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Growth and development of Virginia type groundnutcultivars under Mediterranean conditionsSevgi Caliskan a , M.E. Caliskan a , E. Erturk b , M. Arslan a & H. Arioglu ca Department of Crop Science, Faculty of Agriculture , Mustafa Kemal University , 31040,Hatay, Turkeyb Department of Horticulture, Faculty of Agriculture , Mustafa Kemal University , 31040,Hatay, Turkeyc Department of Crop Science, Faculty of Agriculture , Cukurova University , 01330,Adana, TurkeyPublished online: 13 Dec 2007.
To cite this article: Sevgi Caliskan , M.E. Caliskan , E. Erturk , M. Arslan & H. Arioglu (2008) Growth and development ofVirginia type groundnut cultivars under Mediterranean conditions, Acta Agriculturae Scandinavica, Section B — Soil & PlantScience, 58:2, 105-113, DOI: 10.1080/09064710701312041
To link to this article: http://dx.doi.org/10.1080/09064710701312041
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ORIGINAL ARTICLE
Growth and development of Virginia type groundnut cultivars underMediterranean conditions
SEVGI CALISKAN1, M.E. CALISKAN1, E. ERTURK2, M. ARSLAN1 & H. ARIOGLU3
1Department of Crop Science, Faculty of Agriculture, Mustafa Kemal University, 31040 Hatay, Turkey, 2Department of
Horticulture, Faculty of Agriculture, Mustafa Kemal University, 31040 Hatay, Turkey, and 3Department of Crop Science,
Faculty of Agriculture, Cukurova University, 01330 Adana, Turkey
AbstractThe growth and development of groundnut (Arachis hypogaea L.) are under the influence of complex environmental factors.Understanding of the growth responses of the groundnut to environmental factors may improve the application of bettermanagement practices and develop better cultivars to overcome the problems causing reductions in yield. A two-year fieldexperiment was conducted to determine the growth and development response of groundnut genotypes to environmentalfactors in the eastern Mediterranean region of Turkey in 2001 and 2002. Time from sowing to physiological maturity (R8)ranged from 25138Cd to 25888Cd in 2001 and from 25148Cd to 25738Cd in 2002 while total calendar days varied between147 and 153 and between 156 and 161 depending on genotypes in 2001 and 2002, respectively. Dry matter accumulation ineach part of the plants continued until maturity although accumulation rate differed depending on plant age. Combinationof suitable temperature and photoperiod during the reproductive stages resulted in continuous and abundant reproductiveplant parts, which led to delayed harvest and increased unmarketable pods. The slower growth rate due to the coolerconditions during early stages caused slower biomass accumulation in successive stages indicating the importance of initialcrop growth for final yield. Therefore, the genotypes having high initial growth rate, less reproductive organs, and shortergrowing period should be developed for the Mediterranean conditions by breeders. The management studies shouldalso deal with increased initial growth rate and reduced number of flowers, pegs or pods per plant. Based on ourresults, groundnut has a great yield potential under the Mediterranean conditions. However, further breeding andmanagement studies are needed to improve the yield and profitability and reduce the complications arisen from theMediterranean climate.
Keywords: Dry matter production, groundnut, partitioning, phenological development.
Introduction
Groundnuts (Arachis hypogaea L.) are grown in
several agro-ecological systems and under numerous
socio-economic environments throughout the tropi-
cal and warm temperate regions of the world (Isleib
et al., 1994). The world annual groundnuts produc-
tion is around 35.6 million tonnes from the 26.4
million ha of production area (FAO, 2005). Ground-
nut production in the Mediterranean region is
limited although a long growing period over five
months as well as high yield potential exist in this
region. However, it was foreseen that groundnut
production could be enormously increased in the
Mediterranean basin under irrigated conditions in
the future (Smartt, 1994). Hence, in order to assess
the scope for groundnut production in these types of
environments, it is necessary to understand growth
and development as well as the factors limiting the
yield of groundnut.
Groundnut yield is a product of crop growth rate,
the partitioning of assimilates to reproductive sinks
and the duration of the crop’s reproductive phase
(Duncan et al., 1978). However, each stage is greatly
influenced by the complex environmental factors in
which the genetically controlled characteristics of the
cultivar, weather conditions, soil water regime, and
incidence of insect pests and diseases play an
important role (Kaur and Hundal, 1999). Among
Correspondence: Sevgi Caliskan, Department of Crop Science, Faculty of Agriculture, Mustafa Kemal University, 31040 Hatay, Turkey. Tel: (�90) 326 245
58 26. Fax: (�90) 326 245 58 32. E-mail: [email protected]
Acta Agriculturae Scandinavica Section B � Soil and Plant Science, 2008; 58: 105�113
(Received 23 May 2006; accepted 17 November 2006)
ISSN 0906-4710 print/ISSN 1651-1913 online # 2008 Taylor & Francis
DOI: 10.1080/09064710701312041
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all of these factors, the weather conditions such as
temperature, photoperiod, and irradiance are more
important on growth and development since they
can not be controlled by growers. Hence, many
studies were conducted to understand the growth,
yield, and quality responses of groundnut crop to
the uncontrolled environmental variables such as
soil and air temperature (Leong and Ong, 1983;
Ketring, 1984; Wheeler et al., 1997; Prasad et al.,
2000; Awal & Ikeda, 2002; Craufurd et al., 2002;
Awal & Ikeda 2003) and photoperiod and irradiance
(Witzenberger et al., 1988; Bagnall & King, 1991a,
1991b; Nigam et al., 1994). However, previous
studies also showed that the interaction of these
variables affects the phenological development, dry
matter production, and partitioning of groundnut.
This means that the understanding of the response
of the groundnut to each of these environmental
variables under controlled conditions is not enough
to understand the crop’s response to a certain
environment under field conditions.
The growth analysis of a crop in a certain
environment is important to understand how the
crop growth and development occur and how
production practices should be manipulated. The
quantitative relationships for the allocation of dry
matter among the leaves, stems, roots, and storage
organs are mostly empirical but a rough knowledge
of these relationships is also crucial in the under-
standing of the physiological behaviour of a crop (Tei
et al., 1996). The value of the agricultural experi-
ments could be greatly enhanced if the data related
to the growth and the partitioning of the growth
was available. It would allow better interpretation of
the results within the context of processes and
exploitation of the resource (Royo & Blanco,
1999). Furthermore, the knowledge on the physio-
logical components of yield in a certain environment
could be useful in breeding programs to improve the
yield (Ntare & Williams, 1998). Thus, many studies
for the growth analysis have been conducted to
understand the growth and development processes
in various crops under different environments (Tei
et al., 1996; Royo & Blanco, 1999; Yusuf et al.,
1999; Scholberg et al., 2000). Although some
studies have been reported on the growth and the
development of field-grown groundnut under differ-
ent environmental conditions (Duncan et al., 1978;
Dryer 1982; Bell et al., 1991a, 1991b, 1991c), there
is not such a study conducted under Mediterranean
conditions. The objective of this study was to assess
the growth analysis of groundnut genotypes with
growing degree days based on phenological devel-
opment, dry matter production, and partitioning in a
Mediterranean environment.
Materials and methods
Field experiments were conducted at the Experi-
mental Farm of the Faculty of Agriculture of
Mustafa Kemal University (368 39? N, 368 40? E;
83 m elevation) in the province of Hatay located in
the Eastern Mediterranean region of Turkey in 2001
and 2002. The soil of the experimental site which
was developed from alluvial deposits of river terraces
is typical for the Eastern Mediterranean region of
Turkey and is classified as Vertisol (FAO/UNESCO,
1974) having relatively high clay content with the
predominant clay minerals smectite and kaolinite.
The soil of the experimental plots (0�40 cm depth)
was clayey in texture (38.3% sand, 20.4% silt,
41.2% clay) with low organic matter content
(0.60%) and was slightly alkaline (pH 7.4) in
reaction. The available total nitrogen, available
phosphorus and potassium contents were 0.083%,
122.4 kg/ha, and 690 kg/ha, respectively.
The province of Hatay has typical Mediterranean
climate with hot and dry summers and mild and
rainy winters. The daily climatic data were obtained
from the agro-meteorological station located in a
state farm about 1 km far from the experimental site.
The mean values of the climatic data are presented
in Table 1.
Table 1. Monthly climatical data of experimental area during growing season of 2001 and 2002.
2001 2002
Temperatures (8C) Temperatures (8C)
Months Min. Max. Mean
Rainfall
(mm)
Radiation
(MJm�2day�1) Min. Max. Mean
Rainfall
(mm)
Radiation
(MJm�2day�1)
May 13.3 28.4 20.9 175.6 16.00 13.9 28.3 20.8 13.5 18.26
June 19.2 37.2 26.6 0.0 18.94 19.2 33.4 26.2 2.8 19.51
July 22.9 34.6 28.3 0.0 18.64 22.7 35.5 28.8 0.0 18.50
August 23.7 34.7 28.5 0.0 16.89 22.7 34.0 27.8 0.0 17.06
September 19.9 33.1 25.4 3.3 14.51 18.8 32.7 25.0 13.2 14.00
October 13.5 28.5 20.5 62.1 10.70 13.6 29.6 21.1 14.1 9.82
106 S. Caliskan et al.
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Eight Virginia type groundnut genotypes selected
for their good agronomic performances in the area
(Arioglu et al., 2000) were used in the study: PI
269084, PI 355276, 75/1073, NC-9, Edirne,
Osmaniye 2005, Com, and NC-7. The genotypes
Com and NC-7 are widely grown standard cultivars
in the Mediterranean region of Turkey. The geno-
type Edirne is a local variety and Osmaniye 2005 is a
newly released variety which was developed from
the NC-7�75/1073 cross. The genotypes of PI
269084, PI 355276, 75/1073, and NC-9 are plant
introductions with Zambia, Mexico, Israel, and USA
origin, respectively. The genotypes were grown in a
randomized complete block design with three repli-
cates. The seeds were sown by hand on 20 May in
2001 and 1 May in 2002 in six-row plots. The length
of each plot was 8.0 m. The spacings between the
rows and between plants in each row were 0.7 m and
0.2 m, respectively.
In both years, the groundnut was grown under
irrigated conditions with other cultural inputs ap-
plied consistently with local agronomic practices.
The pre-sowing herbicide, trifluralin, was applied to
the soil with a rate of 2000 mL/ha and the plots were
maintained weed-free by hand-weeding during the
growing period. Plots were fertilized with 60 kg N,
P2O5, K2O per ha before planting and an additional
nitrogen dose of 100 kg per ha was side-dressed at
the pegging stage. Overhead sprinkler irrigation was
applied with approximately two week intervals start-
ing with flowering stage.
The main phenological development stages
(Boote, 1982) such as the appearance of first fully
opened flowers (R1), pegs (R2), pods (R3), and
physiological maturity (R8) were recorded with daily
observations. Then, time from sowing to each
developmental stage was expressed as calendar
days and cumulative growing degree days (GDD)
for each genotype. GDD for each day was calculated
from the mean of the minimum and maximum
temperatures minus base temperature of 108C(Leong & Ong, 1983; Bell et al., 1991c; Craufurd
et al., 2000; Awal and Ikeda, 2002; Awal and Ikeda,
2003). If the mean of the minimum and maximum
temperatures was lower than the base temperature,
the GDD was assumed to be 0.
Six plants per plot were harvested nearly 15 day
intervals starting from 15 days after emergence of all
genotypes giving a total of 10 harvests for growth
analysis. Cumulative GDD from sowing to each
sampling date was also calculated. Harvested plants
were separated into leaves including petiole, stems,
pegs and pods, and dried in a forced air oven at 708Cto a constant weight at least 48 h and dry weights of
all samples were determined. Pod dry weights were
adjusted by multiplying by a factor of 1.65 to allow
for the energy content of oil in the seeds (Duncan
et al., 1978). Harvest index (HI) was calculated
using the pod and the total dry weight at each
sampling. Leaf area was estimated by measuring
green leaf area of a sub-sample with a leaf area meter
(Model MK2, Eijkelkamp Inc., The Netherlands).
Then, leaf area index (LAI) was calculated.
Functional approach for the growth analysis of the
groundnut data obtained from different genotypes
and years was used as suggested by Hunt (1982).
Crop growth is expressed as a function of the
growing degree days rather than calendar time.
Nonlinear regression was used to fit the data of
each genotypes and year for the growth traits of leaf
area index (LAI), total dry weight (TDW), leaf-stem
dry weight (LSDW), peg dry weight (PegDW), and
pod dry weight (PDW) to the asymmetric logistic
peak curve:
y�a=1�eb�cx (1)
where y is the present size of growth trait, x is the
time expressed as growing degree days, a is limiting
growth value which is the horizontal asymptote of
the logistic growth curve, b is the value such that half
life is �log b/c, and c is the rate of decline in relative
growth rate.
First, the data were fit to linear regression using
Eq. [2] by using PROC REG procedure of SAS
statistics software to determine the initial values of
estimated parameters of b and c for the optimization
of PROC NLIN procedure.
ln [(a=y)�1]�b0�b1x (2)
where b0 (intercept) is the initial value of b and b1
(slope) is the initial value of c. Then, the logistic
curves were fit to data using PROC NLIN proce-
dure.
Results and discussion
Wheather and crop phenology
The climatical data during the growing period was
somewhat similar between years and reflected the
long-term average and could be considered as a
typical Mediterranean climate (Table 1). As an
exception, abundant rainfall occurred in early May
in 2001 caused delaying of sowing until late May.
However, no or very little rainfall occurred during
subsequent four months in both years; therefore,
water requirements of the crops were supplied by
using overhead sprinkler irrigation. The heavy rain-
fall after September is not desired by groundnut
growers in the region since it results in delays in the
harvest as well as difficulties for harvest and drying
operations.
Growth and development of Virginia type groundnut cultivars under Mediterranean conditions 107
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The mean temperature was around 208C in May
and increased to around 268C in June and to around
288C in July and August in both years. Then, it
dropped gradually to around 258C and 218C in
September and October, respectively in both years.
The maximum temperature was around 34�358Cduring summer months with an exception of 37.28Crecorded in June of 2001. The reproductive phases
of groundnut genotypes started from mid or late
June in both years. The optimum mean air tempera-
ture range for vegetative growth in groundnut is
between 25 and 308C while the optimum tempera-
ture for the reproductive growth may be similar or
somewhat cooler, i.e. between 22�258C. High tem-
peratures above 358C during the reproductive
phases reduce dry matter accumulation, flower
production, proportion of pegs forming pods, in-
dividual seed mass, and consequently pod yield
(Leong and Ong 1983; Ketring 1984; Wheeler
et al. 1997; Prasad et al. 2000 and 2001).
Time from sowing to each phenological develop-
ment stage was significantly different among geno-
types in both years (Table 2). The genotypes reached
to flowering stage (R1) between 39 days (6158Cd)
and 43 days (6878Cd) in 2001 and between 46 days
(5848Cd) and 49 days (6298Cd) in 2002. The
genotypes of Osmaniye 2005, Com, and NC-7
flowered earlier than other genotypes in both years.
Genotypes reached to R1 with lower cumulative
GDD despite they need longer calendar days in 2002
compared to 2001. The groundnut genotypes were
sown 20 days earlier in 2002; therefore, their early
growth period coincided with relatively cooler per-
iod. This resulted in more calendar days to reach
R1, but less cumulative GDD. Ishag (2000) reported
that time from sowing to first flowering was
significantly affected by growing seasons and geno-
types and those warmer seasons resulted in earlier
flowering.
Late flowering of genotypes was also reflected to
the duration of subsequent growth stages in 2002
and the genotypes reached each stage of longer
duration in calendar day (Table 2). Despite the
periods of R1-R2 and R2-R3 occurred during late
June and July with similar monthly mean tempera-
tures in both years, longer R1-R2 and R2-R3 periods
in 2002 clearly indicated that temperature is not
solely enough to explain phenological development
of groundnut under the field conditions. The com-
bined effects of several environmental factors such as
temperature, photoperiod, water availability, irradia-
tion, soil conditions, pest and diseases are determin-
ing factors. Ishag (2000) also reported significant
variations in respect to duration of different pheno-
logical stages within growing seasons in certain
environments.The thermal times (GDD) for the
entire growth period were not different between
two years (Table 2). Time from sowing to physiolo-
gical maturity (R8) ranged from 25138Cd to
25888Cd in 2001 and from 25148Cd to 25738Cd
Table 2. Time from sowing to main phenological development stages of groundnut genotypes as calendar days and cumulative growing
degree days (8Cd).
R1 R2 R3 R8
Genotypes Days 8Cd Days 8Cd Days 8Cd Days 8Cd
2001
PI 269084 43 692 55 894 72 1225 153 2588
PI 355276 42 666 53 864 70 1199 152 2580
75/1073 41 651 51 832 69 1166 152 2580
NC-9 43 687 53 858 71 1205 151 2569
Edirne 43 681 54 876 72 1225 153 2588
Osmaniye 40 632 50 806 67 1140 147 2513
Com 39 615 50 806 66 1115 152 2580
NC-7 39 615 50 806 66 1109 152 2580
Mean 41 655 52 843 69 1173 151 2572
LSD (0.05) 0,9 16 0,8 13 0,9 18 0,0 0
2002
PI 269084 49 629 64 895 86 1299 158 2544
PI 355276 48 623 62 852 84 1261 159 2548
75/1073 49 635 63 872 85 1286 160 2560
NC-9 49 635 64 889 86 1299 159 2548
Edirne 49 635 64 895 86 1299 161 2573
Osmaniye 46 584 62 846 81 1200 156 2514
Com 47 590 62 852 82 1230 160 2556
NC-7 47 590 62 852 82 1230 161 2573
Mean 48 615 63 869 84 1263 159 2552
LSD (0.05) 0,9 16 0,6 11 0,8 16 0,5 6
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in 2002 while total calendar days varied between 147
and 153 in 2001 and between 156 and 161 in 2002.
The genotype Osmaniye 2005 was recorded as the
earliest genotype in both years. Apparently, the long
duration (as calendar days) of crop growth in the
cultivars in 2002 was mainly due to the longer period
from sowing to first pegging which coincided with
a period of relatively cool temperatures (Table 1).
Similar results were also reported by Banterng et al.
(2003) in Thailand and Ishag (2000) in Sudan with
Virginia type groundnut cultivars.
Leaf area development
The relationship describing the changes in LAI
against thermal time is presented in Figure 1. Also,
the parameters estimating the LAI growth equation
are presented in Table 3. Although some differences
were found among the genotypes for LAI values at
different sampling dates, the curves describing the
changes in LAI with time were similar. The curves
differed between the two years due to the higher
limiting value of LAI which was reached in 2001.
The increase in LAI was relatively slow during the
period of early growth until around 11008Cd. After
this period, the expansion of leaf area increased
with a high rate until around 20008Cd and there-
after, the growth rate slowed down again in both
years. However, LAI values of groundnut genotypes
rarely went down towards maturity in contrast with
most of the annual crops. Less enlargement of the
leaf area during the initial growth could be attributed
to the higher allocation rate of the dry matter to the
roots during this period (Wheeler et al., 1997). The
rapid period of leaf area development coincided with
the onset and initial growth of the pods which also
needed high dry matter accumulation. This was
probably compensated with increasing photosyn-
thetic efficiency as a result of establishing larger
root system and leaf area.
Kiniry et al. (2005) summarized some findings
from eleven previous studies related to LAI. They
noticed that the values of the LAI ranged from 3 to
greater than 8 depending on the experimental
conditions. They also reported that the maximum
LAI values ranged from 5 to 7 at three sites in Texas.
They concluded that LAI values of 5�6 appeared to
be appropriate for groundnut in many regions. In
our study, the LAI values ranged from 7.6 to 9.2 and
from 4.2 to 7.3 depending on cultivars in 2001 and
2002, respectively. Apparently, our findings on the
LAI approximated to the upper limits reported in the
literatures mentioned above. The mean tempera-
tures during most of growing cycle were close to
the optimum for vegetative growth of groundnut in
both years (Table 1). This resulted in abundant and
continuous vegetative growth in groundnut cultivars.
Dry matter production and partitioning
The relationships describing the changes in LSDW,
PegDW, PDW, and TDW as a function of thermal
time fitted to the mean data of each year. Pattern of
the changes in dry weights of different plant parts
from sowing to maturity averaged over genotypes in
2001 and in 2002 are presented in Figure 2 and
Figure 3, respectively. The estimates of the para-
meters of the growth equation for above traits were
presented in Table 3. The curves clearly demon-
strated that the dry matter accumulation to each part
of plants continued until maturity although accu-
mulation rate differed depending on plant age.
The pattern of LSDW accumulation with thermal
time (Figures 2 and 3) was similar to that of LAI
(Figure 1). Likewise, Ma et al. (1992) demonstrated
strong relationship between leaf dry weight and LA
and suggested that LA could be estimated with high
accuracy using leaf dry weight in groundnut. Leaf
production somewhat continued until maturity while
senescence was slow due to suitable temperature
regimes during the growth period in our experi-
ments. Thus, LSDW values did not decline even in
the last period of the growth cycle.
Peg formation started between 800�9008Cd after
sowing depending on genotypes in both years as
discussed earlier. The initial dry weight of peg was
lower in 2002; however, the growth rate increased
more later in this year and curves of each year nearly
overlap after 20008Cd. The increase in PegDW of
the groundnut genotypes continued until maturity in
both years while peg growth rate slowed down after
around 23008Cd (Figures 2 and 3). Actually, onset
of new pegs by genotypes (data not presented)
continued until around 23008Cd and generally
stopped after this time although dry matter accu-
mulation to the formed pegs continued until harvest.
0
1
2
3
4
5
6
7
8
9
10
Growing Degree Days
edniaera
faeL
x 2001
2002
300025002000150010005000
Figure 1. Pattern of the changes in Leaf Area Index, LAI from
sowing to maturity averaged over genotypes. Lines represent the
asymmetric logistic peak curves (Eq. [2]) fit to the observed values
in years.
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Dry matter accumulation of pods also continued
until final harvest in both years (Figures 2 and 3).
The genotype Osmaniye 2005 was the best perform-
ing genotype in respect to PDW at final harvest in
both years. The pod growth rates were slow within
around the first 3008Cd after pod initiation in both
years. The genotypes started with earlier pod growth
and their pod growth continued until the final
harvest with a high rate in 2001 while the pod
growth rate slowed down during the last phase in
2002. These results clearly indicated the importance
of initial crop growth on the subsequent reproduc-
tive growth of the groundnut genotypes.
Bell et al. (1991b) reported that day length was
the primary factor affecting the reproductive devel-
opment and that temperature had less important on
it under field conditions in subtropical Australia. In
contrast, Bagnall & King (1991a, 1991b) reported
that temperature affected the reproductive develop-
ment primarily via its effects on DM accumulation
under controlled conditions. However, both authors
as well as Witzenberger et al. (1988) indicated the
Table 3. Estimates of the parameters of the growth curves for evaluated traits.
2001 2002
Traits Genotypes a b c a b c
TDW
PI 269084 2310.2 6.27 0.0035 2583.2 5.73 0.0027
PI 355276 2911.8 5.51 0.0029 2611.9 5.67 0.0028
75/1073 2617.0 6.91 0.0037 2094.7 6.53 0.0034
NC-9 2683.5 4.64 0.0024 1659.3 8.53 0.0046
Edirne 5255.4 5.25 0.0020 2054.2 5.57 0.0030
Osmaniye 5331.2 5.39 0.0022 2857.6 6.64 0.0031
Com 2604.6 4.69 0.0026 1872.4 5.49 0.0030
NC-7 2930.1 4.42 0.0024 1751.6 5.44 0.0030
overall 3037.5 5.07 0.0025 2162.7 5.91 0.0030
LSDW
PI 269084 1088.9 6.64 0.0045 913.9 5.74 0.0036
PI 355276 1211.6 7.19 0.0048 880.0 4.99 0.0035
75/1073 1190.3 6.27 0.0039 776.6 6.06 0.0041
NC-9 1094.7 6.42 0.0046 881.7 6.18 0.0038
Edirne 1202.7 4.66 0.0028 835.2 7.11 0.0052
Osmaniye 1359.1 4.39 0.0024 1050.4 4.35 0.0022
Com 1156.1 5.01 0.0037 665.5 7.34 0.0056
NC-7 956.1 5.92 0.0048 545.7 5.66 0.0044
overall 1123.0 5.51 0.0038 785.7 5.52 0.0037
GDW
PI 269084 38.0 11.65 0.0072 61.7 8.44 0.0044
PI 355276 51.0 15.75 0.0097 82.7 7.19 0.0037
75/1073 62.9 8.65 0.0051 81.0 9.33 0.0050
NC-9 62.7 8.93 0.0056 62.2 15.78 0.0087
Edirne 149.7 4.92 0.0019 65.5 12.07 0.0069
Osmaniye 163.7 4.89 0.0018 69.5 7.15 0.0039
Com 65.4 7.15 0.0046 58.6 8.34 0.0049
NC-7 119.9 4.34 0.0022 62.1 7.46 0.0044
overall 66.9 6.08 0.0035 67.3 8.57 0.0047
PDW
PI 269084 1066.7 11.82 0.0059 1240.3 12.95 0.0057
PI 355276 2224.9 7.64 0.0031 1277.2 13.91 0.0065
75/1073 1261.6 11.75 0.0058 1079.1 15.20 0.0073
NC-9 1989.0 8.00 0.0031 695.9 15.30 0.0076
Edirne 2197.5 10.02 0.0041 1219.7 9.77 0.0044
Osmaniye 3720.7 6.87 0.0026 1533.9 10.34 0.0049
Com 1311.5 8.07 0.0038 1181.2 8.87 0.0041
NC-7 2603.3 6.16 0.0025 1552.8 6.42 0.0028
overall 1878.3 7.60 0.0033 1209.6 10.17 0.0047
LAI
PI 269084 8.13 7.55 0.0053 6.45 5.33 0.0035
PI 355276 8.58 6.79 0.0044 6.71 5.15 0.0037
75/1073 8.30 6.87 0.0044 5.41 6.34 0.0045
NC-9 7.21 9.63 0.0075 5.27 5.55 0.0039
Edirne 8.55 4.51 0.0028 6.49 7.04 0.0049
Osmaniye 7.55 5.27 0.0033 6.64 4.37 0.0023
Com 8.07 5.23 0.0040 4.37 8.15 0.0065
NC-7 7.54 5.83 0.0048 3.65 5.75 0.0048
overall 7.85 5.92 0.0042 5.43 5.62 0.0040
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importance of interactions between temperature and
photoperiod on the reproductive development of
groundnut and reported that genetic variability
existed for responses to both factors. Shorter day
length with combination of optimum temperature
has stimulative effects on the reproductive growth of
groundnut. Dryer et al. (1981) reported that fruit
formation continued for a longer period under the
cooler soil temperature (238C) conditions. In the
Mediterranean type environments, day length gets
shorter while mean temperature gets lower (�258C)
toward autumn. These conditions results in longer
reproductive growth in groundnut. However, this
growth tendency can cause some problems in
harvest such as delaying of harvest, disparity in pod
maturity, and increasing proportion of unmarketable
pods. Although this response is greatly influenced by
genetic traits reducing new flower production and
shortening the blooming period, appropriate man-
agement systems could be beneficial to overcome
this problem (Cattan & Fleury, 1998).
The total dry matter accumulation also continued
until final harvest as a function of continuous
accumulation to each part as discussed above
(Figures 2 and 3). The delay in phenological
development during the early growth stages due to
cooler conditions in 2002 was reflected in a slower
rate of biomass accumulation in successive stages.
Similar responses of groundnut genotypes were
reported by Banterng et al. (2003) in Thailand.
The slow germination and growth problems due to
cooler conditions at the early sowing dates could be
overcome by the application of mulching. Early
studies showed that polyethylene mulching provided
favorable soil physical environment for early sown
groundnut growth and development to have satisfied
yield (Choi and Chung, 1997; Ghosh et al. 2006;
Khan, 2002; De et al., 2005 and Subrahmaniyan
et al., 2006).
The apparent fractions of current dry matter
above the ground parts of the plant over thermal
time are shown in Figure 4. Although reproductive
development started with the onset of flowering at
around 6008Cd, vegetative parts was still major sink
until around 1100�12008Cd when pod initiation
occurred in groundnut genotypes in both years.
Dry matter of flower and pegs had very small
fraction (2�4%) in total dry matter accumulated
not only in earlier stages but also in later stages. After
the onset of the pods, the fraction of dry matter
accumulated by vegetative plant parts decreased
appreciably. Dry matter accumulation by stems
and leaves accounted for up to 95% of the total
above ground biomass during pod initiation stage
but decreased gradually to around 40% by the end of
the growing season in both years.
Dry matter accumulation by pods (Harvest Index,
HI) increased linearly with thermal time until the
2001
0
250
500
750
1000
1250
1500
1750
2000
2250
2500
2750
0 250 500 750 1000 1250 1500 1750 2000 2250 2500 2750Growing Degree Days
mg(thgie
wyr
D2-) Leaf+stem DW
Peg DWPod DWTotal DW
Figure 2. Pattern of the changes in dry weights of different plant
parts from sowing to maturity averaged over genotypes in 2001.
Lines represent the asymmetric logistic peak curves (Eq. [2]) fit to
the observed values in 2001. DW�dry weight.
2002
0
200
400
600
800
1000
1200
1400
1600
1800
2000
0 250 500 750 1000 1250 1500 1750 2000 2250 2500 2750Growing Degree Days
mg(thgie
wyr
D2 -) Leaf+stem DW
Peg DWPod DWTotal DW
Figure 3. Pattern of the changes in dry weights of different plant
parts from sowing to maturity averaged over genotypes in 2002.
Lines represent the asymmetric logistic peak curves (Eq. [2]) fit to
the observed values in 2002. DW�dry weight.
2001
0,00,10,20,30,40,50,60,70,80,91,0
0 500 1000 1500 2000 2500 3000Growing Degree Days
0 500 1000 1500 2000 2500 3000Growing Degree Days
Leaf+stem DWPod DWPeg DW
2002
0,00,10,20,30,40,50,60,70,80,91,0
Dry
mat
ter
part
ioni
ng c
oeff
icie
nt
Dry
mat
ter
patio
ning
coe
ffic
ient
Leaf+stem DWPod DWPeg DW
Figure 4. Pattern of the changes in dry matter partitioning
coefficient from sowing to maturity averaged over genotypes.
Lines represent the asymmetric logistic peak curves (Eq. [2]) fit to
the observed values in years. DW�dry weight.
Growth and development of Virginia type groundnut cultivars under Mediterranean conditions 111
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end of the growing season in both years. Final HI of
the genotypes ranged from 49.3 to 60.7 and from
29.5 to 67.9 with an average of 53.8 and 54.8 in
2001 and 2002, respectively. Wheeler et al. (1997)
and Craufurd et al. (2002) reported that the pod HI
of the groundnut genotypes increased linearly with
time until maturity irrespective of the growing
temperature but the higher temperatures (�308C)
caused slower HI increasing rate and the lower HI
values at final harvest. Kiniry et al. (2005) reported
that HI values in groundnut varied between 38%
and 62% with an average of 45% in 14 previous
studies from all around the world depending on
environmental conditions and genotypes. They also
reported that HI values ranged from 30% to 58%
depending on the management practices, genotypes,
and locations and suggested that research on the
processes affecting the yield components should
continue to be vigorously pursued to quantify the
differences in HI. Apparently, the mean HI for the
data sets in the present study was relatively high
comparing to several data sets in the literature. This
could be attributed to the stimulating effects of
cooler temperature and shorter day length during
the reproductive growth in the Mediterranean type
environments. However, unmarketable pods, which
were produced toward the maturity, also contributed
to these higher HI values.
Conclusion
In order to obtain high yield, groundnut cultivars
must have high initial growth rate, less reproductive
organs, and shorter growing period under the
Mediterranean conditions. In addition to cultivar
selection, appropriate cultural practices should be
applied to increase initial growth rate and reduce
number of flowers, pegs, or pods per plant.
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