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Agricultural Water Management

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Carbon retention in the soilplant system under different irrigation regimes

Yaosheng Wang a, , Fulai Liu a , Mathias N. Andersen b , Christian R. Jensen a
a b

Department of Agriculture and Ecology, Faculty of Life Sciences, University of Copenhagen, Hjbakkegrd All 13, DK-2630 Taastrup, Denmark Department of Agroecology and Environment, Faculty of Agricultural Sciences, University of Aarhus, P.O. Box 50, DK-8830 Tjele, Denmark

a r t i c l e
Article history: Available online xxx

i n f o

a b s t r a c t
Carbon (C) sequestration through irrigation management is a potential strategy to reduce C emissions from agriculture. Two experiments (Exps. I and II) were conducted to investigate the effects of different irrigation strategies on C retention in the soilplant system in order to evaluate their environmental impacts. Tomato plants (Lycopersicon esculentum L., var. Cedrico) were grown in split-root pots in a climate-controlled glasshouse and were subjected to full irrigation (FI), decit irrigation (DI) and alternate partial root-zone irrigation (PRI) at early fruiting stage. In Exp. I, each plant received 2.0 g chemical nitrogen (N), while in Exp. II, 1.6 g chemical N and maize residue containing 0.4 g organic N were applied into the pot. The results showed that, in both experiments, the concentration and the amount of total C in the soil were lower in FI and PRI as compared to DI, presumably due to a greater microbial activity in the two treatments; particularly the PRI induced drying and wetting cycles of the soils may cause an increase of microbial activities and respiration rate, which could lead to more C losses from the soil. However, in both experiments the total C concentration in the PRI plants was the highest as compared with the FI and DI plants, and this was seemingly due to improved plant N nutrition under the PRI treatment. Consequently, the total amount of C retained in the soilplant system was highest in the FI and was similar, but lower, for the PRI and DI. The different N input in the two experiments might have affected the C retention in the soil and in the plant biomass. Nevertheless, with a same degree of water saving, PRI was superior to DI in terms of enhancing C concentration in the plant biomass, which might have contributed to a better fruit quality in tomatoes as reported by Zegbe et al. (2004, 2006). 2010 Elsevier B.V. All rights reserved.

Keywords: Partial root-zone irrigation Decit irrigation Carbon sequestration

1. Introduction The effects of elevated CO2 concentration in the atmosphere on possible global warming have attracted increased attention into the research for enhancing carbon (C) sequestration in the ecosystems. Increase of the amount of C sequestered in cropland through improved agricultural management has become one important strategy to reduce CO2 emissions to the atmosphere (Lal et al., 1998; Lal and Stewart, 2010). Field management practices such as soil restoration, conservation tillage, cover crops, long-term manure application and crop rotation have shown great potential in enhancing soil C sequestration (Lal, 2004, 2008). In addition to those options, irrigationan essential management tool for sustaining crop production, may also contribute to C retention in the soilcrop system especially in arid areas (Follett, 2001; Gillabel et al., 2007; Lal, 2008). Declining freshwater resources have stimulated research into developing novel irrigation strategies to use the water more ef-

Corresponding author. E-mail address: yawa@life.ku.dk (Y. Wang). 0378-3774/$ see front matter 2010 Elsevier B.V. All rights reserved. doi:10.1016/j.agwat.2010.07.010

ciently. Alternate partial root-zone irrigation (PRI) and decit irrigation (DI) are water-saving irrigation techniques being intensively studied in many regions of the world (Liu et al., 2006). DI is a method that irrigates the entire root zone with an amount of water less then the potential evapotranspiration and the minor stress that develops has minimal effects on the yield (English and Raja, 1996). PRI is a further renement of DI, and the principle behind PRI is to alternately let one part of the root system be exposed to soil drying, while the other part is irrigated, in order to keep the leaves hydrated. PRI has been demonstrated to allow considerable water savingsin some cases almost a doubling of water use efciency (WUE) (Kirda et al., 2007). Besides saving irrigation water, PRI induced drying and wetting cycles of the soils could cause a ush of C and N, and an increase in soil respiration rate (Birch, 1958; Xiang et al., 2008; Butterly et al., 2009). Our recent studies in potato (Solanum tuberosum L.) and tomato (Lycopersicon esculentum L.) have demonstrated that PRI could improve crop N nutrition presumably due to an increased soil mineral N availability caused by an accelerated mineralization rate of soil organic N (Shahnazari et al., 2008; Wang et al., 2009, 2010). Accordingly, it is very possible that PRI may lead to greater C losses from the soils due to the ushes of C and N and an increase in soil respiration. However,

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content (vol.%) of 30.0 and 5.0 at water holding capacity and permanent wilting point, respectively. Before lling the pots, the soil was mixed thoroughly and homogeneously with 2.0 g N as NH4 NO3 in Exp. I, and 1.6 g N as NH4 NO3 together with 25.0 g maize straw, grounded to particle size <1.5 mm, which had a total N content of 16.8 g kg1 and total C content of 391.5 g kg1 in Exp. II. In both experiments, for each pot, 0.87 g pot1 and 1.66 g pot1 of P and K, respectively, were applied to meet the nutrient requirements for plant growth. The average value of soil water content in the soil prole was monitored by time domain reectometry (TDR, TRASE, Soil Moisture Equipment Corp., Santa Barbara, CA, USA) with probes (33 cm in length) installed in the middle of each soil compartment. Before onset of the experiment, the TDR soil moisture measurement has been calibrated by plotting the actual volumetric water content of the soil by weighing the pots against the TDR readings at different pot weight. The climate conditions in the glasshouse were set by an automatic computer controlling system at: 20/17 2 C day/night air temperature, 16 h photoperiod and >500 mol m2 s1 photosynthetic active radiation (PAR) supplied by sunlight plus metal-halide lamps. 2.2. Irrigation treatment Two weeks after transplanting, the plants were subjected to three irrigation treatments: (1) full irrigation (FI) where both soil compartments were watered daily from the surface of the soil at 09.00 h to 30% to compensate the full evapotranspiration water loss during the previous day; (2) partial root-zone drying irrigation (PRI), where one soil compartment was watered to 30% while the other was allowed to dry to 710%, then the irrigation was shifted between the two soil compartments; and (3) decit irrigation (DI), where the same amount of water used for PRI was evenly irrigated into the two soil compartments. The experiment was a completely randomized design with 12 replicates in each treatment. Evapotranspiration (ET) was calculated based on the daily TDR soil moisture measurements. For FI plants, ET at day i was calculated as: ETi = 5.0 [(30.0% N,i ) + (30.0% S,i )] for PRI plants, ET at day i was calculated as: ETi = 5.0 [(30.0% N,i ) + (S,i S,i+1 )] (2) (1)

Fig. 1. The diagram of the split-root setup in tomato pot experiment.

the improved crop N status and WUE under PRI caused increase of plant biomass may thus result in greater C retention in the plant, compensating the C losses from the soil. Thus, in order to evaluate the effects of the irrigation strategies on the total C retention in the soilplant system, both the C economy in the soil and in the plant biomass under different irrigation regimes need to be investigated. Therefore, the objective of the present study was to examine the effects of PRI as compared with DI and full irrigation (FI) on C retention in the soil and the plant biomass. In order to study this, tomato plants were grown in split-root pots in a climate-controlled glasshouse and were subjected to FI, DI and PRI treatments during early fruiting stage in two experiments. In Exp. I, each plant received 2.0 g chemical nitrogen (N), while in Exp. II, 1.6 g chemical N and maize residue containing 0.4 g organic N were applied into the pot. The total C concentrations in the soils and the plants were determined, and the C retention in the soilplant system was analyzed under different irrigation regimes. 2. Materials and methods 2.1. Experimental setup Two experiments (Exps. I and II) were conducted in a glasshouse located at the experimental farm of the Faculty of Life Sciences, University of Copenhagen, Taastrup, Denmark. Tomato (L. esculentum L., var. Cedrico) seedlings were transplanted into 10 L PVC pots (17 cm diameter and 50 cm deep). The pots were evenly divided into two vertical compartments by plastic sheets which were glued to the walls of the pots by silicon sealant such that water exchange between the two compartments was prevented (Fig. 1). The pots were lled with 14.0 kg of soil with a bulk density of 1.36 g dry weight cm3 . Before lling the pots, the soil was sieved by passing it through 2 mm mesh. The soil was developed on moraine deposits from the Weichselian Glaciation, and is classied according to Soil Survey Staff (1992) as an Agrudalf. The soil texture was classied as sandy loam, having a pH of 6.7, with total C of 14.2 g kg1 and total N of 1.6 g kg1 in Exp. I; and with total C of 12.9 g kg1 and total N of 1.4 g kg1 in Exp. II. The soil had a volumetric soil water

when the N (north) soil compartment was being irrigated; or as: ETi = 5.0 [(30.0% S,i ) + (N,i N,i+1 )] when the S (south) soil compartment was being irrigated. For DI plants, ET at day i was calculated as: ETi = 5.0 [(N,i N,i+1 ) + (S,i S,i+1 )] + IPRI IPRI = 5.0 (30.0% N,i ) or 5.0 (30.0% S,i ) where 5.0 is the volume of each soil compartment (L), i is the soil water content (%) at day i of soil compartment just before irrigation. N and S denote the north and south soil compartment of the pots, respectively. IPRI is the amount of irrigation used for the PRI treatment. Plant water use during the treatment period was calculated as the sum of the daily ET. The water used for the irrigation was tap water with negligible concentrations of nutrients. The irrigation treatments lasted 29 days in experiment I and 27 days in experiment II, during which period each soil compartment of the PRI plants had experienced three dry/wet cycles (Fig. 2). (4) (3)

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Fig. 2. Daily average volumetric soil water contents in the pots under FI, DI and PRI treatments in Exps. I and II. PRI-N and PRI-S represent the north and the south soil compartment of the PRI pots, respectively. DAT denotes days after onset of irrigation treatment. Values are means standard error of the means (S.E.) (n = 412).

Fig. 3. Effect of FI, DI and PRI treatments on plant dry biomass of tomato plants in Exps. I and II. DAT denotes days after onset of irrigation treatment. Values are means S.E. Different letters at each sampling date indicate signicant differences between treatments according to Duncans multiple range test at P < 0.05.

Fig. 4. Changes of total C concentration in the plant and soil under FI, DI and PRI treatments in Exps. I and II. DAT denotes days after onset of irrigation treatment. Values are means S.E. Different letters at each sampling date indicate signicant differences between treatments according to Duncans multiple range test at P < 0.05.

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2.3. Measurements and analysis Plant materials and soil samples were collected on 0 (3rd March, 2010), 15, 22 and 29 days in experiment I, and 0 (14th May, 2010), 13, 20 and 27 days in experiment II, after onset of the irrigation treatment (DAT) with four replicates in each treatment at each sampling. During harvest, leaf area was measured with a leaf area meter (model 3050A, Li-Cor Inc., Lincoln, NE, USA). Dry biomass of plant samples was determined after oven drying at 70 C to constant weight. Soil respiration rate was measured in experiment II by a Li-6200 infrared CO2 analyser (Li-Cor, Inc., Lincoln, NE, USA). The plant and soil samples were analyzed for total N and total C using the Dumas dry combustion method in a system consisting of an automated nitrogen carbon analysis unit for solids and liquids (ANCA-SL) Elemental Analyser coupled to a 20-20 Mass Spectrometer (Europa Scientic Ltd. Crewe, UK). Specic leaf N and C contents were calculated as the mass of N and C per cm2 leaf area, respectively.
Fig. 5. Relationship between specic leaf C content and specic leaf N content in the leaves of tomato plants after four weeks FI, DI and PRI treatments in Exp. I (hollow marker) and II (solid marker). **Signicance of the regression line at P < 0.01.

2.4. Statistical analysis The data were analyzed by one-way analysis of variance (ANOVA) using SAS 9.1 (SAS Institute, Inc., 2004). Duncans multiple range test was applied to assess the differences between treat-

Fig. 6. Changes of amount of total C in the plant, soil and C retention in the soilplant system under FI, DI and PRI treatments in Exps. I and II. DAT denotes days after onset of irrigation treatment. Values are means S.E. Different letters at each sampling date indicate signicant differences between treatments according to Duncans multiple range test at P < 0.05.

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ments at a signicant level of 5%. Regression analyses were used to determine the relationships between the measured parameters. 3. Results 3.1. Plant dry biomass The plant dry biomass was the highest in FI, intermediate in PRI, and the lowest in DI during the treatment period in Exp. I (Fig. 3). Nevertheless, in Exp. II, plant dry biomass was similar among treatments before 20 DAT, whereas it was signicantly greater in FI as compared with DI and PRI by the end of the experiment. 3.2. Total C concentration in the plant and soil In both experiments, the total C concentration in the plant was constantly higher in PRI than in FI and DI during most part of the treatment period (Fig. 4). There was a signicant positive correlation between specic leaf C content and specic leaf N content across the irrigation treatments (Fig. 5). However, the total C concentration in the soil showed different pattern. During the period from 0 DAT to 15 DAT in Exp. I, and 0 DAT to 13 DAT in Exp. II, total C concentration in the soils decreased remarkably in all treatments; however, the concentrations of total C in the DI soil increased thereafter, and were slightly higher than those of the FI and PRI soils at the end of the experiments (Fig. 4). 3.3. The amount of total C in the plant and soil, and C retention in the soilplant system The amount of total C in plants was signicantly higher in FI and PRI than in DI on 22 and 29 DAT in Exp. I (Fig. 6). In Exp. II, total C amount in plants was similar among treatments before 20 DAT, whereas it was signicantly greater in FI as compared with DI and PRI on 27 DAT. The total C amount in the soils under all treatments decreased markedly on 15 DAT and 13 DAT in Exps. I and II, respectively, and the total C amount remained constant in FI and PRI thereafter (Fig. 6). In contrast, the amount of total C in the DI soil increased considerably from 15/13 DAT, and was higher than that in FI and PRI at the end of the experiments. In total, the amount of C retained in the soilplant system was the highest in FI, intermediate in PRI, and the lowest in DI during the treatment period of Exp. I (Fig. 6); in Exp. II, the amount of C retention in the soilplant system was slightly higher in FI than in DI and PRI. 4. Discussion Irrigation can affect C balance in several ways. The CO2 xed in plant biomass through photosynthesis can be stored in the soil as organic C by converting plant residue into soil organic matter after the residue has been returned to the soil. Carbon sequestration through irrigation has been estimated to be 50150 kg ha1 yr1 based on increased plant C inputs (Lal et al., 1998). However, irrigation may also decrease soil organic C storage directly by stimulating microbial activity and organic matter decomposition (De Bona et al., 2006). Therefore, the net C sequestration as affected by different irrigation regimes has to be investigated based on both the C retained in the plant biomass and the C stored in the soil. 4.1. Effect of different irrigation regimes on C retention in the plant More C can be retained in the plant by either a greater plant biomass or an increased C concentration in the biomass. In the present study, PRI and DI treatments saved ca. 25% irrigation water,

and the larger irrigation volume applied in the FI resulted in significantly greater plant biomass than those of the PRI and DI (Fig. 3). The great plant biomass in the FI treatment accounted mostly for the greater amount of C retained in the plants (Fig. 6). In the PRI treatment, both the slightly greater dry biomass and signicantly higher total C concentration in the biomass might have contributed to the slightly higher C retained in the plants as compared to the DI treatment, especially in Exp. I. It is notable that the increased plant biomass under PRI as compared with DI was mainly due to a greater water use efciency in the PRI plants, as both irrigation treatments received the same amount of irrigation volume during the treatment period (Wang et al., 2010). However, the reasons for the greater C concentration in the PRI than in the DI plants remain unknown. We speculate that the ability of C utilization in the plant may inuence the C concentration in the plant biomass. It is well known that plant N nutrition plays an important role in regulating C metabolism in plants as N is an essential component for all of the enzymes involving in carbohydrate transport, metabolism and utilization in the plants (Huppe and Turpin, 1994). In the present study, this is clearly illustrated by the signicantly positive linear relationship between specic leaf C content and specic N content observed in the two experiments (Fig. 5), indicating that a higher leaf N content in the PRI plants might have facilitated C xation, transport and metabolism. In addition, the generally higher specic leaf N content in Exp. I than in Exp. II led to greater C concentration in the plant biomass (Figs. 4 and 5), indicating that the status of plant available N in the soil might have inuenced the C economy in tomato plants under the three irrigation treatments. 4.2. Effect of different irrigation regimes on soil C storage It is known that the status of C in the soil is determined by many factors. Among others, soil microbe-mediated C mineralizationimmobilization and turnover processes play an important role in determining soil C dynamics. The CO2 release from the soil into the atmosphere via microbial respiration represents a major pathway of soil C losses (Parkin and Kaspar, 2003; Casals et al., 2009). Many studies have demonstrated that microbial activity is largely dependent on soil moisture conditions (e.g. Magid et al., 1999), and microbial activity has been found to be greatly reduced at low soil water potentials (Fierer and Schimel, 2003); it has also been documented that drying and rewetting cycles of the soils can cause an increase of microbial activities and respiration rate, and thus an increased C loss from the soil (van Gestel et al., 1993; Xiang et al., 2008). We found that the respiration rate was the highest in FI. In the PRI treatment, rewetting the dry soil compartment consistently caused a burst of respiration rate, which was signicantly greater than that of the DI (unpublished data). Consequently, the greater soil respiration under FI and the pulses of respiration induced by the drying and wetting cycles under PRI may lead to more C losses from the soil than under the DI treatment. In accordance with this, in both of the two experiments we observed that the concentration and the amount of total C in the soil was the highest in DI, followed by FI, and the lowest in PRI at the end of the experiments (Figs. 4 and 6). The favorable moisture regime under FI caused high soil microbial activity and great respiration rate, while high moisture content under irrigation may also stabilize C in the soil. Gillabel et al. (2007) found that C storage under center-pivot irrigation was 25% higher than under dryland cultivation in an agricultural system. They postulated that irrigation increased microbial activity resulting in higher C sequestration inside microaggregates. During each irrigation event, the same amount of water for PRI was evenly irrigated onto the surface of the two soil compartments in DI; thus for the latter treatment, the deep soil layer was probably dry, and this might have led to more sloughing off of roots and their decomposition, and thus more root-derived soil organic C

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Y. Wang et al. / Agricultural Water Management xxx (2010) xxxxxx Gillabel, J., Denef, K., Brenner, J., Merckx, R., Paustian, K., 2007. Carbon sequestration and soil aggregation in center-pivot irrigated and dryland cultivated farming systems. Soil Science Society of America Journal 71, 10201028. Huppe, H.C., Turpin, D.H., 1994. Integration of carbon and nitrogen metabolism in plant and algal cells. Annual Review of Plant Physiology and Plant Molecular Biology 45, 577607. Kirda, C., Topcu, S., Cetin, M., Dasgan, H.Y., Kaman, H., Topaloglu, F., Derici, M.R., Ekici, B., 2007. Prospects of partial root zone irrigation for increasing irrigation water use efciency of major crops in the Mediterranean region. Annals of Applied Biology 150, 281291. Kuzyakov, Y., Domanski, G., 2000. Carbon input by plants into the soil. Review. Journal of Plant Nutrition and Soil Science 163, 421431. Lal, R., 2004. Soil carbon sequestration to mitigate climate change. Geoderma 123, 122. Lal, R., 2008. Soil carbon stocks under present and future climate with specic reference to European ecoregions. Nutrient Cycling in Agroecosystems 81, 113127. Lal, R., Stewart, B.A., 2010. Soil Quality and Biofuel Production. Advances in Soil Science. CRC Press, Boca Raton, FL, p. 109. Lal, R., Kimble, J., Follett, R., Cole, C., 1998. The Potential of US Cropland to Sequester C and Mitigate the Greenhouse Effect. Ann Arbor Science Publishers, Chelsea, MI. Liu, F.L., Shahnazari, A., Andersen, M.N., Jacobsen, S.E., Jensen, C.R., 2006. Physiological responses of potato (Solanum tuberosum L.) to partial root-zone drying: ABA signaling, leaf gas exchange, and water use efciency. Journal of Experimental Botany 57, 37273735. Magid, J., Kjrgaard, C., Gorissen, A., Kuikman, P.J., 1999. Drying and rewetting of a loamy sand soil did not increase the turnover of native organic matter, but retarded the decomposition of added 14C-labelled plant material. Soil Biology and Biochemistry 31, 595602. Parkin, T.B., Kaspar, T.C., 2003. Temperature controls on diurnal carbon dioxide ux: implications for estimating soil carbon loss. Soil Science Society of America Journal 67, 17631772. Shahnazari, A., Ahmadi, S.H., Laerke, P.E., Liu, F.L., Plauborg, F., Jacobsen, S.E., Jensen, C.R., Andersen, M.N., 2008. Nitrogen dynamics in the soilplant system under decit and partial root-zone drying irrigation strategies in potatoes. European Journal of Agronomy 28, 6573. Soil Survey Staff, 1992. Keys to Soil Taxonomy, 5th ed. Pocahontas Press, Blacksburg, VA, USA. van Gestel, M., Merckx, R., Vlassak, K., 1993. Microbial biomass responses to soil drying and rewetting: the fate of fast- and slow-growing microorganisms in soils from different climates. Soil Biology and Biochemistry 25, 109123. Wang, H., Liu, F., Andersen, M.N., Jensen, C.R., 2009. Comparative effects of partial root-zone drying and decit irrigation on nitrogen uptake in potatoes (Solanum tuberosum L.). Irrigation Science 27, 443448. Wang, Y., Liu, F., Andersen, M.N., Jensen, C.R., 2010. Improved plant nitrogen nutrition contributes to higher water use efciency in tomatoes under alternate partial root-zone irrigation. Functional Plant Biology 37, 175182. Xiang, S.R., Doyle, A., Holden, P.A., Schimel, J.P., 2008. Drying and rewetting effects on C and N mineralization and microbial activity in surface and subsurface California grassland soils. Soil Biology and Biochemistry 40, 22812289. Zegbe, J.A., Behboudian, M.H., Clother, B.E., 2004. Partial rootzone drying is a feasible option for irrigating processing tomatoes. Agricultural Water Management 68, 195206. Zegbe, J.A., Behboudian, M.H., Clothier, B.E., 2006. Responses of Petopride processing tomato to partial rootzone drying at different phenological stages. Irrigation Science 24, 203210.

(Kuzyakov and Domanski, 2000). Consequently, the concentration and the amount of total C increased considerably after 15 DAT and 13 DAT in experiments I and II, respectively, and was the highest as compared with the FI and PRI soils at the end of the experiments (Figs. 4 and 6). Conclusively, our results obtained in pot experiment showed that both PRI and DI irrigation treatments may lead to a reduced C retention in the soilplant system as compared with the FI treatment, although the reduction was less signicant especially under conditions of organic N input with low chemical N fertilizer. PRI resulted in signicantly higher C concentration in the plant biomass than did the FI and DI treatments; this may attribute to the improved crop quality as reported in the literature in the PRI-treated plants (Zegbe et al., 2004, 2006). However, it is important that more investigations are necessary to further illustrate the effects of different irrigation strategies on C retention in the soilplant system under eld conditions. Acknowledgements This study was partly supported by the European Commission FP7 project: Safe and High Quality Food Production using Low Quality Waters and Improved Irrigation Systems and Management (SAFIR) (FOOD-CT-2005-023168). Technical assistant by Jens Bertelsen is highly acknowledged. References
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