J Agro Crop Sci (2016) ISSN 0931-2250

HEAT STRESS

Response of Mung Bean (Vigna radiata (L.) R. Wilczek) to an

Increasing Natural Temperature Gradient under Different
Crop Management Systems
M. A. P. W. K. Malaviarachchi1,2, W. A. J. M. De Costa2,3, J. B. D. A. P. Kumara2,4,
L. D. B. Suriyagoda2,3 & R. M. Fonseka2,3
1 Field Crops Research and Development Institute, Mahailluppallama, Sri Lanka
2 Postgraduate Institute of Agriculture, University of Peradeniya, Peradeniya, Sri Lanka
3 Department of Crop Science, Faculty of Agriculture, University of Peradeniya, Peradeniya, Sri Lanka
4 Faculty of Agriculture, Sabaragamuwa University of Sri Lanka, Belihul Oya, Sri Lanka

Keywords Abstract
adaptation systems; climate change;
integrated pest management; mulching; Increasing temperatures pose a significant threat to crop production in the tro-
temperature sensitivity pics. A field experiment was conducted with mung bean at three locations in Sri
Lanka representing an increasing temperature gradient (24.4–30.1 °C) during
Correspondence two consecutive seasons to (i) determine the response of mung bean to increasing
W. A. J. M. De Costa
temperature and (ii) test a selected set of crop management practices aimed at
Department of Crop Science
decreasing essential inputs such as water, synthetic pesticides and inorganic nitro-
Faculty of Agriculture
University of Peradeniya gen fertilizer. The control treatment (T1) consisted of standard crop management
Peradeniya 20400 including irrigation, chemical crop protection and inorganic fertilizer applica-
Sri Lanka tion. Adaptation system 1 (T2) included mulching with rice straw at 8 t ha 1
Tel.: +94(0)714430572 with 30 % less irrigation and crop protection and nutrient management as in T1.
Fax: +94(0)812395110 Adaptation system 2 (T3) included crop protection using a pretested integrated
Email: janendrad@gmail.com
pest management package with water and nutrient management as in T2. In
adaptation system 3 (T4), 25 % of the crop’s nitrogen requirement was given as
Accepted March 17, 2015
organic manure (compost) at 0.8 t ha 1 while 75 % was given as inorganic fertil-
doi:10.1111/jac.12131 izer with water management and crop protection as in T3. Durations of both pre-
and post-flowering phases were reduced with increasing temperature. In the
warmer (25.4–30.1 °C) yala season, seed yield (Y) of T1 decreased with increasing
temperature at 366 kg ha 1 °C 1. However, in maha season, Y did not show a
significant relationship across the narrower temperature gradient from 24.4 to
25.8 °C. Pooling the data from both seasons showed a second-order polynomial
response with an optimum temperature of 26.5 °C. In addition to shortened
durations, reduced crop growth rates and reduced pod numbers per plant were
responsible for yield reductions at higher temperatures. In yala, yields of all adap-
tation systems at all locations were on par with yields of the respective controls.
Furthermore, yala yields of T2 and T3 were less sensitive than T1 to increasing
temperatures (265 and 288 kg ha 1 °C 1). In maha, T3 and T4 had greater yields
than the control at the relatively cooler site while having lower yields than the
control at the warmer site. Maha yields of T2 were on par with the control at both
temperature regimes. While demonstrating the significant temperature sensitivity
of mung bean yields, results of the present work showed that components of the
tested adaptation systems could be promoted among smallholder farmers in Asia,
especially in view of their long-term environmental benefits and contributions to
sustainable agriculture in a warmer and drier future climate.

© 2015 Blackwell Verlag GmbH, 202 (2016) 51–68 51

Malaviarachchi et al.
Introduction
Pulse crops provide ample opportunities to diversify crop-
ping systems and to assist in managing the risks involved in
subsistence agriculture in the tropics amidst unpredictable
weather and market patterns (Siddique et al. 2012). Mung
bean, one of the important pulse crops in Asia, occupies a
prominent place in a large number of tropical cropping
systems (Weinberger 2003), because of its importance as a
source of high-quality protein in the cereal-based diets of
many people (Bogahawatte and Kailasapathy 1986, Thi-
rumaran and Seralathan 1988, Tharanathan and Maha-
devamma 2003). It is predominantly grown in the
subhumid and semi-arid tropics in low productive rainfed
farming systems (Lawn and Ahn 1985) where yield varia-
tion is high due to many biotic and abiotic stresses such as
drought (De Costa et al. 1999), a consequence of erratic
rainfall, high temperature (Cooper et al. 2009) and pest,
disease and weed incidences (Allen and Lenn
e 1998, Siddi-
que et al. 2012). Thus, even though mung bean has a high
potential yield, farmers have failed to realize this potential
yield at the field level (Hewavitharana et al. 2010, Siddique
et al. 2012). These stresses are highly likely to be aggravated
in the future with long-term climate change (Tubiello et al.
2007, Lobell and Gourdji 2012). Temperature increases will
affect a wide range of physiological (Prasad et al. 2003,
Porter and Semenov 2005, Hatfield et al. 2011, Lobell and
Gourdji 2012) and phenological (Summerfield and Lawn
1987, Rawson 1992, Summerfield et al. 1997, Lobell and
Gourdji 2012) processes of the crop and increase the fre-
quency of heat stress (Gourdji et al. 2013, Teixeira et al.
2013). In addition, temperature variations will cause
changes in biotic factors such as pest and disease dynamics
(Daugherty et al. 2009, Karuppaiah and Sujayanad 2012).
Furthermore, rates of microbial decomposition of organic
matter would increase with increasing soil temperatures
(Davidson and Janssens 2006) thus influencing soil fertility.
Therefore, development of agronomic management pack-
ages to increase productivity and resilience of tropical crop-
ping systems, which include mung bean, to the adverse
impacts of climate change is of paramount importance
(Siddique et al. 2008).
Even though the environmental control of the flowering
process of mung bean has been studied extensively in con-
trolled environmental conditions (Summerfield and Lawn
1987, Imrie and Lawn 1990) and in the field (Ellis et al.
1994), limited experimental work has been carried out on the
effects of increasing temperature on growth, biomass parti-
tioning and yield of field-grown mung bean in the tropics.
Therefore, the primary objective of this study was to deter-
mine the response of growth, biomass partitioning and yield
of mung bean to increasing growing season temperature. As
a secondary objective, effectiveness of a selected set of
52
agronomic management practices aimed at decreasing essen-
tial inputs in crop management was evaluated at different
growing season temperatures. These included decreasing the
irrigation water input by conserving soil moisture via mulch-
ing, decreasing the use of synthetic pesticides by an integrated
pest management (IPM) package and decreasing the use of
inorganic nitrogen fertilizer by replacing part of the crops’
nitrogen requirement by organic manure. Evaluation of these
modified agronomic practices, referred to as ‘adaptation sys-
tems’ in the present study, becomes relevant because adapta-
tion to climate change in relation to agriculture, especially in
the low-income farming systems in the tropics, necessitates
the adjustment of agronomic practices, agricultural processes
and capital investments in response to climate change threats
(Turner 2004, Easterling et al. 2007, Turner 2008, Siddique
et al. 2012). If effective, these practices could form the core
of an agronomic package of increased resilience to future cli-
mate change to ensure sustainable crop production in
legume-based cropping systems. Introduction of such culti-
vation technology is an urgent need in view of the fact that
yields of legumes in the tropics have not shown sustained
increasing trends during the last few decades despite the
availability of improved varieties and more resource-efficient
agronomic practices (Siddique et al. 2012).
Materials and Methods
Experimental sites
A multilocation field experiment was conducted in the sub-
humid zone of Sri Lanka during two consecutive seasons
from November 2012 to February 2013 (known locally as
the maha season) and from May 2013 to August 2013 (yala
season). The experimental locations were the Experimental
Farm of the University of Peradeniya, Kundasale (KD)
(07.28°N, 80.69°E, altitude 367 m above mean sea level),
the Field Crops Research and Development Institute,
Mahailluppallama (MI) (08.60°N, 80.27°E, 137 m amsl)
and the Regional Agricultural Research and Development
Centre, Kilinochchi (KN) (09.35°N, 80.41°E, 15 m amsl).
The three locations represented three different agro-ecolog-
ical regions of Sri Lanka (Punyawardane 2008), with KD in
IM3c (mid-elevation subhumid zone), MI in DL1b and KN
in DL3 (both in the low-elevation subhumid zone). The
annual average rainfalls of the above agro-ecological
regions are 1100–1200, 900–1100 and 800–1100 mm,
respectively (Punyawardane 2008). The two cropping sea-
sons differed in terms of their rainfall. Maha is the main
cropping season receiving rains from the north-east mon-
soon with the seasonal rainfall ranging from 300 to
800 mm depending on the location within the subhumid
zone of Sri Lanka (Panabokke and Punyawardane 2009). In
contrast, yala is the minor cropping season with a lower
© 2015 Blackwell Verlag GmbH, 202 (2016) 51–68

seasonal rainfall of 100–400 mm from the south-west mon-
soon, which is weakened when it reaches the subhumid
zone. The soils at MI, KN and KD are Rhodustalfs, Eutrus-
tox and Rhodudults, respectively, with a moderately well-
drained sandy clay loam texture (Panabokke 1996). The
initial soil N, P, K and organic matter content at a depth of
0–30 cm were 0.089 %, 24.8, 120 mg kg 1 and 1.14 % at
KD, 0.099 %, 38, 176 mg kg 1 and 0.97 % at MI, and
0.068 %, 29, 292 mg kg 1 and 0.9 % at KN.
Experimental treatments and design
The experimental treatments were four different crop man-
agement systems in a Randomized Complete Block Design
with three replicates at each site. T1, which was the control,
included the current recommended management practices
in terms of irrigation, fertilization and crop protection by
the Department of Agriculture, Sri Lanka. T2, T3 and T4
were different adaptation systems consisting of modified
agronomic packages. Adaptation system 1 (T2) included
mulching with rice straw at 8 t ha 1 for the conservation
of soil moisture with current recommended fertilization
and crop protection practices which are predominantly
based on inorganic fertilizer and synthetic pesticides. Adap-
tation system 2 (T3) included mulching with modified crop
protection to include an IPM package with recommended
nutrient management practices. Adaptation system 3 (T4)
included mulching, modified crop protection (i.e. IPM)
and modified nutrient management. Modified nutrient
management included the addition of 25 % of the crop’s
nitrogen requirement through organic manure in the form
of compost (0.8 t ha 1) while providing 75 % of the nitro-
gen requirement through inorganic fertilizer. Because of
the conserved soil moisture due to mulching, all adaptation
systems received 30 % less irrigation than the control. The
IPM package, designed after preliminary laboratory testing,
included a soil application of a 2 % bleach solution after
crop establishment for eliminating adverse effects of soil-
borne pathogens, weekly foliar applications of a baking
soda (NaHCO3) solution, monthly applications of a talc-
based biopesticide (Bacillus megaterium) to the foliage, a
one-time application of neem seed kernel extract solution
for the control of insect pests and establishment of two
border rows of maize around each plot. The plots contain-
ing the IPM treatments (i.e. T3 and T4) were separated
from plots containing conventional crop protection (T1
and T2) by a 20 m wide barrier of maize rows.
Crop establishment and maintenance
Mung bean (variety MI 6) was established by direct seeding
after disc-ploughing and harrowing the experimental field.
Plots (5 m 9 5 m) were prepared with shallow drains
© 2015 Blackwell Verlag GmbH, 202 (2016) 51–68
Temperature Response of Mung Bean
around them to drain excess water from high-intensity
rains especially during maha. The planting dates in maha
were 4th January 2013, 30th December 2012 at KD and MI
while they were 9th May 2013, 10th May 2013 and 10th
June 2013 in yala at KD, MI and KN, respectively. Even
though crops were established at KN in maha also, they
were destroyed by one event of excessive rainfall at the early
vegetative stage. An inter-row spacing of 30 cm and intra-
row spacing of 10 cm were used. Two seeds per hill were
sown at the beginning and thinned out to have a plant den-
sity of 33.33 m 2. An application of fertilizer (35 kg ha 1
of urea, 100 kg ha 1 of triple super phosphate and
75 kg ha 1 of muriate of potash) at sowing and a top
dressing (30 kg ha 1 of urea) was applied at 50 % flower-
ing in all experiments for treatments 1 (T1), 2 (T2) and 3
(T3). In contrast, 25 % of urea was reduced at each dress-
ing for treatment 4 (T4) while applying 0.8 t ha 1 of com-
post as a replacement for the reduced N from urea. Weed
control was performed manually. Depending on the experi-
mental treatment, pests and diseases were controlled by
periodic spraying of recommended chemicals or by an IPM
package (explained in the following section). Because of
asynchronous pod maturity of mung bean, the crops were
harvested in several picks with the final harvests taken on
15th March 2013 and 10th March 2013 in maha at KD and
MI and on 13th July, 9th July 2013 and 4th August 2013 in
yala at KD, MI and KN, respectively.
Measurements and data analysis
Daily weather data were recorded using an automated
weather station (Watch dog, 2900 ET) located at the exper-
imental sites. A destructive sampling of four randomly
selected plants per plot was taken to measure the leaf area
index at 50 % flowering (LAI50fl), total biomass at 50 %
flowering (TBM50fl) and total biomass at harvest (TBMh)
while an area of 1 m2 from the middle of the plot was used
for measuring seed yield and yield components. LAI50fl was
measured using an automatic leaf area meter (LI 202, CID
Bio-Science Inc., Camas, WA, USA). Dry weights of roots,
stems, leaves and pods were measured by oven drying at
60 °C to a constant weight. Crop growth up to 50 % flow-
ering (CGR50fl) was calculated as the ratio between TBM50fl
and the duration from germination up to 50 % flowering.
Post-flowering crop growth rate (CGRpfl) was calculated as
the ratio between total biomass increment from 50 %
flowering up to final harvest (i.e. TBMh–TBM50fl) and the
duration from 50 % flowering to final harvest.
The count data were subjected to nonparametric analysis
(PROC CATMOD) while parametric data were analysed
using the general linear model (PROC GLM) using the
software, STATISTICAL ANALYSIS SYSTEM (SAS), version 9.0 (Cary,
NC, USA). Since the season was nested within locations
53
Malaviarachchi et al.
and treatments (i.e. crop management systems) were nested
within locations and seasons, it was first tested whether
effects of season and effects of crop management systems
were significant within the nested system using ‘Season
(Location)’ and ‘Treatment (Season Location)’ as error
terms in the analysis of variance. Thereafter, individual sea-
sonal effects, location effects and treatment effects were
examined. Duncan’s new multiple range test (DNMRT)
was used to examine the significance of differences among
treatments at P = 0.05.
The responses of growth and yield parameters to the
temperature gradient represented by the different sites were
quantified by fitting linear or second-order polynomial
functions against the mean location temperature during
each season using regression analysis. To examine the
impacts of temperature extremes, similar regression analy-
ses were performed with mean seasonal minimum and
maximum temperatures as the respective independent vari-
ables. Linear correlation analysis was carried out to exam-
ine the contribution of different yield components to the
observed variation in seed yield across the temperature gra-
dient as represented by different locations and seasons.
Thermal durations for 50 % flowering and maturity were
calculated using daily mean temperatures and a base tem-
perature of 8 °C (Ellis et al. 1994). The dates of final har-
vest were taken for the calculation of thermal durations.
The relationships between leaf area index at 50 % flowering
(LAI50fl) and total biomass at harvest (TBMh) and between
LAI50fl and seed yield across the different locations and sea-
sons were quantified by fitting second-order polynomial
functions with the intercept fixed at zero. The LAI50fl at
which maximum TBMh and seed yield were achieved (i.e.
the optimum LAI50fl) was estimated by taking the first
derivative of the polynomial function and equating it to
zero.
Results
Variation of meteorological conditions during the two
seasons at different locations
In maha 2012/2013, a well-distributed rainfall pattern was
experienced at both sites (Fig. 1a,b). In contrast, yala 2013
was characterized by substantially lower and sporadic rain-
fall. The total rainfall during the first cropping season at
KD and MI was 530 mm and 325 mm with 22 and 27 rain
days, respectively. The corresponding values for the second
season were 2 and 26 mm with 3 and 13 rain days at KD
and MI, respectively. No rainfall was received at KN during
yala 2013. Even though there was substantial variation in
seasonal rainfall among sites in both seasons, supplemen-
tary irrigation ensured that the crops were grown without
any limitation of water. Therefore, the observed variation
54
(a)
(b)
Fig. 1 Seasonal variation of daily rainfall at Kundasale (a) and Mahail-
luppallama (b) during maha 2012/2013 and yala 2013. Kilinochchi did
not receive any rainfall during yala 2013.
of phenology, growth and yield formation of mung bean
crops among the different sites primarily represented the
response of these processes to variation in the site tempera-
ture regimes. KD experienced a lower daily mean tempera-
ture regime than MI and KN in both seasons (Fig. 2a). KN
had a higher temperature regime than MI during the sec-
ond season. The respective seasonal mean temperatures at
KD and MI were 24.4 (range 20.3–27.0 °C) and 25.8 °C
(23.1–28.3 °C) during the maha season while it was 25.4
(22.5–27.9 °C), 28.5 (25.2–30.4 °C) and 30.1 °C (29.8–
31.3 °C), respectively, at KD, MI and KN in the yala sea-
son. The respective mean minimum seasonal temperatures
(Tmin) also were higher in yala than in maha and showed
an increasing trend KD < MI < KN in both seasons
(Fig. 2b). Mean maximum seasonal temperatures (Tmax) in
yala also showed a similar trend among locations (Fig. 2c).
However, in maha, KD had a higher mean seasonal Tmax
(31.3 °C) than MI (30.0 °C).
Except in the yala season at KD, the seasonal totals of
precipitation and seasonal means of daily mean tempera-
tures experienced in the two seasons of the present experi-
ment were within the range of variation shown during the
last decade at the respective locations (Table 1). In com-
parison with the respective decadal means and ranges, the
yala at KD experienced less rainfall, but was cooler.
Daily irradiance of total solar radiation showed substan-
tial fluctuations during the course of both seasons at all
sites (Fig. 3), with the yala season showing a higher level
(10.82–27.47 MJ m 2 day 1) than the maha season (8.98–
23.40 MJ m 2 day 1). Despite the absence (KN) or very
low (KD and MI) rainfall in yala, there were an appreciable
© 2015 Blackwell Verlag GmbH, 202 (2016) 51–68
 Temperature Response of Mung Bean

(a) in yala than in maha at KD and MI. In contrast, the post-

flowering total irradiance was 16 % and 20 % lower in yala
as compared to maha at KD and MI. In comparison with
KD and MI, KN received lower seasonal total radiation in
yala. Both pre- and post-flowering irradiance totals were
also lower at KN than at KD and MI, mainly because of the
shorter durations of both phenological stages (Table 3).

Variation of crop phenology across different locations and

(b)
(c)
Fig. 2 Seasonal variation of daily mean (a), minimum (b) and maximum
(c) temperatures during maha 2012/2013 and yala 2013 at the different
experimental sites.
number of days of relatively lower irradiance
(<15 MJ m 2 day 1) due to cloudiness. Because of the
higher irradiance levels in yala, the seasonal total solar radi-
ation was greater in yala as compared to maha at KD and
MI (Table 2). Notably, total irradiance from sowing to
50 % flowering was, respectively, 16 % and 52 % greater
cropping seasons
Crop management systems and their interaction with sea-
son and location did not show significant variation with
respect to days to 50 % flowering and days to final harvest
whereas there was a significant location effect on these phe-
nological parameters (Table 3). Mung bean at KD reached
50 % flowering in 38 and 36 days in maha and yala,
respectively. In comparison, the same developmental stage
was reached within a shorter period (31 and 33 days) at MI
and 30 at KN. However, the total crop duration was the
same (70 days) at both sites in the first season. In the sec-
ond season, KD reported the longest crop duration
(65 days) while MI and KN recorded lower durations (60
and 55 days, respectively). The post-flowering duration
(i.e. that from 50 % flowering to final harvest) in maha
was greater at MI (39 days) than at KD (32 days) thus
showing an increase with increasing mean location temper-
ature. However, the opposite trend was shown in yala with
the post-flowering duration decreasing from 29 days at KD
to 25 days at KN (Table 3). The respective thermal dura-
tions for 50 % flowering and crop maturity showed only
narrow variations across locations and seasons. The ther-
mal duration for 50 % flowering varied between 493 and
550 °Cd with an average of 521 °Cd in the first season
while it was 575–599 °Cd with an average of 585 °Cd in
the second season (Table 3). On the other hand, the corre-
sponding thermal durations for maturity ranged from
1101 °Cd at KD to 1207 °Cd at MI with an average of

Table 1 Mean and range of seasonal total rainfall and seasonal mean daily mean temperature at the different experimental sites during the period
from 2004 to 2013

Mean seasonal total rainfall (mm) Mean seasonal daily mean temperature (°C)

Site Maha Yala Maha Yala

KD 288.9 (530.0)1 218.5 (2) 24.4 (24.4) 26.1 (25.4)

(74.1–659.2)2 (91.2–430.6) (23.7–24.9) (25.7–26.9)
MI 302.1 (325) 79.1 (26) 26.5 (25.8) 28.8 (28.5)
(85.3–618.4) (11.7–153.1) (25.9–27.2) (28.4–29.2)
KN * 29.3 (0) (0–95.7) * 28.9 (30.1) (29.1–31.1)

KD, Kundasale; MI, Mahailluppallama; KN, Kilinochchi.

1
Value observed in the present experiment is given in italics.
2
Range during the last decade.
*Experiment was abandoned due to heavy rains.

© 2015 Blackwell Verlag GmbH, 202 (2016) 51–68 55

Malaviarachchi et al.
1154 °Cd in the first season while it was 1052–1145 °Cd
with an average of 1108 °Cd in the second season.
Variation of vegetative growth among different crop
management systems in different locations and cropping
seasons
The seasonal effect was significant (P < 0.05) for the
majority of growth and yield parameters (Table 4), with
crop growth rate and total biomass at 50 % flowering
(CGR50fl and TBM50fl) and harvest index (HI) being the
exceptions. The effect of different crop management sys-
tems was significant for all growth and yield parameters
except HI. As the treatment effect was nested within differ-
ent locations and seasons, the observed variation of growth
and yield parameters is presented for each location and sea-
son separately.
Crop growth in different adaptation systems (i.e. T2, T3
and T4) in comparison with the control (i.e. T1) differed
with season and location (Table 5). For example, at KD
during the first season (i.e. maha), T3 treatment (i.e. mulch-
ing + IPM + standard nutrient management) showed
LAI50fl and CGR50fl values which were not significantly dif-
ferent from those of T1, which had the highest among treat-
ments. Both T3 and T4 (i.e. mulching + IPM + 25 % N
from organic source) had substantially greater CGRpfl than
the control in maha at KD. On the other hand, at MI during
the first season, crop growth in T2 (mulching + standard
crop protection and nutrient management) was on par with
Fig. 3 Seasonal variation of daily incident solar radiation during maha
2012/2013 and yala 2013 at the different experimental sites.
2
Table 2 Cumulative totals of incident solar radiation (MJ m
locations and cropping seasons
From sowing to 50 % flowering
Location Season 1 Season 2
Kundasale 646.38 747.66
Mahailluppallama 481.72 731.51
Kilinochchi * 561.43
*Experiment was abandoned due to heavy rains.
Season 1 – Maha 2012/2013; Season 2 – Yala 2013.
56
that of the control in terms of LAI50fl and CGRpfl. During
the second season (i.e. yala), at MI growth parameters of
crops in all adaptation systems were not significantly differ-
ent from those of the control. At KN also, during yala crop
growth in all adaptation systems showed LAI50fl and
CGR50fl which were on par with the control. During the sec-
ond season, at KD also, several adaptation systems showed
growth performances which were not significantly different
from the control. In particular, T4 showed similar growth as
the control in terms of all three growth parameters. More-
over, T3 at KD in yala had similar growth as compared to
the control up to 50 % flowering while T2 had similar
growth during the post-flowering period. In general, during
yala, a majority of crops growing under the three adapta-
tion systems showed growth, which was on par with the
control at their respective locations. Location mean growth
performance as measured by all three parameters was supe-
rior in yala as compared to maha at both KD and MI. Such
a seasonal comparison of crop growth was not possible for
KN as the maha crops were destroyed by excessive rain.
Notably, all three growth parameters showed decreasing
trends at the warmer locations (i.e. MI and KN relative to
KD, Table 4) indicating negative impacts on mung bean
growth with increasing temperature.
Variation of biomass partitioning and yield among
different crop management systems in different locations
and cropping seasons
In the maha season at KD, two adaptation systems (i.e. T3
and T4) produced significantly higher seed yields (Y) than
the control (Table 6). This was primarily due to their
greater total biomass at harvest (TBMh), which was mainly
because of their greater post-flowering crop growth rates
(CGRpfl) (Table 5). Conversely, at MI during maha, the
same two adaptation systems (i.e. T3 and T4) had lower
yields than the control. Here also, lower TBMh brought
about by lower CGRpfl were responsible for the lower Y of
T3 and T4. In contrast to the maha season, during yala,
yields of all adaptation systems at all locations were not sig-
nificantly (P = 0.05) different from the respective control
day 1) during pre- and post-flowering stages of mung bean grown at different
From 50 % flowering to final harvest From sowing to final harvest
Season 1 Season 2 Season 1 Season 2
575.41 486.04 1221.79 1233.69
701.16 561.43 1182.88 1292.94
* 459.64 * 1065.24
© 2015 Blackwell Verlag GmbH, 202 (2016) 51–68
 Temperature Response of Mung Bean

Table 3 Days to 50 % flowering and final harvest of mung bean grown at different locations and cropping seasons

Thermal duration1 for Days to final Thermal duration for

Days to 50 % flowering 50 % flowering (°C days) harvest2 final harvest (°C days)

Location Season 1 Season 2 Season 1 Season 2 Season 1 Season 2 Season 1 Season 2

Kundasale 38 36 549.7 580.8 70 65 1101.4 1051.6

Mahailluppallama 31 33 493.1 598.5 70 60 1207.3 1145.4
Kilinochchi * 30 * 574.7 * 55 * 1125.7

*Experiment was abandoned due to heavy rains.

1
Thermal duration was calculated using a base temperature of 8 °C (Ellis et al. 1994).
2
Final harvest of several picks.

Table 4 Probability levels of significance of the season and treatment effects in the nested treatment structure for the measured growth and yield
parameters of mung bean crops grown under different crop management systems, locations and seasons

Probability

Source of variation LAI50fl TBM50fl TBMh CGR50fl CGRpfl Yield HI

Season (location) 0.0003 0.222 0.034 0.100 0.024 0.048 0.582

Treatment (location season) 0.0012 0.027 0.001 0.004 0.009 0.021 0.601

TBM50fl and TBMh, total crop biomass per unit land area at 50 % flowering and final harvest, respectively; CGR50fl and CGRpfl, respective crop
growth rates from germination up to 50 % flowering from 50 % flowering up to final harvest; HI, harvest index.
Each value is the probability of a given source of variation being due to random variation.

Table 5 Variation of LAI at 50 % flowering (LAI50fl) and crop growth rates up to 50 % flowering (CGR50fl) and during the post-flowering period
(CGRpfl) in mung bean crops growing under different crop management systems at the three experimental locations
2
LAI50 fl CGR50fl (g m day 1) CGRpfl (g m 2
day 1)

Site System Season 1 Season 2 Season 1 Season 2 Season 1 Season 2

Kundasale T1 2.48 a 5.57 a 6.78 a 9.77 ab 7.55 b 23.49 a

T2 1.93 b 2.72 b 5.46 ab 4.53 b 8.13 b 19.91 ab
T3 2.19 ab 5.15 a 7.25 a 11.49 a 13.08 a 12.50 b
T4 1.93 b 3.59 b 4.97 b 8.18 ab 14.76 a 18.42 ab
Mean 2.13 B 4.14 A 6.11 B 8.37 A 10.88 B 18.11 A
CV (%) 9.8 22.7 27.4 33.7 25.4 23.2
Mahailluppallama T1 1.83 a 2.43 a 6.10 a 5.83 a 11.61 ab 14.12 a
T2 1.88 a 2.55 a 4.58 b 5.03 a 13.29 a 11.93 a
T3 1.07 b 2.10 a 3.33 c 6.58 a 7.89 b 13.13 a
T4 1.25 b 2.31 a 4.05 bc 4.94 a 8.54 b 12.66 a
Mean 1.51 B 2.35 A 4.51 B 5.60 A 10.33 B 12.96 A
CV (%) 12.8 13.7 9.6 22.6 21.6 12.5
Kilinochchi T1 * 1.61 a * 4.55 a * 15.02 a
T2 * 1.87 a * 5.44 a * 13.13 b
T3 * 1.72 a * 5.05 a * 6.44 c
T4 * 1.60 a * 4.19 a * 7.89 c
Mean 1.70 4.81 10.79
CV (%) 16.7 16.8 6.3
Rate of change with increasing temperature (°C 1)1 0.45 0.51 1.14 0.80 ns 1.71
R2 0.53 0.58 0.40 0.50 – 0.53

*Experiment was abandoned due to heavy rains.

Within each cropping season and location, treatment (i.e. crop management systems) means with the same lower-case letter are not significantly
different at P = 0.05. Within each location, seasonal means with the same upper-case letter are not significantly different at P = 0.05.
1
Slopes of linear regressions of treatment means against mean seasonal location temperature. A negative slope indicates significant reduction per
unit increase in temperature. ns – No significant relationship. CV - Coefficient of Variation.

© 2015 Blackwell Verlag GmbH, 202 (2016) 51–68 57

Malaviarachchi et al.

Table 6 Variation of seed yield (Y), total biomass at final harvest (TBMh) and harvest index (HI) in mung bean crops growing under different crop
management systems at the three experimental locations

Seed yield (kg ha 1) TBMh (kg ha 1) HI

Site System Season 1 Season 2 Season 1 Season 2 Season 1 Season 2

Kundasale T1 2041 b 3627 a 4992 b 8628 a 0.409 a 0.409 a

T2 1908 b 2969 a 4677 b 7405 a 0.408 a 0.399 a
T3 2998 a 2791 a 6939 a 7762 a 0.433 a 0.360 a
T4 2786 a 3189 a 6611 a 7230 a 0.422 a 0.443 a
Mean 2433 B 3144 A 5805 B 7756 A 0.418 A 0.403 A
CV (%) 14.1 26.4 8.4 16.2 13.3 13.7
Mahailluppallama T1 2113 a 2167 a 6416 a 6723 a 0.326 a 0.325 a
T2 1996 a 2057 a 6604 a 5714 a 0.304 a 0.369 a
T3 1115 b 1752 a 4108 b 6637 a 0.284 a 0.268 a
T4 1232 b 1724 a 4586 b 5933 a 0.271 a 0.290 a
Mean 1614 B 1925 A 5429 B 6252 A 0.296 A 0.313 A
CV (%) 16.1 13.0 16.9 14.0 17.7 21.6
Kilinochchi T1 * 1983 a * 5420 a * 0.368 a
T2 * 1743 a * 4748 ab * 0.370 a
T3 * 1469 a * 4098 ab * 0.365 a
T4 * 1423 a * 3389 b * 0.423 a
Mean 1655 4414 0.382
CV (%) 17.2 21.5 18.8
Temperature sensitivity (°C 1)1 585 327 ns 681 0.087 2

R2 0.45 0.85 0.79 0.93 0.60

*Experiment was abandoned due to heavy rains.

1
See the footnote of Table 4 for an explanation of mean separation symbols and temperature sensitivity.
2
Significant second-order negative polynomial relationship with a minimum estimated HI of 0.333 at 27.8 °C.

yields (Table 6). With the exception of TBMh of T4 at KN,
both TBMh and HI of all adaptation systems at all locations
in yala were on par with their respective controls. Mean
yields across different management systems were signifi-
cantly greater in yala as compared to maha at both KD and
MI. These superior yields in yala were due to the greater
TBMh, which was in turn brought about by greater CGR50fl
and CGRpfl (Table 5). In both seasons, seasonal means of
seed yield, TBMh and HI were lower in the warmer loca-
tions (i.e. MI and KN as compared to KD). Relationships
of these parameters with temperature will be examined in
the next section.
Response of growth, biomass partitioning and yield to the
temperature gradient across locations during different
cropping seasons
Crops growing under currently recommended standard man-
agement practices (T1)
Variation of crop growth and yield among locations in the
control treatment (i.e. T1) was considered to explore the
response of mung bean to a mean seasonal temperature
gradient as soil moisture deficits were avoided through irri-
gation. In the yala season, TBMh and seed yield of T1 had
significant linear relationships with mean seasonal temper-
ature across the experimental sites representing the temper-
ature gradient (P = 0.0044 and P = 0.002, respectively).
TBMh and seed yield decreased by 452 and
366 kg ha 1 °C 1 across the increasing temperature gradi-
ent (Fig. 4a,b). However, when the linear regressions were
carried out across the mean seasonal temperatures of the
two locations in maha, no significant (P = 0.05) relation-
ships were found (Fig. 4a,b) probably because of the very
narrow range of mean seasonal temperatures in maha. On
the other hand, the HI of T1 showed significant (P < 0.05)
decreasing trends with increasing temperature in both sea-
sons (Fig. 4c) at 0.115 and 0.0236 °C 1 in maha and yala,
respectively. Pooling the data from both seasons showed
second-order polynomial response patterns for both Y and
TBMh against mean seasonal temperature (Fig. 4a,b). The
estimated optimum temperatures for Y and TBMh were
26.5 and 27.3 °C, respectively. The estimated maxima for Y
and TBMh at the respective optimum temperatures were
2671 and 7095 kg ha 1. HI also showed a negative second-
order polynomial response with a minimum HI of 0.35
being reached at 28.0 °C (Fig. 4c).
Response of growth, biomass partitioning and yield of crops
growing under different adaptation systems to the tempera-
ture gradient during different cropping seasons
Within a given season, the rates of biomass and yield
reductions of the different adaptation systems (i.e. T2, T3

58 © 2015 Blackwell Verlag GmbH, 202 (2016) 51–68

 Temperature Response of Mung Bean

(a) (b)

(c)

Fig. 4 Relationships between seasonal mean

location temperature and total biomass at har-
vest (TBMh) (a), seed yield (Y) (b) and harvest
index (HI) (c) in mung bean crops grown with
standard recommended management prac-
tices (i.e. the control treatment, T1). ○ – Maha
2012/2013 season’s data; ▲ – Yala 2013 sea-
son’s data. Each data point is a value for a rep-
licate plot at a given location. Solid lines
represent linear temperature response func-
tions of different seasons. Curves represent
polynomial temperature response functions
for pooled data from both seasons.

and T4) in response to increasing temperature were
observed to be within a narrow range (Fig. 5a–d). The
respective rates ranged between 563 and 763 kg ha 1 °C 1
and 265 and 389 kg ha 1 °C 1 for TBMh and yield, respec-
tively, during yala 2013. However, in the maha season,
TBMh and yield of T3 and T4 decreased at comparatively
higher rates (2022, 1446 kg ha 1 °C 1 and 1345,
1110 kg ha 1 °C 1). In contrast, in T2, TBMh increased by
1377 kg ha 1 °C 1 whereas yield did not vary significantly
with temperature. On the other hand, in all adaptation
systems, HI showed significant (P < 0.05) negative second-
order polynomial relationships with mean location temper-
ature in yala (Fig. 5f). The respective temperatures at
which HI was minimum were 28.0, 27.9 and 27.6 °C for
T2, T3 and T4, respectively, with the estimated minimum
HIs being 0.38, 0.31 and 0.27. In contrast to yala, in maha
only T2 showed a significant negative linear relationship
( 0.036 °C 1) with increasing temperature (Fig. 5e).
Relationships between leaf area index at 50 % flowering,
total biomass at harvest and seed yield across different
experimental treatments, locations and seasons
When pooled across experimental treatments and loca-
tions, LAI50fl, one of the principal crop growth parameters
had significant second-order polynomial relationships with
TBMh and seed yield in maha (P = 0.016 and P = 0.0003,
respectively) and in yala (P = 0.0005 and P = 0.0005,
respectively) seasons (Fig. 6 with polynomial relationships
not shown). Furthermore, when the data from both seasons
were pooled, LAI50fl showed highly significant (P < 0.0001)
second-order polynomial relationships with TBMh and
seed yield (Fig. 6a,b). Both TBMh and seed yield reached
their respective maxima of 8137.7 and 3122.0 kg ha 1
when the LAI50fl reached 4.30 and 4.70, respectively.
Yield components and their relationships to yield
All three yield components showed significant variation
between the two seasons (Table 7). On the other hand, the
different crop management systems (i.e. the treatment
effect) had significant effects on the pod number (PN) per
plant only. As the treatment effect was nested within loca-
tions and seasons, variation of yield components in differ-
ent crop management systems is shown separately for
different locations and seasons (Table 8). At KD, PN
showed significant variation among treatments in both sea-
sons. While T3 had significantly greater PN than the con-
trol in maha, PN of the other adaptation systems (i.e. T2
and T4) was not significantly different from that of the con-
trol. In yala at KD, all adaptation systems had PN which
were equal to the control. In contrast to KD, at MI, PN did

© 2015 Blackwell Verlag GmbH, 202 (2016) 51–68 59

Malaviarachchi et al.

(a) (b) (c)

(d) (e) (f)

Fig. 5 Relationships between seasonal mean location temperature and total biomass at harvest (TBMh) (a and b), seed yield (Y) (c and d) and harvest
index (HI) (e and f) in mung bean crops grown with different adaptation systems (□ – T2, ○ – T3, ▲ – T4) in maha 2012/2013 (a, c and e) and yala
2013 (b, d and f) seasons. Each data point is a value for a replicate plot at a given location.

(a) (b)

Fig. 6 Relationships between leaf area index at 50 % flowering (LAI50fl) and total biomass at harvest (TBMh) (a) and seed yield (Y) (b) of mung bean
crops grown with standard crop management (i.e. T1) and in different adaptation systems (i.e. T2, T3 and T4) across the temperature gradient repre-
sented by different locations and seasons. Each data point is a value for a replicate plot at a given location. ○ – Maha 2012/2013 season’s data;
▲ – Yala 2013 season’s data.

not differ significantly among crop management systems in than in maha. In contrast to PN, the seed number (SN) per
both seasons. At KN in yala, T3 and T4 had greater PN than pod did not show significant variation between different
the control. Notably, when averaged across treatments, treatments in all sites and seasons. When averaged across
mean PN of both KD and MI was greater (P < 0.05) in yala treatments, mean SN of maha was greater than that of yala

60 © 2015 Blackwell Verlag GmbH, 202 (2016) 51–68

 Temperature Response of Mung Bean

at MI. However, at KD, mean SN did not differ between (P = 0.05) correlations with Y in both seasons. In addition,
the two seasons. Individual seed weight (ISW) did not dif- several significant negative correlations were observed
fer significantly between treatments at all sites and seasons between different yield components in different seasons.
with the exception of yala at MI, where T4 showed lower These included the negative correlations between PN and
ISW than the control. At both KD and MI, seasonal mean SN and between SN and ISW in the maha season and that
ISW was greater in maha than in yala. Of the three yield between PN and ISW in yala.
components, only PN showed significant declining trends Different yield components showed significant positive
with increasing seasonal mean temperatures across loca- correlations with different vegetative growth parameters.
tions in both seasons (Table 8). These correlations were different in the two seasons. For
The PN showed highly significant (P < 0.0001) positive example, PN was positively correlated to LAI50fl, TBM50fl
correlations with seed yield (Y) in both seasons (Table 9). and TBMh in both seasons (Table 9). On the other hand,
The SN was positively correlated (P < 0.0001) to Y in the SN and ISW were not correlated to any of the above growth
yala season only. The ISW did not have significant parameters in maha. However, in yala, SN showed signifi-
cant positive correlations with LAI50fl and TBM50fl while
ISW was negatively correlated to TBMh.
Table 7 Probability levels of significance of the season and treatment
Similarly, correlations of yield components to the HI
effects in the nested treatment structure for the yield components of
were also different in the two seasons. While none of the
mung bean crops grown under different crop management systems,
locations and seasons yield components were significantly correlated to the HI in
maha, PN and SN had significant positive correlations with
Probability HI in yala (Table 9). In both seasons, HI did not show sig-
Pods per Seeds per Individual seed nificant correlations with any of the growth parameters.
Sources of variation plant pod weight Interestingly, a significant positive correlation was observed
between Y and HI in the yala season only. In contrast, Y
Season (location) 0.006 0.001 <0.0001
showed highly significant positive correlations with TBMh
Treatment (location season) 0.012 0.717 0.531
and all other growth parameters (i.e. LAI50fl and TBM50fl)
Each value is the probability of a given source of variation being due to in both seasons. When the data for both seasons were
random variation. pooled, Y showed a much stronger positive correlation with

Table 8 Variation of pod number per plant (PN), seeds per pod (SN) and individual seed weight (ISW) in mung bean crops growing under different
crop management systems at the three experimental locations
1 1
PN pl Seeds pod ISW (mg)

Site System Season 1 Season 2 Season 1 Season 2 Season 1 Season 2

Kundasale T1 13.1 bc 20.8 ab 8.9 a 9.8 a 59.8 a 51.3 a

T2 12.1 c 21.3 a 8.1 a 8.3 a 61.8 a 51.0 a
T3 18.3 a 16.0 b 9.4 a 10.3 a 58.2 a 50.7 a
T4 16.8 ab 19.3 ab 9.2 a 9.8 a 61.8 a 50.7 a
Mean 15.1 B 19.4 A 8.9 A 9.6 A 60.4 A 50.9 B
CV (%) 13.5 12.2 13.5 14.8 4.1 4.2
Mahailluppallama T1 10.4 a 14.3 a 10.7 a 8.8 a 57.2 a 55.8 a
T2 10.5 a 14.3 a 10.1 a 8.7 a 58.0 a 52.4 ab
T3 7.9 a 11.0 a 10.4 a 9.1 a 58.2 a 52.8 ab
T4 9.3 a 12.3 a 10.3 a 8.5 a 56.8 a 50.4 b
Mean 9.5 B 13.0 A 10.4 A 8.8 B 57.5 A 52.9 B
CV (%) 21.9 13.5 8.7 4.9 1.9 4.2
Kilinochchi T1 * 12.7 a * 9.1 a * 54.5 a
T2 * 12.3 a * 9.3 a * 55.1 a
T3 * 9.3 b * 8.4 a * 56.4 a
T4 * 10.3 b * 9.0 a * 54.7 a
Mean 11.2 9.0 55.2
CV (%) 6.8 10.0 4.2
Temperature sensitivity (°C 1)1 4.0 1.8 1.1 ns 2.0 0.9
R2 0.67 0.85 0.79 – 0.61 0.64

*Experiment was abandoned due to heavy rains.

1
See the footnote of Table 4 for an explanation of mean separation symbols and temperature sensitivity.

© 2015 Blackwell Verlag GmbH, 202 (2016) 51–68 61

Malaviarachchi et al.

Table 9 Linear correlation coefficients between seed yield, yield components, total biomass and harvest index of mung bean crops growing under
standard crop management and different crop adaptation systems across different locations in maha 2012/2013 (above the diagonal) and yala
(below the diagonal) seasons
1 1
Yield HI TBMh LAI50fl TBM50fl Pods pl Seeds pod ISW

Yield – 0.053ns 0.774*** 0.678*** 0.651*** 0.896*** 0.232ns 0.113ns

HI 0.580*** – 0.207ns 0.043ns 0.075ns 0.107ns 0.219ns 0.083ns
TBMh 0.783*** 0.042ns – 0.488* 0.365ns 0.563** 0.076ns 0.104ns
LAI50fl 0.554*** 0.174ns 0.572*** – 0.825*** 0.592** 0.390ns 0.096ns
TBM50fl 0.532*** 0.163ns 0.592*** 0.950*** – 0.651*** 0.374ns 0.117ns
Pods pl 1 0.877*** 0.522** 0.703*** 0.370* 0.351* – 0.479* 0.175ns
Seeds pod 1
0.620*** 0.392* 0.289ns 0.588*** 0.550*** 0.295ns – 0.584**
ISW 0.315ns 0.157ns 0.499** 0.261ns 0.250ns 0.489** 0.051ns –

HI, harvest index; TBMh and TBM50fl, total biomass at harvest and 50 % flowering; LAI50fl, leaf area index at 50 % flowering; ISW, mean individual
seed weight.
Each correlation had 24 and 36 data points in maha and yala, respectively. ns – Non-significant at P = 0.05; * – Significant at P < 0.05;
** – Significant at P < 0.01; *** – Significant at P < 0.001.

TBMh (r = 0.778 with P < 0.0001) than with HI
(r = 0.284 with P = 0.028).
Effects of seasonal temperature extremes
Effects of minimum and maximum temperatures were
examined on mung bean crops growing under standard
crop management (i.e. T1) (Table 10). When the data for
both seasons were pooled, both TBMh and Y showed signif-
icant second-order polynomial relationships with mean
seasonal minimum temperature (Tmin) across different
locations. The respective optimum Tmin for maximum
TBMh and Y were 23.0 and 23.4 °C. Across the two
seasons, the HI also showed a negative second-order poly-
nomial response to mean seasonal Tmin with the minimum
HI at 24.6 °C. When the responses to mean seasonal Tmin
were examined separately for the two seasons, negative lin-
ear relationships were shown for Y and HI. Seed yield (Y)
showed a greater sensitivity (as indicated by the slope of
the fitted linear function) to increasing Tmin during the
warmer yala season with a yield reduction of
422 kg ha 1 °C 1 as compared to the cooler maha season
( 26 kg ha 1 °C 1). However, the HI showed a greater
sensitivity to increasing Tmin during the maha season. In
contrast to Y and HI, TBMh showed a second-order poly-
nomial response to increasing Tmin during yala while show-
ing a positive response in maha.
In contrast to the response to Tmin, TBMh and Y showed
negative linear responses to increasing mean seasonal maxi-
mum temperatures (Tmax), for both seasons separately and
when the data for the two seasons were pooled (Table 10).
The sensitivity to increasing mean seasonal Tmax was
greater in yala for Y. However, the opposite was observed
for TBMh. The HI showed a positive response to increasing
Tmax in maha, but a negative response in yala. When the
data for the two seasons were pooled, HI did not show a
significant variation across the range of mean seasonal
Tmax.
Discussion
The principal objective of the present work was to assess
the sensitivity of mung bean yields to increasing tempera-
ture across different locations representing a natural tem-
perature gradient over two contrasting cropping seasons.
In terms of the seasonal mean temperature, the first season
(i.e. maha 2012/2013) had a narrower temperature range of
1.4 °C (from 24.4 to 25.8 °C) across the different locations.
In contrast, crops in the second season (yala 2013) experi-
enced a broader temperature range of 4.7 °C (25.4–
30.1 °C). Comparison with the meteorological data at the
locations during the last decade (Table 1) showed that the
two cropping seasons were largely representative of the cli-
matic conditions experienced in these locations. A primary
assumption in this multilocational experimental set-up was
that temperature would be the principal environmental fac-
tor controlling growth, biomass partitioning and yield of
mung bean crops. Impacts of precipitation variation
among locations were largely eliminated by irrigating the
crops. Excess water from high-intensity rainfall was
removed by drains between plots. Any effects on crop
growth and yield formation due to soil variation among
sites were eliminated by providing recommended doses of
nutrients as inorganic (T1, T2 and T3) or a combination of
inorganic and organic (T4) fertilizer. Any variation among
sites in pest and disease incidence was controlled via con-
ventional chemical-based (T1 and T2) or IPM (T3 and T4)
methods. Therefore, it was assumed that the effects of any
uncontrolled variations among different sites would not
override and obscure the principal response to the

62 © 2015 Blackwell Verlag GmbH, 202 (2016) 51–68

 Temperature Response of Mung Bean

Table 10 Fitted relationships of total biomass at harvest (TBMh), seed yield (Y) and harvest index (HI) with mean minimum and maximum seasonal
temperatures (Tmin and Tmax) for mung bean crops growing under standard crop management (T1) across three experimental locations and two crop-
ping seasons

Season TBMh (kg ha 1) Y (kg ha 1) HI

Tmin
Maha TBMh = (234*Tmin) + 556 (R2 = 0.348) Y=( 26*Tmin) + 1547 (R2 = 0.008) Y = ( 0.059*Tmin) + 1.603 (R2 = 0.497)
Yala TBMh = ( 512*Tmin2
) + (24034*Tmin) + 273395 Y=( 422*Tmin) + 12993 Y = ( 0.029*Tmin) + 1.151
(R = 0.644) (Optimum Tmin = 23.47 °C)
2
(R2 = 0.533) (R2 = 0.446)
Both1 TBMh = ( 121*Tmin2
) + (5580*Tmin) + 57277 Y=( 2
74*Tmin ) + (3336*Tmin) + 34492 Y = (0.0025*Tmin
2
) (0.1228*Tmin) + 1.91
(R2 = 0.402) (Optimum Tmin = 23.01 °C) (R2 = 0.227) (Optimum Tmin = 23.42 °C) (R2 = 0.08)
(Tmin for minimum HI = 24.56 °C)
Tmax
Maha Y = ( 488*Tmax) + 20273 (R2 = 0.348) Y = ( 54*Tmax) + 3743 (R2 = 0.008) Y = (0.122*Tmax) 3.326
(R2 = 0.497)
Yala Y = ( 416*Tmax) + 19874 (R2 = 0.572) Y = ( 341*Tmax) + 13469 Y = ( 0.022*Tmax) + 1.156
(R2 = 0.508) (R2 = 0.390)
Both1 Y = ( 207*Tmax) + 12543 (R2 = 0.112) Y = ( 234*Tmax) + 9738 (R2 = 0.24) Y = ( 0.0082*Tmax) + 0.6935
(R2 = 0.025)
1
Data from both seasons pooled.

temperature gradient. Accordingly, results of the present
study are especially relevant in view of the conclusion of
Lobell and Burke (2008) that uncertainty in the sensitivity
of crop yields to temperature variation rather than to pre-
cipitation variation constitutes a greater proportion to the
uncertainty of estimated impacts of climate change on crop
yields.
Our results, especially those of the second season where
the crops experienced a broader temperature range, show
that despite being a warm-season grain legume that is
adapted to higher temperature regimes (Rachie and Rob-
erts 1974), mung bean yields are reduced significantly when
growing temperatures increased from ca. 25–30 °C
(Fig. 3). It is notable that this range of mean seasonal tem-
peratures is well within the temperature range across which
mung bean is currently grown in different parts of the
world (Lawn and Ahn 1985). Furthermore, the current
growing season temperatures in many parts of the tropics
where mung bean is grown exceed the estimated optimum
temperature of 26.5 °C for maximum yield (Christensen
et al. 2007, Lobell et al. 2008). Therefore, it is highly likely
that future increases in air temperature due to the
enhanced greenhouse effect (IPCC 2013) would have nega-
tive impacts on future mung been yields in Sri Lanka and
South-East Asia. This is in agreement with the findings of
many studies in which negative impacts of future warming
on the yields of several major food crops (e.g. wheat, maize,
rice and soya bean) have been shown both at the global
(Lobell and Field 2007, Lobell and Gourdji 2012, Teixeira
et al. 2013) and regional (Lobell and Asner 2003, Lobell
et al. 2008, Cooper et al. 2009, Schlenker and Roberts
2009, Schlenker and Lobell 2010, Hatfield et al. 2011)
scales. As most of the published studies have focused on
major food crops of global importance, to our knowledge,
the present work constitutes the first published report on
the sensitivity of mung bean yields to seasonal warming
across a multilocational temperature gradient. The esti-
mated temperature optimum of 26.5 °C for mung bean in
the present work is higher than the ranges of 22–23 °C esti-
mated for soya bean (Hatfield et al. 2011, Lobell and Gour-
dji 2012) and 23–24 °C for peanut (Prasad et al. 2003) and
red kidney bean, that is Phaseolus vulgaris (Laing et al.
1984, Prasad et al. 2002).
Analysis of yield responses to seasonal means of mini-
mum (Tmin) and maximum (Tmax) temperatures
(Table 10) showed that yields of mung bean (Y) are more
sensitive to Tmax than Tmin. This is in agreement with the
results of Lobell and Field (2007) who found greater
impacts of increasing temperatures on yields of wheat and
maize when Tmax and Tmin were used instead of seasonal
means of daily mean temperatures (Tavg). This is demon-
strated by the lower optimum for Tmin (i.e. 23.4 °C) as
compared to the optimum for Tavg (26.5 °C) and by the
negative linear response of Y to Tmax. Our analysis also
showed that mung bean yields are more sensitive to
increasing Tmin and Tmax in the warmer yala season than in
the relatively cooler maha.
Results of the present experiment show that high tem-
perature-induced yield reductions in mung bean occur due
to reductions in both total biomass at harvest (TBMh) and
HI. This was shown especially in the second season when a
wider temperature range was experienced. This indicates
that higher temperatures influence both vegetative and
reproductive growth and also both developmental and

© 2015 Blackwell Verlag GmbH, 202 (2016) 51–68 63

Malaviarachchi et al.
growth processes. These observations agree with the
expected responses based on the effects of increasing tem-
peratures on basic physiological processes (Hatfield et al.
2011, Lobell and Gourdji 2012). The influence of tempera-
ture variation on developmental processes could be
observed in the variation of durations to 50 % flowering
and maturity, with increasing temperatures clearly hasten-
ing the flower initiation and maturity. These shortened
durations clearly contributed to the yield reductions
observed at higher temperatures. This is in agreement with
the conclusions of reviews of previous experimental work
by Hatfield et al. (2011) and Lobell and Gourdji (2012)
and the specific predictions of Cooper et al. (2009) for
yields of peanut and pigeon pea at higher temperatures.
The estimated thermal durations for flowering and matu-
rity in the present work were comparable to those of other
work on mung bean. For example, Chauhan et al. (2010)
estimated the thermal time for flowering and maturity for a
typical mung bean variety (Emarald) grown in Australia to
be 595 and 1200 °Cd, respectively, at a base temperature of
7.5 °C, which are within the range of our values, which
were estimated at a base temperature of 8 °C (Table 3).
The small seasonal differences in the thermal durations
observed in the present study might have been caused by
photothermal variations as even slight variations in photo-
period can alter the initiation of flowers in mung bean
(Summerfield et al. 1997). Similar photothermal sensitivity
of flowering has been shown in other grain legumes such as
soya bean (Hatfield et al. 2011) and for annual crops in
general (Craufurd and Wheeler 2009).
The increasing temperatures across the locations clearly
reduced vegetative growth as shown by the decreasing sea-
sonal means of LAI50fl, CGR50fl, CGRpfl (Table 5) and
TBMh (Table 6). These reductions reflect the negative
impacts of increasing temperatures on the processes of leaf
expansion, radiation interception, photosynthesis and bio-
mass accumulation (Squire 1990, Porter and Semenov
2005, Hatfield et al. 2011, Lobell and Gourdji 2012) indi-
cating that the temperature range experienced across loca-
tions in the present study exceeded the respective optimum
temperatures for the above processes. While the high tem-
perature-induced reductions in LAI50fl and TBMh were
partly caused by the shortened durations of the pre- and
post-flowering stages (Table 3), the reductions in CGR50fl
and CGRpfl showed that the rates of key physiological pro-
cesses responsible for growth were also reduced at the
supra-optimal temperatures experienced by crops of the
present work. However, it is notable that within a given
location, all the above growth parameters were higher in
the warmer yala season as compared to the cooler maha
season (Tables 4 and 5). This difference between the two
seasons was most probably caused by the greater irradiance
levels in the yala season (Table 2), especially during the
64
period from planting up to 50 % flowering, which allowed
greater radiation interception and biomass production.
The observed relationships of LAI50fl with TBMh (Fig. 6a)
and seed yield (Fig. 6b) demonstrate the importance of
early vegetative growth in yield determination of mung
bean. Therefore, greater irradiance levels during the vegeta-
tive phase enabled the yala crops to achieve greater yields
than in maha despite the warmer temperatures. This find-
ing indicates that, if irrigation is available, growing crops
during periods of greater irradiance, especially during the
vegetative stage, enable achievement of higher yields at
higher temperatures.
The stronger correlations between yield and TBMh as
compared to those between yield and HI (Table 9) indi-
cate that increasing temperatures have a greater influence
on processes responsible for biomass production than on
processes determining biomass partitioning to yield. How-
ever, this does not mean that the reproductive processes
of mung bean are not sensitive to increasing temperatures.
Observed variations in the duration from germination to
50 % flowering (Table 3) showed that flower initiation
was sensitive to even the narrow temperature range
between locations in the first season. The clear reductions
in PN per plant with increasing temperatures across loca-
tions (Table 8) indicate the possible adverse impacts of
heat stress on the number of flowers formed, their fertil-
ization and retention of the fertilized flowers and young
pods. These results of the present study affirm the high
sensitivity of crop reproductive processes of legumes to
higher temperatures (Gross and Kigel 1994, Prasad et al.
2000, 2001, 2002, 2003, Lobell and Field 2007, Hatfield
et al. 2011). The highly significant positive correlations
that were observed between seed yield and PN in both sea-
sons showed that PN is a key determinant of yield in
mung bean. Thus, increasing temperatures exert a key
negative impact on mung been yields by decreasing PN
(Table 8). Furthermore, the significant positive correla-
tions of PN with all growth parameters (i.e. LAI50fl,
TBM50fl and TBMh) (Table 9) indicate that the magnitude
of vegetative growth partially influences the number of
pods retained probably by controlling the assimilate
supply to developing pods. In addition to PN, SN per pod
(SN) also exerted a significant influence on yield determi-
nation when the crops were subjected to a higher temper-
ature range in the yala season (Table 9). This was
probably because of adverse impacts of higher tempera-
tures on fertilization and seed formation (Prasad et al.
2001, 2003, Hatfield et al. 2011). The significant negative
correlations that were observed between different yield
components in the two seasons (Table 9) indicated that
the yield formation process of mung bean possesses a cer-
tain degree of flexibility of adapting to higher tempera-
tures by increasing SN and ISW when PN is decreased.
© 2015 Blackwell Verlag GmbH, 202 (2016) 51–68

Performance of adaptation systems
The ‘adaptation systems’ that were tested in the present
study were aimed at reducing the inputs of water, synthetic
pesticides and inorganic nitrogen fertilizer. As such the spe-
cific adaptations tested here are components of ecosystem-
based approaches and conservation agriculture (Siddique
et al. 2012), which are part of innovative cultivation tech-
nologies especially aimed at resource-limited smallholder
farmers in the tropics. A summary of seed yields of the
adaptation systems in comparison with the respective con-
trols (i.e. the standard crop management practices)
(Table 6) show that with the exception of the maha season
at MI, performances of the three adaptation systems were
either superior (e.g. T3 and T4 at KD in maha) to or on par
with the control (e.g. T2, T3 and T4 at all locations in yala).
When the reduced cost of inputs and the long-term bene-
fits to the soil (through mulching in T2 and organic matter
addition in T4) and the environment (through reduced
application of synthetic pesticides and inorganic nitrogen
fertilizers) are taken in to consideration, the modified agro-
nomic and crop protection practices which formed the
adaptation systems can be recommended to upland crop-
ping systems in the tropics.
The success of IPM in the present work in either obtain-
ing higher yields over chemical-based pest and disease con-
trol or maintaining the same yield levels is a notable
achievement, especially in view of the pesticide-related
environmental and health hazards that have been reported
in Sri Lanka (Van der Hoek et al. 1998, Aponso et al. 2003,
Marasinghe et al. 2011). Similar success stories of IPM in
legumes have been reported for chickpea in India (Singh
et al. 2003) and northern Bangladesh (Harris et al. 2008)
and soya bean in Brazil (Bueno et al. 2011). Adaptation
systems T3 and T4 at MI in maha represented the only situ-
ation where the performance of an adaptation system was
inferior to that of the control (Table 6). This was because
of the higher incidence of the mung bean pod borer (Maru-
ca vitrata), which could not be controlled by the IPM pack-
age practised in these two adaptation systems. This is in
agreement with the general consensus about IPM that its
success is not universal and could be location and season
specific (Landa et al. 2004, Van den Berg 2004).
The effectiveness of mulching to conserve soil moisture
and reduce the irrigation requirement was demonstrated
by the observation in the present study that seed yield of
the T2 treatment (i.e. the adaptation system in which
mulching and reduced water input was the only agronomic
modification from the control treatment) was always on
par with that of the respective control (T1) (Table 6). This
finding is especially relevant in the subhumid zone of Sri
Lanka where soil water deficits constitute a major con-
straint to increasing the productivity of upland cropping
systems (Abeyrathna 1956).
© 2015 Blackwell Verlag GmbH, 202 (2016) 51–68
Temperature Response of Mung Bean
The estimated yield reductions per unit of increased tem-
perature (Figs 4b and 5c,d) indicated the sensitivity of dif-
ferent crop management systems to increasing
temperature. In the warmer yala season, which experienced
a wider temperature range, yields of T2 and T3 showed
lower temperature sensitivity (Fig. 5d) than the control
(Fig. 4b) while the sensitivity of T4 was on par with that of
the control. In the cooler maha season, which experienced
a narrower temperature range, T2 showed a substantial
positive yield response to increasing temperatures (Fig. 5c).
The beneficial effects of mulching in making a cropping
system more resilient to increasing temperature are in
agreement with the simulated beneficial impacts of mulch-
ing on crops in the semi-arid tropics (Cooper et al. 2009).
In contrast, yields of T3 and T4 showed substantially greater
temperature sensitivity than the control. This was because
of the significantly lower yields of T3 and T4 at the warmer
site (i.e. MI) in maha (Table 6). Despite this, on the whole,
results of the present work indicate the likelihood that these
adaptation systems have a greater resilience to the expected
temperature increases in a future climate.
Characters of mung bean varieties suitable for a warmer and
drier future climate
Temperature responses of the measured growth parameters
and yield components and their interrelationships provide
indications to plant breeders about characters to be incor-
porated in future mung bean varieties to achieve higher
yields in a warmer and drier climate. Varieties with higher
crop growth rates, both during pre- and post-flowering
stages need to be developed to compensate for reduced
crop durations. For this, varieties which are able to achieve
the optimum LAI (leaf area index) of ca. 4.5 (Fig. 6) within
the shortest possible duration and maintain it for a maxi-
mum duration need to be bred. The strong positive corre-
lation between seed yield and PN means that future
varieties should be capable of maximum pod initiation and
retention in warmer temperatures.
Acknowledgements
This work was funded by the Higher Education for Twenty
First Century (HETC) Quality and Innovation Grant Win-
dow 3 (QIG-3) project. The Integrated Pest Management
(IPM) package was formulated by Dr. Devika M. De Costa
with assistance from Nadeeka Dharmadasa.
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