Hasan 2008

Morphological variability of greater yam
(Dioscorea alata L.) in Malaysia

Sayed M. Zain Hasan
1
*, Andrew A. Ngadin
1
, Ramisah M. Shah
1
and Norizan Mohamad
2
1
Faculty of Agro-technology and Food Science and
2
Faculty of Science and Technology,
Kolej Universiti Sains dan Teknologi Malaysia (KUSTEM), 21030 Mengabang Telipot,
Kuala Terengganu, Malaysia
Received 19 September 2006; Accepted 30 April 2007
Abstract
An assessment of morphological variation among 70 accessions of greater yam (Dioscorea
alata L.) collected throughout Malaysia was made. Data of 47 morphological variables
measured from the accessions were subjected to multivariate analysis using principal com-
ponent (PCA) and cluster analysis (CA). The PCA results indicated that the characters contribu-
ting largely to the species variability were those related to the shape, size and esh colour of
underground tubers; shape and colour of aerial tubers; position, shape, size and vein colour of
the leaves; petiole colour; shoot growth rate; and number of days for shoots to germinate. The
two-dimensional plot of the rst two PCs showed a separation between accessions of purple
tuber groups and those of white tuber groups, but was unable to distinguish accessions
according to tuber shape groups, i.e. irregular, oblong and round, as revealed by visual obser-
vation. The dendrogram of CA revealed four major groups of D. alata in Malaysia, which
supported the PCA grouping. This study demonstrated that D. alata in Malaysia consists of
numerous genotypes revealing wide inter- and intra-group variability.
Keywords: Dioscorea alata L; morphological traits; multivariate analysis; variability
Introduction
Dioscorea alata L. (greater yam or ubi badak) is a mono-
cotyledonous tuber crop of the Dioscoreaceae family and
is reported to be an old crop species native to South-East
Asia (Burkill, 1951; Coursey, 1967). Its tubers have high
nutritional value and were probably the main source of
sustenance for the people of South-East Asia for several
decades before the introduction of rice (Burkill, 1966;
Martin, 1976). Today, it is recognized as an important
food crop in many parts of the tropical and sub-tropical
regions of the world (Purseglove, 1983).
D. alata is the most polymorphic species in the genus
Dioscorea (Martin and Rhodes, 1973, 1977; Martin and
Delpin, 1978). Typically, D. alata can easily be distin-
guished from the other species in the genus Dioscorea
by its quadrangular winged stems. However, much
confusion exists among the cultivated varieties (cultivars)
of the species (Onwueme and Ganga, 1996), perhaps due
to the presence of hundreds of variants carrying numer-
ous complex characteristics that overlap. Determining
the level of variation and identifying the variants within
the species is invaluable for genetic improvement and
conservation of the crop (Okoli, 1988).
Several morphological variability studies on D. alata
have been conducted elsewhere. In New Caledonia,
Lebot et al. (1998) undertook a variability study on the cul-
tivars of D. alata and classied the cultivars into four major
groups, while the study by Velayudhan et al. (1989) on the
local cultivars inIndia found15 groups of D. alatacultivars.
Meanwhile, Sastrapradja (1982) reported the variability of
D. alata collected in Java, Indonesia, suggesting that
D. alata in Java was represented by six groups of cultivars.
In Malaysia, D. alata is not considered a major crop,
and thus there is limited information on the species, * Corresponding author. E-mail: sayed@kustem.edu.my
q NIAB 2008
ISSN 1479-2621
Plant Genetic Resources: Characterization and Utilization 6(1); 5261
DOI: 10.1017/S1479262108920050
particularly with regard to the variability, evaluation and
classication of the cultivars. Nevertheless, numerous local
cultivars or landraces of the crop were identied by local
farmers and residents (Martin and Delpin, 1978). According
tothe tuber eshcolours, the species cansimply be differen-
tiated into white and red-purple cultivar groups (Onwueme
and Ganga, 1996), known vernacularly as ubi nasi and ubi
kedudok respectively. We found that the cultivars were
also distinguished by the local residents according to tuber
shape, i.e. irregular, oblong and round shapes, and named
locally as ubi badak or ubi dinding, ubi torok and ubi
tempayan, respectively. Based on such observations,
cultivated D. alata in Malaysia could be differentiated into
six morphological groups, namely irregular-white, oblong-
white, round-white, irregular-purple, oblong-purple and
round-purple. Such groupings, however, are very ambigu-
ous where each group consists of a range of variability that
causes confusion when identifying the cultivars (see Fig. 1).
In the present work, multivariates of principal com-
ponents and cluster analysis based on the morphological
traits were carried out to determine the level of variabil-
ity, the relationship among accessions, and to identify
groups of cultivars present in D. alata collected in Malay-
sia. The result would be very useful for conservation
planning and genetic improvement of the crop.
Materials and methods
Plant material
A total of 70 accessions of various cultivated groups of
Dioscorea alata (see Fig. 1) were collected from nume-
rous locations in Peninsular Malaysia, Sabah and Sarawak
(Table 1). The collections were made in 2003 and 2004
during the growing season (October to December).
Each collection, consisting of 35 tubers, was given an
accession number and simply grouped into six typical
cultivar groups, i.e. irregular-white, irregular-purple,
oblong-white, oblong-purple, round-white and round-
purple groups (Table 1). All accessions were maintained
as a living collection in the experimental plot at KUSTEM,
Terengganu, Malaysia.
Planting the materials
Material was planted in the eld in March and April, after
4 months of tuber dormancy. Minisets of tubers with a
size of 400500 cm
3
were cut and used as the planting
materials. Prior to planting, the minisets were disinfected
with a 2.0% (w/v) solution of fungicide (thiuram) for
Fig. 1. Photograph showing various size, shape and branching patterns of the tubers from three groups of D. alata used
in this study.
Morphological variability of Dioscorea alata L. in Malaysia 53
Table 1. Accession numbers, collecting sites and cultivar groups of yam (Dioscorea alata) used in the present study
Accession no.
(KUSTEM/DA-)
Location
(village, district/division, state)
Cultivar group
(typical tuber characteristics)
-01 Kg. Cetok, Tanah Merah, Kelantan Irregular-white
-02* Kg. Cetok, Tanah Merah, Kelantan Irregular-purple
-03 Kg. Gaung, Kuala Berang, Terengganu Oblong-white
-04* Kg. Gaung, Kuala Berang, Terengganu Irregular-white
-05* Kg. Gaung, Kuala Berang, Terengganu Irregular-purple
-06* Kg. Gaung, Kuala Berang, Terengganu Oblong-purple
-07* Kg. Paloh, Tanah Merah, Kelantan Irregular-white
-08 Kg. Paloh, Tanah Merah, Kelantan Irregular-white
-09 Kg. Paloh, Tanah Merah, Kelantan Round-white
-11* Kg. Paloh, Tanah Merah, Kelantan Irregular-purple
-12* Kg. Paloh, Tanah Merah, Kelantan Oblong-purple
-15 Kg. Paloh, Tanah Merah, Kelantan Oblong-white
-18* Kg. Paloh, Tanah Merah, Kelantan Round-purple
-21 Kg. Paloh, Tanah Merah, Kelantan Irregular-purple
-22* Kg. Pasir Mas, Kelantan Irregular-white
-23* Kg. Pasir Mas, Kelantan Round-white
-24* Kg. Bunut Sarang Burung, Tumpat, Kelantan Irregular-white
-25* Market Kota Bahru, Kelantan Irregular-white
-27* Market Kota Bahru, Kelantan Oblong-white
-29 Kg. Kedai Menanti, Pasir Putih, Kelantan Irregular-white
-30* Kg. Kedai Menanti, Pasir Putih, Kelantan Oblong-purple
-31 Kg. Kedai Menanti, Pasir Putih, Kelantan Irregular-white
-32* Kg. Seri Nilam, Kuala Terengganu, Terengganu Oblong-purple
-33* Manir, Marang, Terengganu Irregular-white
-34* Manir, Marang, Terengganu Round-white
-35 Manir, Marang, Terengganu Round-purple
-36 Manir, Marang, Terengganu Irregular-white
-38* Manir, Marang, Terengganu Round-purple
-39* Manir, Marang, Terengganu Oblong-white
-40 Manir, Marang, Terengganu Oblong-purple
-44 Manir, Marang, Terengganu Round-white
-47 Manir, Marang, Terengganu Round-purple
-48 Manir, Marang, Terengganu Oblong-white
-50 Manir, Marang, Terengganu Oblong-white
-51* Manir, Marang, Terengganu Round-purple
-52* Manir, Marang, Terengganu Round-white
-56* Manir, Marang, Terengganu Irregular-purple
-57* Kg. Kapimpinan, Papar, Sabah Irregular-purple
-58* Kg. Kapimpinan, Papar, Sabah Irregular-white
-59* Kg. Siburan, Kuching, Sarawak Irregular-purple
-60* Kg. Kapimpinan, Papar, Sabah Irregular-white
-61* Kg. Kapimpinan, Papar, Sabah Oblong-purple
-62 Kg. Bangol, Sik, Kedah Oblong-purple
-63 Kg. Bangol, Sik, Kedah Irregular-purple
-64* Kg. Relau Berdiri, Batu Kurau, Perak Oblong-white
-65 Kg. Sempeneh Seberang, Batu Kurau, Perak Oblong-white
-66* Felda Jengka 6, Maran, Pahang Round-white
-67 Felda Jengka 6, Maran, Pahang Oblong-purple
-68 Kg. Kahang, Kluang, Johor Irregular-white
-69 Kg. Batu Merah, Kangsar, Perlis Oblong-purple
-70 Kg. Batu Merah, Kangsar, Perlis Oblong-purple
Kg., Kampung or village.
S. M. Zain Hasan et al. 54
5 min. The minisets were pre-sprouted in a nursery bed
before being transferred to the eld.
The minisets were transplanted into holes c. 10 cm
deep on a 2 m 1 m ridge. The spacing within and
between the ridges was 0.25 m. The young plants were
shaded with dry leaves for the rst 20 d after transplan-
ting. Plots were fertilized with 200 kg/ha of 12 : 12 : 15
NPK at 3 and 6 months after planting. Plants were irri-
gated when necessary by using a eld sprinkler and
tube droplet water system. Weeds were controlled manu-
ally by hand-weeding.
Morphological data recording
The morphological measurements were conducted on
the living plants under eld conditions by using 47
agro-morphological traits (Table 2). The characters used
and methods of data recording were according to the
International Plant Genetic Resources Institutes (IPGRI)
descriptors for yam (Dioscorea spp.) with some modi-
cation (IPGRI, 1997). Only those that discriminated
between our samples were used for the present analysis.
The characters were measured on at least three different
healthy plants and the data were then averaged for analy-
sis. The data were recorded either directly from the
measurement, using a 19 scale or as a binary recording
(1 present and 0 absent). All data were standardized
and subjected to multivariate principal component and
cluster analysis.
Principle component analysis
Principle component analysis (PCA) was performed using
SPSS software program version 12.0 (Norman, 2004). In
the PCA, the data were used to generate eigen values,
percentage of the variation accumulated by PCA and
load coefcient values between original characters and
respective PCA. Those principal components (PC) with
eigen values .1.0 were selected and those characters
with load coefcient values $0.6 were considered
as highly relevant characters scored for that PC and
supposed to be a useful trait for distinguishing the taxa
( Jeffers, 1967). The rst two principal components
which accounted for the higher proportion of the total
variations were used to plot the two-dimensional dis-
persion or scatter diagram of the accessions.
Cluster analysis
For cluster analysis (CA), the same standardizeddata matrix
was used to generate a morphological distance among
accessions using Euclidean dissimilarity coefcient, and
then used to generate a hierarchical dendrogram through
an unweighted pair-group method average (UPGMA)
(Sokal and Michener, 1958) using the software package
NTSYSpc programme (Rohlf, 2000). This analysis is used
tostudy patterns of variance andrelationshipamong acces-
sions, where accessions with close genetic distances are
placed in close proximity in the dendrogram.
Results
Morphological observation of tuber variability
Tubers of D. alata show a wide range of morphological
variation. Visual observation, based on the tuber character-
istics, enabled the separation of D. alata accessions into
three main groups of cultivars, namely irregular, oblong
and round or ovate groups (Fig. 1). Each group contained
tubers ranging from 10 to 60 cm in length, 1550 cm
in width, and weighing between 0.1 and 5 kg/tuber. Most
tubers developed branches or lobes with various shapes
and sizes, although some tubers developed without
having any branch or lobe and thus formed a single
oblong, round or ovate tuber. The tuber esh colour also
varied, displaying white, yellowish white, purple, red-
purple and mixed colours.
Principal component analysis
The result of the multivariate analysis based on the 47
morphological characters revealed a great diversity
among the 70 accessions of D. alata cultivars used in
this study. The rst ten principal components (PCs)
gave eigen values greater than 1.0 and together explained
70.2% of the total variance in the data set (Table 3).
Scores on the rst principal component (PC-1), which
accounted for 19.5% of the total variation, were highly
correlated (correlation coefcient .0.6) with the charac-
ters related to the tuber esh colours (FCCS, FCLP and
UFCCS) and petiole colour (PC) (Table 4). The second
principal component (PC-2) explained 10.7% of the
total variation and was highly associated with the aerial
tuber characteristics (EAT, ATS, ATD, SCAT and FCAT).
The third principal component (PC-3), which
explained 8.7% of the variation, was mainly loaded by
the characters related to the underground tuber shapes
(DL, TS, TTB) and weight (WGT). The fourth component
(PC-4) explained 6.2% of the variation and was deter-
mined mainly by the leaf and petiole length measure-
ments (LML2, LMW1 and PLM). The PC-5, PC-6, PC-7,
PC-8, PC-9 and PC-10 explained an additional 5.2, 5.0,
4.5, 3.8, 3.4 and 3.0%, respectively, of the total variation.
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h
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t
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PC-5 was dominated by leaf vein colour variables (LVCUS
and LVCLS), PC-6 by the leaf length variables (LML1 and
TLM), PC-7 by the leaf shape variable (LS), PC-8 by
number of days to shoot emergence (DE) and stem
length (SL) variables, PC-9 by leaf position differences
(PL) and PC-10 by root trait variables (RTS and PRT).
The eleventh and subsequent PCs were considered less
important as their contribution percentages to the total
variation were small. Therefore they are not used in the
discussion (Table 4).
The scatter diagram of the rst two PCs (Fig. 2) shows
the phenotypic variation among accessions of D. alata
collected in Malaysia. The accessions are widely dis-
persed along the both PCs axes. The dispersion slightly
supported the delimitation of the accessions based on
observations of the esh colour. The accessions of
purple tuber gathered on the negative side and white
accessions on the positive side along the second PC
axis (Fig. 2). However, there was an undened pattern
of clustering with overlap dispersion among accessions
of irregular, oblong and round tuber groups. This
suggests that tuber character alone may not be sufcient
for delimiting taxonomic units in D. alata.
The accessions of the three observedgroups are scattered
and intermixed among the four large groups, i.e. A, B, Cand
D, formed in the scatter diagram (Fig. 2). Accessions of
the oblong group (nos 3, 6, 12, 32, 39, 48, 64, 65, 69 and
70), round group (nos 9, 23, 35, 44, 52 and 66) and irregular
group (nos 1, 2, 4, 5, 7, 8, 25, 28, 29, 33, 36, 41, 42, 45, 56,
58, 59, 60 and 68) are mixed in sub-group A1. They are
characterized by the presence of aerial tuber, fast germinat-
ing tuber with less than 1 week for shoot emergence, fast
growing shoot that can reach up to 80cm length within
20d, large and branching tubers (up to 5 kg/tuber) and
elongated leaves.
Accessions 14 and 20 of irregular-purple and acces-
sions 30 and 40 of oblong-purple groups are mixed in
sub-group A2 (Fig. 2) and distinguishable by the presence
of numerous roots on the tuber surface. Meanwhile,
accessions of irregular-white (24, 46, 49, 54 and 55) and
oblong-white (43 and 53) formed another sub-group
(A3) in the diagram and differed from the others by
having a moderately lobed tuber.
Three accessions of oblong-purple (nos 61, 62 and 67)
and one accession of irregular-purple (no. 57) were
together in group B (Fig. 2) and they are differentiated
from the others by the presence of single unbranched
tubers with the smallest size (815 cm length and
68 cm width). Accessions 11 and 21 of irregular-purple
and accessions 18, 38, 47 and 51 of round-purple are
mixed in group C, which possess plants with moderate
shoot growth that can reach 2040 cm length within
20 d, single tuber, moderate tuber size and long sagittate
leaf. The two groups B and C displayed a wide range of
within-group variation in terms of tuber size, shape,
branching and lobes.
The scatter diagram (Fig. 2) also displayed a close link
with group D, comprising one accession (no. 34) of
round-white, three accessions (nos 15, 27 and 50) of
oblong-white and ten accessions (nos 10, 13, 16, 17, 19,
22, 26, 31 and 37) of irregular-white cultivars. Plants of
this group are characterized by a slow growing habit,
producing new shoots at 24 weeks after sowing and
shoot elongation of 20 cm length within 20 d. This
group is distinguishable by the presence of large lobes
and branches on the tubers.
Cluster analysis
Hierarchical cluster analysis showing the dendrogram of
taxa relationships is presented in Fig. 3. The analysis
separated the 70 accessions as different genotypes with
Euclidean dissimilarity distance ranging from 0.88
(between accessions 41 and 42) to 8.42 (between clusters
A and B grouping with remaining accessions). At the dis-
similarity distance 6.54, the dendrogram identied
four major clusters, A, B, C and D, containing between
4 and 47 accessions per cluster. At dissimilarity distance
<4.6, cluster A is separated into three sub-clusters, A1,
A2 and A3, containing 35, 4 and 7 accessions, respec-
tively. The hierarchical cluster analysis supports the
result of PCA where clustering patterns in the dendro-
gram (Fig. 3) conrms the grouping pattern revealed by
the scatter diagram (Fig. 2).
Discussion
Variability was obtained in all 47 characteristics, indica-
ting a high degree of morphological polymorphism
Table 3. Eigenvalue, variability percentage and accumu-
lated variation explained by each component of the rst
ten principal components (PCs)
Principal
component
(PC) Eigen-values
Variation
of each
component
(%)
Accumulated
variation
(%)
1 9.156 19.482 19.482
2 5.072 10.792 30.274
3 4.115 8.755 39.029
4 2.920 6.213 45.241
5 2.426 5.162 50.403
6 2.358 5.017 55.420
7 2.118 4.506 59.926
8 1.796 3.822 63.748
9 1.621 3.449 67.197
10 1.396 2.971 70.168
T
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present in the species of D. alata in Malaysia. This shows
that there are diverse variable arrangements at the indi-
vidual genotype level, indicating ample possibilities for
obtaining desirable trait combinations in specic
materials to meet the demand requested by researchers,
farmers and consumers.
The substantial morphological variation within and
between cultivar groups of D. alata may associate with
cross-pollination and sexual recombination, and perhaps
mutation, followed by intensive selection by isolated
human communities in diverse environments (Martin,
1976). Our record indicated that the male : female sex
ratio of the germplasm was 7.15 : 2.85, which is con-
sidered substantial for creating the variation within
the species. It also likely that continuous vegetative
propagation and selection have contributed to the
wide phenotypic diversity observed in this species
(Velayudhan et al., 1989).
Despite showing a great phenotypic variation among
accessions, this investigation has also found characters
that are useful as markers for classifying the cultivars
and for genetic improvement of the crop. Traits that
best discriminate between the accessions are under-
ground tuber size and esh colour, aerial tuber shape
and colour, leaf position, shape and size, leaf
vein colour, petiole colour, shoot growth rate, and
number of days for tuber miniset to germinate. This
is in agreement with Sastrapradja (1982), Martin
(1976), Onwueme (1978), and Martin and Rhodes
(1973, 1977), conrming that the most effectively used
characters in classication of D. alata are tuber, leaf
and growth characteristics.
In the present study we found that the multivariate
analysis based on the 47 agro-morphological descriptors
differentiated the accessions of D. alata in Malaysia into
four major cultivar groups, identied as groups A, B, C
and D, which possess high inter- and intra-group variabil-
ity (Figs 2 and 3). This result was in line with the result
obtained by Lebot et al. (1998) in New Caledonia,
where four major groups of D. alata morphotypes were
also identied.
Conclusions
This work contributes to increasing knowledge regarding
D. alata genetic resources available in Malaysia. In many
cases, the resources are threatened with loss by genetic
erosion due to social and economic change in this
country. Thus, genetic conservation and improvement
based on the selected materials should be encouraged
as they could lead to a reversal of genetic resource
erosion in this species. Such activities would also
maximize the use of genetic resources and provide the
opportunity to explore the potential use of this species
for other purposes.
Fig. 2. Two-dimensional plot of the rst two principal components (PC-1 and PC-2). Accessions that are encircled by the
dashed lines and marked as groups A, B, C and D corresponded with the grouping in the cluster analysis (Fig. 3).
Acknowledgements
The authors wish to thank for En Hassan for his
technical assistance. This study was funded by the
KUSTEM fundamental research grant, via the allocation
no. 57 002.
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Hasan 2008

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Morphological variability of greater yam

(Dioscorea alata L.) in Malaysia

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