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CSIRO PUBLISHING

www.publish.csiro.au/journals/ajar Australian Journal of Agricultural Research, 2005, 56, 971–981
Growth and seed yield of lentil (Lens culinaris Medikus) genotypes
of West Asian and South Asian origin and crossbreds between
the two under rainfed conditions in Nepal
R. Shrestha
A,B,D
, K. H. M. Siddique
A
, N. C. Turner
A,C
, D. W. Turner
B
, and J. D. Berger
A
A
Centre for Legumes in Mediterranean Agriculture (CLIMA), Faculty of Natural and Agricultural Sciences,
The University of Western Australia, 35 Stirling Highway, Crawley, WA 6009, Australia.
B
School of Plant Biology, Faculty of Natural and Agricultural Sciences, The University of Western Australia,
35 Stirling Highway, Crawley, WA 6009, Australia.
C
CSIRO, Plant Industry, Private Bag No. 5, Wembley, WA 6913, Australia.
D
Corresponding author. Email: renuka shrestha@hotmail.com
Abstract. Nineteen diverse lentil genotypes, 8 originating from South Asia, 6 from West Asia, and 5 crossbreds
using parents from South Asia and West Asia (or other Mediterranean environments), were evaluated for growth,
phenology, yield, and yield components at Khumaltar in the mid-hill region of Nepal. Additionally, dry matter
production, partitioning, root growth and water use of 8 selected genotypes from the 3 groups were measured at
key phenological stages. The seed yield of the West Asian genotypes was only 330 kg/ha, whereas the South Asian
genotypes produced a mean seed yield of 1270 kg/ha. The crossbreds had a significantly (P=0.05) greater seed yield
(1550 kg/ha) than the South Asian genotypes. The high seed yield of both the South Asian and crossbred genotypes
was associated with rapid ground cover, early flowering and maturity, a long reproductive period, a greater number
of seeds and pods, high total dry matter, greater harvest index, and high water use efficiency. West Asian genotypes,
on the other hand, flowered 43 days later, matured 15 days later, and had a shorter reproductive period (by 22 days)
than the crossbred and South Asian genotypes. The 23% greater seed yield in the crossbreds compared with the
South Asian genotypes was the result of a similar increase in seed size (weight per seed).
There were no significant differences in total root length (mean 4.7 km/m
2
), root dry matter (mean 95.5 g/m
2
),
or water use among the 3 groups during the major part of the growing period. There was a significant difference
in total water use due to the longer growing season of the West Asian genotype ILL 7983 and its ability to use
late-season rainfall. Maximum water use efficiencies for seed yield of 7.0 kg/ha.mm and for above-ground dry
matter of 18.9 kg/ha.mm were comparable with those reported in India and the Mediterranean environments of
south-western Australia and Syria.
Additional keywords: genetic variation, photoperiodic sensitivity, earliness, pilosae, residual soil water.
Introduction
Lentil (Lens culinaris Medikus) is the most important
grain legume in Nepal. Although initially only cultivated
on the plains (terai) of southern Nepal, recently lentils
have gained popularity in the hill and mountain regions
of Nepal, where it now contributes 5% of the country’s
area and production of lentil (ABPSD 2004). Lentil is
grown during the winter growing period (October–March)
as a post-rainy season crop. It is usually grown
in rotation with rice (Oryza sativa L.) and/or as a
mixed crop with wheat (Triticum aestivum L.), barley
(Hordeum vulgare L.), mustard (Brassica juncea L.), linseed
(Linum usitatissimum L.), grasspea (Lathyrus sativus L.), or
field pea (PisumsativumL.). Lentil is sown into the wet paddy
before the rice is harvested (the utera or relay system) or
immediately after the rice has been harvested and the soil
cultivated (the paira or post-rice system). In both cropping
systems, lentil depends predominantly on residual soil water
from the preceding rainy-season crop for its growth and
development. Soil water deficit duringcropestablishment and
in the post-flowering period is the major constraint affecting
productivity of the crop. The large year-to-year variability
in winter (October–January) rainfall from 11 to 204 mm has
led to highly variable seed yields of 220 to 1800 kg/ha in
experimental plots (Shrestha 1997, 1998).
The lentils currently grown in Nepal are the small-
seeded pilosae type of landraces belonging to the subspecies
microsperma (Bahl et al. 1991) that have limited genetic
© CSIRO 2005 10.1071/AR05050 0004-9409/05/090971
972 Australian Journal of Agricultural Research R. Shrestha et al.
variation (Erskine 1985; Erskine and Saxena 1991; Erskine
et al. 1998). In Nepal, lentil improvement activity began
in 1977, but the systematic genetic improvement of lentil
began only in 1985 with the establishment of the National
Grain Legume Research Program (NGLRP) (Bharati and
Neupane 1991). Systematic evaluation of the local landraces
and introductions from neighbouring countries (India,
Bangladesh, Pakistan) and the International Center for
Agricultural Research in Dry Areas (ICARDA), Syria, began
in 1985. However, none of the direct introductions of large-
seeded West Asian genotypes has performed well. Under
Nepalese conditions these West Asian genotypes flower
when the South Asian genotypes are maturing, suggesting
that photoperiod or vernalisation requirements limit the
adaptation of germplasm from the Mediterranean region
(Sinha 1977; Erskine and Hawtin 1983; Summerfield et al.
1984, 1985; Erskine 1997). In 1981, ICARDA initiated
an extensive crossing program of small-seeded pilosae
lentils from South Asia with germplasm from other regions,
particularly large-seeded germplasm from West Asia and
Ethiopia, to widen the genetic diversity in South Asian
lentils (Erskine and Hawtin 1983; Erskine et al. 1998). The
incorporation of desirable traits, particularly early flowering,
from the South Asian landraces into the large-seeded
Mediterranean germplasm through hybridisation and direct
introduction from other short-day growing environments
(cv. Precoz, Argentina) has provided an opportunity to select
lentil genotypes with improved seed size and higher yield
(Erskine 1983).
Since the establishment of the NGLRP in Nepal, several
exotic germplasms including both small and large-seeded
types were introduced from ICARDA and tested under
various agro-ecological zones (Bharati and Neupane 1991).
It was only during the early 1990s that the introduction and
field evaluation of genotypes from ICARDA that were bred
for targetted environments of South Asia, led to identification
of a number of promising genotypes (Shrestha 1996, 1997;
Shrestha and Neupane 2000). Furthermore, recent studies in
Nepal have also shown the superior performance of these
genotypes in terms of seed yield and disease resistance under
both the post-rice and rice relay system(Francis and Siddique
2001; CLIMA 2005). However, to date no systematic studies
with regard to phenology, dry matter production/partitioning,
water use, and root growth of lentils have been conducted
in Nepal, particularly with the newly developed crossbreds
between the South Asian and West Asian (or Mediterranean)
genotypes. Hence, a subset of the promising genotypes
from the Nepal lentil improvement program, including the
large-seeded types from ICARDA and the standard local
cultivars, was selected for detailed studies under a rainfed
environment. The main objective of the study was to evaluate
the genotypic variation in the adaptation of a range of
lentil germplasms originating from South Asia, West Asia
(or from Mediterranean-type environments), and crossbreds
between the 2 sources, and to identify the morphological and
phenological traits associated with high dry matter and seed
yield under the rainfed environment of the mid-hills region
of Nepal. We hypothesise that there will be more variation in
terms of phenology, cropdevelopment andseedyieldbetween
lentil groups (West Asia, South Asia, and crossbreds) than
within groups.
Materials and methods
Site description
A field experiment was conducted during the 2001–02 season
at Khumaltar (85.20

E, 27.40

N; 1360 m above sea level) in the
Kathmandu Valley, Nepal. The Kathmandu Valley is representative of
the warm temperate mid-hill region of Nepal.
The soil in Khumaltar is a deep, silt loamto silt clay loamdeveloped
on depositional alluvial terraces (Sherchan 1997). It contains 1.9%
organic matter (OM) with a pH of 4.3 and bulk density of 1.3 g/cm
3
and 1.7 g/cm
3
at 0–15 cm and 15–60 cm soil depth, respectively. The
soil texture is 16% sand, 62% silt, and 22% clay, with 0.09% nitrogen
(N), 27.4 mg phosphorus (P)/kg, and 85.0 mg potassium (K)/kg. Soil
analyses were done by the Soil Science Division, Khumaltar, Nepal,
using the hydrometer method and USDA textural classification for
soil textural classes, pH in 1 : 1 soil : water (Jackson 1958), OM by
the Walkley-Black method (Hesse 1972), total N by the Kjeldahl
method (Hesse 1972), available P by the Olsen method (Olsen et al.
1954), and exchangeable K by using an ammonium acetate extractant
(ADAS 1981).
Maximum and minimum temperatures, rainfall, and Class A pan
evaporation at the experimental site were recorded daily at 0845 hours
and 1745 hours during the experiment. Pan evaporation data were not
available for the period September–November.
Genotypes and agronomic practices
Nineteen lentil genotypes, 8 locally adapted landraces from South Asia,
6 from West Asia, and 5 crossbreds (where one of the parents was
from South Asia and the other from West Asia or from Mediterranean-
type environments), were grown in a completely randomised block
design with 4 replicates (Table 1). Each plot was 5.5 m long and
4 m wide (16 rows). The row spacing was 25 cm and plant-to-plant
spacing within a row was 2–5 cm. Border rows were maintained at
both ends of each block and border strips, 0.5 m on either side of the
plots, were excluded from the measurements. Before sowing, the land
was ploughed, harrowed, and levelled. The previous crop was soybean
(Glycine max L.).
Seeds were hand-sown at a depth of 3–5 cm on 21 October at a
density of 200 plants/m
2
. Seeds were treated with Bavistin fungicide
(2 g/kg seed). Rhizobium inoculum (Rhizobium leguminosarum) was
mixed with moist soil and sprinkled over the rows. At sowing,
20 kg N/ha (diammonium phosphate: 18% N, urea: 46% N), 40 kg P/ha
(diammonium phosphate: 20% P), and 20 kg K/ha (potassium chloride:
50% K) were applied. Carbofuran 3% CG insecticide was broadcast at
2 kg/ha at sowing. Seedlings began to emerge 6 days after sowing (DAS)
and reached 75–90% emergence at 8–10 DAS. Crops were harvested
from 25 March to 24 April. The net area for harvest was 3.0 m by 2.5 m
(12 rows of 2.5 m length).
Seedling establishment
Plant number was counted 28 DAS, and at maturity, using a 1-m length
of row at 4 random positions, and at dry matter harvests throughout the
season, using 2 quadrats of 0.5 m by 0.5 m per plot.
Adaptation of diverse lentil genotypes in the mid-hills of Nepal Australian Journal of Agricultural Research 973
Table 1. Lentil genotypes used in the study, their seed weight (g/100 seeds) at sowing, origin, and pedigree
ILL, International Legume Lentil; FLIP, Food Legume Improvement Program
Accession number Cultivar name Seed Group Origin Pedigree
weight
ILL 7957 (FLIP96-25L) 3.5 West Asia ICARDA ILL 5883 (Syria) ×ILL 6246 (Syria)
ILL 7978 (FLIP96-46L) 4.3 West Asia ICARDA 91S 82379
ILL 7983 (FLIP96-51L) 2.3 West Asia ICARDA 91S 88607
ILL 8006A 3.9 West Asia ICARDA Not known
ILL 8621 (FLIP2002-20L) 3.6 West Asia ICARDA ILL 5883 (Syria) ×ILL 7149 (Syria)
ILL 8633 (FLIP2001-12L) 3.0 West Asia ICARDA ILL 5883 (Syria) ×ILL 7005 (Syria)
ILL 3512 (LG 7) Simal 1.6 South Asia India Landrace
ILL 2573 (PL 639) Khajura 2 1.7 South Asia India L 9-12 ×T8
ILL 2580 (L 1278) Shital 1.6 South Asia India Landrace
ILL 7346 (LG 198) Khajura 1 2.0 South Asia India Landrace
ILL 7200B (FLIP92-35L) 1.7 South Asia ICARDA Not known
ILL 4402 1.7 South Asia Pakistan Landrace
ILL 7723 2.7 South Asia Pakistan 89503, landrace
ILL 8010 Sindur 1.6 South Asia Nepal Landrace
ILL 6829 (FLIP89-71L) 1.9 Crossbred
A
ICARDA ILL 4407 (Pakistan) ×ILL 4605 (Argentina)
ILL 7537R (FLIP93-36L) 2.2 Crossbred
A
ICARDA 90S 30799
ILL 7979 (FLIP96-47L) 2.4 Crossbred
A
ICARDA 91S 87270
ILL 7982 (FLIP96-50L) 2.1 Crossbred
A
ICARDA 91S 88526
ILL 7986 (FLIP96-54L) 2.9 Crossbred
A
ICARDA ILL 5748 (Syria) ×ILL 2578 (India)
A
Crosses between genotypes from South Asia and West Asia (or other Mediterranean-type environments).
Phenology and growth parameters
Times from sowing to 50% flowering (at least 1 fully open flower
in 50% of the plants), first pod (50% of plants with their first pod
visible), last flower (90% of the plants with no flowers), 50% podding
(50% of pods in a plot turned yellow), and physiological maturity
(90%of plants in a plot turned golden brown with fully filled pods) were
recorded on a whole-plot basis. Flowering and pod filling durations
were calculated as time from 50% flowering to days to last flowering,
and time from first podding to maturity, respectively. The reproductive
growth period was defined as the duration from 50% flowering to
physiological maturity.
At physiological maturity, 10 plants were selected at random from
the harvest area to measure plant height and primary branch number
(branches subtending from the main stem), number of pods, and seeds
per plant. Plant height was measured fromground level to the last visible
node on the main stem (just below the apical bud).
Morphological characters
Assessment of leaf morphology and seed coat colour was made
according to the International Board for Plant Genetic Resources
(IBPGR 1985). Stem pigmentation was recorded at 44 DAS, leaf colour
at 44 and 116 DAS, and leaflet size at 104 DAS. Leaf hair density was
estimated using a magnifying glass at 116 DAS.
Ground cover
Ground cover was recorded at the vegetative, flowering, and podding
stages. The proportion of ground area (%) covered by the crop canopy
was estimated visually between 1000 and 1300 hours (4 estimates
per plot).
Dry matter production and partitioning
Above-ground dry matter was measured at 58 DAS (vegetative),
50% flowering, 50% podding, and maturity. Plants were cut at the
base from a 0.25-m
2
area (0.5 m by 0.5 m quadrat) at 2 positions
per plot. Plant samples were oven-dried at 70

C for 48 h before
weighing. Senescent and fallen leaves were collected and included in the
dry weight.
Five plants were sampled and separated into leaves, stems, flowers,
and pods. Projected green area was measured using a Systronics leaf area
meter (Model 211, Naroda, Ahmedabad, India). Green area included the
area of leaves, stems, and pods. Plant samples were then oven-dried at
70

C to a constant weight.
Yield and yield components
At maturity, plants were harvested by cutting stems at ground level.
Seeds were separated from the straw by hand-threshing. The straw
(leaves, stems, and pod walls) was oven-dried at 70

Cto constant weight.
Two to four sun-dried, pre-weighed subsamples of seed, each weighing
about 100–200 g, were oven-dried at 70

C for 48 h to estimate the seed
yield on an oven-dry basis. Five hundred seeds were counted, oven-
dried at 70

C for 48 h, and weighed to measure seed size (expressed as
100-seed weight). Harvest index (HI) was calculated as the proportion
of seed weight to the total above-ground dry matter.
Soil water content
Soil water content from 0 to 60 cm depth was measured gravimetrically
in 10-cm sections of a 50-mm-diameter soil core. Measurements were
made before sowing, at 58 DAS (vegetative stage), 50% flowering, 50%
podding, physiological maturity, and immediately after the final crop
harvest. One sample (1 core) per plot was taken before sowing, and the
subsequent samplings (after crop establishment) consisted of 2 cores,
one centered over the cut stems of plants within a row and the other
between the rows at 12.5 cm from the lentil plants. All the genotypes
(4 replicates) were sampled for soil water content before sowing and at
harvest, and the soil sampling at different growth stages was done in
7 contrasting lentil genotypes, 2 from West Asia (accessions ILL 7978,
ILL 7983), 3 from South Asia (cvv. Simal, Khajura 2, and Sindur) and
2 crossbreds (accessions ILL 7979, ILL 7982).
974 Australian Journal of Agricultural Research R. Shrestha et al.
The fresh soil samples were weighed and subsequently oven-dried
at 100

C to constant weight. Volumetric water content was calculated
using the measured bulk density for the corresponding soil depth. Mean
soil water content was obtained from within-row and between-row
measurements. Bulk density was measured down to 60 cm depth, at
15-cm intervals using a core size of 7.1 cm height and 7.5 cm diameter
(2 samples per depth).
Crop water use is the water transpired by the plants plus water
lost via soil evaporation (evapotranspiration). Total crop water use was
determined by summing the change in water content in the soil between
0 and 60 cm over the period considered, and the rainfall. Drainage and
run-off were assumed to be negligible, a reasonable assumption in this
season. Water-use efficiencies were calculated as the ratios of above-
ground dry matter (WUE
DM
) or seed yield (WUE
grain
) to total crop
water use from sowing to final harvest.
Root length and dry matter
Roots were sampled at the same time as the above-ground dry matter and
soil water contents. Root measurements were made on the same 7 lentil
genotypes as sampled for water use. Root sampling was limited to the
inner 12 rows. Root samples were collected in 10-cm increments to the
maximum depth below which no more roots were visible. Roots were
sampled using a 50-mm-diameter tube auger. One sample consisted of
2 cores, 1 core centered over the cut stems of the plants and the other
core at 12.5 cm from the lentil plants.
The soil samples with intact roots were then soaked in Calgon
(sodiumhexametaphosphate/0.1%sodiumtri-polyphosphate) overnight
to disperse the clay, and roots were washed free of soil over a 1-mmsieve.
Dead (black) roots were removed. Root length was estimated using the
modified line intersect method as described by Tennant (1975). The roots
were then oven-dried at 70

C for 48 h and weighed. Total root length
per unit area of land (km/m
2
) was calculated by totalling root length
density (cm/cm
3
) to the measured depth.
Data analysis
Dry matter, seed yield, and yield components were analysed using
one-way ANOVA and linear mixed models [for estimating variance
components by the method of residual or restricted maximum
likelihood (REML)], with genotypes nested within origin or group
(West Asia, South Asia, and crossbreds) in Genstat 6th edition (Lawes
Agricultural Trust, Rothamsted Experimental Station, Hempstead,
Hert, UK). Random effect included genotypes within the group.
Least significant differences (l.s.d., P=0.05) were calculated (Genstat
2002). Morphological traits, expressed on a qualitative scale (higher
value indicating the higher intensity of the character), were analysed
using hierarchical cluster analysis in SPSS, version 11.5 (SPSS
Inc., Wacker Drive, Chicago, IL, USA) (SPSS 2002). Correlation
coefficients among various parameters were estimated using the Pearson
correlation (SPSS 2002).
Results
Weather
The mean monthly temperature in September was 23

C
and reached the lowest mean temperature of 10.0

C in
January (Fig. 1). The minimum temperature ranged from
−1.0 to 1.4

C from the third week of December to the
second week of January and temperatures slowly increased
from February onwards. Mean daily evaporation (class A
pan) ranged from 1.5 mm/day (January) to 3.4 mm/day
(April). The rainfall during the autumn months of September
and October was 158 mm, followed by a period without rain
–25 0 25 50 75 100 125 150 175
–5
0
5
10
15
20
25
30
35
Sep Oct Nov Dec Jan Feb Mar Apr
0
10
20
30
40
0
1
2
3
4
5
6
(a)
(b)
–50
R
a
i
n
f
a
l
l

(
m
m
/
d
a
y
)
T
e
m
p
e
r
a
t
u
r
e

°
C
E
v
a
p
o
r
a
t
i
o
n

(
m
m
/
d
a
y
)
Days after sowing
Fig. 1. (a) Daily maximum (—) and minimum (– –) air temperatures,
and (b) pan evaporation (. . .) and rainfall (histograms) during the
growing season at the experimental site (2001–02).
from November to mid-January. Rainfall of 159 mm was
recorded fromthe second week of January to the first week of
April (86–164 DAS), coinciding with increasing evaporation.
The rainfall from January to April was 112% higher than the
long-termaverage for the same period. The 87 mmof rainfall
received after 147 DASmay have been too late to benefit yield
in the early-maturing genotypes.
Seedling establishment and canopy development
The initial number of plants at 28 DAS (160–164/m
2
) did
not vary among groups or genotypes within groups and
the mean emergence was 82% of total seed sown. Plant
numbers declined after podding due to death by vascular
wilt (Fusarium oxysporum f. sp. lentis), particularly in
the large-seeded genotypes. Plant numbers ranged from
80–151 plants/m
2
among lentil genotypes at harvest, with
a 19% lower stand (compared with the initial stand) in the
West Asian genotypes compared with 1% in the South Asian
and crossbred genotypes.
At 58 DAS (vegetative phase), the percentage ground
cover differed significantly among groups (P<0.001), but
not within groups: South Asian and crossbred lentils had
25% greater ground cover than West Asian lentils (Fig. 2).
Ground cover increased to a maximum of 87% at flowering
(133 DAS) for West Asian genotypes, compared with 97% at
podding(154DAS) for SouthAsianandcrossbredgenotypes.
Adaptation of diverse lentil genotypes in the mid-hills of Nepal Australian Journal of Agricultural Research 975
Days after sowing
25 50 75 100 125 150 175
G
r
o
u
n
d

c
o
v
e
r

(
%
)
0
20
40
60
80
100
Fig. 2. Percent ground cover of lentil genotypes from West Asia (

),
South Asia (ᮀ), and crossbreds (N ), measured at different
growth phases. Fitted curves for genotypes from West Asia (—),
South Asia (. . .), and crossbreds (– –).
An erect plant type, low plant density, and fewer branch
numbers in the West Asian genotypes (except ILL 7983)
contributed to the lower canopy cover than in the South Asian
and crossbred genotypes (data not presented).
Morphology and phenology
Morphologically, the crossbred genotypes were similar to
the South Asian genotypes in terms of leaf and stem
pigmentation, testa pattern (speckled) and cotyledon colour
(red), whereas the leaflet size, colour, and pubescence density
of the crossbreds were intermediate between the South Asian
and West Asian genotypes (data not shown).
In the Kathmandu Valley, the South Asian and crossbred
lentils reached 50% flowering in mid-January when the
photoperiod was 10.6 h (1200 degree days). This occurred
about 1.5 months earlier than for the West Asian genotypes in
early March, which flowered when the daylength was 11.6 h
(1700 degree days). The West Asian genotypes were also
considerably later to pod and mature than the South Asian and
crossbred genotypes (Fig. 3). The nested ANOVA revealed
significant differences between both groups and genotypes
within groups in the time to 50% flowering, 50% podding,
maturity, and the reproductive growth period. However,
REML demonstrated that variance components for these
traits were 3–7 times higher between groups compared
with genotypes within groups. The late flowering of the
West Asian genotypes was also associated with a short
flowering duration (mean 30 days) and pod filling duration
(mean 33 days) compared with the mean duration of 58 days
Days after sowing
0 40 80 120 160 200
ILL 7986
ILL 7982
ILL 7979
ILL 7537R
ILL 6829
Sindur
ILL 7723
ILL 4402
ILL 7200B
Khajura 1
ILL 2580
Khajura 2
Simal
ILL 8633
ILL 8621
ILL 8006A
ILL 7983
ILL 7978
ILL 7957
Flowering Sowing Podding Maturity
West Asia
Crossbreds
South Asia
Fig. 3. Duration from sowing to 50% flowering (solid bars), first
pod (open bars), and physiological maturity (grey bars), for 19 lentil
genotypes.
and 40 days, respectively, in the South Asian and crossbred
genotypes (data not presented). There were no significant
differences in phenology between the crossbreds and the
South Asian genotypes.
Green area index (GAI), dry matter production,
and partitioning
There were no differences in GAI at the vegetative phase
(58 DAS) but group differences emerged throughout the
growing season. The West Asian genotypes had the highest
GAI (4.1) at 133 DAS, whereas the South Asian genotypes
attained the maximum GAI (3.5) at podding (154 DAS)
(Fig. 4a).
Total above-ground dry matter at maturity differed
significantly between groups (P<0.001) and between
genotypes within groups (P=0.003), with 83% of the
variance explained by the groups. At maturity the highest
dry matter was recorded in the crossbreds, followed by the
South Asian genotypes, but the West Asian genotypes had
a 35–40% lower dry matter at maturity than South Asian
genotypes and crossbreds (Fig. 4b). As a result, the
South Asian and crossbred genotypes had double the average
rate of dry matter accumulation at 4.2 g/day compared with
the West Asian genotypes at 2.2 g/day.
There was significant variation in the partitioning of dry
matter (Fig. 5), particularly between groups (64–94% of
variance in dry matter components for leaf, stem, seed, and
pod wall). At the vegetative stage (58 DAS), leaf dry matter
was 70% of the total, but decreased to 20% in the South
Asian and crossbred genotypes and 27% in the West Asian
genotypes by maturity. At maturity, the pod wall accounted
for about 19% of total dry matter in the South Asian and the
crossbred genotypes, significantly higher than the West Asian
976 Australian Journal of Agricultural Research R. Shrestha et al.
0
100
200
300
400
500
0
1
2
3
4
5
6
50 75 100 1 25 150 175
0
1
2
3
4
50 75 100 1 25 150 1 75
50
100
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300
Days after sowing
G
r
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a
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i
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t
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s
h
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r
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a
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u
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(
m
m
)
A
b
o
v
e
-
g
r
o
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d
r
y

m
a
t
t
e
r

(
g
/
m
2
)
(a) GAI
(c) Root-to-shoot ratio
(b) Dry matter
(d) Cummulative water use
Fig. 4. (a) Green area index; (b) total above-ground dry matter; (c) root-to-shoot ratio
for lentil genotypes from West Asia (circles), South Asia (squares), and crossbreds
(triangles); and (d) cumulative water use (0–60 cm soil depth) in genotypes from
West Asia: ILL 7983 (

), crossbred: ILL 7979 (N ), and South Asia: Simal (ᮀ) and
Khajura 2 (᭿) during the growing period. Bars indicate standard errors.
(a) ILL 7978
0
100
200
300
400
500 (b) Simal
Days after sowing
(c) ILL 7979
0 50 100 150 0 50 100 150 0 50 100 150
D
r
y

m
a
t
t
e
r

(
g
/
m
2
)
Fig. 5. Dry matter partitioning of 3 representative lentil genotypes from(a) West Asia, (b) South Asia, and (c) crossbreds.
Legends: leaf (no fill), stem (hatched), pod wall (grey), and seed (black).
genotypes (14%). The crossbreds had the highest dry matter
in the seeds (36%) at maturity, followed by the South Asian
genotypes (30%) and then the West Asian genotypes (14%).
The ratio of leaf to stem dry matter in the crossbred and
South Asian groups was 19% higher than in the West Asian
group; however, variation within groups was 3 times greater
than between groups. The crossbreds had a significantly
(P<0.001) higher ratio of seed to pod dry matter than the
West Asian and the South Asian group, and this ratio was the
least in the West Asian group.
Root growth
Lentil groups and genotypes within groups did not differ
significantly in total root dry matter (75.7–128.6 g/m
2
) or
Adaptation of diverse lentil genotypes in the mid-hills of Nepal Australian Journal of Agricultural Research 977
total root length (3.9–5.6 km/m
2
). There was an increase in
total root dry matter up to about 130 DAS and an increase
in root length to about 150 DAS, followed by a decrease to
maturity caused by root death (data not shown).
There was a significant variation among groups in root-to-
shoot ratio (50–97% of variance) during the growing season,
with the highest values recorded in West Asian material
(Fig. 4c). There was a dramatic decrease in the root-to-shoot
ratio in all groups over the growing season from 2.0–3.3 at
58 DAS to 0.16–0.38 at maturity. At maturity there were
no significant differences among the groups in the root-to-
shoot ratio.
Root length density, measured within the row,
reflected the root dry matter pattern. Root length
density was greater in the top 10 cm than at deeper
soil depths (Fig. 6) and decreased with increasing lateral
distance from the plants (data not presented). Variance
analysis showed that root length density only varied
significantly among the groups of genotypes at podding,
when South Asian genotypes >crossbreds >West Asian
genotypes (Fig. 6).
Water use
There were no significant differences in water use among
the 3 groups of lentils during the major part of the growing
season (Fig. 4d). The total seasonal water use ranged from
194 mm in the early South Asian genotypes to 278 mm
in the long-season West Asian genotypes, because the
West Asian genotypes were able to use the late-season
rainfall (Fig. 1).
Root length density (cm/cm
3
)
0 1 2 3 4 5
20
60
40
80
S
o
i
l

d
e
p
t
h

(
c
m
)
Fig. 6. Distribution of mean root length density (cm/cm
3
) with
depth measured within the row for lentil genotypes from West Asia:
ILL 7978 (

), ILL 7983 (

); South Asia: Sindur (᭿), Simal (ᮀ); and
crossbreds: ILL 7982 (N ), ILL 7979 () at podding. Standard errors
were smaller than the symbols.
Growth, yield, and yield components
Seed yield varied from 130 to 1800 kg/ha among the
genotypes, with significantly (P<0.001) different mean
seed yields of 1550 kg/ha, 1270 kg/ha, and 330 kg/ha
for crossbreds, South Asian, and West Asian genotypes,
respectively (Table 2). However, the mean straw yield
(average 2400 kg/ha) did not vary significantly among the
3 groups. The HI varied significantly between genotypes
and groups, ranging from 0.08 to 0.41, with mean values
of 0.38, 0.33, and 0.14 for the crossbreds, South Asian, and
West Asian genotypes, respectively (Table 2).
The mean plant height at maturity (39 cm) was similar for
the 3 lentil groups. The South Asian and crossbred genotypes
produced 1 more primary branch/plant than the West Asian
genotypes, which had 4 branches/plant. The number of
filled pods, seeds/plant, 100-seed weight, seed yield, and
final above-ground dry matter showed significant differences
among genotypes and groups (P<0.001). The mean number
of pods/plant was 82, 91, and 41 in crossbreds, South
Asian genotypes, and West Asian genotypes, respectively,
and was significantly different (P<0.001) among groups.
Genotypes from South Asia and the crossbreds had
100–120% more filled pods than the West Asian genotypes
that had much higher numbers of unfilled pods. The
South Asian genotypes produced significantly higher number
of seeds/plant (170) than the crossbreds (158), whereas the
West Asian genotypes produced the fewest seeds/plant (56).
Hundred-seed weight was 1.6–2.2 g in crossbreds, 1.3–1.8 g
in the South Asian genotypes, but 1.4–2.2 g in the West
Asian genotypes (Table 2). On average, the seed size in the
crossbreds was 22%higher (P=0.05) thaninthe SouthAsian
genotypes. The mean seed size at final harvest was 38–61%
smaller in the West Asian genotypes than their original seed
size at sowing, whereas the reductions were 3–33% and
13–28% in crossbreds and South Asian genotypes,
respectively (Table 1). The mean seed size of West Asian
and crossbred groups was significantly (P<0.001) greater
than of the South Asian group; however, 64% of variation in
seed size was within the group.
Using all 19 genotypes, seed yield was negatively
correlated with time to flowering (r =−0.85**), time to
maturity (r =−0.64**), and the number of unfilled pods
(r =−0.66**), but positively correlated with the number of
filledpods (r =0.57**) andseeds (r =0.61**), above-ground
dry matter (r =0.87**), and HI (r =0.91**). Also, there was
a significant positive correlation between number of pods and
the number of primary branches (r =0.58**).
Water use efficiency (WUE)
The mean WUE of 18.3 kg dry matter (DM)/ha.mm
and 6.3 kg grain/ha.mm for selected genotypes from
South Asia and the crossbreds were significantly higher
than the 9.6 kg DM/kg.mm and 1.9 grain/ha.mm for selected
978 Australian Journal of Agricultural Research R. Shrestha et al.
Table 2. Seed yield, yield components, and water use efficiencies (WUE) for dry matter (DM) and grain at maturity of lentil
genotypes grown at Khumaltar
Genotypes Pods/plant Seeds/plant 100-seed Seed yield Total dry HI WUE
DM
WUE
grain
weight (kg/ha) matter (kg/ha.mm)
(g) (kg/ha)
West Asia
ILL 7957 37 40 1.95 242 2847 0.08 – –
ILL 7978 58 84 2.21 435 1789 0.24 8.2 2.0
ILL 7983 53 91 1.37 507 3057 0.16 11.0 1.8
ILL 8006A 27 28 1.51 129 1659 0.08 – –
ILL 8621 38 47 2.03 340 2673 0.12 – –
ILL 8633 33 47 1.85 337 2631 0.13 – –
South Asia
Simal 90 166 1.53 1440 4173 0.35 18.2 6.3
Khajura 2 88 175 1.41 1220 3585 0.34 18.7 6.3
ILL 2580 91 169 1.52 1351 3909 0.34 – –
Khajura 1 96 175 1.35 1094 3452 0.32 – –
ILL 7200B 112 220 1.43 1152 3612 0.31 – –
ILL 4402 105 195 1.42 1346 4062 0.33 – –
ILL 7723 68 129 1.82 1125 3762 0.30 – –
Sindur 73 134 1.55 1392 4082 0.35 18.9 5.9
Crossbreds
ILL 6829 96 179 1.65 1715 4245 0.40 – –
ILL 7537R 84 168 1.72 1795 4520 0.40 – –
ILL 7979 86 163 1.72 1374 4270 0.32 18.5 5.9
ILL 7982 88 186 1.82 1477 3640 0.41 17.2 7.0
ILL 7986 54 95 2.24 1398 4036 0.35 – –
P value <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001
l.s.d. (=0.05) 29 60 0.19 330 50 0.06 3.4 1.4
West Asian genotypes (Table 2). There was no significant
difference in WUE between the crossbred and South Asian
genotypes (Table 2).
Analysis of the seed yields against crop water use
using the French and Schultz (1984) method and an
estimated soil evaporation (intercept) of 115 mm derived
for Mediterranean-type environments of southern Australia
(Siddique et al. 1998, 2001) showedthat the WUE(drymatter
produced per unit of water lost through evapotranspiration)
of the South Asian and crossbred lentils was similar at
10–15 kg grain/ha.mm to those determined in Western
Australia, India, Jordon, and Syria (Yusuf et al. 1979; Sharma
and Prasad 1984; Silim et al. 1987, 1993; Siddique et al.
1998, 2001; Zhang et al. 2000), whereas the West Asian
lentils had a WUE for grain of less than 5 kg grain/ha.mm
(Fig. 7a). The WUE for selected South Asian and
crossbred genotypes was about 40 kg DM/ha.mm, whereas
the West Asian genotypes had only half of this value (Fig. 7b).
Discussion
Rainfed lentil production is increasing in the mid-hill region
of Nepal. In this region, the present study has shown that
lentil genotypes fromSouth Asia and the crossbreds between
South and West Asian parents showed better adaptation than
West Asian genotypes. The high seed yield is related to
their early flowering, early maturity, large number of filled
pods and seeds, high above-ground dry matter, and high
dry matter partitioning to seeds. The times from sowing
to 50% flowering and maturity were particularly important
phenological traits in the adaptation. During adaptation and
acclimatisation in the lower latitudes away from their centre
of origin in West Asia, the South Asian genotypes acquired
early flowering and the ability to mature early (Erskine
1997), thereby showing less responsiveness to photoperiod
or vernalisation, combined with a greater sensitivity to
temperature near flowering (Summerfield et al. 1984, 1985;
Erskine et al. 1990, 1994).
These evolutionary changes were strongly evident in
the present study. In the mid-hills region of Kathmandu
Valley, the South Asian and crossbred lentils flowered at a
shorter daylength (10.6 h) than the West Asian genotypes
(11.6 h). Similar observations were reported when the
West Asian genotypes were evaluated in India and Pakistan
(Erskine 1983; Waldia et al. 1988; Ceccarelli et al. 1994).
For example, the large-seeded Syrian local lentil took
46 days longer when grown in Pakistan (photoperiod 10.9 h)
compared with northern Syria (photoperiod 12.2 h) (Erskine
1983). As a result the flowering and particularly pod filling
periods were truncated in West Asian material by the
increasing temperatures and increasing water deficits in
Adaptation of diverse lentil genotypes in the mid-hills of Nepal Australian Journal of Agricultural Research 979
0
500
1000
1500
2000
2500
0
1500
3000
4500
6000
(a) (b)
0 400 300 200 100 0 400 300 200 100
Total water use (mm)
T
D
M

(
k
g
/
h
a
)
S
e
e
d

y
i
e
l
d

(
k
g
/
h
a
)
Fig. 7. Relationship between (a) seed yield, and (b) above-ground dry matter and cumulative water use
for selected lentil genotypes fromWest Asia (

), South Asia (᭿), and crossbreds (N ). Lines are based on
an intercept of 115 mm and water use efficiencies for (a) 15 (– –), 10 (· · · ·), and 5 (—) kg grain/ha.mm,
and (b) 40 kg/ha.mm total above-ground dry matter. Data also plotted from other studies in India (ᮀ)
(Yusuf et al. 1979; Sharma and Prasad 1984); Tel Hadya, Syria (

) (Zhang et al. 2000); Breda,
Syria (

) (Silim et al. 1987, 1993); Jordon () (Badarneh and Ghawi 1994); and
Western Australia () (Siddique et al. 1998, 2001).
spring–early summer, resulting in few filled pods, a large
number of unfilled pods, poor dry matter accumulation, and
hence low HI and very poor seed yields. The implication
of this poor adaptation is that the limited genetic diversity in
South Asian lentils will not be overcome by farmers passively
adopting West Asian germplasmbecause there is no incentive
for them to do so. At present, only the early maturing, large-
seeded cultivar Precoz (ILL 4605) is being used in the lentil
breeding programs in Nepal. Active steps are required in the
lentil breeding program to widen the genetic base of released
cultivars. These include a crossing programbased on artificial
long-day conditions, or applying vernalisation so that a wide
range of West Asian parental germplasm can be exploited
by breeders.
Indeed, our results confirm the importance of early
phenology in breaking the South Asian bottleneck by
crossing with West Asian material (Erskine 1997). We
demonstrate a yield advantage in the crossbred material
over local landraces and cultivars, which is associated with
rapid growth, early flowering, and early maturity from the
South Asian parents, and seed size traits fromthe West Asian
parents. Moreover, the increase in seed size in crossbred
material makes it a more marketable proposition than
South Asian landraces and cultivars (Sharma et al. 1991), and
the fact that straw yield was not compromised is important
in regions such as Nepal where straw is a highly valued stock
feed. The total dry matter for the South Asian genotypes
in this study was very similar to that under farmers’ fields
(Maskey et al. 2001).
Large-seeded genotypes, particularly of West Asian
origin, were not adapted in Nepal because of their
photoperiod sensitivity, and also greater susceptibility to
boron deficiency in traditional plain areas (Bharati and
Neupane 1991; Erskine et al. 1998; Srivastava et al. 2000).
The crossbred ILL 7986 was the first large-seeded genotype
to produce more than 1 t/ha under the mid-hill rainfed
environments of the Kathmandu Valley of Nepal due to
its reduced photoperiod sensitivity (flowered in 83 days)
trait derived from the large-seeded parent ILL 4605 from
Argentina (Mediterranean-type environment) (Khanna-
Chopra and Sinha 1987), its long reproductive growth period
of 82 days, and its early maturity of 165 days compared
with the late flowering (132 days) and maturity (178 days)
in the large-seeded West Asian genotypes. Moreover, the
large leaflets, lack of pubescence, and large white flowers
of ILL 7986 make it clearly distinguishable from the local
landraces or cultivars in farmers’ fields, thereby preventing
the sale of its seed under a false name. Thus, the large-
seeded trait of the Mediterranean types can be transferred
to South Asian genotypes without detrimentally affecting
the adaptation of the South Asian genotypes. Also, the large
variance between rather than within groups suggests the
greater benefit of crossing between the groups, rather than
within the groups.
In contrast to the phenological and morphological traits
discussed above, the crop water use (evapotranspiration)
and root length density did not vary among the groups. The
high seasonal crop water use (278 mm) in the West Asian
genotype ILL 7983 was largely associated with a longer
period of growth and ability of this late West Asian
genotype to use the late-season rainfall in the year of study.
Nevertheless, the West Asian genotypes were not able to
take advantage of this late rainfall, still producing very low
yields. The measured seasonal crop water use (194–278 mm)
980 Australian Journal of Agricultural Research R. Shrestha et al.
was similar to that reported in Syria (213–326 mm, Silim
et al. 1987); India (115–228 mm under a range of water
supplies, Saraf and Baitha 1985), and Western Australia
(174–273 mm, Siddique et al. 2001), giving us confidence
that the measured crop water use captured most of the crop
water use and was not confounded by losses through deep
drainage or run-off. Recognising that the soil evaporation
in this study in the post-rainy season may be less than in a
Mediterranean environment, it is interesting to note that the
WUE
DM
values estimated for the South Asian and crossbred
genotypes (8.2–18.9 kg/ha.mm) were comparable with
those reported by Zhang et al. (2000) (9.2–18.1 kg/ha.mm),
Silim et al. (1993) (7.9–13.8 kg/ha.mm) and Siddique et al.
(2001) (8.5–16.7 kg/ha.mm), and the WUE
grain
for the
high-yielding genotypes (5.9–7.0 kg/ha.mm) was similar to
or slightly lower than those reported by Silim et al. (1993)
(6.3 kg/ha.mm), Silim et al. (1987) (9.05 kg/ha.mm), and
Yusuf et al. (1979) (8.2 kg/ha.mm) under adequate water
supply, Siddique et al. (2001) (2.4–7.2 kg/ha.mm) under
rainfed Mediterranean-type environments. West Asian
lentils, on the other hand, had markedly lower water use
efficiencies even for dry matter production in the mid-hill
environment of Nepal compared with the South Asian and
crossbred genotypes.
Conclusions
This study has shown the superior adaptation of the crossbred
and South Asian lentil genotypes compared with the
West Asian genotypes to the rainfed mid-hill environments
of the Kathmandu Valley of Nepal. The small number
of pods and seeds per plant and poor seed yield in the
West Asian genotypes are attributed to late flowering and
maturity under the short days of the Kathmandu Valley.
Morphologically, the crossbred genotypes are closer to
South Asian genotypes than the West Asian genotypes.
The crossbreds had significantly higher yields than the
South Asian genotypes. They inherited the rapid ground
cover, high dry matter, and early flowering traits from the
South Asian parents while retaining the larger seed size
of the parental genotypes from West Asia. The importance
of phenological adaptation coupled with greater seed size,
provides an explanation for the superior performance of
the crossbreds under the mid-hill environment of Nepal.
Although this paper reports a single year’s results, the fact that
the poor yield of the West Asian lentils was a consequence of
their inappropriate phenology, and not other adaptive traits,
suggests that the results will apply generally and not vary
with season. Future targetted crossing of superior parental
genotypes between South and West Asian germplasms and
early generation selection and evaluation under a range of
environments in Nepal could lead to further improvements
in the adaptation and seed yield of this important grain
legume crop.
Acknowledgments
Ms R. Shrestha gratefully acknowledges the support of
a John Allwright Fellowship from the Australian Centre
for International Agricultural Research (ACIAR). We thank
Dr D. Tennant for valuable guidance on soil water and root
length measurements, and Dr S. Asseng and Dr H. Zhang
for their advice on root growth and water use data.
Ms R. Shrestha records her gratitude to Prof. C. M. Francis
of CLIMA, The University of Western Australia, for his
encouragement and support, ICARDA (Syria) for providing
the large-seeded and crossbred lentil accessions, and the
Nepal Agricultural Research Council (NARC) for the
experimental facilities.
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