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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

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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: scaliskan@mku.edu.tr

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

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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.

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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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