Springer 2006

Genetic Resources and Crop Evolution (2006) 00: 111

DOI 10.1007/s10722-005-1884-6

-1

Genetic diversity in barley (Hordeum vulgare L. ssp. vulgare) landraces from

Uttaranchal Himalaya of India
T. Manjunatha, I.S. Bisht*, K.V. Bhat and B.P. Singh
National Bureau of Plant Genetic Resources, Pusa Campus, New Delhi-110 012, India; *Author for correspondence (e-mail: bishtis@redimail.com/bishtis@nbpgr.ernet.in
Received 22 March 2005; accepted in revised form 29 July 2005

Key words: Barley landraces, Genetic diversity, Hordeum vulgare spp. vulgare, Uttaranchal Himalaya

Abstract
In the high altitude areas of western Himalaya, barley is a crop of marginal, low input drought stressed
environments. The landraces grown in these areas are favoured for their quality, both as grain and straw.
However, area under the naked barley landraces, during the last three to four decades, has declined
considerably and their ex situ and in situ conservation requires attention. Morphological and RAPD
descriptors of a collection of 70 landraces from the higher Himalayan ranges of Uttaranchal state were used
to analyse levels of genetic diversity. A wide range of variation was recorded for various morphological
characters in univariate analysis. The multivariate analysis based on six quantitative traits distinguished
accessions from dierent geographical areas in the region but failed to separate naked from covered
barleys. Clustering based on qualitative traits, however, clearly distinguished naked and covered forms.
RAPD proles eciently dierentiated naked barleys from covered forms, but could not dierentiate
between oriental and occidental covered forms. A set of 11 predominant landraces were subjected to
detailed population genetic analysis. More diversity was observed in covered barleys than the naked forms,
both for morphological and RAPD markers. The low diversity in naked barley populations was attributed
to either genetic drift or to a founder eect, while the high diversity in covered barley populations was
attributed to their large-scale cultivation for animal feed and brewing purposes. Allelic combinations were
not randomly distributed, as a geographic trend was closely related to covered and naked barleys. Since
naked barleys are poorly represented in the national collections, a systematic collection from the entire
region is suggested. The genetic dierences between covered and naked barleys may be relevant to breeding
programmes since the variability created through hybridisation between the contrasting forms could be
exploited.

Introduction
Cultivated barley (Hordeum vulgare ssp. vulgare) is
one of the oldest of cultivated plants (Harlan 1968)
and has been grown in India since ancient times.
The origin of cultivated barley is disputed. Aberg
(1940) postulated that the six-rowed wild barley
(H. vulgare ssp. agriocrithon) found in Tibet was

the progenitor of cultivated barley, but Harlan

(1976) maintained that barley was domesticated in
Southwest Asia from a two-rowed wild barley,
H. vulgare ssp. spontaneum. Freisleben (1940)
(elaborated by Takahashi 1955) proposed a
diphyletic origin, where the six-rowed cultivated
barley in the Oriental region was derived from ssp.
agriocrithon, and the two-rowed cultivated barley

2
in Southwest Asia originated from ssp. spontaneum. In a recent study, Badr et al. (2000) demonstrated that barley was rst brought into culture in
the Fertile Crescent, and that the Himalayas is a
diversication region of domesticated barley.
Cultivated barleys are classied as hulled, in which
the lemma and palea are fused to the pericarp, and
naked, in which the cha is easily separated from
the grain. Where barley forms a major part of the
human diet, naked types are preferred. The widespread distribution of naked barley makes the
hulled or naked caryopsis character a key trait to
follow the origin and domestication process of
barley (Harlan 1995; Salamini et al. 2002). A recent study (Taketa et al. 2004) indicates that
naked barley has a monophyletic origin, probably
in southwestern Iran, and all naked barleys are
likely to share a common ancestor.
In India, barley is cultivated over ca. 3 million ha, mainly in the north Indian plains but also
in the hilly regions of the Himalayas, up to an
elevation of around 4000 m. In the Himalayas, the
six-rowed covered types are common, and tworowed types, both covered and naked, are grown
only to a limited extent. Cultivation of the sixrowed naked forms is conned to the higher
Himalayan ranges. In the high altitude areas of the
western Himalaya, barley is generally a crop of
marginal, low input drought stressed environments. The landraces grown in these areas are favoured by farmers for their quality, both as grain
and straw. However, the area under the naked
barley landraces, which were very popular and
widely grown in higher Himalayan ranges three to
four decades ago, has declined considerably and
their ex-situ and in-situ conservation now requires
attention.
Although some information is available with
regard to the phylogeny and evolution of Himalayan barleys, there is only limited knowledge of
the genetic diversity present in the traditional
landraces. The present study was carried out in
order to:
1. Estimate the extent of the genetic diversity in
landraces using morphological and molecular
characterisation data.
2. Assess the extent of diversity within and between populations with a view to devise strategies
for their ex situ and in situ (on-farm) conservation.

Materials and methods

General diversity analysis
Morphological characterisation
The experimental material comprised 70 barley
landraces assembled from dierent parts of northwestern Himalayas in Uttaranchal state and represented all agro-ecological conditions of the state.
The material was collected from almost all diversity rich areas of the state during 20002001. The
material included covered (64) and naked (6)
barleys (Table 1). Landraces with specic local
adaptations for distinct microclimatic niches were
included in the study. The material was grown in a
randomised block design with two replications at
two locations, namely NBPGR Regional Station,
Bhowali (temperate) and NBPGR, New Delhi
(sub-tropical to semi-arid) during the rabi (winter)
season of 200203. The accessions were grown in
four 2-m row plots, with a between-row spacing of
25 cm, and a within-row spacing of 10 cm. Five
commercial varieties (DL-36, DL-85, Jyoti, VLB-1
and VLB-60) at the Delhi location and two (VLB1, VLB-60) at Bhowali were included as controls.
Recommended agronomic practices were followed
through various stages of crop growth at both the
locations. Data were recorded on 10 randomly
chosen competitive plants per plot for 16 descriptors (Table 2) following the IBPGR (1982)
descriptor list. Six quantitative characters (days to
owering, days to maturity, plant height, number
Table 1. Passport information of barley landraces studied.
Districta

No. of
accessions

Hulled
(covered)

Hull-less
(naked)

Elevational
ranges (m)

Almora
Bageshwar
Chamolib
Champawat
Nainital
Pithoragarhb
Rudraprayag
Tehrib
Uttarkashi
Total

8
1
5
1
3
25
10
9
8
70

8
1
4
1
3
20
10
9
8
64

13001800
1400
15002000
1500
15001800
13004500
13001600
15002000
15002000

The districts are parts of Uttaranchal State in Indian Himalaya

falling between 2870 N to 3140 N latitude and 7740 E to
8102 E longitude.
b
Eight landraces from Pithoragarh, two from Tehri and one
from Chamoli districts were studied for population genetic
parameters.

3
Table 2. Descriptors recorded for morphological characterisation of barley landraces.
1.
2.
3.
4.
5.
6.
7.
8.
9.

10.
11.
12.
13.
14.
15.
16.

Growth class (seasonality): 1=Winter; 2=Intermediate; 3=Spring

Plant height (cm): Height of the plant at maturity
Number of productive tillers
Days to ower: Counted as the number of days from sowing to 50% of the plants in ower
Days to maturity: Counted as the number of days from sowing to 80% of the plants in physiological maturity
Row number or lateral orets: 1=six rowed, 2=two rowed, large or small sterile lateral orets; 3=two rowed, rudimentary
sterile lateral
Spike density: A visual measure of spike density; 1=lax (rachis internode length >4 mm); 2=intermediate; 3=dense (rachis
internode length <2 mm)
Spikelet groups (triplet): 1=few (<20); 2=average (2030); 3=many (>30)
Hoodedness/awnedness: 1=sessile hoods; 2=elevated hoods; 3=awnless, or awnleted (<2 mm) on all rows; 4=awned (on
external rows only for two rowed forms and on all six rows for six rowed forms); 5=awned on central rows only, lateral
rows awnless or awnleted (for six rowed forms only)
Awn roughness: 1=smooth; 2=rough
Length of rachilla hairs: 1=short; 2=long
Kernel covering: whether lemma and palea adhere the caryopsis: 1=naked grains; 2=covered grains
Lemma colour: 1=white/brown; 2=purple/black
Grain (pericarp) colour: 1=white; 2=blue; 3=black; 4=Others
100-seed weight (g)
Yield per plant (g)

of tillers per plant, 100-seed weight and yield per

plant) were subjected to statistical analysis by
ANOVA.
Classication (cluster analysis) and ordination
(principal component analysis) analyses were
performed. Skewed data on quantitative traits
were transformed before multivariate analysis.
Wards minimum variance clustering method was
used to classify accessions in discrete clusters
(Sneath and Sokal 1973), whereas the Weighted
Average Linkage technique using Jaccards similarity index was used for qualitative traits. The
scores for various character states of dierent
accessions were converted to a binary code before analysis using qualitative traits. Principal
components analysis was performed using quantitative traits.
RAPD analysis
Young leaves from eld-grown plants were used
for extraction of total genomic DNA following the
CTAB procedure detailed in Saghai-Maroof et al.
(1984). Leaves from 15 to 20 plants per accession
were bulked to represent a DNA sample for each
accession. PCR conditions were optimised by
determining the most appropriate concentrations
of template DNA, Taq DNA polymerase and Mg
concentration required to generate repeatable
PCR amplication proles. Eighty six random
10mer primers from the sets OPA, OPB, OPC,
OPD, OPK (Operon Technologies, USA) were

used to amplify four templates, and from this

screen, 20 primers were chosen for the study, on the
basis of good amplication and clarity of bands.
For the genotype screen, each 25 lL PCR contained 1 reaction buer (10 mM TrisHCl, pH 8.3
and 50 mM KCl), 2.5 mM MgCl2, 1 U of Taq DNA
polymerase; 200 lM each of dATP, dTTP, dCTP
and dGTP (all reagents from Promega, USA);
0.6 lM primer and 20 ng of template DNA. The
amplication programme consisted of a denaturation (94 C for 3 min), followed by 40 cycles 94 C
(1 min), 32 C (1 min) and 72 C (1 min), ending
with a single step of 72 C for 5 min. The amplication products were electrophoresed in 1.8%
agarose gel and stained with ethidium bromide.
Scoring and data analysis
The two-way data matrix of varieties X amplicons
was used to calculate pair-wise Jaccards similarity
coecients. This matrix of similarity coecients
was subjected to UPGMA to generate a dendrogram using the average linkage procedure. The
data matrix was used to calculate correlations
among variables. These correlations were subjected to eigenvector analysis and the rst three
most informative principal components were
extracted to study the pattern of variations
observed among the accessions. All the numerical
taxonomic analyses were conducted using the
computer programme NTSYS-pc, version 1.80
(Exeter Software, New York).

4
Table 3. Range of variation for various morphological traits in barley landraces.
Descriptors

Min.

Max.

Kurtosis

Skewness

Mean

SD

SE

CV

Quantitative traits
Delhi location
1. Days to owering
2. Days to maturity
3. Number of tillers
4. Plant height
5. 100-seed wt.
6. Yield/plant

78.0
121.0
7.0
83.0
2.1
5.3

110.0
139.0
38.0
128.0
4.3
47.5

2.25a
0.42
1.09

0.55
0.50
2.09a

1.36a

0.61a
1.03a
0.27

0.15
1.19a

98.11
129.77
16.41
109.20
3.35
18.77

6.24
3.27
5.61
9.02
0.39
6.68

0.51
0.27
0.46
0.74
0.03
0.54

6.36
2.51
34.19
8.25
11.66
35.62

Bhowali location
1. Days to owering
2. Days to maturity
3. Number of tillers
4. Plant height
5. 100-seed wt.
6. Yield/plant

124.0
163.0
3.0
66.8
1.6
2.5

151.0
185.0
15.0
95.5
5.2
13.8

1.77a
1.26a
1.09

0.04

0.13
0.18

0.75a
0.10
1.03a
0.03
0.33
0.39

134.56
169.79
7.41
81.84
3.01
7.34

4.61
3.73
2.87
6.90
0.71
2.37

0.38
0.31
0.46
0.57
0.06
0.19

3.43
2.20
38.73
8.44
23.57
32.28

Qualitative traits
7. Growth class: All spring types (100%)
8. Row number or lateral orets: All six rowed (100%)
9. Spike density:1 (lax)=45.3%; 2 (intermediate)=41.3%; 3 (dense)=13.4%
10. Spikelet groups (triplet) per spike: 1 (<20):6.7%; 2 (2030)=77.3%; 3(>30)=16%
11. Hoodedness/awnedness: 4 (awns on all rows)=92%; 5 (awns on central rows only)=8%
12. Awn roughness: Predominantly smooth (97%)
13. Length of rachilla hairs: 1 (short)=46.7%; 2 (long)=53.3%
14. Kernel covering: 1 (naked grains)=8%; 2 (covered grains)=92%
15. Lemma colour: 1 (white/brown)=88%; 2 (purple/black)=12%
16. Grain (pericarp) colour: 1 (white)=88%; 2 (blue)=1.3%; 3 (black)=10.7%
a

Signicant at p 0.01.

Table 4. Means of quantitative variables in dierent clusters of

barley landraces.
Descriptors

Cluster I

Cluster II

Cluster III

Delhi location
Days to owering
Days to maturity
Plant height
No. of tillers
100-seed weight
Yield/plant
No. of accessions

91.22
126.28
107.31
19.35
3.27
18.16
23

100.44
131.17
106.96
14.15
3.53
22.23
32

102.33
131.17
106.96
15.22
3.13
13.93
20

Bhowali location
Days to owering
Days to maturity
Plant height
No. of tillers
100-seed weight
Yield/plant
No. of accessions

131.52
167.13
85.43
19.35
3.27
6.45
28

135.14
170.91
79.20
14.15
2.52
7.30
35

140.08
173.04
80.49
15.22
3.70
9.47
12

Detailed population genetic studies

Detailed population genetic studies were carried
out to assess within and between population

variation among prominent local landraces with a

view to select suitable sites for on-farm conservation. Eleven landraces were selected for the study
of population genetic parameters. Of these, eight
widely grown landraces (six hulled and two hullless) were selected from Munsyari valley in
Pithoragarh district, a supposedly diversity-rich
area adjoining Tibet in Kumaon region of Uttaranchal Himalaya, and three important landraces
(two hulled and one hull-less from a relatively
distant area in Garhwal region of Uttaranchal
state, for comparison. Ten individual plants per
accession were characterised by RAPD proling
with four primers (OPN-1, OPN-19, OPO-9 and
OPO-20). The primer selection was dictated by the
availability of the primers as this study was carried
out at a later stage to investigate the reasons for
low polymorphism between populations. The genetic diversity of the populations was studied by:
1. Per cent polymorphic loci (Pp)
2. Number of alleles per locus (Ap)

Figure 1. Wards minimum variance dendrogram of 75 barley accessions based on quantitative traits at Delhi location (*landraces
selected for population genetic studies, N = naked (hull-less) landraces).

6
Table 5. Principal components analysis using quantitative
traits of barley landraces.
PC axes

Total variation
explained
%

Characters
weightings

with

high

Cumulative

Delhi location
I
38.68

38.68

II

25.33

64.10

III

18.06

82.16

Bhowali location
I
41.54

41.54

II

23.48

65.03

III

15.84

80.88

Days to owering (0.93),

days to maturity (0.93)
100-seed weight (
0.80),
yield/plant (
0.77)
Plant height (
0.88),
Number of tillers/plant
(
0.59)
Days to owering (0.83),
days to maturity (0.78)
100-seed weight (
0.83),
yield/plant (
0.62)
Yield/plant
(
0.58),
Number of tillers/plant
(0.49)

3. Genetic diversity for each locus and population

(Hep)
4. Diversity in the overall populations (Ht)

Results
Morphological characterisation
Univariate analysis
The ANOVA revealed signicant dierences for
all the quantitative variables. All accessions owered and matured early at Delhi, and produced
more yield per plant and a greater 100-seed weight
compared to those at Bhowali (Table 3). GE
interaction was signicant, with signicant dierences between locations being observed for days to
owering, days to maturity and 100-seed weight.
Therefore, no pooled analysis was possible for
these characters. Pooled analysis was performed
for plant height and yield per plant, and this revealed that the performance of the landraces differed across the locations. The distribution of yield
per plant, number of tillers and 100-seed weight
among the quantitative traits was skewed, in both
the positive and negative directions. For the
qualitative traits, diversity was revealed for spike
density, spikelet groups (triplets) per spike, length

of rachilla hairs, awnedness, kernel covering,

lemma colour and grain colour. Not much dierence was observed in growth class, row number
and awn roughness (Table 3).
Multivariate analysis
Cluster analysis classied the accessions into three
major groups at both locations. Cluster II comprised 32 accessions, cluster I 23 and cluster III 20
accessions at Delhi. Similarly at Bhowali, cluster II
comprised 35 accessions, cluster I 28 and cluster III
12 accessions (Table 4). In Delhi, cluster I consisted
mainly of early owering/maturing accessions with
high tillering capacity and moderate to high yield
potential. These accessions were collected from
lower elevation zones (10001500 m asl) of Uttaranchal Himalaya. Cluster II comprised accessions
with high yield potential from mid elevation zones
(15002000 m asl) and cluster III comprised accessions with late owering/maturity, low yield per
plant and low 100-seed weight, assembled mainly
from high elevation sites (more than 2000 m asl) of
Pithoragarh district. The Wards minimum variance dendrogram of clustering pattern using Delhi
location data is presented in Figure 1. In Bhowali,
cluster I comprised early owering accessions with
greater plant height and low yield per plant, accessions assembled mainly from mid elevation zones.
Cluster II comprised accessions with low 100-seed
weight and plant height mainly from higher elevations. Cluster III mainly comprised accessions with
late owering/maturity, high 100-seed weight and
high yield per plant from lower elevations.
The rst three most informative principal components accounted for, respectively, 82% and 81%
of the variation at Delhi and Bhowali (Table 5).
The characters contributing the greatest weightings in the rst PC axis were days to owering and
days to maturity. For the second PC axis, 100-seed
weight and yield per plant were the most important traits, while for the third axis, plant height
and number of tillers per plant at Delhi, and yield
per plant and number of tillers/plant at Bhowali
dominated. The principal component analysis
largely conrmed the groupings of the accessions
obtained through cluster analysis.
The clustering pattern based on qualitative traits
revealed that the majority of the accessions
grouped into two major clusters comprising 35
accessions in cluster I and 25 accessions in cluster
II, followed by 7 accessions in cluster III, 4

Figure 2. Weighted average linkage dendrogram of 75 barley accessions (including 5 check varieties) based on qualitative traits.

accessions in cluster IV and 1 accession each in

cluster V and cluster VI (Figure 2). Here also, a
fair association between genetic diversity and

geographical diversity was revealed. Cluster IV

comprised only naked barleys, and the distance
between these clusters shows that naked barleys

8
Table 6. Comparison of bands generated with RAPD primers.
Primers with DNA sequence

Total number of bands

Number of polymorphic loci

Percent polymorphic loci

OPN-16 (AAGCGACCTG)
OPN-17 (CATTGGGGAG)
OPN-19 (GTCCGTACTG)
OPO-7 (CAGCACTGAC)
OPO-11 (GACAGGAGGT)
OPO-18 (CTCGCTATCC)
Total

4
4
5
5
4
4
26

2
1
1
3
3
3
13

50
25
20
60
75
75
50.0

Table 7. Properties of RAPD amplicons generated by 4 primers in 11 selected barley landraces.

Primer

Total number of bands

Number of polymorphic loci

Percent polymorphic loci

OPO-9
OPO-20
OPN-19
OPN-1
Total

7
10
8
11
36

4
9
7
10
30

57.10
90.00
87.50
90.91
83.33

are the most easily discriminated based on qualitative traits. Cluster III mainly comprised covered
barleys with black lemma/grain colour from high
altitude areas of Pithoragarh district. The majority
of the accessions in cluster I were from higher
elevation zones of Uttaranchal state, whereas in
cluster II majority of the accessions belonged to
mid elevation zones.

RAPD proling
Of the 20 decamer primers selected, six were
informative (Table 6), generating 26 bands of size
range 2001800 bp, of which 13 were polymorphic. Percent polymorphism ranged from 20 to
75% with two primers (OPO-11 and OPO-18)
displaying 75% polymorphism. Genetic similarities between the accessions is presented as a dendrogram (Figure 3). Cluster I comprised 68
accessions, mainly covered barleys. Cluster II
comprised 4 naked barleys. One naked barley (IC356114) and two check varieties VLB-1 and DL-85
did not cluster in either groups and were the most
diverse. One naked barley accession from Chamoli
clustered with covered barleys in cluster I. Bootstrap analysis of RAPD data indicated less than
50% support for the major nodes of the clusters.
This indicated that the branches were not strongly
supported and there is a need to add more number

of markers. No distinct association between clustering pattern and area of collection was found,
but brittle and non-brittle rachis were clearly separated (Figure 4). The diversity among accessions
in general appears to be low as indicated by the
similarity co-ecient values between various
accessions.
Four primers, two each from OPO and OPN
series, were used to detect within population
diversity. Of the 36 bands generated, 30 were
polymorphic. The percent polymorphism ranged
from 57.1 to 87.5% (Table 7). The number of alleles per locus was estimated by simple counting.
The number of alleles per locus ranged from 1.05
(IC-260864) to 1.45 (IC-356093) with overall
population average of 1.25 alleles per locus. The
overall population genetic diversity was greater in
the eight landraces from Munsyari valley in
Pithoragarh district than in the three landraces
from other areas (one from Chamoli and two from
Tehri district). Covered barleys were more diverse
than naked barleys (Table 8).

Discussion
The study revealed variation among the barley
landraces for all six quantitative traits studied. The
naked barleys were generally poor yielding at both
test locations. The set of accessions under study

Figure 3. UPGMA dendrogram of 75 barley accessions (including 5 check varieties) based on RAPD data.

did not perform uniformly with regard to most of

the traits across locations, indicating the importance of genotype environment interaction. As

the provenance of most of these accessions is from

the high Himalayan ranges, their relatively good
performance at Delhi suggests wide adaptability of

10
Table 8. Summary diversity analysis of selected barley landraces.
Diversity parameters

Munsyari populations

Other populations

Covered barleys

Naked barleys

Whole population

No. of alleles per locus

Genetic diversity

1.262
0.097

1.1997
0.085

1.293
0.106

1.114
0.060

1.250
0.094

Figure 4. UPGMA dendrogram of 11 selected barley landraces based on RAPD data.

Himalayan barleys. The quantitative traits used

for general diversity analysis in the present study,
however, failed to discriminate covered and naked
barleys. Within the qualitative traits, accessions
from higher elevations had mainly brittle or lax
rachis as against intermediate to dense spikes in
accessions collected from relatively lower elevations. The naked barleys invariably had brittle
rachis. Variation was also recorded for lemma and
grain colour, even in covered barleys, but white/
brown seed colour was more predominant than
purple/black types. White/brown seed colour type
barleys are most preferred for cultivation with
specic uses in Himalayas as animal feed and
brewing. There were also dierences in awn type.
Populations with awns only on the central rows,
and awnless or awnleted lateral rows were characteristic of the blue/black grain covered and
naked barleys. The variation seems to be regionally distributed in clustering pattern based on
quantitative traits (Figure 1). Such regional variability could be due to geographic isolation,
founder eects and microclimatic dierences
between regions.
Cluster analysis using qualitative traits
clearly discriminated naked and covered barleys
(Figure 2). Naked barleys were far more distinct
than covered barley for morphological traits.
RAPD proling was also eective in discriminating between naked and covered barleys. Most of

the naked barleys were grouped together and were

distinct from the covered barleys (Figure 3).
Murphy and Witcombe (1986) and Witcombe and
Murphy (1986) have demonstrated that covered
and naked barleys from the Himalayas dier signicantly from each other in a multivariate sense.
A higher diversity, and the grouping of black/blue
lemma covered barleys with naked barleys in one
cluster, shows that both occidental and oriental
types coexist in the Himalayas and that blue
aleurone and purple/black caryopsis are peculiar
to Himalayan barleys of exclusively oriental origin. Murphy and Witcombe (1986) have suggested
that naked barleys arose by a single gene mutation
from covered barleys a long time ago, and that
thereafter cultural and adaptive selection pressures
have caused these forms to diverge.
A systematic survey of the entire Himalaya region is needed to safeguard further erosion of the
existing landraces. More detailed diversity analyses from selected sites in the Himalayas will identify potential sites for in situ conservation. The
importance of these genetic resources is underlined
by the surprising observation that both the naked
and covered landraces were competitive with
commercial cultivars at Delhi, so that the Himalayan barleys can be viewed as a source of useful
genetic variation for a range of production traits.
Of particular interest to breeding programmes is
that the genetic dierences uncovered between

11
covered and naked barleys would have potential to
unlock novel variation following the hybridisation
of contrasting forms.

Acknowledgements
The authors thank Dr. B.S. Dhillon, Director,
NBPGR for providing facilities for the work.
Thanks are also due to the Ocer-in-Charge and
other scientists working on barley germplasm at
NBPGR Regional Station, Bhowali (Uttaranchal)
for helping in eld evaluation of barley landraces
at Bhowali location. Financial support in the form
of scholarship received from Indian Council of
Agricultural Research for M.Sc. degree is duly
acknowledged by the senior author.

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