within a plant had no eþ ect on glycoalkaloid concentrations in tubers of similar size.
An inverse relationship between total glycoalkaloid concentration and tuber fresh weight of individual
tubers was found for both cultivars and small tubers of Pentland Dell (20–40 g fresh weight) exceeded
the safety limit of 20 mg per 100 g fresh weight. The pattern of glycoalkaloid accumulation diþ ered
between genotypes and appeared related to tuber growth. The implications of these results in terms of
food safety are discussed.
( 1999 Society of Chemical Industry

Keywords : glycoalkaloid ; a-solanine ; a-chaconine ; potato; tuber size ; Solanum tuberosum

INTRODUCTION diþerent cultivars.1 Variation of glycoalkaloid con-

Tubers of the cultivated potato (Solanum tuberosum
L) contain two naturally occurring toxins, the steroi-
dal glycoalkaloids, a-solanine and a-chaconine, which
comprise over 95% of the total glycoalkaloids present
in the potato plant.1 Both a-solanine and a-chaconine
are concentrated in the peripheral layers of the
potato tuber so are present at low levels in peeled
tubers and usually do not present any food safety
problems for the consumer.2,3 However, undesirable
high and toxic levels have been found and many
cases of acute poisoning and even deaths have been
reported in humans after consumption of potatoes
with excessive glycoalkaloid levels.4h6 Since early
this century, levels of less than 20 mg per 100 g of
tuber fresh weight (FW) have been considered safe
for human consumption.7 Concern about this safety
limit has been expressed due to the small safety
margin between potentially toxic levels and the
glycoalkaloid levels found in marketable potatoes.8
Several factors during growth, harvesting and
post-harvest treatment can aþect glycoalkaloid accu-
mulation.9,10 Genotype has a major eþect and tuber
glycoalkaloid levels can vary considerably between
centration has also been reported between tubers
from the same experimental plot11 and between indi-
vidual tubers within the same commercial package.12
Causes of such variation have not been deüned
clearly. Small tubers within a plant are expected to
have high glycoalkaloid concentrations due to the
higher skin to ýesh ratio whereas larger tubers have
lower glycoalkaloid concentrations.13 However, the
relationship between tuber size and TGA concentra-
tion is still unclear and may diþer between geno-
types. Physiological factors within a plant such as the
tuber position might also be important. In the
present study the variation in the glycoalkaloid
content of tubers of individual plants was assessed
and the relationships between tuber size and glyco-
alkaloid content were investigated using two culti-
vars diþering in maturity group. These were Estima,
which accounted for 40% of the total area of second
early maturing cultivars grown in Great Britain in
1994 and 199514 and Pentland Dell, which accounted
for 9% of the area of maincrop cultivars grown in
Great Britain and is a very important maincrop culti-
var in Northern Ireland.

¹ Pres ent addres s : Karagiannopoulou 125, Levadia Viotias , 32100 Belfas t BT9 5PX, UK
Greece.
* Corres pondence to : Barbara MR Harvey, Department of Applied (Received 15 September 1997 ; revis ed vers ion received 2 March
Plant Science, The Queen’s Univers ity of Belfas t, Newforge Lane, 1998 ; accepted 14 April 1998 )

( 1999 Society of Chemical Industry. J Sci Food Agric 0022–5142/99/$17.50 32


EXPERIMENTAL
Plant material
Pentland Dell and Estima were grown as part of 1994
and 1995 cultivar trials at the Northern Ireland Plant
Testing Station, Crossnacreevy (Department of
Agriculture for Northern Ireland). Two days before
planting the experimental areas (medium clay loam)
were fertilised with 125 kg N, 225 kg P (P O ) and
2 5
250 kg K (K O) and the trials were laid out in four
2
blocks with cultivar plots randomised within the
blocks. Each block consisted of four rows with 76 cm
between rows and there were 25 tubers in each row
with 30 cm between tubers. Summer weather condi-
tions diþered considerably between the years ; mean
monthly values (J une to August inclusive) for rainfall
were 69 mm in 1994 and 47 mm in 1995, temperature
was 13.1¡C and 15.6¡C in 1994 and 1995 respectively
and irradiance was 402 MJ m~2 in 1994 and
502 MJ m~2 in 1995.
Sampling was performed on 11 October 1994 (160
days after planting) and on 10 October in 1995 (170
days after planting). This was 4 weeks after haulm
destruction using Reglone (ICI, plc). Two plants of
each cultivar were randomly selected from the centre
of two of the four blocks in 1994 and from all four
blocks in the 1995 trial. Individual tubers were har-
vested carefully by hand and the position of each
tuber within each plant was recorded. The presence
or absence of an attached stolon was also recorded
for each tuber. Any green or damaged tubers were
discarded. Each tuber was placed in a labeled plastic
bag and stored in a paper box to avoid any exposure
to light. The tubers were kept at 4¡C for 24 h and
then individually processed and analysed for glyco-
alkaloids.
Sample preparation
Tubers were washed, weighed and frozen with liquid
nitrogen. Each tuber was then freeze-dried, weighed
and ground into a üne powder through a 0.5 mm
cyclone mill (Retsch). The dried powder was packed
in a plastic container and stored at [ 20¡C for up to
3 months before glycoalkaloid analysis.
Extraction and purification
Extraction followed the procedure suggested by Dale
et al15 with some modiücations. A sample of 0.48 g
of freeze-dried potato powder was mixed thoroughly
with extraction solution (water/acetic acid/sodium
bisulphite, 100 : 5 : 0.5 (v/v/w)) using mortar and
pestle. The mixture was diluted to a total of 12 ml of
extraction solution in 15 ml centrifuge tubes. The
tubes were placed in a tube rotator for 30 min and
then centrifuged at 2000 ] g, 4¡C for 30 min. The
clariüed aliquot (10 ml) was puriüed using the clean
up procedure of HellenaŽ s et al.16
Chemical analysis
A high-performance liquid chromatography (HPLC)
method based on HellenaŽ s et al16 was used. The ana-
J Sci Food Agric 79 :32–36 (1999)
Variation in glycoalkaloid concentration of potato tubers
lytical column (250 ] 4.6 mm id) was packed with
ODS2, Spherisorb, 5 lm, C18 phase (HPLC Tech-
nology Ltd, UK) and the mobile phase was
acetonitrile/0.01 M potassium phosphate buþer, pH
7.05, 50 : 50 (v/v). The ýow rate was 1.5 ml min~1
with injection volume 50 ll and the column effluent
was monitored for UV absorption at 202 nm. The
analysis equipment consisted of an HP1050 series
quaternary pump, autosampler and multiwavelength
detector (Hewlett Packard, USA). The concentra-
tions of a-solanine and a-chaconine were quantiüed
using standard curves produced from known
amounts of pure standards and results were
expressed as content (mg) or concentration (mg per
100 g FW) in individual tubers. Total glycoalkaloid
(TGA) was calculated as the sum of a-solanine and
a-chaconine.
Statistical analysis
Experimental data were analysed using Instat 2
(Graphpad Software Inc) and Genstat V17 was used
for two-way analysis of variance (ANOVA) and
linear regressions. The linear regression equations of
glycoalkaloid content per tuber versus tuber FW
were converted to inverse equations between concen-
tration and tuber FW. From these equations the
tuber FW at which the glycoalkaloid concentration
was half the overall safety limit (ie 10 mg per 100 g
FW) was calculated for each plant and compared
between cultivars by ANOVA. A probability of
P O 0.05 was accepted as signiücant in all analyses.
RESULTS
Variation between and within plants
Mean tuber glycoalkaloid concentration per plant of
cultivars Pentland Dell and Estima sampled in 1994
and 1995 is shown (Fig 1) by box and whisker
plots.18 Plants of cultivar Pentland Dell generally
had higher mean tuber glycoalkaloid concentration
than plants of Estima in both years. For Pentland
Dell the mean TGA concentration per plant was cal-
culated to be 10.0 ^ 1.09 and 16.3 ^ 1.25 mg per
100 g fresh weight in 1994 and 1995, respectively,
and for Estima the values were 4.0 ^ 0.79 and
4.3 ^ 0.23 mg per 100 g fresh weight in 1994 and
1995, respectively. However, the variation between
plants expressed as coefficient of variation was
similar for Estima and Pentland Dell in both years.
All values were between 22% and 25%. Considering
variation within plants for each cultivar, the standard
deviation in tuber glycoalkaloid levels per plant was
small for Estima in 1994 and 1995 and the range
between minimum and maximum tuber glycoalkaloid
levels was also small (Fig 1). Larger variation within
plants was observed for cultivar Pentland Dell than
cultivar Estima for both years but it was signiücantly
diþerent only in 1995.
33
F Papathanasiou, SH Mitchell, BMR Harvey

Figure 2. Total glycoalkaloid concentration (mg per 100 g FW) in

individual tubers of mature plants of cultivar (a) Pentland Dell and
(b) Es tima harves ted in 1994. Tubers and s tolons are drawn to
approximate s cale, the s cale bar indicates 50 mm. Stolons are
s hown as a broken line where the tuber was no longer connected
to the parent plant. * \ Pooled s ample.
Figure 1. Tuber TGA concentration (mg per 100 g FW) in plants of
cultivars Pentland Dell and Es tima harves ted in (a) 1994 and (b)
1995. Box and whis ker plots dis play the mean (=), s tandard
deviation (boxes ) and the range from minimum to maximum
for Pentland Dell was observed in tubers of less than
values for individual tubers of each plant (vertical line). 2 g FW which had over 70 mg per 100 g FW.
Although most of the large tubers of Pentland Dell
were well below the safety limit of 20 mg per 100 g
FW, tubers in the range of 20–40 g FW had TGA
Effect of tuber position within a plant concentrations above or just below this. In contrast
The eþect of tuber position within a plant on glyco-
alkaloid concentration was investigated for mature
plants of cultivars Pentland Dell and Estima har-
vested in the 1994 year crop and positional diagrams
were produced. TGA concentration in each tuber of
individual plants is shown in positional diagrams
(Fig 2) for Pentland Dell and Estima harvested in
1994. There was no obvious eþect of the position of
the tuber within each plant on either tuber size or
glycoalkaloid concentration. Tubers of similar size
were present in the top and the bottom of the plant
proüle and had similar glycoalkaloid levels.
However, tuber size and TGA concentration were
related as generally the smaller tubers had the higher
concentrations of TGA for both cultivars (Fig 2).
Stolons were detached from most tubers of cultivar
Estima whereas only a small number of tubers within
each plant of Pentland Dell were detached from their
stolons.

Relationship with tuber size

TGA concentration in both cultivars decreased as
the tubers enlarged up to about 50 g FW (Fig 3(a)).
There was little change with further increase in size,
with concentrations remaining higher for Pentland
Dell than Estima. The highest TGA concentration
Figure 3. Plots of (a) TGA concentration (mg per 100 g FW) and
(b) TGA content per tuber (mg per tuber) vers us tuber FW for
individual tubers of cultivars Pentland Dell (…) and Es tima (>)
harves ted in 1995. Each point repres ents one tuber.

34 J Sci Food Agric 79 :32–36 (1999)


Table 1. Tuber fres h weight (g) at which the concentration of
a-s olanine, a-chaconine and TGA is half the overall s afety limit
(ie 10 mg per 100 g FW) for cultivars Pentland Dell and Es tima
harves ted in 1994 and 1995
Harves t year Cultivar SE
Pentland Dell Es tima
1994
a-Solanine 10.2 3.8 1.6
a-Chaconine 23.7 9.3 3.9
TGA 51.4 14.3 6.7
1995
a-Solanine 15.2 3.8 0.9
a-Chaconine 46.8 9.9 1.3
TGA 113.2 15.1 7.1
a NS, not s ignificant ; ** P O 0.01 ; *** P O 0.001.
to Pentland Dell, even the smallest tubers of Estima
had TGA concentrations well below the safety limit.
Patterns were very similar in 1994 (data not shown)
and 1995. The tuber FW at which the a-solanine,
a-chaconine and TGA concentration was half the
overall safety limit (ie 10 mg per 100 g FW) appeared
to diþer between the cultivars in both years but the
diþerence was statistically signiücant only in the
1995 harvest (Table 1).
When glycoalkaloid content per tuber was plotted
against tuber size for each cultivar, a steady increase
of TGA content with increasing tuber size was
evident (Fig 3(b)). A cultivar diþerence in the
relationship between tuber FW and TGA content
was observed in both harvest years but regression
analysis showed a statistically signiücant diþerence
only in the 1995 sampling.
DISCUSSION AND CONCLUSIONS
Mean tuber glycoalkaloid concentration was much
higher in plants of cultivar Pentland Dell than in
plants of cultivar Estima in both 1994 and 1995. Dif-
ferences in glycoalkaloid levels of Pentland Dell and
Estima have already been shown not only in mature
tubers19 but also in immature tubers.20 Concentra-
tions for Estima were similar in 1994 and 1995 but
Pentland Dell had higher TGA concentration
(averaged across all tubers per plant) in 1995 than in
1994. The cause of this diþerence between years is
not known but could have been the unusually hot
and dry conditions in 1995. The experimental trial
was not irrigated and consequently the plants fre-
quently showed symptoms of stress. Eþects of stress
on TGA levels in Pentland Dell but not in Estima
would be consistent with previous reports that culti-
vars with genetically determined high levels of gly-
coalkaloids are proportionally responsive to adverse
conditions and accumulate excessive levels in their
tubers more easily than do cultivars with low levels.1
Pentland Dell also showed greater within plant varia-
tion in glycoalkaloid concentration than Estima,
J Sci Food Agric 79 :32–36 (1999)
Variation in glycoalkaloid concentration of potato tubers
especially in 1995. The tuber size range (2–300 g
FW) was similar for the two cultivars in 1995 but
Pentland Dell had considerable numbers of very
small tubers with very high TGA concentrations
(Fig 3(a)). The box and whisker plots showed that
Pa
six of the eight Pentland Dell plants sampled in 1995
had tubers with TGA levels of more than twice the
safety limit of 20 mg per 100 g FW.
NS The position of the tuber within a plant could have
NS contributed to the within plant variation observed as
NS upper tubers might be exposed to diþused sunlight
through loose soil and thus have higher glycoalkaloid
levels than the tubers at the bottom of the plant
**
*** proüle. There was no relationship found between
** glycoalkaloid concentration of individual tubers and
their position within plants of cultivars Pentland
Dell and Estima. This contrasts with a previous
report21 of diþerences in tuber glycoalkaloid levels
relating to depth of soil cover. The authors suggested
that greening of the upper tubers due to sunlight
exposure was the main reason for the glycoalkaloid
diþerences. However, in the current study plants
were all earthed up and tubers with any green dis-
colouration were discarded before glycoalkaloid
determination. Glycoalkaloid levels could have
increased without any visible greening of the tuber
but that was not found to be the case. From the
above it seems that the position of the tuber within a
plant is not an important factor for tuber glycoalka-
loid levels as long as tubers are protected from sun-
light by sufficient soil cover to prevent any greening.
Small size and immaturity of tubers have often
been associated with higher glycoalkaloid levels.5,13
The individual tubers in this investigation were har-
vested late in the season, so that immaturity eþects
would be minimised and tuber size eþects could be
evaluated. An inverse relationship was found
between tuber size of individual tubers and TGA
concentration for both Pentland Dell and Estima; as
tuber size increased the TGA concentration of the
individual tubers generally decreased and then
leveled oþ. Similar patterns of decrease were report-
ed for samples taken from commercial packs of culti-
var Bintje12 and of several other commercial
cultivars.22 Although the higher TGA concentration
in the very small tubers should be expected consider-
ing that glycoalkaloids are concentrated mostly in the
peripheral layers,3 reviewers have stated that gly-
coalkaloid accumulation occurs only in the early
stages of tuber development and that the glycoalka-
loids are diluted as tuber size increases.5,23 However,
the approximately constant TGA concentrations in
larger tubers indicate that the changes in glycoalka-
loid levels of individual tubers are not due to a
simple dilution eþect. This is supported by the plot
of tuber fresh weight against TGA content (Fig 3(b))
which demonstrated that glycoalkaloid accumulation
generally continued as tuber size increased. Trends
of TGA accumulation were diþerent between the
two cultivars studied. Estima showed a gradual
35
F Papathanasiou, SH Mitchell, BMR Harvey
increase in glycoalkaloid content with increasing
tuber size whereas Pentland Dell had a much steeper
increase. Similarly, Wolf and Duggar24 reported a
rise of TGA content per tuber with increase in tuber
size for cultivar White Rural but only up to a certain
weight. Cultivars with only gradual accumulation of
TGA content as tuber size increases or a complete
cessation of accumulation after a certain weight are
desirable especially when tubers are harvested imma-
ture.
Considering that a need to lower the recommend-
ed safety limit of 20 mg per 100 g FW has been
expressed,4,8 the tuber size at which the TGA con-
centration was half the safety limit (ie 10 mg per
100 g FW) was calculated. For Pentland Dell, tubers
below 52 g FW in 1994 and 114 g FW in 1995 had
TGA levels above 10 mg per 100 g FW, whereas for
cultivar Estima only tubers below 15 g FW had TGA
levels above half the safety limit. These results show
that if lower safety limits are recommended market-
able tubers of Estima would pose no danger to the
consumer, even when eaten with the skin but, for
Pentland Dell, only big tubers would be considered
safe for consumption. Furthermore, for cultivar
Pentland Dell individual tubers below 40 g FW had
TGA concentrations above or just below 20 mg per
100 g FW. Consumption of such small tubers (20–
40 g FW) as ‘baby’ potatoes, which may be eaten
unpeeled could lead to the exposure of the consumer
to relatively high glycoalkaloid levels.
The results presented here indicate that TGA con-
centration of individual tubers is dependent both on
cultivar and tuber size. A relationship between tuber
size and TGA concentration was found and it was
shown that the decrease of glycoalkaloid concentra-
tion as tubers enlarge is not simply a dilution eþect,
as accumulation appears to continue with increasing
tuber size. Since tubers in this study were harvested
at maturity, further work is required to determine
patterns of glycoalkaloid accumulation during tuber
development.
ACKNOWLEDGEMENTS
The authors thank Dr Ethel White and Mrs Carol
Hall at the Plant Testing Station, Department of
Agriculture for Northern Ireland for advice and
access to the cultivar trials.
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