Euphytica (2012) 183:119131

DOI 10.1007/s10681-011-0520-0

Genotypic variability and genotype by environment

interactions for inulin content of Jerusalem artichoke
germplasm
Ratchanee Puttha Sanun Jogloy
Preeya P. Wangsomnuk Supalax Srijaranai

Thawan Kesmala Aran Patanothai

Received: 19 May 2011 / Accepted: 18 August 2011 / Published online: 3 September 2011
Springer Science+Business Media B.V. 2011

Abstract Genetic variations in germplasm and
genotype 9 environment interaction (G 9 E) are
important for crop improvement. The objectives were
to explore genetic variation in Jerusalem artichoke
germplasm and to evaluate G 9 E interaction for
inulin content. Seventy-nine accessions of Jerusalem
artichoke were evaluated in a randomized complete
block design with two replications for three seasons.
Significant variation in inulin content (55.374.0%
dry weight) was observed and the genotypes with
high inulin content could be identified although there
was intermediate G 9 E interaction. Genotypes were
also significantly different for days to maturity, fresh
tuber yield, biomass and harvest index and G 9 E
interactions for these traits were also significant.
The correlation between inulin content and days to
maturity was not significant (r = -0.20), whereas
inulin content and fresh tuber yield were significantly
R. Puttha  S. Jogloy (&)  T. Kesmala  A. Patanothai
Department of Plant Science and Agricultural Resources,
Faculty of Agriculture, Khon Kaen University,
Khon Kaen 40002, Thailand
e-mail: sanun@kku.ac.th
P. P. Wangsomnuk
Department of Biology, Faculty of Science,
Khon Kaen University, Khon Kaen 40002, Thailand
S. Srijaranai
Department of Chemistry, Faculty of Science,
Khon Kaen University, Khon Kaen 40002, Thailand
associated (r = 0.22). JA 37 and CN 52867 are
promising for high yield and inulin content.
Keywords Biomass  Days to maturity 
Helianthus tuberosus L.  Tuber yield
Introduction
Inulin, a highly water-soluble carbohydrate consisting
of oligomer of 2?1-linked b-D-fructose ending in an
a-D-glucose unit (Shu 1998; Roberfroid 1999; Hellw-
ege et al. 2000), is found at high concentration in the
roots or tubers of various crop in Asteraceae family
(Moerman et al. 2004; Muir et al. 2007) and it is also
found in a traceable amount in various fruit and
vegetable crops (Niness 1999; Muir et al. 2007).
Inulin is largely utilized as dietary fiber in food
(Niness 1999; Stevens et al. 2001; Cherbut 2002).
Degree of polymerization (DP) of inulin varies from
2 to 60 depending on plant species (Prosky and
Hoebregs 1999), and the degree of polymerization is
about 10 in Jerusalem artichoke (Raccuia and Melilli
2010). Inulin is most applicable as fat replacer
(Kocsis et al. 2007b). In the large intestine, inulin is
fermented by the intestinal microflora and stimu-
lates growth of the bifidobacteria and lactobacillus
(Kleessen et al. 2007; Roberfroid 2007a). Moreover,
a large number of animal studies (Zaky 2009) and
preliminary human (Davidson and Maki 1999) data
show that dietary inulin reduce serum cholesterol and

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the risk of heart disease. Because of its health
promoting properties, inulin is important as func-
tional food (Milner 1999; Roberfroid 2007b), food
industries (Azis et al. 1999; Danilcenko et al. 2008),
nutritional composites (Coussement 1999) and med-
icines in food and pharmaceutical industries (Monti
et al. 2005; Valluru and Van den Ende 2008).
Jerusalem artichoke (Helianthus tuberosus L.) is a
valuable source of inulin (Azis et al. 1999; Muir et al.
2007). Its tubers contain 20.431.9% of dry matter,
from which carbohydrates are the main component.
Most of carbohydrates consist of water-soluble inulin,
reaching 5075% of dry matter or 730% of fresh
mass of the tubers (Danilcenko et al. 2008; Kays and
Nottingham 2008). Inulin is primarily stored in the
tubers, but temporary storage also occurs in the stems
prior to tuber filling (Monti et al. 2005; Kays and
Nottingham 2008). The two enzymes: sucrose:
sucrose 1-fructosyltransferase (1-SST) and fructan:
fructan 1-fructosyltransferase (1-FFT) involve in
inulin bio-synthesis (Hellwege et al. 2000; Valluru
and Van den Ende 2008).
High content of inulin and a peculiarly high tuber
yield allow Jerusalem artichoke to be designated as a
fructan crop with a bright future (Kocsis et al.
2007b), and it is a promising candidate for inulin
production in the tropical climates (Baldini et al.
2004). Breeding program has initiated at Khon Kaen
University and focused on the improvement of tuber
yield (Pimsaen et al. 2010). In a pioneer work of the
project, high interactions between genotype and
environment (G 9 E) were observed for tuber yield,
tuber number and tuber size and multi-location trials
are necessary to identify the best genotypes (Pimsaen
et al. 2010).
Other works carried out under temperate climates
also reported genotypic differences and significant
G 9 E interactions for tuber yield, tuber number,
tuber size (Berenji and Sikora 2001), nutrients in the
forage of Jerusalem artichoke (Seiler and Campbell
2004, 2006), maturity, female fertility, chlorophyll
intensity and disease susceptibility in hydroponics
(Serieys et al. 2010). Terzic and Atlagic (2009) found
variability in Jerusalem artichoke tubers for nitrogen
and sugar content (inulin 80%).
Harvest time affected inulin content and degree
of polymerization of Jerusalem artichoke (Kocsis
et al. 2007a). Kocsis et al. (2007b, 2008) found
that cultivars, amount of rainfall, rain distribution,
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Euphytica (2012) 183:119131
weather and irrigation also affected stalk dry weight,
tuber dry weight, sugar yield, inulin yield in Jerusa-
lem artichoke grown in semi-arid areas.
The information on the variation in inulin content
in a large number of Jerusalem artichoke accessions
is rather limited and the extent to which the cultivars
interact with the environments has not been ade-
quately researched. The objectives of this study were
to explore genetic variability for inulin content in a
large number of Jerusalem artichoke germplasm and
to evaluate genotype 9 environment interaction for
inulin content. The information would enable breed-
ers to make informed decisions about suitable parents
for high inulin content in Jerusalem artichoke breed-
ing program.
Materials and methods
Soil properties and weather data
Soils were taken from ten positions in each site at
the depths of 015 and 1530 cm from soil surface.
The soils were analyzed for chemical and physical
properties. Rainfall and maximum and minimum air
temperatures during the three seasons were recorded
daily from planting until harvest by a meteorological
station located near the experimental site.
Jerusalem artichoke gemplasm and crop
management
Seventy-nine accessions of Jerusalem artichoke used
in this study were kindly donated from three institu-
tions. They included one accession from the North
Central Regional Plant Introduction Station (NCR-
PIS), USA, 25 accessions from the Leibniz Institute
of Plant Genetics and Crop Plant Research (IPK) of
Germany and 53 accessions from the Plant Gene
Resource of Canada (PGRC) (Table 1).
The experiment was undertaken at the experimen-
tal farm of the Faculty of Agriculture, Khon Kaen
University, Khon Kaen, Thailand (Lat 16270 N, Long
102480 W, 179 masl). The 79 accessions were laid
out in a randomized complete block design with two
replications for three seasons in two late rainy
seasons (SeptemberDecember 2008 and 2009) and
one early rainy season (MarchJuly 2009). The plot
size was four-row plot with 5 m long with spacing of
Euphytica (2012) 183:119131 121

Table 1 Jerusalem
Entry no. Accession no. Name of accession Origin Genetic resources
artichoke accessions,
sources of origin and 1 JA 1 7305 Canada PGRC
genetic resources
2 JA 2 7306 Canada PGRC
3 JA 3 7307 Canada PGRC
4 JA 4 7308 Canada PGRC
5 JA 5 7309 Canada PGRC
6 JA 6 7310 Canada PGRC
7 JA 7 7312 Canada PGRC
8 JA 8 7512 Canada PGRC
9 JA 9 7513 Canada PGRC
10 JA 10 HM Hybrid A Canada PGRC
11 JA 11 HM Hybrid B Canada PGRC
12 JA 12 HM Hybrid C Canada PGRC
13 JA 13 HM-2 Canada PGRC
14 JA 14 HM-3 Canada PGRC
15 JA 15 HM-5 Canada PGRC
16 JA 16 HM-7 Canada PGRC
17 JA 18 HM-9 Canada PGRC
18 JA 20 HM-11 Canada PGRC
19 JA 21 HM-12 Canada PGRC
20 JA 23 DHM-3 Canada PGRC
21 JA 25 DHM-5 Canada PGRC
22 JA 30 DHM-16 Canada PGRC
23 JA 35 W-97 Canada PGRC
24 JA 36 W-106 Canada PGRC
25 JA 46 DHM-14-3 Canada PGRC
26 JA 47 DHM-14-6 Canada PGRC
27 JA 55 USA PGRC
28 JA 58 Intress USSR PGRC
29 JA 59 Volzskij-2 USSR PGRC
30 JA 60 Jamcovskij krashyj USSR PGRC
31 JA 61 Vadim USSR PGRC
32 JA 69 TUB-346 USD-ARS-SR USA PGRC
33 JA 70 TUB-365 USD-ARS-SR USA PGRC
34 JA 71 TUB-675 USD-ARS-SR USA PGRC
35 JA 72 TUB-676 USD-ARS-SR USA PGRC
36 JA 75 #2 Canada PGRC
37 JA 76 #4 Canada PGRC
38 JA 77 #5 Canada PGRC
39 JA 92 Industrie USSR PGRC
40 JA 93 Leningradskii (NC10-65) USSR PGRC
41 JA 97 D19-63-340 France PGRC
42 JA 108 83-001-3 (37 9 6) Canada PGRC
43 JA 109 83-001-4 (37 9 6) Canada PGRC
44 JA 114 83-001-9 (37 9 6) Canada PGRC
45 JA 120 83-003-1 (6 9 20) Canada PGRC

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122 Euphytica (2012) 183:119131

Table 1 continued
Entry no. Accession no. Name of accession Origin Genetic resources

46 JA 122 83-004-2 (6 9 20) Canada PGRC

47 JA 132 83-007-2 (69 9 3) Canada PGRC
48 CN 52867 PGR-2367 USSR PGRC
49 JA 37 Comber Canada PGRC
50 JA 38 B.C. #1 Canada PGRC
51 JA 67 Oregon White USA PGRC
52 JA 89 Waldspindel France PGRC
53 JA 102 073-87 Germany PGRC
54 HEL 53 Germany IPK
55 HEL 61 Tambovskij Krasnyi Russian Federation IPK
56 HEL 62 Sachalinskij Krasnyi Russian Federation IPK
57 HEL 65 Sejanec 19 Russian Federation IPK
58 HEL 66 Kievskij Belyj Ukraine IPK
59 HEL 68 Unknown IPK
60 HEL 69 Unknown IPK
61 HEL 231 Germany IPK
62 HEL 335 Unknown IPK
63 Ames 2729 TUB-49 South Dakota NCRPRIS
64 HEL 243 Bianka Germany IPK
65 HEL 246 Unknown IPK
66 HEL 248 Rote Zonenkugel Germany IPK
67 HEL 250 Medius France IPK
68 HEL 253 Unknown IPK
69 HEL 256 Unknown IPK
70 HEL 265 BT4 Hungry IPK
71 HEL 272 D19-63-340 France IPK
72 HEL 278 Voelkenroder Spindel Unknown IPK
Kays and Nottingham 73 HEL 280 BS-83-22 Unknown IPK
(2008); NCRPIS the North 74 HEL 288 RAI Poland IPK
Central Regional Plant 75 HEL 293 RA9 Poland IPK
Introduction; IPK the
Leibniz Institute of Plant 76 HEL 308 Unknown IPK
Genetics and Crop Plant 77 HEL 315 Unknown IPK
Research of Germany; 78 HEL 316 Unknown IPK
PGRC the Plant Gene
79 HEL 317 Unknown IPK
Resource of Canada

0.5 m between rows and 0.5 m between plants within
rows.
Soil was ploughed once using a 3-disc tractor and
twice using a 7-disc tractor and ridged at a distance of
1 m. Then, the ridges were leveled to make soil beds.
Sprouted seed tubers were used as planting materials.
To prepare the sprouted seed tubers, the tubers were
cut into small pieces each of which had 23 buds/
piece. The tuber pieces were incubated in plastic bags
containing moistened coconut peat at the bottom and
the top of the bags about 7 days under ambient
conditions. The plastic bags were kept opened for
good aeration. The tuber pieces with active buds and
roots were further transferred to plug plastic trays
containing a mixture 1:1 soil:burnt rice husk medium
about 7 days for germination. The two-leaf sprouted
seedlings were then suitable for transplanting in the
field. The crop was well managed for optimum
growth and yield. Weeds were controlled with single
manual weeding at 1 month after transplanting and

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Euphytica (2012) 183:119131
when it grown out. Supplementary fertilizer was
applied at 1 month after transplanting at the rate of
156.3 kg/ha of NP2O5K2O formula 151515. shaken well, kept at ambient conditions to reduce
A Terraclor (quintozene 24% W/V EC) was applied
monthly for 3 months after transplanting at the rate
25 ml/20 l of water for control of stem rot (Sclero-
tium rolfsii). Experimental fields were uniformly
sprinkler-irrigated.
Data collection
Maturity was determined by the browning of the
stems and defoliation. This is a sign indicating the
complete translocation of reserved assimilates to
the tubers and appropriate time for harvest, and days
to maturity was recorded. At harvest, plants at the end
of the rows and border rows were discarded. Then,
five plants in each plot were sampled randomly and
used for determination of tuber yield, biomass,
harvest index (HI) and inulin content.
The stems were cut at the crowns, tubers were dug
and roots were discarded. Fresh shoot and tuber
weight were recorded in the field. A random shoot
sample of 100 g and fresh tuber sample of 200 g for
each plot was taken, oven dried at 80C for 48 h or
until constant weight, and weighed. Biomass was
calculated from shoot and tuber dry weight. Harvest
index (HI) was calculated as the ratio of tuber dry
weight divided by biomass.
Sample preparation and analysis of inulin content
At harvest, five plants for each accession were
randomly collected and all fresh tubers were washed
to eliminate soil. The sample preparation and inulin
analysis has been reported previously (Saengkanuk
et al. 2011). Briefly, the tubers were sliced longitu-
dinally into thin pieces at the middle of the tubers.
Fifty grams of sliced tuber were soaked in 100 ml
absolute ethanol at 10C for 24 h to prevent enzy-
matic degradation of inulin in the tubers. Ethanol
solution was discarded and the samples were imme-
diately stored at -20C in sealable plastic bags until
analysis.
The samples were oven-dried at 60C for 10 h or
until constant weight. The dried samples were milled
and kept in desiccators until inulin extraction. For
inulin analysis, the sample of each plot was divided
into two grams for three replications, and inulin was
123
extracted by reverse osmosis water in a 25 ml flask at
80C for 20 min on a water bath. The samples were
heat and then treated by filtering the solution through
a Whatman no. 1 filter circle followed by immediate
analysis. The extract of 500 ll was transferred to a
25 ml volumetric flask, hydrolyzed with 750 ll of
hydrochloric acid (3% v/v HCl). The volume was
adjusted with reverse osmosis water to 25 ml and
heated at 97 2C for 45 min on a hot plate and
allowed to cool to room temperature before analysis.
If analysis could not be undertaken immediately, the
samples were stored at 4C in plastic bottles. Fructose
was determined by spectrophotometry using perio-
date reaction (Saengkanuk et al. 2011). Inulin content
was calculated based on fructose, ignoring trace
amount of glucose and reducing free fructose,
glucose and sucrose. The inulin analysis results are
expressed as a percentage of inulin content on dry
weight basis.
Statistical analysis
Analysis of variance was performed for individual
seasons and error variances were tested for homoge-
neity (Gomez and Gomez 1984). Combined analysis
of variance of three-season data was performed for
days to maturity, fresh tuber yield, biomass, HI, and
inulin content in 79 Jerusalem artichoke accessions.
Although error variances were homogenous, there
were significant genotype 9 environment interac-
tions for all characters, and data of individual seasons
were reported. Duncans multiple range test (DMRT)
was used to compare mean differences. All calcula-
tions were done using computer software MSTAT-C
(Bricker 1989). The relationships among traits were
calculated using simple correlation by using STA-
TISTIX8 software program (Statistix8 2003).
Results and discussion
Soil properties and weather data
The soils in the experiment were sandy and infertile.
The chemical and physical properties were slightly
different among experimental sites (Table 2). The
soils in the late rainy season 2008 and 2009 trials
were higher in electrical conductivity (EC) and
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124 Euphytica (2012) 183:119131

Table 2 Chemical and physical characteristics of the soil in the experimental fields
Fields Depth pH EC CEC OM Total Available Exchangeable Particle size, mun (USDA) Texture
(cm) (1:1 H2O) (1:5 H2O) (c mol/kg) (%) N (%) P (mg/kg) class
(dS/m) K Ca Sand: Silt: Clay:
(mg/kg) (mg/kg) 2.00.05 0.050.002 \0.002
(%) (%) (%)

A9 015 6.8 0.03 4.40 0.53 0.029 35.3 41.2 375 87.9 8.1 4.1 Sand
1530 6.9 0.03 4.37 0.49 0.027 35.6 40.2 410 86.9 7.9 5.2 Sand
A 13 015 7.6 0.11 3.48 0.65 0.036 89.4 57.2 500 88.8 7.9 3.4 Sand
1530 7.7 0.08 3.45 0.55 0.030 67.7 41.7 430 87.7 8.0 4.3 Sand

A 9 is the experimental field for early rainy season 2009 and A 13 is the experimental field for late rainy season 2008 and 2009

available P than the soil in the early rainy season
2009 trial. However, the EC values in all seasons
were in lower range (B0.2 dS/m) (Geng-Mao et al.
2008), indicating that there are not enough nutrients
available for optimal plant growth, whereas available
P values were sufficient ([15 mg/kg) in all seasons
(Lebot 2009). Potassium values were intermediate
and nitrogen values were low. As the nutrient values
felled into the same ranges, the differences in
nutrients among seasons would not cause significant
differences in inulin content and other parameters.
Maximum temperatures (T-max) and minimum
temperatures (T-min) were slightly different among
seasons. T-max in late rainy seasons were 30.5C in
2008 and 32.5C in 2009 (Fig. 1a, c) and T-max in
the early rainy season was 33.9C (Fig. 1b). T-min
values in late rainy seasons were 21.7C in 2008 and
22.0C in 2009, and T-min in the early rainy season
was 25.3C. As Jerusalem artichoke in this study was
grown in the tropics, the growing temperatures were
much higher than optimum temperatures reported
previously, most Jerusalem artichoke cultivars
require average annual temperatures of between 6
and 26C, within a growing season of at least 125
frost-free days (Kays and Nottingham 2008).
Jerusalem artichoke can grow under tropical
conditions, but yields are often suboptimal (Kays
and Nottingham 2008). Plants are much smaller and
mature earlier, yielding smaller and fewer tubers than
in temperate regions (Kays and Nottingham 2008).
Tuber yield of the accession BT4 (HEL 265 in this
study) could be as high as 3590 g/plant under
temperate conditions (Berenji and Sikora 2001), but
its yield was much lower (258 g/plant) under tropical
conditions (data not reported). Days to maturity were
similar in ranks for Jerusalem grown under tropical
and temperate conditions. HEL 243 (Bianka) was
early mature and JA 89 (Waldspindel) was rather late
(Kocsis et al. 2007a). In this study, HEL 243 was
early mature (107 days), whereas JA89 was rather
late mature (110 days) (data not reported). The
maturity range reported for these Jerusalem artichoke
accessions was 104116 days in this study.
Rainfalls in the late rainy seasons were 619.7 mm
in 2008 and 204.1 mm in 2009 (Fig. 1a, c), and
rainfall in the early rainy season 2009 was 601.1 mm
(Fig. 1b). Rain distribution in the late rainy season
2008 extended to mid growing season, whereas rain
distribution in the late rainy season 2009 was limited
to early growth stages. However, drought was not a
problem because irrigation was available throughout
the crop cycle. In contrast, the rainfall well distrib-
uted throughout the crop cycle in the early rainy
season 2009.
Under rained conditions, rainfall and rain distri-
bution greatly affect growth and yield of Jerusalem
artichoke (Kocsis et al. 2008). However, photoperi-
ods and temperatures also play an important role in
the differences in yield, biomass, harvest index, and
inulin content in Jerusalem artichoke (Kays and
Nottingham 2008; Raccuia and Melilli 2010).
Genotypic variability
and genotype 9 environment interactions
Seasons were significantly different for days to
maturity, fresh tuber yield, biomass and harvest
index (HI) except for inulin content (Table 3). This
indicated that inulin content was more stable than
other characters across seasons. This information is
very useful for selection of parents in Jerusalem
artichoke breeding for high inulin content.

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Euphytica (2012) 183:119131 125

Fig. 1 Rainfall (mm),

maximum temperature
(T-max) and minimum
temperature (T-min) (8C)
during the crop growth
period of 79 Jerusalem
artichoke accessions in late
rainy season 2008 (a), early
rainy season 2009 (b) and
late rainy season 2009 (c)

Tuber yield and harvest index were higher in the
late rainy seasons for both years, whereas days to
maturity and biomass were higher in the early rainy
season in 2009 (Table 4). The variations in genotype,
season, location, drought stress, harvest date (Kocsis
et al. 2007a, b, 2008), soil properties (Geng-Mao
et al. 2008), temperatures (Raccuia and Melilli 2010),
salt tolerance (Xiao-Hua et al. 2010) and cropping
practices (Rodrigues et al. 2007) could affect these
agronomic traits and inulin content.
Jerusalem artichoke accessions were significantly
different for all characters, and there were significant
genotype 9 environment interactions for these char-
acters (Table 3). The results showed genetic vari-
ability for these characters and also indicated the
need for multi-location trials to identify superior

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126 Euphytica (2012) 183:119131

Table 3 Mean squares and proportion of sum of squares to artichoke accessions in late rainy season 2008, early rainy
total sum of squares for days to maturity, fresh tuber yield, season 2009 and late rainy season 2009
biomass, harvest index (HI) and inulin content of 79 Jerusalem
Source of variation df Days to maturity Fresh tuber yield Biomass HI Inulin content

Environment (E) 2 14647.6** (0.72) 217425.1** (0.04) 278406.6** (0.21) 8.22** (0.66) 302.3ns (0.05)
Rep within environment 3 59.3 (0.00) 1378.2 (0.00) 710.6 (0.00) 0.03 (0.00) 87.4 (0.02)
Genotypes (G) 78 83.9** (0.16) 78993.1** (0.54) 17143.4** (0.51) 0.04** (0.14) 92.7** (0.56)
G9E 156 27.9** (0.11) 25380.8** (0.35) 3471.4** (0.21) 0.02** (0.15) 20.9** (0.25)
Error 234 0.4 (0.00) 3231.0 (0.07) 708.8 (0.06) 0.01 (0.06) 6.7 (0.12)
CV (%) 0.6 20.2 19.0 14.0 4.0
Numbers within the parentheses are proportion of sum squares to total sum of squares
ns, ** Non significant and highly significant at P \ 0.01 probability levels, respectively

Table 4 Days to maturity, fresh tuber yield, biomass, harvest index (HI) and inulin content for averaged over 79 Jerusalem artichoke
accessions
Seasons Days to maturity Fresh tuber Biomass HI Inulin content
(days) yield (g/plant) (g/plant) (% dry wt)

Late rainy season 2008 103 3.3b 296.7 109.7a 115.3 43.0b 0.71 0.1a 67.0 4.5
Early rainy season 2009 118 6.2a 239.0 169.3b 188.4 83.3a 0.30 0.2b 65.3 4.5
Late rainy season 2009 100 4.5b 308.2 155.5a 116.1 57.1b 0.67 0.1a 64.3 5.2
F-test ** ** ** ** ns
Data are presented as mean SD (n = 3), values with different letters within the same column are significantly different at P \ 0.05
by DMRT
** Significant at P \ 0.01 probability level

genotypes. It is clear that environment contributed to
the small portion of variations in fresh tuber yield and
inulin content (45%), whereas genotype contributed
a larger portion to the variations (5456%). A large
estimate of genetic variance indicates that selection
and breeding initiatives can proceed immediately
(Ordonez et al. 2005). The contribution of geno-
type 9 environment interaction was higher than that
of environment but it was still lower than the
contribution of genotypic differences in fresh tuber
yield and inulin content (Table 3).
In contrast to this study, the environment contrib-
uted to a large portion of variations in tuber fresh
weight, whereas genotype and genotype 9 location
contributed a smaller portion to variations (Pimsaen
et al. 2010). The environment, accession and geno-
type 9 environment interaction were highly signifi-
cant for tuber yield per plant (Berenji and Sikora
2001; Baldini et al. 2004). The results both supported
and contrasted to those in previous studies possibly
largely due to materials used and the differences in
environments.
The variations in biomass as affected by environ-
ment and genotype 9 environment differences were
significant but smaller than the variation of genotype.
However, days to maturity and HI were mainly
affected by the variations caused by season.
Figure 2 showed patterns of genotype 9 environ-
ment interactions of high and low groups of Jerusa-
lem artichoke accessions for inulin content. The high
and low groups were separated to reduce confounding
effect in genotypes with intermediate performance.
The interactions within high or low groups were high,
whereas the interactions between groups were rather
low. Therefore, high and low groups were clearly
separated except for few accessions, but identification
of Jerusalem artichoke accessions for high or low
inulin content within high or low groups was difficult.
Previous study in 37 cultivars of Jerusalem artichoke
could identify the genotypes with high fructose
content (Grando, Kharkov, C-122, Meillo and Dubo)
(Ben Chekroun et al. 1996). However, there acces-
sions were not included in this study. Inulin contents
in different maturity classes were also studied in

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Euphytica (2012) 183:119131
Fig. 2 Selected eighteen accessions ranked in the top low and
high inulin content for late rainy season 2008, early rainy
season 2009 and late rainy season 2009
Jerusalem artichoke (Kocsis et al. 2007b, 2008) and
two accessions (Bianka and Waldspindel) are com-
mon to this study (HEL 243 and JA 89, respectively)
(Kocsis et al. 2007b). Inulin content in Bianka
was moderate (63.5%) but it was low (48.7%) in
Waldspindel (Kocsis et al. 2007b). Inulin content
(65.3%) in HEL 243 was similar to that reported
previously, whereas higher inulin content (69.3%)
was observed in JA 89.
As the objective of this study was to identify
Jerusalem artichoke genotypes with high or low
inulin content, the accessions were then divided into
two extreme groups each of which has nine acces-
sions (Table 5). The high group consisted of JA 37,
JA 6 JA 15, JA 67, JA 70, JA 59, CN 52867, JA 97
and JA 30 and the values of inulin content ranged
from 70.1 to 74.0%. The low group comprised of JA
11, JA 35, JA 55, JA 14, HEL 288, HEL 335, HEL
62, JA 58 and JA 4 with the range of 55.3 to 59.9%.
The ranks in the two groups were not overlapped
and statistically different, and, therefore, the geno-
types with low and high inulin content were readily
identified.
Days to maturity ranged between 104 and 116 days
(Table 5). The ranges between high and low groups
were overlapped. The lack of significant correlation
127
(0.20, P [ 0.05) between these characters was
observed (Table 6). The relationship between days to
maturity and inulin content was not significant, indicat-
ing independent segregation of these traits. Days to
maturity in this study (104116 days) were much
shorter than those in other studies (Ordonez et al.
2005; Kocsis et al. 2007b). The difference in days to
maturity among different studies indicated the differ-
ential response of Jerusalem artichoke to different
environments.
A wide range of tuber yield was observed among
Jerusalem artichoke accessions, ranging from 77.9 to
730.5 g/plant (Table 5). However, the ranks of high
and low groups were overlapped, showing the inde-
pendency between these traits. The correlation
between these characters was low although it was
significant (0.22, P B 0.05). High variation was
observed for tuber yield and selection for this character
is possible. Jerusalem artichoke tubers are also variable
in size, color and shape (Vasic et al. 2002). Tuber size
and tuber number are important components that affect
tuber yield (Pimsaen et al. 2010).
Similar to tuber yield, a wide range of biomass
was also observed, ranging from 39.5 to 277.1 g/plant
(Table 5). The ranks of high and low groups were
overlapped, indicating the independency between
these traits. The correlation between these traits was
also not significant (0.15, P [ 0.05). Variation in
biomass was high and selection for this character is
possible. The high variation of biomass was respon-
sible to variation of tuber yield. Growth, yield and
compositional characteristics of Jerusalem artichoke
have tendentious influence to biomass production
(Kiru and Nasenko 2010).
Harvest index ranged from 0.3 to 0.7, and the
values were overlapped between low and high
groups, showing some extent of independency
between these traits (Table 5). The correlation coef-
ficient between inulin content and harvest index was
low although it was significant (0.22, P B 0.05)
(Table 6). Harvest index and days to maturity were
negatively associated (-0.43, P B 0.01), whereas
harvest index and fresh tuber yield were positively
associated (0.31, P B 0.01). However, the correlation
between harvest index and biomass was not signif-
icant (-0.07, P [ 0.05). The high HI indicated the
ability of accession to produce tubers. Jerusalem
artichoke is a crop with exceptionally high harvest
index (0.640.78) and shows high ability to produce
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 128

123
Table 5 Selected eighteen accessions ranked in the top nine high and low for inulin content, days to maturity, fresh tuber yield, biomass and harvest index (HI) for 79 Jerusalem
artichoke accessions averaged over three seasons
Groups No. Accession no. Inulin content (% dry wt) Days to maturity (days) Fresh tuber yield (g/plant) Biomass (g/plant) HI

High 1 JA 37 74.0 1.0a 104 7.9t 474.7 70.6bcd 203.7 87.6cf 0.7 0.2abc
2 JA 6 73.6 0.3ab 104 7.9t 199.5 49.2vb0 116.1 66.7ry 0.5 0.2ek
3 JA 15 71.7 2.0abc 108 13.6klm 228.8 57.2sy 153.8 113.7ir 0.5 0.2fl
4 JA 67 71.6 5.6abc 111 2.0ef 386.1 294.9ej 212.5 35.3cd 0.5 0.4hm
5 JA 70 71.0 4.2ad 104 7.8st 135.7 50.7a0 e0 88 25.3xe0 0.5 0.3fl
6 JA 59 70.8 7.2ae 104 7.8st 214.6 64.0ua0 102.8 51.6uc0 0.5 0.3fl
7 CN 52867 70.8 0.4ae 107 5.8nop 730.5 157.2a 277.1 45.8a 0.7 0.2ae
8 JA 97 70.6 3.2bf 104 7.8st 331.6 17.4lp 131.6 28.1pv 0.6 0.1ae
0 0 0
9 JA 30 70.1 5.4cg 104 7.9t 82.2 90.2vb 54.2 30.3e i 0.5 0.4jm
Low 1 JA 11 59.9 3.2xa0 104 7.8st 85.0 91.5c0 d0 e0 49.9 32.2f0 i0 0.4 0.3lmn
2 JA 35 58.9 1.3yb0 104 7.8st 190.3 100.3a0 e0 93.0 20.2yf0 0.6 0.3gm
3 JA 55 58.6 6.4zc0 104 7.8st 151.7 99.8ye0 43.2 9.4g0 h0 i0 0.6 0.3ek
4 JA 14 58.3 2.1zc0 104 7.9t 155.2 128.6yd0 41.8 21.3h0 i0 0.5 0.3kn
5 HEL 288 57.8 0.2 zc0 110 15.9ghi 344.4 7.4 hn 207.2 102.3cf 0.5 0.2jm
6 HEL 335 56.9 3.1a0 b0 c0 113 13.5c 307.6 120.4js 213.3 79.7c 0.4 0.3jm
7 HEL 62 56.8 3.4a0 b0 c0 115 10.6a 209.5 45.2ua0 138.0 68.6nu 0.5 0.2kn
8 JA 58 56.0 2.4b0 c0 104 7.7st 121.8 56.2b0 e0 63.3 12.7d0 i0 0.5 0.3ek
0
9 JA 4 55.3 3.9c 108 13.7lmn 257.4 259.8pw 147.8 150.0ls 0.5 0.2fl
Max 74.0 116 730.5 277.1 0.7
Min 55.3 104 77.9 39.5 0.3
Mean 65.5 3.9 107 3.7 281.3 114.7 139.9 53.5 0.6 0.1
F-test ** ** ** ** **
Data are presented as mean SD (n = 3), minimum, maximum and mean values were calculated from 79 accessions, values with different letters within the same column are
significantly different at P \ 0.05 by DMRT
** Significant at P \ 0.01 probability level
Euphytica (2012) 183:119131
Euphytica (2012) 183:119131 129

Table 6 Correlation coefficient between days to maturity,

fresh tuber yield, biomass, harvest index (HI) and inulin con-
tent of 79 Jerusalem artichoke accessions in late rainy season
2008, early rainy season 2009 and late rainy season 2009
Characters Days to Fresh tuber Biomass HI
maturity yield

Fresh tuber 0.26*

yield
Biomass 0.54** 0.85**
HI -0.43** 0.31** -0.07 ns
Inulin -0.20 ns 0.22* 0.15 ns 0.22*
content
ns, *, ** Non significant and significant at P B 0.05 and 0.01
probability levels, respectively

tubers (Baldini et al. 2004), but it still has much lower

harvest indices than sugarbeet (HI = 0.8) (Schitten-
helm 1999).
The low harvest index in the early rainy season
2009 compared to the late rainy seasons 2008 and
2009 was possibly due to higher temperature
(25.734.0C) and day lengths (Fig. 1) that promoted
vegetative growth and flowering. Furthermore, the
plants produced prolific rhizomes rather than tubers
(Kays and Nottingham 2008).
Days to maturity was well associated with fresh
tuber yield (0.26, P B 0.05) and biomass (0.54,
P B 0.01), showing dependence of days to maturity
on fresh tuber yield and biomass. Fresh tuber yield was Fig. 3 Relationships between inulin content and days to
well associated with biomass (0.85, P B 0.01), indi- maturity (a), inulin content and fresh tuber yield (b) and fresh
cating that a fresh tuber yield might play a significant tuber yield and days to maturity (c) of 79 Jerusalem artichoke
accessions
contribution to biomass in most accessions.
Figure 3 showed the relationships of inulin content
with days to maturity, fresh tuber yield and the fresh tuber yield and high inulin content. The
relationship between fresh tuber yield and days to genotypes CN 52867, JA 37 and JA 67 are promising
maturity evaluated from 79 Jerusalem artichoke because they had high inulin content and high fresh
accessions grown in three environments. The overall tuber yield.
relationship between inulin content and days to The relationship between fresh tuber yield and
maturity was negative and not significant (-0.20, days to maturity was positive and significant (0.26,
P [ 0.05), indicating that genotypes with early P B 0.05). The results suggested that genotypes with
maturity tended to have higher inulin content than high fresh tuber yield tended to have late maturity.
did genotypes with late maturity. JA 37, JA 6 and JA CN 52867 was interesting because it had intermediate
70 were interesting because they had high inulin maturity and high fresh tuber yield. Low correlation
content and early maturity. also suggested the possibility to select genotypes with
The relationship between inulin content and fresh early maturity and high fresh tuber yield. HEL 278,
tuber yield were positive and significant (0.22, JA 37 and JA 108 meet these criteria. JA 37 was the
P B 0.05). The results indicated that there was most interesting accession because it had high inulin
possibility to select Jerusalem artichoke with high content, high fresh tuber yield and early maturity.

123
130
In conclusion, Jerusalem artichoke accessions
have high variations in fresh tuber yield, biomass
and inulin content and selection for these characters
is possible among these accessions. Selection for
early maturity and HI may be difficult because
of low variations for these characters. Correlation
between fresh tuber yield and inulin content was
positive and significant; suggesting that simultaneous
selection for these traits is possible. JA 37 and CN
52867 were identified as early maturity, high bio-
mass, high harvest index and high yielding with high
inulin content, respectively. These data would enable
breeders to make informed decisions about suitable
parents for Jerusalem artichoke breeding programs.
Acknowledgments The study was funded under the Strategic
Scholarship Program for Frontier Research Network for
the Join Ph.D. Program Thai Doctoral Degree from the
Office of the Higher Education Commission, Thailand.
Grateful acknowledgments are made to the Khon Kaen
University Senior Research Scholar Project of Assoc. Prof.
Dr. Sanun Jogloy under Khon Kaen University Fund. Miss
Araya Saengkanuk is acknowledged for her assistance in inlin
analysis. Department of Chemistry, Faculty of Science,
Khon Kaen University, Thailand is acknowledged for
providing laboratory facilities. The North Central Regional
Plant Introduction Station, USA, the Leibniz Institute of Plant
Genetics and Crop Plant Research, Germany and the Plant
Gene Resource of Canada are acknowledged for their donation
of Jerusalem artichoke germplasm.
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