ORIGINAL ARTICLE

Photosynthetic characteristics of groundnut (Arachis hypogaea L.)

under water deficit stress

K. A. Kalariya • A. L. Singh • K. Chakraborty •

P. V. Zala • C. B. Patel

Received: 25 January 2012 / Accepted: 13 March 2013

� Indian Society for Plant Physiology 2013

Abstract In a field experiment during summer season,

the chlorophyll fluorescence and net photosynthetic rate

were studied in six Spanish groundnut cultivars under

water deficit (WD) condition between beginning bloom to

beginning seed (WD I) and beginning seed to beginning

maturity (WD II), by withholding irrigation 31–62 DAS

and 62–87 DAS, respectively. The severity of drought was

so that the soil moisture content of the stressed field

depleted to 9.4 and 6.2 % at 0–15 cm soil depth in WD I

and WD II at 62 and 87 DAS, respectively, as against 17

and 19 % in control. The mean RWC of leaves decreased

from 92 in control to 88 in WDI and from 91 in control to

84 in WD II with the least decrease in ICGS 44 under WD I

and in TAG 24 under WD II. The WD condition signifi-

cantly increased non-photochemical quenching but

decreased the maximum quantum yield of PS II (Fv/Fm)

from 0.81 in control to 0.77 under WD I condition which

was again resumed to 0.80 after 48 h of withdrawal of

stress. The rate of photosynthesis which was 29 and

36 lmol m-2 s-1 in well irrigated plots decreased to 26

and 28 lmol m-2 s-1 with a deduction of 11 and 30 %

under WD I and WD II, respectively. Cultivar TAG 24

showed better stress recovering capacity with high photo-

synthesis under both control as well as WD condition

whereas, data on chlorophyll fluorescence parameters

showed that cultivar ICGS 44 was least affected to damage

via photoinhibitory action.

Keywords Chlorophyll fluorescence � Groundnut �

Leaf temperature � Photosynthesis � Water deficit

stress

Introduction

Groundnut (Arachis hypogaea L.) is an important food

legume and oilseed crop of the world grown on about 24

million ha of land in about 120 countries mostly in tropics

and subtropics of arid and semi-arid regions where the

availability of water is a major constraint on yield and crop

frequently suffers drought of various spells and intensities.

As a result, the groundnut productivity is less than

1,000 kg ha-1 in more than 50 % of groundnut growing

countries. In India, the groundnut is cultivated on about 6

million ha with productivity of about 1,400 kg ha-1. Of all

the abiotic stresses, drought is most damaging one in

groundnut and it has a broad physiological spectrum

affecting many metabolic processes and it is difficult to

assess the contribution of individual processes to the final

damage done to the plants. Drought affects the groundnut

plants at all the stages, with varied sensitivity and crop

cultivars that show differential responses (Singh 2004,

2011; Nautiyal et al. 2012).

The photosynthetic rate (PN) of leaves decreases as

relative water content (RWC) and water potential (W)

decreases and limitation of net photosynthetic rate in low

moisture stressed plant is mainly through stomatal closure

(Cornic and Massacci 1996; Singh 2004) and/or by meta-

bolic impairment. The relative magnitude of stomatal and

non-stomatal factors limiting photosynthesis depends on

the severity of drought. Photosynthesis, the most important

process influencing crop production, is inhibited by

drought stress and high PN is one of the most important

K. A. Kalariya (&) � A. L. Singh � K. Chakraborty �

P. V. Zala � C. B. Patel

Directorate of Groundnut Research, PB 5, Junagadh 362 001,

Gujarat, India

e-mail: kuldeep_ka@yahoo.co.in

123

Ind J Plant Physiol.

DOI 10.1007/s40502-013-0027-x

breeding strategies for crop improvement (Richards 2000).

Chlorophyll fluorescence is a very sensitive probe of

physiological status of leaves and plant performance in a

wide range of situations and successfully applied in crop

improvement programs by carefully selecting and analys-

ing fluorescence parameters (Daniele et al. 2006). Though

the initial phase of drought stress in plants due to stomatal

limitation of photosynthesis is not perceived by the fluo-

rescence parameters that can only monitor the efficiency of

primary photochemical processes of photosynthesis. Araus

et al. (1998) proposed chlorophyll fluorescence as a

selection criteria for improvement of grain yield of durum

wheat in Mediterranean conditions, where frequent drought

accompanied by high temperatures is common. Since then

chlorophyll fluorescence continues in mainstay in studies

of photosynthetic regulation and plant responses to the

environment due to its sensitivity, convenience, and non-

intrusive quality. In recent years, the chlorophyll fluores-

cence technique has become ubiquitous in plant eco-

physiological studies.

Therefore, a study was aimed at unravelling the changes

in chlorophyll fluorescence and net photosynthetic rate

under water deficit condition at different phenophases in

groundnut cultivars of different maturity period.

Materials and methods

A field experiment was conducted during summer 2011

(January–June) using six Spanish groundnut varieties at the

research farm of Directorate of Groundnut Research,

Junagadh, Gujarat (lat 21.0 310N, Long 70.0360E), India in

Vertic Ustochrept (pH 7.5) soil. The experiment was laid in

a split plot design taking three irrigation treatments viz.

control, water deficit (WDI) and WD II in main plot, and

six varieties in sub plots. The net plot size was 4 9 3 m

with 9 rows per plot at 45 cm row to row and 10 cm plant

to plant spacing. To synchronize phenophases the

groundnut cultivars SG 99, ICGS 44 and ICGV 86031 of

130 days duration were sown during last week of January

and cultivars TAG 24, AK 159 and DRG 1 of 120 days

duration were sown during first week of February.

The water deficit condition was created by with-holding

irrigation from beginning bloom to beginning of seed i.e.

31–61 DAS (WD I) and from beginning seed to beginning

maturity, i.e. 62–87 DAS (WD II) while adequate moisture

was maintained in the control. The RWC, chlorophyll

fluorescence, photosynthesis and leaf temperature were

recorded from the third leaf at 62 and 87 DAS in WD I,

WD II, respectively and in control and 48 h after relief of

water stress. Maximum efficiency of PSII (Fv/Fm) of the

dark adapted leaves were recorded after 30 min dark

adaptation by leaf clips and actual quantum yield of PSII

and photosynthesis (at 1,650 PPFD artificial light) were

recorded between 08:00 and 10:00 h by LI-COR 6400,

Portable Photosynthesis system (LI-COR Inc. Lincon, NE,

USA) with modulated fluorescence measurement. This

system also simultaneously measures the net rate of pho-

tosynthesis (PN), stomatal conductance (gs) leaf tempera-

ture (Tleaf) and transpiration rate (E) etc. The RWC was

calculated using formula: RWC = 100*((FW - DW)/

(TW - DW)), Where, FW is fresh weight, TW is turgid

weight and DW is dry weight.

Results and discussion

Soil water plant relations

The soil moisture content of the field at 0–15 cm soil depth

was 17.0 and 19.5 % in control plots which, by with-holding

irrigation decreased to 9.4 and 6.2 % under water deficit

condition at the end of WD I and WD II, respectively.

However, the soil moisture content in 15–30 cm soil depth

was 18.0 and 19.0 % in control plots which decreased to 10.1

and 9.4 % in WD I and WD II, respectively. The soil mois-

ture content depleted by 45 and 44 % in WD I and 68 and

51 % in WD II as compared with control at 0–15 cm and

15–30 cm depth, respectively. The RWC of groundnut

leaves which was 93 and 91 % in control decreased to 88 and

84 % by imposition of water deficit at the end of WD I and

WD II, respectively with the highest value in SG 99

(Table 1). Stomatal conductance (gs) and transpiration rate

also decreased under water deficit condition (Table 1). The

gs in control at 62 and 87 DAS was 0.40 and 0.67 m s-1

which decreased to 0.32 and 0.30 m s-1 in WD I andWD II,

respectively which after relieving of stress by irrigation

further increased gs to 0.35 and 0.40 mol m-2 s-1, respec-

tively. Highest stomatal conductance was found in cultivar

ICGS 44 under WD I and in DRG 1 under WD II. The

varieties with minimum transpiration were SG 99, AK 159

and DRG 1 because of corresponding reduction in stomatal

conductance whereas, ICGV 86031 and TAG 24 showed

higher transpiration rates.

The RWC, leaf water potential, stomatal resistance, rate

of transpiration, leaf temperature and canopy temperature

are important parameters that influence water relations in

groundnut (Nautiyal et al. 1995, 2002, 2012; Singh 2004).

The RWC of leaves is reported to be higher during the

initial stages of development and declines as the dry matter

accumulates and leaf matures (Jain et al. 1997). In non-

stressed groundnut plants the RWC in leaves ranges from

85 to 90 %, while under drought stress it goes down

heavily (Babu and Rao 1983). Daniele et al. (2006) found

that in groundnut, the genotypic discrimination of RWC

trait depends on the water regime and genetic background

Ind J Plant Physiol.

123

and this trait has no value as selection criteria but serves to

characterize finely the water status of plants. Water uptake

was reported to be maximised by increasing root depth.

Allen et al. (1976) measured soil water extraction during

water stress and found that groundnut roots effectively

extracted soil water to a depth of at least 180 cm in fine

sand soil as roots in lower depths continue to grow deeper

even though vegetative growth appears to stop.

Simmonds and Ong (1987) found that the cultivar Robut

33–1 extracted water more rapidly from deeper layers

when grown at high vapour pressure deficits than grown in

more humid air. Devries et al. (1989) reported that cultivar

Florunner had higher root length density in deeper layers

(60–150 cm) during drought periods. All these traits con-

tribute to groundnut’s ability to avoid drought stress in this

study. Drought stressed plants transpire less than unstressed

plants, as a result decreased rate of transpiration was

observed in this study. The same pattern was also observed

for stomatal conductance. Subramaniam and Maheswari

(1990) reported that leaf water potential, transpiration rate

and photosynthetic rate decreased progressively with

increasing duration of water stress in groundnut.

Maximum efficiency of PS II (Fv/Fm) and quantum

yield of PS II (UPSII)

Maximum efficiency of PS II (Fv/Fm) decreased signifi-

cantly from 0.81 in control to 0.77 under WD condition and

again reached to 0.80 after 48 h of withdrawal of stress

showing a good stress recovery indication under WD I

(Table 2). The recovery was highest in cultivar ICGS 44

and AK 159. Under WD II also the Fv/Fm decreased from

Table 1 Leaf relative water content (RWC), stomatal conductance (gs), transpiration rate and leaf temperature (Tleaf) in groundnut varieties at

62 DAS in WD I and 87 DAS in WD II as affected by water deficit conditions during summer 2011

Varieties Relative leaf water content (%) gs (m s-1)

62 DAS 87 DAS 62 DAS 87 DAS

Control WDI Mean Control WDII Mean Control WDI SR Mean Control WDII SR Mean

AK 159 91.3 87.7 89.5 91.0 83.9 87.5 0.41 0.30 0.36 0.35 0.68 0.25 0.41 0.45

DRG 1 91.8 86.4 89.1 92.4 82.6 87.5 0.42 0.28 0.41 0.37 0.66 0.43 0.42 0.50

TAG 24 91.2 88.1 89.6 90.5 83.1 86.8 0.41 0.34 0.36 0.37 0.62 0.38 0.50 0.50

ICGS 44 90.9 90.3 90.6 92.0 83.9 87.9 0.42 0.38 0.34 0.38 0.62 0.22 0.40 0.41

ICGV 86031 91.7 85.8 88.7 91.2 84.1 87.6 0.39 0.31 0.32 0.34 0.70 0.24 0.34 0.43

SG 99 94.2 91.8 93.0 90.8 86.5 88.6 0.38 0.31 0.35 0.34 0.72 0.29 0.33 0.45

Mean 91.8 88.4 90.1 91.3 84.0 87.7 0.40 0.32 0.35 0.36 0.67 0.30 0.40 0.46

CD (p = 0.05)

Water deficit 1.7 2.09 0.02 0.09

Varieties 2.4 NS NS NS

Interactions NS NS 0.06 0.12

Varieties Transpiration rate (mmol m-2 s-1) Tleaf (�C)

62 DAS 87 DAS 62 DAS 87 DAS

Control WDI SR Mean Control WDII SR Mean Control WDI SR Mean Control WDII SR Mean

AK 159 10.6 5.6 10.1 8.8 9.9 5.4 7.9 7.7 32.1 34.7 31.9 32.9 33.0 35.5 34.0 34.2

DRG 1 10.6 5.8 11.2 9.2 10.5 7.4 7.4 8.4 32.4 37.9 32.8 34.4 32.7 38.7 33.6 35.0

TAG 24 10.1 7.5 9.7 9.1 11.0 7.2 8.1 8.8 31.2 37.0 32.3 33.5 31.9 37.8 34.3 34.6

ICGS 44 10.4 6.0 9.2 8.5 10.6 5.7 7.9 8.1 32.4 36.9 32.8 34.0 32.9 37.2 34.1 34.7

ICGV 86031 10.1 7.8 9.8 9.3 12.3 6.6 7.1 8.7 32.9 36.6 32.8 34.1 33.2 38.5 34.6 35.5

SG 99 10.5 5.2 10.1 8.6 9.7 6.1 7.8 7.8 32.1 36.4 32.2 33.6 32.3 37.2 33.8 34.4

Mean 10.4 6.3 10.0 8.9 10.9 6.4 7.7 8.3 32.2 36.6 32.5 33.7 32.7 37.5 34.1 34.73

CD (p = 0.05)

Water deficit 0.74 1.60 1.4 1.3

Varieties NS NS NS NS

Interactions 1.57 1.14 NS NS

SR represents 48 h after stress relief

Ind J Plant Physiol.

123

0.84 in control to 0.80 in WD condition which recovered

after 48 h of withdrawal of stress. Significant varietal dif-

ference for UPSII with the high value in TAG 24, ICGS 44

and SG 99 was observed under WD I. Also, there was

significant interaction of varieties and treatment for UPSII

where the reduction in these parameters was lower in TAG

24, ICGS 44 and SG 99 than other varieties under WD I.

This was also reflected in terms of higher rate of photo-

synthesis in these varieties under WD I. After 48 h of stress

recovery, increase in the proportion of absorbed energy

utilised for photochemistry (UPSII) with the highest value in

cultivar TAG 24 and SG 99 showed the better stress

recovering capacity. Groundnut has the ability to recover

from prolonged desiccation indicative of moisture stress

endurance (Babu and Rao 1983). Bogalea et al. (2011)

showed that the F0 has increased while variable fluores-

cence (Fv), maximum fluorescence (Fm) and optimum

quantum yield fluorescence (Fv/Fm) decreased under water

deficit condition in durum wheat. The increase in F0 is a

characteristic of PSII inactivation whereas a decline in Fv

under stress may indicate the increase in non-photochem-

ical quenching process at or close to reaction centres.

Similarly, the increase in F0 and the decrease in Fm under

water deficit with concomitant decrease in Fv/Fm, indicate

the occurrence of chronic photo inhibition due to photo

inactivation of PSII centres (Baker and Horton 1987).

Leaf temperature (Tleaf) and non-photochemical

quenching (NPQ)

There was increase in leaf temperature by 4.4 and 4.8 �C

due to WD I and WD II, respectively compared to control

but, there was no significant difference between Tleaf

among groundnut cultivars (Table 1). The correlation

(Table 3) between transpiration and Tleaf being r = -0.87

and -0.71 under WD I and WD II, respectively shows the

cooling effect of leaf under higher transpiration. The NPQ

also significantly increased under water deficit condition.

The correlation between Tleaf and NPQ was r = 0.80 under

WD I and r = 0.71 under WD II. Likewise increase in

UPSII with a corresponding decrease in NPQ have been

observed for stress recovered plant also indicates that after

stress recovery, energy was not utilised for heat dissipation,

but for photochemistry. The highest decrease in NPQ in

TAG 24 variety again confirms its better stress recovering

capacity. Most of the plants adapt themselves to water

stress by dissipating the excess excitation energy thermally

with the down regulation of PSII activity to protect pho-

tosynthetic apparatus from photo-damaging effect under

water stress often coinciding with high leaf temperature

(Bilger and Bjorkman 1990). Shahenshah and Isoda (2010)

reported that under green-house condition, Tleaf in

groundnut was increased due to water deficit stress. The

increase in volume of NPQ coupled with a negative effect

on rate of photosynthesis (r = -0.78 and -0.80 at WD I

and WD II, respectively) confirms that the incident energy

was diversified towards heat dissipation instead of photo-

chemistry under water deficit stress (Table 3). Excess light

is harmlessly dissipated in the antennae complexes of PSII

as heat is involved in a process of xanthophyll cycle, a

reversible conversion of the carotenoid violaxanthin (V) to

zeaxanthin (Z) via antherozanthin (A) as well as low thy-

lakoid pH. The xanthophyll cycle pigments Z and A are

formed from V during conditions of excess light and are

both thought to be involved in the photo-protective dissi-

pation process (Bjorkman and Demmig-Adams 1994).

At the leaf level, the dissipation of the excitation energy

through processes other than photosynthetic carbon

metabolism is an important defence mechanism under

water stress conditions and is accompanied by down-reg-

ulation of photochemistry in the long term carbon metab-

olism (Chaves et al. 2002). Also the energy dissipation in

closed stomata can occur via ATP and NADPH, which are

used for other metabolic processes, and are obviously

important mechanisms of tolerance and protection against

water stress and photo-oxidative damage. Daniele et al.

(2006) reported that fluorescence parameters showed

strong sensitivity to drought and good genotypic discrim-

ination in groundnut genotypes.

Net photosynthetic rate (PN)

The rate of photosynthesis significantly decreased due to

water deficit condition at both the stress periods (Table 2).

The mean PN was 29 and 36 lmol m-2 s-1 which due to

water stress decreased to 26 and 28 lmol m-2 s-1 at WD I

and WD II, respectively. However, the relief of stress

further increased PN to 28 and 29 lmol m-2 s-1, respec-

tively. The decrease in PN compared to control under WD I

and WD II by 11 and 30 %, respectively showed that the

growth period as defined by Boote (1982) between R5 and

R7 is more susceptible to water deficit condition for carbon

partitioning. The highest decrease was in cultivar DRG 1

under WD I, but interestingly the least decrease also in the

same cultivar under WD II shows stage-specificity in tol-

erance and susceptibility to water deficit condition. Canopy

photosynthesis is reduced by moisture stress mainly due to

reduction in stomatal conductance and leaf area, and as

moisture stress increases, stomata start closing as a

mechanism to reduce transpiration. As a consequence, the

entry of carbon dioxide is also reduced (Singh 2011). The

decrease in conductance of mesophyll cells due to moisture

stress results in low conductance of carbon dioxide and a

reduction in photosynthesis. During pod development

stage, both the vegetative and reproductive sinks

operate concurrently in groundnut because of relatively

Ind J Plant Physiol.

123

Table 2 Maximum efficiency yield of PS II (Fv/Fm), quantum yield of PS II (UPSII), non-photochemical quenching (NPQ) and photosynthetic rate

(PN) in groundnut varieties at 62 DAS in WD I and 87 DAS in WD II as affected by water deficit condition during summer 2011

Varieties Fv/Fm PS II (UPSII)

62 DAS 87 DAS 62 DAS 87 DAS

Control WDI SR Mean Control WDII SR Mean Control WDI SR Mean Control WDII SR Mean

AK 159 0.800 0.730 0.810 0.780 0.830 0.800 0.850 0.827 0.38 0.15 0.41 0.31 0.37 0.24 0.32 0.31

DRG 1 0.820 0.740 0.790 0.783 0.850 0.800 0.850 0.833 0.39 0.11 0.38 0.29 0.48 0.25 0.32 0.35

TAG 24 0.820 0.790 0.800 0.803 0.840 0.790 0.830 0.820 0.42 0.31 0.43 0.38 0.39 0.32 0.33 0.35

ICGS 44 0.810 0.790 0.810 0.803 0.830 0.800 0.840 0.823 0.40 0.30 0.39 0.36 0.37 0.31 0.33 0.33

ICGV

86031

0.820 0.760 0.800 0.793 0.840 0.800 0.830 0.823 0.43 0.13 0.34 0.30 0.44 0.32 0.34 0.37

SG 99 0.810 0.790 0.800 0.800 0.840 0.790 0.840 0.823 0.42 0.21 0.43 0.35 0.42 0.34 0.29 0.35

Mean 0.813 0.767 0.802 0.794 0.838 0.797 0.840 0.825 0.41 0.20 0.40 0.33 0.41 0.29 0.32 0.34

CD (p = 0.05)

Water

deficit

0.007 0.004 0.05 0.05

Varieties 0.014 0.006 0.04 NS

Interactions 0.025 0.011 0.07 NS

Varieties NPQ

62 DAS 87 DAS

Control WDI SR Mean Control WDII SR Mean

AK 159 1.66 2.04 1.95 1.88 1.23 1.91 1.55 1.56

DRG 1 1.39 2.25 1.63 1.76 1.03 1.74 1.5 1.42

TAG 24 1.66 1.99 1.67 1.77 1.23 1.53 1.19 1.32

ICGS 44 1.66 2.06 1.79 1.84 1.17 1.68 1.34 1.40

ICGV 86031 1.53 2.16 1.99 1.89 1.00 1.58 1.3 1.29

SG 99 1.44 2.14 1.62 1.73 1.14 1.67 1.43 1.41

Mean 1.56 2.11 1.78 1.81 1.13 1.69 1.39 1.40

CD (p = 0.05)

Water deficit 0.18 0.20

Varieties NS 0.10

Interactions NS 0.18

Varieties PN (lmol m-2 s-1)

62 DAS 87 DAS

Control WDI SR Mean Control WDII SR Mean

AK 159 30.4 25.7 26.8 27.6 33.8 25.7 31.4 30.3

DRG 1 27.9 23.3 29.6 26.9 34.0 28.1 28.4 30.2

TAG 24 29.0 26.6 27.9 27.8 38.0 27.7 31.0 32.2

ICGS 44 27.5 25.2 26.7 26.5 36.1 29.0 27.3 30.8

ICGV 86031 29.1 26.6 27.6 27.8 38.8 28.1 28.7 31.9

SG 99 28.3 25.7 27.5 27.2 35.5 27.3 29.0 30.6

Mean 28.7 25.5 27.7 27.3 36.0 27.7 29.3 31.0

CD (p = 0.05)

Water deficit 1.6 2.3

Varieties NS 1.4

Interactions 2.2 2.4

SR represents 48 h after stress relief

Ind J Plant Physiol.

123

indeterminate growth habit. Under such situation, parti-

tioning of photosynthates into developing pods is most

important physiological trait in yield determination. Single

leaf PN has been studied for several groundnut genotypes

and genetic variation is reported (Nautiyal et al. 1995;

Collino et al. 2001). The photosynthesis is fundamental in

both biomass accumulation and productivity and it could

be best utilized in identifying the efficient cultivars and to

understand the physiological traits of productivity both

under normal and stress conditions. Under increasing

moisture deficit stress, the low SLA type groundnut

genotypes were able to maintain higher RWC, PN and gs(Nautiyal et al. 2002). Nautiyal et al. (2012) reported

variation in PN and gs and based on average PN groundnut

cultivars were divided into high, medium and low groups

based on the cropping seasons and accessed the influence

of environment on PN, gs and leaf temperature. They also

reported that photosynthetic rate and associated traits dur-

ing different reproductive growth stages in summer season

showed wide genetic variability in their response to meet

the extra demand of photosynthates by the developing pods

and cultivar TAG 24 showed higher PN during all the

growth stages and maintained least increase in leaf tem-

perature (1.0 �C), which makes it most suitable for culti-

vation in summer season. Bhagsari et al. (1976) observed

large reductions in photosynthesis and stomatal conduc-

tance as the relative water content of groundnut leaves

decreased from 80 to 75 within 3 days of withholding

water in potted plants. Stomatal responses are more closely

linked to soil moisture content than to leaf water status, as

stomata are responding to chemical signals (ABA) pro-

duced by dehydrating roots (Davies and Zhang 1991).

Stage specificity in tolerance and susceptibility to water

deficit condition in this study clearly indicated that pod

development stage between 62 and 87 DAS (WD II) was

more susceptible to water stress than growth period

between 31 and 62 DAS (WD I) and variety TAG 24

showed better stress recovering capacity with high photo-

synthesis under WD condition. The data on chlorophyll

fluorescence parameters showed that cultivar ICGS 44 was

least affected to damage via photo-inhibitory action.

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Table 3 Correlation matrix between various parameters in groundnut varieties at 62 DAS in WD I and 87 DAS in WDII as affected by water

deficit condition during summer 2011

Fv/Fm NPQ PN Transpiration Tleaf

WD I WD II WD I WD II WD I WD II WD I WD II WD I WD II

NPQ -0.71** -0.73**

PN 0.66** 0.62** -0.78** -0.80**

Transpiration 0.71** 0.67** -0.83** -0.87** 0.84** 0.94**

Tleaf -0.67** -0.87** 0.80** 0.71** -0.77** -0.70** -0.87** -0.71**

gs 0.74** 0.64** -0.80** -0.84** 0.76** 0.88** 0.73** 0.93** -0.64** -0.73**

* significant at 5 % level

** significant at 1 % level

Ind J Plant Physiol.

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