Senthilkumaran 2007

Genet Resour Crop Evol (2008) 55:287–302
DOI 10.1007/s10722-007-9234-5
RESEARCH ARTICLE
Diversity in buckwheat (Fagopyrum spp.) landrace

populations from north-western Indian Himalayas
R. Senthilkumaran Æ I. S. Bisht Æ K. V. Bhat Æ
J. C. Rana
Received: 3 October 2006 / Accepted: 12 March 2007 / Published online: 13 June 2007
Springer Science+Business Media B.V. 2007
Abstract Buckwheat (Fagopyrum spp.) is widely wheat landraces. Inter-population genetic parameters
distributed in Indian Himalayas in varying frequencies. indicated that accessions from Uttaranchal were more
The environmental heterogeneity of Indian Himalayas variable than Himachal Pradesh accessions. The spe-
offers a great deal of variability in various traits of cies-wise population data on two cultivated and one
economic importance to cultivated buckwheat species. wild species [F. cymosum (Trev.) Meisn.] indicated
In the present study buckwheat landrace populations, that F. tataricum accessions were more diverse than F.
comprising both the cultivated species, Fagopyrum esculentum accessions. Expected heterozygosity was
esculentum Moench and F. tataricum (L.) Gaertn., more for F. esculentum because of its out-crossing
were characterized for morphological and molecular nature. The estimated FST-value indicated low differ-
diversity. A total of 46 populations, 30 of F. esculen- entiation among populations of individual species
tum and 16 of F. tataricum, from the north-western zone-wise. Differentiation between species is, how-
Himalayan ranges of three administrative regions, ever, relatively strong. Understanding the population
Himachal Pradesh, Jammu and Kashmir and Uttaran- structure of landraces is important for assessing the
chal were studied for morphological characterization level of diversity among buckwheat landrace popula-
in separate field trials. The material was characterized tions and also for in situ management of farmers’
for a set of 23 descriptors (14 quantitative and 9 landraces on-farm. Furthermore, the genetic differ-
qualitative). Wide range of variations was recorded for ences between landraces are potentially relevant to
most of the quantitative traits studied. Distinct regional breeding programmes in that the variability created
variations for some important quantitative traits were through hybridization of the contrasting forms could be
evident in buckwheat landrace populations. Molecular exploited.
characterization using Random Amplified Polymor-
phic DNA markers generated enough polymorphism to
differentiate the inter-population diversity of 51 buck- Keywords Buckwheat (Fagopyrum spp.) F.
cymosum F. esculentum F. tataricum Genetic
diversity Molecular characterization North-western
R. Senthilkumaran I. S. Bisht (&) K. V. Bhat
National Bureau of Plant Genetic Resources, Pusa Himalayas
Campus, New Delhi 110 012, India
e-mail: bishtis@rediffmail.com; bishtis@nbpgr.ernet.in Introduction
J. C. Rana
National Bureau of Plant Genetic Resources, Regional Increased reliance on major food crops has been
Station, Shimla, Himachal Pradesh 171 004, India accompanied by a shrinking of the food basket
123
288 Genet Resour Crop Evol (2008) 55:287–302
which humankind has been relying upon for tions led to a uniform frequency of allozymes.
generations (Prescott-Allen and Prescott-Allen Migration could have taken place along several
1990). The shrinking of agricultural biodiversity trade routes through this area from southern China
has reduced both the intra- and inter-specific and Tibet. Ohnishi (1988) found that the places
diversity of crops, increasing the level of vulnera- where differentiation occurred were those where a
bility among users, particularly the poorer sections, sufficient genetic source for migration had been
for whom diversity in crops is a necessity for lacking, that is the margin of the distribution.
survival rather than a choice. Many of the under- Buckwheat grain is grown mainly for human
utilized crops have been included in worldwide consumption and as animal feed, although it can also
plans of action after having successfully raised the be used as a green vegetable, a green manure cover
interest of decision-makers particularly during the crop and a smother crop to crowd-out weed and as a
last 5-10 years. Buckwheat (Fagopyrum spp.) is one source of buckwheat honey. It is highly nutritive and
such underutilized crop, which holds tremendous unlike common cereals, which are deficient in lysine,
agronomic and nutritional benefits and chosen for buckwheat has excellent protein quality in terms of
the present study primarily for assessing the extent essential amino acid composition. The flowers and
and distribution of genetic diversity in north- green leaves are used for extraction of Rutin (glyco-
western Indian Himalayas. side) used in medicine and the flower produces honey
Common buckwheat (Fagopyrum esculentum of good quality. The other importance of buckwheat
Moench) is a self-incompatible species. The main as a food is its great effect in reducing the risks to
producers of common buckwheat are China, Rus- person exposed to high radiation hazards in high
sian Federation, Ukraine and Kazakhstan (Campbell mountain areas.
1995). Major exporters are China, Brazil, France, About 1,000 accessions of buckwheat (F. escu-
the USA and Canada. Japan accounts for almost all lentum and F. tataricum) have been collected from
of the world’s buckwheat imports. The other Indian Himalayas and are maintained as active
cultivated species, F. tataricum (L.) Gaertn. or collection at the National Bureau of Plant Genetic
Tartary buckwheat, is also produced in many areas Resources (NBPGR), Regional Station, Shimla
of the world but generally is consumed or traded (Himachal Pradesh) and also represented as base
locally. F. tataricum, because of its frost tolerance collection in the Indian National Gene Bank at
potential, is mainly cultivated in the high altitude NBPGR, New Delhi. Much of this diversity (*570
mountainous areas of Asia and to a lesser extent accessions) was recently collected under the Na-
elsewhere. In many areas the trend is for replace- tional Agricultural Technology Project on Plant
ment of common buckwheat, which has lower Biodiversity operative at the NBPGR during 1999-
yielding ability and lacks frost tolerance, with other 2005. Himalayas offer a wealth of diversity in
crops (Bisht et al. 2006, 2007). At these altitudes terms of landrace populations of buckwheat since
Tartary buckwheat frequently outyields common the crop is cultivated in large stretch of land of
buckwheat. Therefore, in some areas of India, higher hills of Himachal Pradesh, Uttaranchal,
Nepal and China common buckwheat production is Jammu and Kashmir, Sikkim and North-eastern
declining but Tartary buckwheat is remaining stable hills for centuries, and this extent of diversity
or increasing. Ohnishi (1988) found that cultivation would suggest need for their continuous collection
of common buckwheat in the Himalayas was and devising strategies for ex situ and in situ (on-
limited to approximately 500-2,500 m altitude. farm) conservation. It would also suggest possibility
Above 2,500 m, Tartary buckwheat replaces it in of using landraces for improved yield and better
cropping patterns. He found a considerable unifor- utilization in agriculture and industry. The present
mity of allelic frequencies of isozyme loci and study was therefore undertaken with the objectives
suggests that the recent cultivation of buckwheat in of estimating genetic diversity in buckwheat land-
this area is a possible interpretation. An alternative races using morphological and molecular character-
and may be more plausible interpretation is that the ization data and identifying useful genetic diversity
history of cultivation in this area is quite long, but in landrace populations for promoting its use and
that the repeated migration from adjacent popula- in situ conservation on-farm.
123
Genet Resour Crop Evol (2008) 55:287–302 289
Materials and methods The material was grown in randomized block

design with two replications at NBPGR, Regional
Morphological characterization Station, Shimla (latitude 318050 N, longitude 778050 E,
altitude 2,176 MASL) during 2005 cropping season
The experimental material comprised 42 buckwheat in separate trials for each species. The accessions
landraces of both the cultivated species, F. esculen- were grown in four 2 m row plots, with a between-
tum (28 accessions) and F. tataricum (14 accessions) row spacing of 30 cm, and a within-row spacing of
assembled from north-western Himalayas represent- 20 cm. Four commercial varieties, two of F.
ing different agroecological niches. The passport data esculentum (PRB1 and VL7) and two of F. tataricum
showing the origin of accessions are given in Table 1. (Himpriya and Shimla B1) were included as controls.
Table 1 List of accessions used for morphological and molecular characterization

Serial number Accessions for morphological Accessions used for molecular Species State
characterization characterization
1 – IC-258232 F. esculentum HP
2 IC-258237 IC-258237 F. tataricum HP
4 – IC-274424 F. tataricum HP
10 IC-278972 IC-278972 F. esculentum J&K
11 IC-279059 IC-279059 F. esculentum J&K
12 – IC-280349 F. esculentum J&K
17 IC-329199 IC-329199 F. esculentum HP
20 IC-338640 IC-338640 F. esculentum UA
26 IC-356112 IC-356112 F. tataricum UA
28 IC-360829 – F. esculentum UA
123
Table 1 continued
Serial number Accessions for morphological Accessions used for molecular Species State
characterization characterization

45 – IC-383595 F. esculentum UA
49 IC-423431 IC-423431 F. esculentum HP
55 Himpriya (control) Himpriya (control) F. tataricum HP
56 PRB1 (control) PRB1 (control) F. esculentum HP
57 Shimla B1 (control) – F. tataricum HP
58 VL7 (control) VL7 (control) F. esculentum UA
59 – Wild F. cymosum1 HP
60 – Wild F. cymosum2 HP
HP: Himachal Pradesh, J&K: Jammu and Kashmir, UA: Uttaranchal
Optimal agronomic practices were followed through statistical package developed at the INDOSTAT
various stages of crop growth. Data were recorded for Services (Hyderabad, Andhra Pradesh, India).
23 descriptors, 14 quantitative and 9 qualitative Classification (cluster analysis) and ordination
(Table 2). Five competitive plants were randomly (principal component analysis) analyses were also
chosen per plot for recording observations on these performed. Skewed data on quantitative traits were
descriptors (IPGRI 1994). transformed before multivariate analysis. Ward’s
Data on quantitative characters were subjected to minimum variance clustering method was used to
statistical analysis and analysis of variance (ANO- classify accessions on discrete clusters (Sneath and
VA) was carried out. The frequency distribution and Sokal 1973) using quantitative traits.
comparative summary statistics of buckwheat acces-
sions for quantitative and qualitative traits adminis- Molecular characterization (RAPD analysis)
trative region/state-wise (Himachal Pradesh, Jammu
and Kashmir and Uttaranchal,) were computed. The A total of 51 accessions of three Fagopyrum species
statistical analysis was carried out using INDOSTAT were included for molecular characterization using
123
Table 2 List of descriptors used for germplasm characterization

Serial number Descriptor
Quantitative descriptor
1 Days to 50% flowering
2 Leaf length (cm)
3 Leaf width (cm)
4 Number of leaves
5 Number of internodes
6 Petiole length (cm)
7 Number of primary branches
8 Number of inflorescence/plant
9 Length of cyme (cm)
10 Plant height (cm)
11 Days to 80% maturity
12 Yield/plant (g)
13 1,000-seed wt (g)
14 Number of seeds per inflorescence
Qualitative descriptor
15 Early plant vigour (3: Poor; 5: Good; 7: Very Good)
16 Plant growth habit (1: Erect; 2: Semi-erect; 3: Spreading)
17 Leaf colour (1: Green; 2: Pink; 3: Red; 4: Others)
18 Leaf margin colour (1: Green; 2: Pink; 3: Red; 4: others)
19 Leaf blade shape (1: Ovate; 2: Hastate; 3: Sagittate; 4: Cordate; 5: Others)
20 Stem colour (1: Green; 2: Pink; 3: Red; 4: Others)
21 Seed shattering (0: Non-shattering; 3: Low; 5: Moderate; 7; high)
22 Seed shape (1: Triangular; 2: Ovate; 3: Conical; 4: Others)
23 Seed colour (1: Grey; 2: Brown, 3: Black; 4: Mottled; 5: Others)
Random Amplified Polymorphic DNA (RAPD) anal- mining the most appropriate concentrations of tem-
ysis. These include 29 populations of F. esculentum, plate DNA, Taq polymerase and Mg concentration
20 accessions of F. tataricum and 2 accessions of required to generate repeatable PCR amplification
wild F. cymosum [F. acutatum (Lehm.) Mansf. ex profiles. One hundred random primers of 10 bp long
Hammer]. The passport information on these acces- each (10 mers) of OPA, OPB, OPC, OPD, OPM
sions is presented in Table 1. A total of 37 accessions (Operon Technologies, Alameda, CA USA) series
including both cultivated types were common for were used to amplify genomic DNA of three
morphological and molecular characterization. randomly chosen genotypes, included in the study.
Healthy seeds cleaned with distilled water were kept Out of these, twenty primers that gave satisfactory
on the germination paper soaked in water. The rolled amplification and band resolution were chosen for the
towels were incubated at 278C in the incubator for study.
germination and watered once in 2 days. Following the experiments for optimization of
Young leaves from field-grown plants were used component concentrations, PCR amplification was
for extraction of total genomic DNA following the carried out with 10 ng/ml of genomic DNA, 2 mM
CTAB procedure detailed in Saghai-Maroof et al. MgCl2, 1 U Taq DNA polymerase, 1· PCR buffer
(1984). Leaves from 15 to 20 plants per accession with 15 mM MgCl2 0.6 mM decamer primers and
were bulked to represent a DNA sample for each 0.2 mM of dNTP mix. The volume was made up to
accession. PCR conditions were optimized by deter- 25 ml with sterile water. PCR tubes containing the
123
above components were capped and given a pulse
of seeds/
Number
spin to allow proper settling of the reaction mixture.
0.00
832.27*** 169.24*** 115.12*** 28.92*** 1.03
0.00
0.51
infl.
PCR amplification was carried out in GA9600
Thermalcycler (ABI, Systems, USA). The amplifica-
7.848*
weight
tion programme consisted of a denaturation (948C for
1.40
1,000
15.04
seed
(g)
3 min), followed by 40 cycles 948C (1 min), 328C
(1 min) and 728C (1 min), ending with a single step of
728C for 5 min. The amplification products were
1.99
0.65
56.94
Yield/
plant
electrophoresed in 1.8% agarose gel and stained with
ethidium bromide.
maturity
Days to
3.19
7.62
86.98
Scoring and data analysis
80%
The two-way data matrix of varieties · amplicons
433.54
16.28
49.20
height
was used to calculate pair-wise Jaccard’s similarity
Plant
(cm)
coefficients. This matrix of similarity coefficients was
subjected to UPGMA to generate a dendrogram using
of cyme
83.73*** 4.76***
Length
the average linkage procedure. All the numerical
(cm)
0.51
0.37
2.53
taxonomic analyses were conducted using the com-
puter programme NTSYS-pc, Version 1.80 (Exeter
Number
of infl./
Software, New York, NY, USA; Rohlf 1993).
0.94
1.26
41.81
plant
Bootsrapping
Number of
branches
5.72*** 0.88***
primary
The reliability of the node of UPGMA tree was tested
0.04
0.11
0.49
by bootstrap analysis using 1000 pseudoreplicates.
Bootstrapping methods are general set of methods for
Petiole
length
Table 3 ANOVA summary for quantitative traits of F. esculentum landraces
creating pseudoreplicate data sets in situations where

leaves internodes (cm)
0.17
0.43
3.02
true resampling is impractical or impossible. The
pseudoreplicate samples are created using numerical
3.06*** 3.044*** 9.21*** 10.03***
Number Number
simulation rather than resampling. The frequency

0.27
0.78
5.32
with which a given branch is found upon analysis of

of
these pseudoreplicate data sets is recorded as the

bootstrap proportion. These proportions can be used
1.50
0.80
4.95
of
to assess the reliability of individual branches in the

optimal tree. The analysis was carried out using Free
width
Tree software.
(cm)
Leaf
0.34
0.63
1.81
length
flowering (cm)
Leaf
0.43
0.61
1.81
Results
Treatment 29.00 30.47***
Morphological characterization
Days to
* (p<0.05), *** (p<0.01)

0.02
0.29
59.00 15.12
50%
Analysis of variance (ANOVA) and mean perfor-

1.00
29.00
mance of 46 buckwheat landraces (including four

DF
control varieties) for quantitative traits are presented

Replicates
in Tables 3 and 4, respectively, for the trials

conducted on F. esculentum (30 accessions) and F.
Total
Error
tataricum (16 accessions), separately.
123
Highly significant differences for all quantitative

of seeds/
Number
variables were recorded for F. esculentum landraces.
0.00
0.80
0.00
0.39
weight (g) infl. In case of F. tataricum, no significant difference for
traits like leaf length, petiole length, number of
primary branches and number of inflorescence/plant
26.32 9.901*
1006.16** 365.04*** 457.91 3.994*
Yield/ 1,000
plant seed
9.45 1.51
226.99 2.98
were recorded. None of the accessions of F. escu-
lentum and F. tataricum outyielded the best local
control varieties VL7 and Shimla B1, respectively.
The landraces with other important desired traits such
as early flowering/maturity, determinate growth
maturity
Days to
55.06
243.10
211.11
habit, 1,000-seed weight, etc. could, however, be

80%
identified.
The range of variation for 30 F. esculentum
1231.94*
236.16
640.86
landraces revealed maximum variation for yield/

height
Length of Plant
(cm)
plant, number of inflorescence/plant, length of cyme,

number of internodes and number of primary
branches. The range of variation for 16 F. tataricum
3.152*
cyme
landraces revealed that the important characters for

(cm)
2.97
1.10
2.15
greater variability were yield/plant, length of cyme,

plant height, number of inflorescence/plant, petiole
Number
of infl./
length and leaf width. Species-wise variations and

4.13
38.72
31.50
29.27
plant
regional variations within species are presented in

Table 5. Significant differences were obtained for
Number of
days to flowering, number of primary branches and

branches
primary
days to maturity in F. esculentum. Significant differ-

0.00
0.10
0.07
0.08
ences were obtained for number of leaves, number of

internodes, petiole length and number of inflores-
cence/plant in F. tataricum. Except seed shape and
Petiole
length
Table 4 ANOVA summary for quantitative traits of F. tataricum landraces
internodes (cm)
0.02
3.51
3.30
3.30
seed colour, not much variations for qualitative

characters were revealed between F. esculentum
and F. tataricum landraces.
Number Number
2.99*** 3.85***
2.820*
Common buckwheat accessions from Uttaranchal

0.45
2.17
of
Himalaya were early flowering but indeterminate in

growth habit. Relatively greater determinate growth
2.531*
leaves
0.42
1.73
habit was recorded in two accessions from Jammu

of
and Kashmir. Accessions from Himachal Pradesh had

relatively greater yield potential. F. tataricum acces-
width
4.58*
(cm)
Leaf
0.05
1.46
2.93
sions from Himachal Pradesh were relatively early

* (p<0.05), ** (p<0.02), *** (p<0.01)
flowering/maturing. Uttaranchal accessions were

length
invariably determinate in growth habit. Seed shatter-

(cm)
Leaf
2.00
1.79
1.04
1.43
ing was more in F. esculentum accessions.

Cluster analysis using Ward’s minimum variance
flowering
Treatment 15 58.53***
DF Days to
technique classified the 30 F. esculentum landraces

1.53
0.80
31 28.76
50%
into four major groups. Cluster II comprised 12

accessions followed by 9 accessions in cluster IV
15
1
and 8 accessions in cluster I (Fig. 1). Cluster III

Replicates
comprised only one accession. A fair degree of

association between geographical origin and genetic
Total
Error
diversity was revealed. Cluster I was characterized
123
Table 5 Comparative performance of different quantitative traits species-wise and administrative region-wise (number of accessions
in parenthesis) of buckwheat landraces
Serial number Descriptors F. esculentum* F. tataricum**
HP (4) J&K (2) UA (24) Mean (30) HP (11) J&K (–) UA(5) Mean (16)
Quantitative characters
1 Days to 50% flowering 58.8 60.3 52.2 53.4 56.9 – 59.6 57.8
2 Leaf length (cm) 7.4 7.9 7.8 7.8 6.8 – 6.9 6.8
3 Leaf width (cm) 7.7 8.6 8.4 8.4 8.1 – 8.2 8.1
4 Number of leaves 15.8 14.6 15.8 15.8 13.2 – 14.5 13.6
5 Number of internodes 15.1 14.2 15.9 15.7 12.6 – 14.2 13.1
6 Petiole length (cm) 7.9 8.1 7.7 7.7 7.3 – 8.9 7.8
7 Number of primary branches 3.7 2.3 3.4 3.3 3.7 – 3.5 3.6
8 Number of inflorescence/plant 23.1 25.8 25.8 25.6 20.1 – 24.4 21.4
9 Length of cyme (cm) 5.9 5.9 6.7 6.6 4.5 – 4.0 4.3
10 Plant height (cm) 118.0 147.0 112.8 115.6 79.9 – 85.5 81.7
11 Days to 80% maturity 123.2 111.6 127.2 125.8 106.3 – 100.7 104.6
12 Yield/plant 17.2 13.1 15.2 15.3 18.9 – 16.0 17.9
13 1,000-seed weight (g) 21.2 22.1 21.9 21.8 17.3 – 16.3 16.9
14 Number of seeds/inflorescence 4.3 4.0 5.1 4.9 5.0 – 4.80 5.00
Qualitative characters
15 Early plant vigour Predominantly very good Predominantly very good
16 Plant growth habit Predominantly erect Predominantly erect
17 Leaf colour Predominantly green Predominantly green
18 Leaf margin colour Predominantly pink Predominantly pink
19 Leaf blade shape Predominantly hastate Predominantly hastate
20 Stem colour Uniform frequency of green and red Predominantly green
21 Seed shattering Low to medium Predominantly low
22 Seed shape Uniform frequency of triangular and ovate Predominantly ovate
23 Seed colour Uniform frequency of brown to black Predominantly brown
HP: Himachal Pradesh, J&K: Jammu and Kashmir, UA: Uttaranchal
*Significant differences were obtained for days to flowering, number of primary branches and days to maturity
**Significant differences were obtained for number of leaves, number of internodes, petiole length, and number of inflorescence/plant
with accessions late in flowering, more number of The 16 F. tataricum landraces could be classified
leaves, more number of inflorescence/plant and into three distinct clusters (Fig. 2), cluster II
determinate growth habit. Cluster II was character- comprising maximum (13) accessions followed by
ized with accessions late in maturity, greater plant cluster I with two and cluster III with only one
height and highly indeterminate growth habit. accession (control variety Shimla B1). The cluster II
Cluster IV was characterized with accessions low was characterized with accession early in flowering/
in plant height and low 1,000-seed weight. The maturity, greater petiole length, more number of
cluster III with only one accession was character- leaves, more number of internodes and determinate
ized with early flowering, more yield/plant, more growth habit. Cluster I was characterized with
number of inflorescence/plant, low number of accession low in yield/plant, low plant height, low
internodes and determinate growth habit. Inter- number of inflorescence/plant and low leaf length/
cluster distances were more between Cluster III width.
and other clusters. Intra-cluster distances were more Principal component analysis performed on quan-
for cluster I. titative traits of F. esculentum accessions revealed
123
Fig 1 Ward’s minimum

variance dendrogram of 30 J&K IC-278972
F. esculentum landraces J&K IC-279059
based on quantitative traits HP

UA
IC-329199
IC-338640
I
UA IC-340872
UA IC-340876
UA IC-340879
UA IC-340881
UA IC-340888
UA IC-360826
UA IC-360829
UA IC-360846
UA IC-360847
UA IC-361326 II
UA IC-361359
UA IC-362233
UA IC-362255
UA IC-382680
UA IC-383449
UA
UA
IC-383593
IC-383648
III
UA IC-391566
UA IC-392556
HP IC-423431
UA IC-444150
UA IC-444175
UA IC-444178 IV
UA IC-444186
HP PRB-1
UA VL-7
Fig 2 Ward’s minimum

variance dendrogram of 16 I
HP IC-258237
F. tataricum landraces HP IC-258243
based on quantitative traits HP IC-329195
HP IC-329401
HP IC-329407
UA IC-356112
UA IC-361316
UA IC-361317
UA IC-362253 II
UA IC-362260
HP IC-381048
HP IC-381052
HP IC-382270
HP IC-423485 III
HP Himpriya
HP Shimla B1
that the first three most informative components leaf width and petiole length. The principal compo-
accounted for 59.97% variance. It also presented the nents analysis in general confirmed the groupings
characters with greater weightings in each of the obtained through cluster analysis. The two-dimen-
three principal component axes. Important characters sional scatterplot of PCA ordination using the first
with greater weightings in principal component axis I two most informative PC axes is presented in Fig. 3.
included number of internodes, number of leaves, Principal component analysis performed on quan-
number of inflorescence/plant and yield/plant. Impor- titative traits of 16 F. tataricum accessions revealed
tant characters with greater weightings in principal that the first three most informative components
component axis II included leaf length, plant height accounted for 74.36% variance. Important characters
and petiole length. Important characters with greater with greater weightings in principal component axis I
weightings in principal component axis III included included leaf length, number of leaves, plant height,
123
Fig 3 Two-dimensional
principal components
ordination of 30 F.
esculentum landraces
number of inflorescence/plant, number of internodes Molecular characterization (RAPD analysis)

and petiole length. Important characters with greater
weightings in principal component axis II included Primer screening
yield/plant, days to 50% flowering and days to
maturity. Important characters with greater weigh- Of the 100 primers surveyed, 20 were selected for the
tings in principal component axis III included days to analysis as others gave suboptimal, indistinct or
maturity, length of cyme and leaf width. The monomorphic amplification products. A total of 138
principal components analysis in general confirmed amplification products were scored with the selected
the groupings obtained through cluster analysis. The primers, which exhibited overall 92.85% polymor-
two-dimensional scatterplot of PCA ordination using phism. A representative RAPD profile obtained with
the first two most informative PC axes is presented in the primer OPB 17 is presented in Fig. 5 indicating
Fig. 4. the extent of polymorphism observed among the
Fig 4 Two-dimensional
principal components
ordination of 16 F.
tataricum landraces
123
Fig 5 RAPD profiles of 51

buckwheat landrace
populations generated by
primer OPB 17
buckwheat populations. The average number of and the range was between 200 and 4,000 kb. In
amplification products generated was 6.9 per primer general, the extent of polymorphism observed was
with a maximum of eight with five primers and a high. Ten of 20 primers showed 100% polymorphism
minimum of six with seven primers. The size of the and only two primers showed less than 85%
amplification products varied with the primer used polymorphism. The RAPD data obtained were sub-
Fig. 6 UPGMA
dendrogram of 51
buckwheat accessions
123
jected to UPGMA clustering to find out the relation- The zone-wise population data on 49 accessions of
ship between the accessions analysed. two cultivated species indicated that Uttaranchal
accessions were more variable followed by acces-
sions from Himachal Pradesh, Jammu and Kashmir
Diversity analysis
and Zonal checks (Table 6). Information on acces-
sions from Jammu and Kashmir (three populations)
Figure 6 shows the clustering pattern obtained from
and the three zonal check varieties is, however, less
the UPGMA analysis of the data. Reliability of
informative because of low number of accessions.
clustering pattern was tested by correlating cophe-
The species-wise population data of 51 accessions of
netic values with Jaccard’s similarity coefficients
two cultivated and one wild species indicated that F.
(Mantel 1967) and the correlation coefficient of 0.893
tataricum populations were more variable than F.
indicated very good fit. The dendrogram obtained
esculentum accessions (Table 6). Information on F.
was rerooted with F. cymosum (two accessions) as the
cymosum accessions (two populations) is not much
out-group since this species is morphologically most
informative because of low number of accessions.
distinct and distantly related wild species of the other
Expected heterozygosity is more for F. esculentum.
two cultivated species. The tree was rerooted with F.
Zone-wise and species-wise population structure is
cymosum using ‘‘PHYLIP’’ Version 3.60 (Felsen-
presented in Table 7. The estimated FST-value of 0.04
stein 1993).
that falls within the upper and lower 95% limit
The UPGMA dendrogram clearly revealed three
indicates that there is low differentiation among the
major groups (clusters). Cluster I comprised 25
populations of the Fagopyrum species zone-wise. The
accessions, predominantly of F. tataricum. Cluster
differentiation among populations species-wise is,
II comprised 24 accessions, predominantly of F.
however, strong as the calculated FST-value of 0.23
esculentum. The FST-value of 0.23 indicated good
falls above the upper 95% limit. Nei’s genetic
differentiation at the species level.
distances as well as FST revealed that F. cymosum
is relatively more close to F. esculentum than F.
Population structure and species differentiation tataricum (Table 8).
Statistics of genetic diversity and population struc-

ture were computed, after estimating allele frequen- Discussion
cies, following strictly the treatment of Lynch and
Milligan (1994). A total of 189 fragments were Morphological characterization
recorded. The mean number of fragments per
population recorded were 72.2%. The total number Buckwheat varies a great deal in many characteristics
of segregating fragments were 100%. such as seed size and shape, pericarp colour, plant
Table 6 Population data (zone-wise and species-wise) of buckwheat populations

Zone/species Number Number Number of Proportion of Expected heterozygosity
of accessions of loci polymorphic loci polymorphic loci or Nei’s gene diversity
Population data zone-wise of 49 cultivated accessions

Himachal Pradesh 21 189 153 81.0 0.32 ± 0.01
Jammu and Kashmir 3 189 121 64.0 0.38 ± 0.02
Uttaranchal 22 189 166 87.8 0.36 ± 0.01
Controls 3 189 105 55.6 0.33 ± 0.02
Population data species-wise of 51 buckwheat accessions
F. cymosum 2 189 33 17.5 0.13 ± 0.02
F. esculentum 29 189 168 88.9 0.36 ± 0.01
F. tataricum 20 189 176 93.1 0.32 ± 0.01
F. cymosum populations were not included in zone-wise population data analysis
123
Table 7 Population structure of buckwheat accessions (Lynch and Milligan method)

n HT HW HB FST Lower 95% limit Upper 95% limit
Population structure zone-wise

4 0.36 0.35 ± 0.02 0.014 ± 0.01 0.04 ± 0.44 0.01 0.19
Population structure species-wise
3 0.35 0.27 ± 0.07 0.09 ± 0.00 0.23 ± 0.26 0.002 0.22
HT, the total gene diversity, i.e. expected heterozygosity or gene diversity in the overall samples zone-wise and species-wise
HW, the average gene diversity within zones and within species
HB, the average gene diversity among populations in excess of that observed between zones and between species
Table 8 Nei’s genetic distances (upper diagonal, after Lynch yield/plant, length of cyme, plant height, number of
and Milligan 1994, Phylip format) and pair-wise FST between inflorescence/plant, petiole length and leaf width.
populations (below diagonal, Phylip format) species-wise
Common buckwheat accessions were largely inde-
Species F. cymosum F. esculentum F. tataricum terminate in growth habit particularly accessions
from Uttaranchal Himalaya. A few accessions from
F. cymosum 0.0000 0.1343 0.1681
Jammu and Kashmir included in the study were
F. esculentum 0.2950 0.0000 0.0395
relatively determinate types. More such accessions
F. tataricum 0.3644 0.0735 0.0000
are required to be collected and used in crop
improvement. Report on morphological characteriza-
height, leaf size and shape and many others. Highly tion of 408 accessions of buckwheat from India (Joshi
significant differences for all quantitative variables and Paroda 1991), 280 accessions from Korea (Choi
were recorded for F. esculentum landraces. In case of et al. 1995), 507 accessions from Nepal (Baniya et al.
F. tataricum, however, no significant differences for 1995), 1,221 accessions from China (Yan et al. 1992)
traits like leaf length, petiole length, number of and 217 accessions from Japan (Ujihara 1983) has
primary branches and number of inflorescence/plant been documented (Campbell 1997). Yoshida et al.
were recorded. None of the accessions of F. esculen- (1995) reported on 234 strains of Tartary buckwheat
tum and F. tataricum outperformed in yield the best collected from main production areas of the world.
local controls VL 7 and Shimla B1, respectively. The Most of these reports, however, lack evaluation data
landraces with other important desired traits could, on various desired traits. Buckwheat genetic re-
however, be identified. Many cultivars of buckwheat sources have also been evaluated and characterized
have been developed by many countries during the for morphological and quantitative traits in DPR,
past several decades. Many of these have not been Korea (Kim and Li 1992). Information on genetic
documented to date except in a few instances, such as resources of buckwheat emanating from several other
by Joshi and Paroda (1991). These authors review studies has also been well documented (Kortov 1976;
some of the known new cultivars that have been Kubeizek and Komendra 1980; Nikitin 1980; Paush-
developed over the past two to three decades (Camp- eva and Dontsova 1980; Oshini 1986).
bell 1997). These new cultivars developed mainly Although there are many collections of common
from local germplasm contain many economically and Tartary buckwheat, there appears to have been
important traits that have been combined during the little agreement on important traits to be documented
breeding process and are therefore very important until recently. To date, in most evaluations, seed size
from conservation perspective. and density as well as plant height and lodging ability
The range of variation for F. esculentum landraces have been reported. Days to flowering and maturity
was maximum for yield/plant, number of inflores- also have been recorded on many of the collections.
cence/plant, length of cyme, number of internodes There appears to be little agreement between the
and number of primary branches. The range of breeding programmes worldwide on the most impor-
variation for F. tataricum landraces revealed that the tant traits for emphasis in crop improvement, other
important characters for greater variability were than yielding ability. Breeding programmes have had
123
to select traits and attempt to stabilize them as in therefore need of much larger effort in collecting this
many cases they do not exist or are hard to find in species, compared to common buckwheat, to effec-
collections. tively sample the existing germplasm. It must be
It would appear that the lack of frost resistance by recognized, however, that the areas in which this
common buckwheat or the frost resistance found in collection should take place are in mountainous areas
Tartary buckwheat is having an effect on grower of the Himalaya where it is often difficult to obtain
preferences in many marginal areas of production. ready access.
Common buckwheat production is declining while In situ conservation of buckwheat landraces with
Tartary buckwheat production is staying almost specific adaptations from diversity rich areas and
stationary. Unfortunately to date there has been no their conservation on-farm is a priority activity in
reported finding of frost resistance, a very favourable marginal areas especially for Tartary buckwheat
trait, in common buckwheat. As a result, breeding which is an important staple food in higher Himala-
programmes have been looking at means of intro- yan ranges. In situ conservation and crop improve-
ducing this trait from other species. Perhaps a more ment can complement one another in such areas.
extensive evaluation of germplasm from very moun- Breeding programmes that evaluate landraces and use
tainous regions may find variability in this trait them in local improvement efforts are expected to
(Campbell 1997). As evaluations of these kinds of produce material of direct value for marginal agro-
traits are expensive and appear to have limited climatic zones as well as achieve significant local
success, most breeding programmes have concen- conservation (Harlan 1992; Brush 1999; Almekinders
trated on yield parameters and other agronomic traits and Elings 2001). The ‘grassroot breeding’ can build
such as determinate growth habit for inclusion into upon existing knowledge and skills of farmers and
new varieties. Although frost tolerance is a very link farmers from different regions through the
favourable characteristic of Tartary buckwheat there exchange of information and landraces (Iwanaga
has been little evaluation of this trait and therefore the 1995; Brush and Meng 1998; Berthaud et al. 2001;
distribution of variability of this trait is not known. Cooper et al. 2001).
Common buckwheat is a self-incompatible species
and this therefore dictates the breeding patterns most Molecular characterization
used on it. Owing to its out-crossing characteristics,
all lines that are being developed must be kept in Molecular characterization using RAPD markers
isolation, either spatial or in cages, from each other. generated higher polymorphism to differentiate the
Tartary buckwheat crop improvement programmes diversity among 51 buckwheat populations. The
can be handled in the same manner as other self- bootstrap values fully supported the resolution shown
pollinating crops. It is thus much easier to generate in UPGMA dendrogram. In one cluster (Fig. 6), a few
and maintain the high numbers of segregating populations of F. esculentum are mixed-up with F.
progeny and advanced lines that are required in a tataricum. As these are two distinct species, this
plant breeding programme. mixing-up is difficult to explain and substantiate. It
The collection of Tartary buckwheat has lagged may, however, be emphasized at the moment that
behind that of common buckwheat as can be seen under farmers’ dynamic management, when new
from the listing of buckwheat collections where F. genetic combinations arise through different means
tataricum accessions are far less (approximately one- (introgression, natural hybridization or mutation),
tenth) than F. esculentum. This is in large part there is considerable evidence that farmers are likely
because common buckwheat is the most economi- to recognize, maintain and use the new genetic
cally important species (Campbell 1997). In north- combinations. New genetic combinations may be
western Himalayas, the common buckwheat is grown desirable for performance but may raise concerns for
intercropped with potato and area under its cultiva- conservationists. These are of particular relevance in
tion has reduced considerably (Bisht et al. 2006, thinking about the setting and design of an on-farm
2007). However, in higher Himalayan ranges (above conservation programme. These accessions are pres-
2,500 m), Tartary buckwheat is extremely important ently being investigated further for variation in ITS
as a food stuff for indigenous population. There is sequences.
123
Inter-population genetic parameters indicated that Acknowledgements The authors are thankful to the Director,
accessions from Uttaranchal were more variable NBPGR for providing facilities for this study. The authors also
thank all NBPGR scientists associated in assembling the
than Himachal Pradesh accessions. The species-wise buckwheat landraces from north-western Himalayas. The
population data on two cultivated and one wild scholarship provided to the senior author (RS) by the Indian
species (F. cymosum) indicated that F. tataricum Council of Agricultural Research for Master’s degree from
were relatively more polymorphic than F. esculen- Indian Agricultural Research Institute, New Delhi is gratefully
acknowledged.
tum accessions. Expected heterozygosity was more
for F. esculentum because of its out-crossing nature.
The estimated FST-value indicated low differentia- References
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Genet Resour Crop Evol (2008) 55:287–302

Diversity in buckwheat (Fagopyrum spp.) landrace

Materials and methods The material was grown in randomized block

Table 1 List of accessions used for morphological and molecular characterization

33 IC-361326 IC-361326 F. esculentum UA

Table 2 List of descriptors used for germplasm characterization

above components were capped and given a pulse

creating pseudoreplicate data sets in situations where

simulation rather than resampling. The frequency

with which a given branch is found upon analysis of

these pseudoreplicate data sets is recorded as the

to assess the reliability of individual branches in the

* (p<0.05), *** (p<0.01)

Analysis of variance (ANOVA) and mean perfor-

mance of 46 buckwheat landraces (including four

control varieties) for quantitative traits are presented

in Tables 3 and 4, respectively, for the trials

tataricum (16 accessions), separately.

Highly significant differences for all quantitative

habit, 1,000-seed weight, etc. could, however, be

landraces revealed maximum variation for yield/

plant, number of inflorescence/plant, length of cyme,

landraces revealed that the important characters for

greater variability were yield/plant, length of cyme,

length and leaf width. Species-wise variations and

regional variations within species are presented in

days to flowering, number of primary branches and

days to maturity in F. esculentum. Significant differ-

ences were obtained for number of leaves, number of

seed colour, not much variations for qualitative

Common buckwheat accessions from Uttaranchal

Himalaya were early flowering but indeterminate in

habit was recorded in two accessions from Jammu

and Kashmir. Accessions from Himachal Pradesh had

sions from Himachal Pradesh were relatively early

flowering/maturing. Uttaranchal accessions were

invariably determinate in growth habit. Seed shatter-

ing was more in F. esculentum accessions.

technique classified the 30 F. esculentum landraces

into four major groups. Cluster II comprised 12

and 8 accessions in cluster I (Fig. 1). Cluster III

comprised only one accession. A fair degree of

diversity was revealed. Cluster I was characterized

Fig 1 Ward’s minimum

based on quantitative traits HP

Fig 2 Ward’s minimum

number of inflorescence/plant, number of internodes Molecular characterization (RAPD analysis)

Fig 5 RAPD profiles of 51

Statistics of genetic diversity and population struc-

Table 6 Population data (zone-wise and species-wise) of buckwheat populations

Population data zone-wise of 49 cultivated accessions

Table 7 Population structure of buckwheat accessions (Lynch and Milligan method)

Population structure zone-wise

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