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Indian J. Agric. Res., 49 (4) 2015: 299-307Print ISSN:0367-8245 / Online ISSN:0976-058X

Physiological responses of tomato (Lycopersicon esculentum Mill ) cv. ArkaAshish to elevated atmospheric CO2 under water limiting conditions

H. Mamatha*, N.K. Srinivasa Rao and T. Vijayalakshmi1

Division of Plant Physiology and Biochemistry,Indian Institute of Horticultural Research, Bangalore-560 089, India.Received: 15-01-2015 Accepted: 08-06-2015 DOI: 10.5958/0976-058X.2015.00055.4

ABSTRACTThe study was conducted to understand the physiological responses of tomato cv. Arka Ashish to 550 ppm elevated CO2

(EC) and to what extent EC alleviates the adverse effects of water stress at peak flowering stage. The plants grown at EC hadsignificantly higher PN with decreased stomatal conductance (gs) and leaf transpiration rate (E) compared to plants grown at380 ppm ambient CO2 (AC), irrespective of water supply conditions. Plants grown at EC recorded a lower number ofstomata on both adaxial and abaxial surfaces. The plants at EC maintained higher water potential(w), instantaneous wateruse efficiency (iWUE) and lower osmotic potential (s) at irrigated and water stress condition. Higher SOD and GR activitywas observed at EC compared to the plants grown at AC. Increased number of fruits and fruit weight per plant were observedat EC both at control and water stress condition.

Key words: Fruit yield, Gas exchange characteristics, Intercellular CO2, Stomatal density, Water relations.

*Corresponding autor’s e-mail: [email protected] for Environment Jawaharlal Nehru Technological University, Hyderabad-500 085, India.

INTRODUCTIONThe anthropogenic activities have caused increased

emission of CO2 into the atmosphere and the levels havereached 399.65ppm (March, 2014) from the pre-industriallevels of 280 ppm (Anonymous, 2014). As per theRepresentative Concentration Pathways (RCPs) used in theIPCC3 AR5 it is projected that the prescribed CO2

concentrations reach 421 ppm (RCP2.6), 538 ppm (RCP4.5),670 ppm (RCP6.0), and 936 ppm (RCP 8.5) by the year 2100(IPCC, 2013). The crop plants under elevated atmosphericCO2 would be benefited due to the fact that high CO2 aroundRubisco-enzyme accelerates the carboxylation reaction, whilesuppressing the competing oxygenation reaction andsubsequently reducing CO2 loss and energy costs associatedwith photorespiration (Leakey et al., 2009; Long et al., 2004).

Studies under controlled conditions have shown thatCO2 fertigation enhances PN and yield in both C3 and C4 crops(Kimball et al., 2002; Reddy et al., 2010).The doubling ofCO2 from 35 to 70 Pa, stimulates the PN by 25 to 75 % (Stitt,1991). Elevated CO2 increases net photosynthetic rate andalso above and/or below ground biomass in C3 species(Bowes, 1996; Drake et al., 1997). Plants grown at elevatedCO2 maintain high internal CO2 and hence increased PN,resulting in better growth and yield. Increased plant wateruse efficiency (WUE) and decreased transpiration rate at

elevated CO2 have been observed (Robredo et al., 2007).

Tomato, being a C3 plant, responds positively to elevatedCO2. Higher PN, total biomass and yield and reductions intranspiration rate (E) and stomatal conductance (gs) have beenobserved in tomato plants grown at elevated CO2 (Yelle etal., 1990; Behboudian, 1994). It is observed that the elevatedCO2 caused reduction in stomatal density in seven-year-oldScots pine seedlings grown at 750 ppm (Lin et al., 2001) andalso in egg plants grown at 700 ppm elevated CO2 along withwater stress (Bikash and Michihiro, 2011) .

The increased global temperature is likely to alterprecipitation patterns and also cause high evapotranspirationin crop canopies. This situation could put both natural andagricultural vegetation to greater risks of more severe andprolonged water deficiency (Ellsworth, 1999; Houghton etal., 2001). Tomato is very sensitive to water stress at floweringand fruit enlargement stages (Sibomana et al., 2013). Tomatohas highest demand for water during flowering stage(Doorenbos and Kassam, 1979). Water deficit causesreduction in number of flowers, fruits, fruit size and weightand ultimately low marketable yields (Rahman et al.,1999;Veitkohler et al., 1999; Adams, 1990).

Studies to elucidate the extent to which elevated CO2

influences physiology under water stress conditions have

















































































































































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shown that the plants grown under elevated CO2 dry moreslowly with reduced stomatal conductance and transpirationrate (Bunce, 1998; Robredo et al., 2007). This Indicatesthat, the plants grown at elevated CO2 conditions could usewater more efficiently and tolerate drought (Vu et al., 1998).The reduced plant water use delays the onset and severity ofwater deficit (Robredo et al., 2007). In tomato,increased CO2

ameliorated the detrimental effects of drought stress for fivedays (Paez et al., 1984). Consistent increase in gas exchange,dry matter production and yield of potato was observed underelevated CO2 at various levels of water stress as compared toambient CO2 (Fleisher et al., 2008). Barley plants grown underwater stress with elevated CO2 showed less depletion of soiland plant water content (Robredo et al., 2010). Thus, theelevated CO2 enables crop plants to overcome the adverseeffects encountered under water scarce conditions.

Studies have shown increased activities ofantioxidant enzymes, SOD and CAT at elevated CO2 duringinduction of water stress in wheat (Lin and Wang, 2002).Very high activities of antioxidant enzymes ascorbateperoxidase (APX), catalase (CAT), glutathione S-transferase(GST) were observed in wheat plants grown at elevated CO2

with low water supply. Elevated CO2 had favourable effectson the development and stress tolerance in crops (Benczeet al., 2014).

The present experiment was conducted to study theresponse of tomato plants under elevated CO2 with waterlimiting conditions and the extent to which the elevated CO2

alleviates the adverse effects of water stress through itsultimate influence on yield formation.

MATERIALS AND METHODSExperimental conditions and plant materials: The studywas conducted in the open top chambers (OTCs) duringDecember-2009 to March - 2010 at Indian Institute ofHorticultural Research, Bangalore, India. Thirty day oldseedlings of tomato cv. Arka Ashish were transplanted at aspacing of 50 X 50 cm in four OTCs having dimension of3m x 3m x 2.4 m covered with polyvinylchloride (PVC) sheet150 micron thickness with 95 percent light transmission. Fiverows and five plants per row were maintained in each OTC.Among the four OTC’s two were maintained with ambientCO2 (380 ppm) and the remaining two with 550 ppm elevatedCO2. The CO2 levels inside the OTCs were continuouslymonitored by non-dispersive infrared (NDIR) CO2 analyser(Lambda T, ADC Bioscientific Ltd., U.K.). The air samplefrom each OTC to the CO2 analyser was drawn at the crop -canopy height and the CO2 concentration was measured bythe analyzer. When the CO2 3levels inside the OTC dropped

below the set concentration, solenoid valve opened to supplyCO2 from the gas cylinder and remained open until the setconcentration was attained. As soon as the CO2 level crossedset concentration, the valve closed. Thus, desired level ofCO2 was maintained inside the OTC. Elevated CO2 level of550 ± 50 ppm was maintained daily for seven hours between09:00-16:00hrs. (IST).

Out of the four OTCs which were maintained atambient and elevated CO2 levels, two OTCs from each CO2

level, received continuous irrigation and another two wereimposed with water stress at flowering stage for three weeks,between 49 to 70 days after transplanting (DAT). Thus makingfour treatment combinations, ambient CO2 (380ppm) +control watersupply, ambient CO2 (380ppm) + water stress,elevated CO2 (550 ppm) + control watersupply and elevatedCO2 (550 ppm) + water stress. After releasing water stresson 70 DAT, plants received regular water supply till the endof the experiment.

Gas exchange characteristics: Gas exchange characteristicswere recorded from the uppermost fully expanded leafbetween 9:30 h to 11:30 hrs using portable photosynthesissystem (LI 6400 XT, LI – COR, Lincoln, USA) duringflowering stage at 49 and 70 DAT. A total of fivemeasurements were taken from five plants in each treatment.Cuvette temperature, light intensity and humidity were set tomean atmospheric conditions viz., 33°C, 1,200 µmol m-2s-1

and 60 %, respectively. Leaf was held in the chamber for tenminutes to obtain stable readings. While recording gasexchange characteristics, the CO2 levels in the leaf chamberwere maintained at 550 ppm and 380 ppm in respective CO2level treatments. PN, gs, and E were calculated by the softwareoperating in LI 6400 XT, using the Von Caemmerer andFarquhar (1981) equations. PN, gs, and E were expressed asµmol(CO2) m

-2 s-1, mol(H2O) m-2 s-1 and mol(H2O) m-2 s-1 ,respectively.Instantaneous water useefficiency (iWUE)wascalculated from gas exchange measurements on individualplants, as the ratio of carbon assimilation(PN) to leaftranspiration rate (E) and expressed as mmol CO2 mol-1 H2O(Dang et al., 1991; Zhang and Marshall, 1994).

Stomatal density: The epidermal layer of the leaves used forrecording gas exchange characteristics was carefully removedand mounted on a slide. The stomatal number was countedusing a compound fluorescence microscope (OLYMPUSBX60., India) at 40X magnification. The microscopic fieldarea was measured using stage and ocular micrometer and thenumber of stomata per mm2 was worked out.

Water and osmotic potential: The w and s were recordedat 49 and 70 DAT from five fully expanded leaves. The w

was recorded using a pressure chamber (ARIMAD-3000,









































































































































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Israel).Third fully expanded leaf from top was excised andthe petiole was inserted into the pressure chamberimmediately. The values are expressed in mega Pascal (-MPa).Leaf s was measured using a vapour pressure osmometer(VAPro, WESCOR, USA). Leaf cell sap was squeezedmanually from third fully expanded leaf onto filter paper andplaced in the osmometer sample chamber to read the s inmega Pascal (-MPa).

Yield characteristics: The total fruit weight (g plant-1) andnumber of fruits per plant were quantified by pooling thefruit weight and number of fruits harvested at regular intervals.

Assay of antioxidant enzyme activity: Activity of superoxide dismutase (SOD) and glutathione reductase (GR) wasestimated in the fully expanded leaves. Leaf tissue (0.5 g)was ground to a fine powder with liquid nitrogen using pestleand mortar and extracted at a ratio1:4 (w/v) fresh weight inextraction buffer (50mM potassium phosphate buffer pH 7containing 1mM EDTA, 2mM ascorbate and 1% solublePVP). The homogenate was centrifuged at 12,000g for 15min at 4°C and the supernatant was used for the assays. Theactivity of SOD was determined according to the method ofGiannopolitis and Chloride (1977) by following the photo-reduction of nitroblue tetrazolium (NBT) with somemodification. The reaction buffer (3 ml) contained 50 mMpotassium phosphate buffer (pH 7.8), 0.1 mM EDTA, 13 mMmethionine, 75µM NBT, 2mM riboflavin and 50 µl of theenzyme extract. Test tubes containing the reaction mixturewere irradiated under fluorescent light for 15 min. Theabsorbance of irradiated and non-irradiated solution wasmeasured at 560 nm. One unit of SOD activity was assignedas the amount of enzyme that would inhibit 50 % of reaction.GR (Glutathione reductase) activity was assayed as thedecline in the absorbance of NADPH at 340 nm. The reactionmixture consisted of 50 mM phosphate buffer (pH 7.0), 0.5mM oxidized glutathione (GSSG), 0.1 mM NADPH and 100µl of enzyme extract (Jiang and Huang, 2001).

RESULTS AND DISCUSSIONThe plants grown at 550 ppm elevated CO2 (EC)

recorded significantly higher PN compared to plants grown at

380 ppm ambient CO2 (AC) irrespective of water supplyconditions. Though, water stress caused reduction in PN ofthe plants grown at both EC and AC, the reduction was lowerat EC. Plants at AC showed 13.05 % reduction in PN at theend of three week water stress. However, plants grown at ECshowed only 5.78 % reduction in PN after three week waterstress (Table 1).

Several studies across C3 and C4 crops have shownsignificant positive influence of elevated CO2 on plant growthand yield. Mamatha et al. (2014) recorded significantly higherPN , decreased leaf transpiration and stomatal conductance inplants grown at elevated CO2 (700 and 550ppm) as comparedto ambient CO2 (380 ppm). Behboudian (1994) observed asignificant increase in PN in tomato plants grown at 1000ppm and 700 ppm elevated CO2 levels as compared to theplants grown at 340 ppm ambient CO2. Higher netphotosynthesis rates across eight tomato genotypes (Trippet al., 1991) and two species, ‘Vedettos’ (Lycopersiconesculentum) and LA1028 (Lycopersicon chmielewskii) (Yelleet al.,1990) have also been reported. Vanaja et al. (2011)found that elevated CO2 significantly improved the PN in bothwell watered and moisture stressed sunflower plants. Underwater stress condition the sunflower plants grown at elevatedCO2 recorded significantly higher PN (60%) as compared withambient CO2 (32%). Similar results were observed by Wardet al. (1999) in C3 and C4 plants.

The plants grown under EC with well irrigated aswell as water stress had lower gs. The imposition of waterstress caused 86.95% and 33.33% reduction in gs in the plantsgrown at EC and AC, respectively (Table 2).Concomitant tothe reductions in gs, the E also decreased at EC (Table 3).

The plants grown at EC with increased PN and low Ehad higher instantaneous water use efficiency (iWUE) ascompared to plants grown at AC. The higher iWUE was observedat well irrigated as well as under water stress. After the thirdweek of water stress 6.03% increase in iWUE was observed atEC as compared to EC with control. However there was only0.94% increased iWUE was recorded at AC with water stress ascompared to AC with well irrigated condition(Table 4).

TABLE 1: Effect of elevated CO2 and water stress levels on net photosynthetic rate of tomato cv. Arka Ashish

Treatments CO2 levels Mean (T)550 ppm (C1) 380 ppm(C2)

Irrigated (T1) 25.44 23.21 24.71Water stress(T2) 23.97 20.18 21.69Mean (C) 24.33 22.07

F-Test S.Em.+ CD (P=0.05)Treatments (T) * 0.41 1.38CO2 levels (C) * 0.41 1.38TxC * 0.59 1.96

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The plants grown at EC showed reductions in gswithboth well irrigated as well as water stress conditions. Studieshave reported that in the herbaceous plants, reduction in gs

under elevated CO2 might be between 27 and 40% (Field etal., 1995). Vanaja et al. (2011) also recorded similar resultsin sunflower and maize. We observed the duration of waterstress up to three weeks and recorded a substantial reductionsin gs especially at EC. Concomitant to the reductions in gs,the E also decreased at EC and water stress caused, furtherreductions. Studies have shown that in tomato, in addition toincrease in PN, elevated CO2 caused decrease in gs (Yelle etal., 1990) and transpiration rate (Behboudian, 1994). Duringwater stress, the decreased gs and E are due to partial closureof stomata and plants reduce water loss by decreasingconductance to water vapour and thus decreased transpiration(Rao et al., 2000). In our study at EC increased PN and low Erecorded higher instantaneous water use efficiency (iWUE)as compared to plants grown at AC. Peach (Prunus persica)seedlings grown at EC (700 ppm) recorded 51, 63% higher

TABLE 2: Effect of elevated CO2 and water stress levels on stomatal conduction of tomato cv. Arka Ashish



F-Test S.Em+ C.D.@5%Treatments (T) * 0.01 0.02CO2 levels (C) * 0.01 0.02TxC * 0.01 0.03

TABLE 3: Effect of elevated CO2 and water stress levels on leaf transpiration rate of tomato cv. Arka Ashish

Treatments CO2 levels Mean (T)

550 ppm (C1) 380 ppm(C2)Irrigated (T1) 8.15 11.05 9.60Water stress(T2) 6.71 9.36 8.04Mean (C) 7.43 10.21

F-Test S.Em+ C.D.@5%Treatments (T) * 0.34 1.01CO2 levels (C) * 0.34 1.01TxC NS 0.48 1.43

TABLE 4: Effect of elevated CO2 and water stress levels on instantaneous water use efficiency (iWUE) of tomato cv. Arka AshishTreatments CO2 levels Mean (T)

550 ppm (C1) 380 ppm(C2)


F-Test S.Em+ C.D.@5%Treatments (T) NS 0.12 0.35CO2 levels (C) * 0.12 0.35TxC NS 0.16 0.49

WUE in well watered and water stress condition, respectively,than WUE of comparable seedlings grown in AC (350ppm)(Centritto et al., 2002).

Plants grown at EC had significantly lower stomataldensity and a reduction of 24%(39.05) on adaxial and29%(100) on abaxial surfaces was observed as compared toAC (Table 5). The significant reduction in stomatal densityof both adaxial and abaxial surfaces at EC was observed inthe present study.The adaptation of stomatal properties toelevated CO2 concentration was observed in eggplants, wherestomatal density was 24% lower on adaxial surface and 5%lower on abaxial surface under elevated CO2 than in ambientCO2concentration(Bikash and Michihiro, 2011).

Imposition of water stress for a period of three weekscaused decrease in s at both EC and AC. Lower (morenegative) s (1.23)was observed at EC compared to plantsgrown at AC(1.02) (Table 7). In comparision to plants grownat AC, EC maintained higher (less negative) w at well













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TABLE 5: Effect of elevated CO2 stomatal density of tomatocv. Arka Ashish

Treatments Adaxial (/mm2) Abaxial (/mm2)T1 39.05 100.00T2 51.43 140.00F-Test * *S.Em+ 1.41 7.30CD@5% 4.24 21.92

TABLE 6: Effect of elevated CO2 and water stress levels on leaf water potential of tomato cv. Arka Ashish



F-Test S.Em+ C.D.@5%Treatments (T) S 0.04 0.11CO2 levels (C) S 0.04 0.11TxC NS 0.05 0.16

TABLE 7: Effect of elevated CO2 and water stress levels on osmotic potential of tomato cv. Arka Ashish




irrigated as well as water stress durations (Table 6).The resultsof w values are in conformity with the results of Vanaja etal.,(2011).Tomato plants grown at elevated CO2 recordedhigher (less negative) w and Lower (more negative) s

(Mamatha et al., 2014). The lower (more negative) osmoticpotential with better osmoregulation was observed in wheatplants grown under 1000 ppm CO2 imposed with water stressas compared to control plants (Sionit et al., 1981). In ourstudy elevated CO2 counteracted the water stress effect bymaintaining high water potential, decreased gs and E, withhigher PN; which is in conformity with the results of Robredoet al., (2007). Higher leaf water potential and lower osmoticpotential helped plants to maintain higher turgorpressure(Sionit et al.,1981). Elevated CO2 improves soil andwater relations under drought conditions by maintaininghigher WUE. The decline in PN with more negative waterpotential might be the result of a reduction in gs. In barley,the higher CO2 concentration enabled plants to offset therestriction imposed by stomata (Robredo et al., 2007). Twotomato cultivars grown at low (350 ppm) and high (675 ppm)CO2 recorded decreased total leaf water potential and osmotic

potential when water was withheld for five days at thirteendays after emergence of tomato plants, but the decrease wasless rapid in high CO2 than in low (Paez et al., 1984). Holcuslanatus plants exposed to 700 ppm CO2 concentration withwater stress recorded higher water potential as compared toplants raised at ambient CO2 with water stress (Clark et al.,1999). Xiao et al. (2005) suggest that elevated CO2 mayenhance drought avoidance and improved water relations,thus weakening the effect of drought stress on growth ofCaragana intermedia seedlings.

Higher SOD activity was recorded in plants grownat EC with water stress(459.370) as compared to all othertreatments. However lower SOD activity(283.753) wasrecorded in plants grown at AC with water stress (Table 9).The activity of GR also was higher(0.194) in plants grown atEC imposed with three weeks water stress compared to allother treatments (Table 8). Higher SOD activity was observedin the plants grown at EC irrespective of water supply andactivity increased as the water stress progressed. The activityof GR also was higher in plants grown at EC imposed withthree weeks water stress compared to plants grown at ACwith water stress.Similar results were recorded by Schwanzet al., (1996), where Pendunculate oak (Quercus robur) andmaritime pine (Pinus pinaster) seedlings grown at 700 ppmrecorded significant enhancements of the enzymatic activityin response to drought. Further they also observed that SODactivity reflects the need for detoxification of O2

-, the rate ofO2

- production increased in plants grown with drought stresswith elevated CO2. Perhaps plants grown at an elevated CO2,

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-, the rate ofO2


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-, the rate ofO2



TABLE 8: Effect of elevated CO2 and water stress levels on glutathione reductase(GR) activity of tomato cv. Arka Ashish




Table 9. Effect of elevated CO2 and water stress levels on Super oxide dismutase (SOD) activity of tomato cv. Arka Ashish




concentration contain increased metabolic reserves and, thus,might have an enhanced metabolic flexibility to respond tooxidative stress compared with plants grown at ambient CO2

concentrations. Higher antioxidant enzyme activity under highCO2 with stress indicate the better free radical scavengingmechanism due to higher WUE, compared to ambient CO2

grown plants under stress condition. Stimulation of theantioxidant enzymes APX, CAT, GST were recorded by Benczeet al. (2014) in wheat plants grown at elevated CO2 and drought.

Increased number of fruits and fruit weight per plantwere observed irrespective of water supply. Plants exposedto EC with well irrigated condition recorded a 80 % increasein the number of fruits (Fig.1) and 128% increase in fruit

FIG 1: Effect of elevated CO2 on number of fruits per plant oftomato cv. Arka Ashish grown at 550 ppm elevated CO2(EC) +

control water supply (C1T1), 380 ppm ambient CO2 (AC) +control water supply (C2T1), EC + water stress (C1T2), AC +

water stress (C2T2). The values are means ± SE (n = 4).

FIG 2: Effect of elevated CO2 on yield (gram plant-1) of tomatocv. Arka Ashish grown at 550 ppm elevated CO2(EC) + control

water supply (C1T1), 380 ppm ambient CO2 (AC) + control watersupply (C2T1), EC + water stress (C1T2), AC + water

stress(C2T2).The values are means ± SE (n = 4).

weight per plant over AC (Fig.2). However yield reductionwas less (44%) at EC with water stress over EC with controlas compared to AC with water stress over AC with control(50%).The number of fruits increased by 129% and fruitweight increased by 159% at EC under water stress for threeweeks as compared to the plants grown at AC. In the presentstudy at elevated CO2 higher number of fruits per plant withincreased yield irrespective of water stress was observed.Studies have suggested that CO2 enrichment increases sinkstrength more than source strength in tomato, and during fruitdevelopment carbohydrates may be partitioned to the fruits,leading to higher yields (Peet et al., 1991). Mamatha et al.(2014) recorded maximum number of fruits and fruit weight

































































































































































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REFERENCESAdams, P. (1990). Effects of watering on the yield, quality and composition of tomatoes grown in bags of peat.

J.Hortic.Sci.,65:667-674.Anonymous. 2014. www.esrl.noaa.gov/gmd/ccgg/trends/ NOAA/ESRL:Recent monthly average Mauna Loa CO2, March

2014.Behboudian, H.M. (1994). Carbon Dioxide Enrichment in ‘Virosa’ Tomato Plant: Responses to Enrichment Duration and to

Temperature. HortScience., 29:1456-1459.Bencze, S.; Bamberger, Z.; Janda, K.; Balla, B.; Varga, Z.; Bedo. and Veisz, O. (2014). Physiological response of wheat

varieties to elevated atmospheric CO2 and low water supply levels. Photosynthetica., 52:71-82.Bikash, C.; Sarker and Michihirohara. (2011). Effects of elevated CO2 and water stress on the adaptation of stomata and gas

exchange in leaves of eggplants (Solanumm elongena L.). Bangladesh J. Botany., 40:1-8.Bowes, G. (1996). Photosynthetic responses to changing atmospheric carbon dioxide concentration. In: Baker N.R.,(1996)

ed. Photosynthesis and the environment. Advances in photosynthesis, 5, Dordrecht, The Netherlands: KluwerAcademic Publishers, Pp. 387-407.

Bunce, J.A. (1998). Effect of humidity on short-term responses of stomatal conductance to an increase in carbon dioxideconcentration. Plant Cell Environ., 21:115-120.

Centritto, M.; Lucas, E.M. and Jarvis, P.G. (2002). Gas exchange, biomass, whole-plant water-use efficiency and wateruptake of peach (Prunus persica) seedlings in response to elevated carbon dioxide concentration and wateravailability. Tree Physiol., 22:699-706.

Clark, H.; Newton, P.C.D and Barker D.J, (1999). Physiological and morphological responses to elevated CO2 and a soilmoisture deficit of temperate pasture species growing in an established plant community. J. Exp. Bot., 50:233-242.

Dang, Q.L.; Lieffers, V.J.; Rothwell, R.L.; S.E. and Mcdonald. (1991). Diurnal variation and interrelations of ecophysiologicalparameters in the three peatland woody species under different weather and soil moisture conditions. Oecologia.,88:317-324.

Doorenbos, J. and Kassam, A.H. (1979). Yield response to water. Food and Agriculture Organization irrigation and drainagepaper, 33, Food and Agriculture Organization, Rome. 157pp.

Drake, B.G.; Gonzalez-Meler, M.A. and Long, S.P. (1997). More efficient plants: a consequence of rising atmosphericCO2? Annual Review of Plant Physiology and Plant Molecular Biology, 48:609-639.

Ellsworth, D.S. (1999). CO2 enrichment in a maturing pine forest: are CO2 exchange and water stress in the canopy affected?Plant Cell Environ., 22:461- 472.

per plant in tomato plants exposed to elevated CO2 (700 and550ppm), percent increase in fruit yield was 125 and 54 at700 and 550ppm, respectively as compared to control plants.An increase in yields by 80% in the early (first three weeksof harvest) and 21.5% in total yield was observed in tomatocv.Vedettos (Yelle et al., 1990). Plants exposed to 900 ppmrecorded seven percent increase in cumulative yields ofmarketable tomato than plants exposed to 350 ppm (AlejandroFierro et al., 1994). Richard et al. (1997) recorded 22-41percent increased cumulative tomato yield at increasing CO2concentrations from 450-675 ppm. Islam et al. (2006) recordeda significantly larger fruits in tomato cv. ‘Momotaro’, ‘Ladyfirst’, and ‘minicarol’ grown at elevated CO2 (850 + 50)compared to ambient (350 + 50) CO2 concentration. Yieldincrease was also due to good osmotic adjustment by anincreased concentration of dissolved solutes in leaf cells (Sionitet al., 1981). Thus, the loss in total number of fruits and fruit

weight per plant due to water stress at flowering stage from 49to 70 DAT could be alleviated by elevated CO2.

CONCLUSIONSIn the present study, we observed a significant

increase in PN at EC irrespective of water supply conditions.Concomitantly the EC caused reductions in gs and E at bothwell irrigated and water stress condition. Plants adapted toEC with lower stomatal density on both adaxial and abaxialsurfaces. Overall, the EC counteracted the water stress effectby maintaining high water potential with good osmoticadjustment by maintaining high dissolved solutes, decreasedgs and E, and higher PN. Higher activity of SOD and GR atEC indicated better free radical scavenging mechanism.TheEC alleviated the adverse effects of water stress for threeweeks imposed at flowering stage with plants recording highernumber of fruits per plant with increased yield.

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Field, C.B.; Jackson, R.B. and Mooney, H.A. (1995). Stomatal responses to increased CO2 - implications from the plant tothe global scale. Plant Cell Environ., 18:1214-1225.

Fierro, A.; Tremblay, N. and Andre, G. (1994). Supplemental Carbon Dioxide and Light Improved Tomato and PepperSeedling Growth and Yield. Hortscience., 29:152-154.

Fleisher, David H., Timlin. Dennis J. and Reddy. V.R. (2008) Elevated carbon dioxide and water stress effects on potatocanopy gas exchange, water use, and productivity. Agr. Forest. Meteorol. 148:1109-1122.

Giannopolitis, C.N. and Chloride, S.K. (1977). Superoxide dismutase. I. occurrence in higher plants. Plant Physiol., 59:309-314.

IPCC, 2013: Summary for Policymakers. In: Climate Change 2013: The Physical Science Basis. Contribution of WorkingGroup I to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change [Stocker, T.F., D. Qin,G.-K. Plattner, M. Tignor, S.K. Allen, J. Boschung, A. Nauels, Y. Xia, V. Bex and P.M. Midgley (eds.)]. CambridgeUniversity Press, Cambridge, United Kingdom and New York, NY, USA.

Islam, S.; Khan, S.; James, O. and Garner. (2006). Elevated atmospheric CO2 concentration enhances carbohydrate metabolismin developing Lycopersicon esculentum mill. Cultivars. Int. J. Agri. Biol., 8:157-161.

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Volume 49, Issue 4, 2015 307














Volume 49, Issue 4, 2015 307














Volume 49, Issue 4, 2015 307














Volume 49, Issue 4, 2015 307














Volume 49, Issue 4, 2015 307














Volume 49, Issue 4, 2015 307














Volume 49, Issue 4, 2015 307














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