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E, 27.40
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, owers,
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
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
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 (200102).
from November to mid-January. Rainfall of 159 mm was
recorded fromthe second week of January to the rst week of
April (86164 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 benet yield
in the early-maturing genotypes.
Seedling establishment and canopy development
The initial number of plants at 28 DAS (160164/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
80151 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 signicantly 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 owering
(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 leaet 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% owering 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 owered 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
signicant differences between both groups and genotypes
within groups in the time to 50% owering, 50% podding,
maturity, and the reproductive growth period. However,
REML demonstrated that variance components for these
traits were 37 times higher between groups compared
with genotypes within groups. The late owering of the
West Asian genotypes was also associated with a short
owering duration (mean 30 days) and pod lling 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% owering (solid bars), rst
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 signicant
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
signicantly 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 3540% 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 signicant variation in the partitioning of dry
matter (Fig. 5), particularly between groups (6494% 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, signicantly 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
150
200
250
300
Days after sowing
G
r
e
e
n
a
r
e
a
i
n
d
e
x
R
o
o
t
t
o
s
h
o
o
t
r
a
t
i
o
W
a
t
e
r
u
s
e
(
m
m
)
A
b
o
v
e
-
g
r
o
u
n
d
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 (060 cm soil depth) in genotypes from
West Asia: ILL 7983 (
), ILL 7983 (
) (Silim et al. 1987, 1993); Jordon () (Badarneh and Ghawi 1994); and
Western Australia () (Siddique et al. 1998, 2001).
springearly summer, resulting in few lled pods, a large
number of unlled 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 articial
long-day conditions, or applying vernalisation so that a wide
range of West Asian parental germplasm can be exploited
by breeders.
Indeed, our results conrm 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 owering, 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 elds
(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 deciency in traditional plain areas (Bharati and
Neupane 1991; Erskine et al. 1998; Srivastava et al. 2000).
The crossbred ILL 7986 was the rst 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 (owered 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 owering (132 days) and maturity (178 days)
in the large-seeded West Asian genotypes. Moreover, the
large leaets, lack of pubescence, and large white owers
of ILL 7986 make it clearly distinguishable from the local
landraces or cultivars in farmers elds, 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 benet 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 (194278 mm)
980 Australian Journal of Agricultural Research R. Shrestha et al.
was similar to that reported in Syria (213326 mm, Silim
et al. 1987); India (115228 mm under a range of water
supplies, Saraf and Baitha 1985), and Western Australia
(174273 mm, Siddique et al. 2001), giving us condence
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 (8.218.9 kg/ha.mm) were comparable with
those reported by Zhang et al. (2000) (9.218.1 kg/ha.mm),
Silim et al. (1993) (7.913.8 kg/ha.mm) and Siddique et al.
(2001) (8.516.7 kg/ha.mm), and the WUE
grain
for the
high-yielding genotypes (5.97.0 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) (2.47.2 kg/ha.mm) under
rainfed Mediterranean-type environments. West Asian
lentils, on the other hand, had markedly lower water use
efciencies 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 owering 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 signicantly higher yields than the
South Asian genotypes. They inherited the rapid ground
cover, high dry matter, and early owering 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 years 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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Manuscript received 11 February 2005, accepted 23 June 2005
http://www.publish.csiro.au/journals/ajar