Partial root-zone drying by International Potato Center

Adolfo Posadas, International Potato Center (CIP), Guliver Rojas, Escuela de Post Grado, UNALM, Lima, Perú Miguel Málaga, Escuela de Post Grado, UNALM, Lima, Perú Víctor Mares, International Potato Center (CIP) Roberto A. Quiroz, International Potato Center (CIP)

2008-2 Working Paper

Partial root-zone drying: An alternative irrigation management to improve the water use efficiency of potato crops

ISBN 978-92-9060-360-3 Production Systems and the Environment Division Working Paper No. 2008-2

Working Paper

3 International Potato Center • Working Paper 1

Partial root-zone drying: An alternative irrigation management to improve the water use efficiency of potato crops

The Production Systems and the Environment Division Working Paper Series comprises preliminary research results published to encourage debate and exchange of ideas. The series also includes documentation for research methods, simulation models, databases and other software. The views expressed in this series are those of the author(s) and do not necessarily reflect the official position of the International Potato Center.

Comments are invited. This series is available on the internet at www.cipotato.org

Partial root-zone drying: An alternative irrigation management to improve the water use efficiency of potato crops

International Potato Center • Working Paper 1

4 © International Potato Center (CIP), 2008 ISBN 978-92-9060-360-3 CIP publications contribute important development information to the public arena. Readers are encouraged to quote or reproduce material from them in their own publications. As copyright holder CIP requests acknowledgement, and a copy of the publication where the citation or material appears. Please send a copy to the Communication and Public Awareness Department at the address below. International Potato Center P.O.Box 1558, Lima 12, Peru cip@cgiar.org • www.cipotato.org Produced by the CIP Communication and Public Awareness Department (CPAD) Production Coordinator Cecilia Lafosse Design and Layout Elena Taipe and contributions from Graphic Arts Printed in Peru by Comercial Gráfica Sucre Press run: 200 November 2008

Table of Contents Abstract ............................................................................................................................................................................iv Introduction.................................................................................................................................................................... 1 Physiological aspects ............................................................................................................................................ 2 Partial root-zone drying ....................................................................................................................................... 2 The water factor in potato................................................................................................................................... 3 Materials and methods ............................................................................................................................................... 4 Results and discussion................................................................................................................................................. 6 In-field PRD applicability ..................................................................................................................................... 6 Production effects.................................................................................................................................................. 7 Yield and WUE......................................................................................................................................................... 9 Quality........................................................................................................................................................................ 9 Growth parameters ............................................................................................................................................. 10 Conclusions................................................................................................................................................................... 10 References ..................................................................................................................................................................... 10

iii

Abstract Drought is a severe environmental stress that limits agricultural production. Vegetable crops, including potato, have high water requirements and in most countries full or supplemental irrigation is necessary for successful vegetable production. However, water availability for agriculture is being reduced as a consequence of global climate change, environmental pollution and growing demand for other uses. Therefore, great emphasis is placed on crop management for dry conditions with the aim of increasing water use efficiency. To see how restricted irrigation systems affect water use efficiency and yield of potato, an experiment was conducted in an arid area in coastal Peru at the International Potato Center in Lima. Partial Root-Zone Drying (PRD), an innovative irrigation system in which both halves of the root system are alternately dried and well watered, was compared to conventional irrigation (CI) on an early potato cultivar (4 months) grown in furrows in randomized plots. Plants were fully and uniformly irrigated for 60 days following planting (pre-experimental period) and then treatments were applied up to the harvest time. For CI every furrow was irrigated during each watering. The PRD system consisted of alternately irrigating one of the two neighboring furrows during consecutive watering. CI and PRD were further divided into two treatments with different watering amounts, resulting in a total of four irrigation treatments, ranging from CI1 with 100% of the water typically applied to the potato crop in Lima, according to crop requirements; CI1/2 (50% of the amount of water applied to CI1); PRD1 (same amount of water as CI1/2); and PRD1/2 that received half of the water applied to PRD1. Fresh tuber yield was significantly higher for CI1 (45.1 t ha-1), followed by PRD1 (36.2 t ha-1), CI1/2 (33.9 t.ha-1) and PRD1/2 (31.0 t.ha-1). Water use efficiency (WUE) calculated for total water use (pre-experimental and experimental periods) was similar for PRD1/2 (2.6 kg DM ha-1. m-3), PRD1 (2.4 kg DMha-1. m-3), CI1 (2.3 kg DM.ha-1. m-3), and CI1/2 (2.2 kg DM.ha-1. m-3). However, WUE calculated for water used during the experimental period showed larger differences as it was higher for PRD1/2 (14.6 kg DM ha-1 m-3) followed by PRD1 (8.1 kg DM ha-1 m-3), CI1/2 (7.5 kg DM ha-1 m3

), and CI1 (4.9 kg DM ha-1 m-3). Our results suggest that the PRD irrigation system might become

an alternative in large potato-producing areas in the world, where water is limiting and where salinity might become a problem. Keywords: PRD, irrigation system, WUE, potato.

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Partial root-zone drying: An alternative irrigation management to improve the water use efficiency of potato crops INTRODUCTION Drought is a severe environmental stress that limits agricultural production in many agroecosystems worldwide. Many vegetable crops, including potato, have high water requirements and in most countries supplemental irrigation is necessary for successful vegetable production. However, in many countries water availability for agriculture is being reduced as a consequence of global climate change, environmental pollution and growing demand for other uses. Therefore, great emphasis is placed on water management for dry conditions based on plant and crop physiology, with the aim of increasing water use efficiency by major crops. Regulated deficit irrigation (RDI) and partial root-zone drying (PRD) are two irrigation methods that attempt to decrease the agricultural demand for water. PRD is an irrigation technique whereby half of the root zone is irrigated while the other half is allowed to dry out. The treatment is then cyclically reversed allowing the previously well-watered side of the root system to dry down while fully irrigating the previously dried side. The PRD technique is rather simple, requiring only the adaptation of irrigation systems to allow alternate wetting and drying of parts of the root zone (Loveys et al., 2000, Stikic et al., 2003). However, important issues such as the growth stage at which PRD should be applied to the potato crop to improve WUE without yield reductions remain to be addressed (Liu, 2006a). When PRD irrigation is applied to a crop, the normal root to shoot signaling system that operates in water-deficient soils is altered, causing the drying half of the root system to release abscisic acid (ABA) thus reducing stomatal aperture, whereas the fully hydrated roots maintain a favorable water status throughout the aboveground parts of the plant. In other words, PRD uncouples the biochemical signal in response to water stress from the hydraulic signal and physical effects of reduced water availability (Bacon, 2003). This mixed root signals causes a limited closure of stomata to restrict water vapor loss without a severe restriction of CO2 entrance. The outcome is reasonably good yields with considerable water savings and higher water use efficiency (WUE), which is of paramount importance in areas where water resources are limiting. PRD has been successfully used in fruit-producing crops such as tomatoes, grapes, oranges, olive trees, tomato, corn, cotton and others, but no extensive research has been conducted in root and tuber crops, particularly in semi-arid environments where the water resource is scarce. The results P A R T I A L

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in the former crops demonstrated that PRD has no major negative effect on the yield but improves fruit quality with a reduction of more than 50% of the consumption of water (Loveys et al., 2001).

Physiological aspects The first physiological response of plants to water deficit is stomatal closure, which slows down the fall in plant water potential. Stomatal closure results from the regulation of osmotic pressure in the guard cells, mediated by ABA released by roots in drying soil. Recent studies on progressive root drying in potatoes have shown that root-sourced ABA reduces stomatal conductance (Liu et al., 2005). This kind of communication is known as non-hydraulic or chemical signaling, which differs from hydraulic signals, which are based on changes in the xylem sap tension (Stikic et al., 2003). When the tips of young roots come into contact with dry soil, the release and high concentration of ABA in the xylem prompts stomatal closure to reduce water loss and bud growth, and prevent wilting (Zhang et al., 1989; Zhang & Outlaw, 2001; Khalil & Grace, 1993; Jia et al., 1996). Besides the reduction of water vapor loss, stomatal closure brings about a series of physiologic and metabolic adjustments that includes, among others, the decrease of photosynthesis rate and alterations in translocation and distribution of photosynthates (Hanson & Hitz, 1982; Kaiser, 1987). Waggoner in 1969 (cited by Harris, 1992) considered the possibility of manipulating stomatal opening in potato in order to increase the yield per unit volume of transpired water, although this increment would also mean a reduction in the yield per unit area. Interestingly, Xu et al., (1998) stated that ABA stimulates tuber formation in potato whereas Jackson (1999) has suggested that ABA participates in the control of tuber formation although its direct effect is not totally clear yet. The effect of partial root-zone drying in tuber formation has not yet been elucidated.

Partial root-zone drying PRD is based in the theoretical assumption that a small narrowing of stomatal opening may reduce water loss substantially with a minimum effect on CO2 uptake and photosynthesis (Jones, 1992). It is known that when the root system is exposed to dry soil, it responds by sending ABAmediated chemical signals to the leaves to close stomata and reduce the water loss (Davies & Zhang, 1991). On the other hand, plants with a good watering regime usually keep turgor and their stomata wide open in response to hydraulic signal through xylem water pressure. Therefore, it is expected that contradictory root signals brought about by PRD would cause a slight reduction of the stomatal opening that would decrease the water loss substantially with only a small effect on the photosynthesis rate, provided plant turgor is maintained by the watered 2

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fraction of the root system. If this is achieved in practice, the WUE in terms of the carbon gained per unit of water lost will be increased at a minimum cost of CO2 uptake and yield reduction. In grapevine and other fruit-crops it has been demonstrated that the PRD has no effect on yield, but it can improve quality, with a reduction of more than 50% of the consumption of water (Loveys et al., 2001). Besides the continuous chemical signaling that PRD stimulates (Stoll et al., 2000), this practice induces the growth of secondary roots, which reduces the vulnerability to drought (Zhang & Tardieu, 1996). A root system more widely distributed in the soil volume as a result of the lateral dry-wet cycle can result in an improved uptake of nutrients and water by the root system (Kang et al., 1998). Implementation of PRD irrigation requires that the watering system allow a wet and dry cycle in different areas of the root, independently if it is flood or pressurized irrigation (Loveys et al., 2001). The cycling is essential for maintaining a constant emission of signals from the root to the foliage, because a drought-primed root is not able to sustain its production of ABA for long periods of time (Davies and Hartung, 2004). The alternating frequency is determined according to the crop, soil type and environmental factors. The soil is a most important factor because texture and structure influence the water infiltration rate and high levels of salts increase the effect of water stress in plants (Kriedemann & Goodwin, 2004). Contrary to the conventional deficit irrigation techniques, in which the watering schedule depends mainly on potential evapotranspiration (ET), in PRD more emphasis is given to direct measures of the soil water content in the root area. The frequency of irrigation in the PRD system varies according to environmental conditions, but the watering volumes depend on the soil type and root depth, without adjustments for environmental conditions. For the calculation of the real ET in the PRD system, it is necessary to make an adjustment of the crop coefficient (kc), since, like in all methods of deficit irrigation, this value is not similar to that of a regime of normal watering.

The water factor in potato Potato is a crop that is very sensitive to water deficit. Even in normal conditions of watering, water stress happens during the noontime due to high transpiration rates (Harris, 1978; Kumar et al., 2003) and short periods of water stress are usually caused by inadequate watering practices. Although the high incidence of pests and diseases are partly responsible for low yields, the main restrictive factor of yield and quality is water stress. It is considered that the world yield average (20 t/ha) could be increased by approximately 50% by optimizing the water supply to the crop (Kumar et al., 2003). This sensitivity to water stress makes potato a water-demanding crop, P A R T I A L

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requiring from 400 to 600 liters of water to produce 1 kilogram of tuber dry matter (Beukema & Van der Zaag, 1979). Under field conditions, the water requirements vary between 350 to 500 mm over the growing season, depending on the crop period, environmental conditions, soil type and cultivar (Sood & Singh., 2003). Potato plants can respond with increments of up to 2 t/ha for each 2 cm of water lamina (Harris, 1978). The optimal yield is highly dependent on well-planned watering with low volume and high frequency (Vayda, 1994; Wright & Stark, 1990). The potato’s limited tolerance to drought is due to its comparatively shallow root system (50-60 cm) and the stomatal tendency to close (Harris, 1992; Kleinkopf & Westermann, 1981; and Bailey, 2000), which reduce leaf extension rates (Haverkort & MacKerron, 2000). Stomatal closure also reduces CO2 uptake and photosynthetic activity, increases leaf temperature and photorespiration, and is therefore negative for crop production (Egúsquiza, 2000). The longer the reduction of stomatal opening lasts, the higher the reduction in yield (Martínez y Huamán 1993). When the water stress is short, most of cells recover; but if it lingers, the plant withers (Beukema & Van der Zaag, 1979). Thus, PRD may reduce water stress by decreasing vegetative development. The critical period to water deficit in potato is during tuber development; achieving high yields requires an adequate water supply from tuber initiation to maturity (Salter & Goode, 1967; Jensen et al., 2000 and Egúsquiza, 2000) and even short episodes of water stress during this period can cause significant reductions in yield and quality (Miller & Martin, 1987; Kumar et al., 2003) causing chained, hollow and small tubers (Jensen et al., 2000). Deficit irrigation techniques, intended to save water, produce results that are generally nonprofitable, since the reduction in the amount of applied water in the total root zone does not compensate for the economic loss due to the reduction in yield and quality (Shock & Feibert, 2002). However, PRD represents an alternative to deficit irrigation techniques, and more experimental tests on the potato crop are needed. Therefore, the objective of this initial study was to test the effects of PRD on WUE and tuber production as compared to full irrigation, and investigate its effect on morphological and physiological characteristics of the potato crop.

MATERIALS AND METHODS A field experiment with the potato variety Única was conducted in an arid area in coastal Peru, at the International Potato Center in Lima, where the average rainfall is 23 mm y-1. Soils were sandy loams with good drainage. A complete randomized block design with five replicates was used to compare four watering treatments (See Figure 1). The distance among plants was 0.30 m and 4

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among furrows, 0.9 m. The conventional furrow irrigation scheme was used. The water was siphoned into the furrows from the irrigation canals (see Figure 2) and the amount of water per day-furrow was measured. In the PRD treatments, which consisted of alternately irrigating one of the two neighboring furrows during consecutive waterings, the furrows with irrigation were alternated each week. All plots were irrigated in the conventional way for 42 days after emergence (60 days after planting), the stage at which the corresponding treatments (experimental phase) were applied. We estimated the percentage of tubers formed during the pre-experimental phase, when all plots were equally irrigated. This estimation was used for the attribution of tuberization to each treatment and the calculation of WUE. The experiment had four irrigation treatments, as follows: CI1 was the conventional way where every furrow was irrigated during each watering cycle with 100% of the water typically applied to the crop in Lima according with crop requirements; CI1/2 was similar to CI1 but received 50% of the water lamina applied to CI1; PRD1 received the same watering lamina as CI1/2 in the alternate way described above; and PRD1/2, which received half of the water applied to PRD1 in the same alternate way. Figure 1. Experimental techniques for testing the PRD irrigation on potato. In A, by using a plastic membrane. In B, by dividing manually the root system in two joined pots. In C, using PVC siphons in field plots.

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RESULTS AND DISCUSSION In-field PRD applicability Scientists at the International Potato Center have been evaluating different procedures for applying the PRD technique on potato since 2003. Plastic membranes were placed vertically underground (figure 1A) before crop planting, so that the roots would grow in both sides of the membrane; however, results were not as expected as the membrane impeded the fertilization work and caused high temperatures that damaged the roots. In another experiment, emerged plants were transplanted into adjacent pots with half of the root system in each pot (figure 1B); however potato roots are very fragile and were damaged by manipulation. The conclusion was that the simplest way to apply and assess PRD on potato is by furrow cultivation as in normal field conditions, and the use of PVC siphons (figure 1C) to accurately control the volume of applied water on each treatment. Although the division of the root system or the applied water is not as exact as in pots, PRD in the field creates a gradient of humidity, in which the roots that are in touch with the driest area of the gradient are expected to produce the chemical signals. This assumption is supported by fieldwork with PRD in corn (Kang & Zhang, 2004). Figure 2 shows the water status of the soil in adjacent furrows during our experimental phase. The peaks in the curves indicate the watering dates. In the CI treatments the humidity of the soil was similar in all furrows as they simultaneously received the same amount of water. However, in the PRD treatments the alternate furrow irrigation caused a distinct spatial and temporal pattern of soil humidity among furrows. An important difference between PRD in the field as compared to the response under controlled conditions in which roots are divided in adjacent pots is that in pots the root half that is well irrigated maintains constant a high water status of the plant (Stikic et al., 2003; Davies et al., 2000 and Dry et al., 2000) whereas PRD in the field causes periodic symptoms of water stress in the plant. This seems to be associated to the unavoidable temporary water stress that happens in potato during sunny days as pointed out by Kumar et al., (2003), and to the likelihood that PRD brings about some degree of water stress due to the fact that the amount of water applied is determined by relative soil water content and not by actual soil and leaf water potential. Kriedemann & Goodwin (2004) pointed out that in PRD, as in any of water deficit irrigation techniques, an adjustment of the crop coefficient (Kc) should be made.

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Figure 2. Water content of the soil in adjacent furrows during the experimental phase.

Soil Water Content in Adjacent Furrows CI½

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Soil Water Content in Adjacent Furrows PRD1

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Soil Water Content in Adjacent Furrows PRD½

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Production effects Table 1 shows the results of all the response variables. The letters beside the numbers indicate the statistical differences according to the Waller-Duncan test.

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Table 1: Potato responses under CI and PRD watering regimes. VARIABLE

CI1

CI1/2

PRD1

PRD1/2

Signif.

Fresh Tuber Yield (t/ha)

45.1 a

33.9 bc

36.2 b

31.0 c

< 0.05

2.3 a

2.2 a

2.4 a

2.6 a

5.5 a

5.1 a

6.1 a

6.5 a

WUEi (kgdm/m /ha) Experimental period 3 WUEet (kgdm/m /ha) Experimental period Tuber growth (t/ha) during exp period 3 Adjusted WUEi (kgdm/m /ha) exp period % Dry Matter in tubers

4.9 8.6 38.3 4.2 22.1 b

7.5 9.1 28.8 6.4 22.5 b

8.1 11.9 30.8 6.9 22.9 ab

14.6 14.2 26.4 12.5 24.1 a

< 0.05

% CT weight

98.1 a

96.3 a

97.5 a

96.4 a

Chip color (grades del 1 al 5)

1.75

1.50

1.25

nsad

Chip Oil (%)

34 a

32 a

29 a

Number of stems per plant

1.5

1.8

1.7

nsad

Number of stolons per plant

22.0 a

22.2 a

20.4 a

22.3 a

Number of CT per plant

8.1 a

8.0 a

7.7 a

7.4 a

WUEi (kgdm/m /ha) Total growth period 3

WUEet (kgdm/m /ha) Total growth period 3

Number of NCT per plant

3.5 a

2.7 a

3.8 a

4.0 a

Leaf dried weight (g/plant)

29.6 a

22.5 b

22.6 b

22.5 b

< 0.05

Steam dried weight (g/plant)

21.3 a

18.5 a

20.9 a

18.0 a

Stolon dried weight (g/plant)

4.4 b

4.8 b

5.8 a

4.7 b

< 0.05

Root dried weight (g/plant)

0.8 a

0.6 b

0.5 b

< 0.01

Tuber dried weight (g/plant)

274.9 a

200.4 b

204.8 b

182.5 b

< 0.01

Relation A/S

0.32 a

0.27 b

0.29 ab

0.30 ab

0.66 b

0.74 a

0.70 ab

0.72 a

0.34 a

0.28 b

0.31 ab

0.32 ab

Fresh Foliage weight at harvest (kg/plant)

220.0 a

166.2 a

189.8 a

182.2 a

5.1

9.5

11.2

11.1

nsad

Root Expansion (dm )

19.8

22.8

23.2

24.8

nsad

EC soil (dS/100g)

0.67

0.34

0.62

0.32

nsad

Root concentration (dm ) 2

WUEi: Water use efficiency at irrigation technique level (ratio between total dry weight in tubers and the volumen of applied water); WUEet: Water use efficiency at crop level (ratio between total dry weight in tubers and the evapotranspiration demand); CT: commercial Tubers; NCT: Non commercial tubers; HI: harvest index (ratio between tubers dry weight and plant dry weight); PI: Production Index (ratio between stems and leaves dry weight and plant dry weight); A/S: Aerial and subterranean relation (ratio between stems and leaves dry weight and stolons, roots and tubers dry weight); EC: electric Conductivity; nsad: no statistical analysis done; ns: no significant.

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Yield and WUE Significant differences in tuber yield were found among treatments. Highest fresh tuber yield was obtained under CI1, which was significantly superior to all other treatments. However, it is interesting to see that yields under CI1/2 and PRD1/2, which received the same total amount of water (pre-experimental irrigation + experimental irrigation), did not differ. This seems to indicate that PRD (applied during the experimental phase) had no negative effect on tuber growth as yield in both cases were determined by total water applied, not by the watering regime. This is further corroborated by the fact that the treatment with the lowest yield was PRD1/2, which received the least water, both as total as well as during the experimental phase. These results are in agreement with reports that water stress slows the vegetative development and reduces tuber yield (Harris, 1992; Kumar et al., 2003; Jensen et al., 2000; Miller & Martin, 1987; Salter & Goode, 1967; Wright & Stark, 1990). However, Liu et al. (2006b) found no difference in potato tuber yield between full irrigation and PRD (70% of water applied to full irrigation from tuber initiation to maturity) in a field experiment, which suggest that PRD could be an effective strategy to improve WAE while sustaining yields provided PRD is optimized in terms of the timing of application and shifting and volume of irrigation water (Shahnazari et al., 2008) No differences in WUE were evident, either calculated on the basis of applied water (WUEi) or evapotranspired water (WUEet) when the assessment was based on total water applied. However, when the same analysis was based on water applied during the experimental phase, differences between treatments in WUEi and WUEet were evident, PRD producing increased efficiencies inversely related to water volume used.

Quality Although it is well known that water stress increases the number of small tubers (Kumar et al., 2003), in the present work, the weight percentage of CT was near 100% in all treatments with no significant differences among treatments. Also, no differences were found in the quality for industry variables tested, although the PRD1/2 treatment produced tubers with higher % DM that led to processed chips of better coloration and less absorbed oil. It is known that water stress tends to improve the quality of chips due to the higher % DM in tubers that gives chips a clearer and more uniform color (Kumar et al., 2003; Jensen et al., 2000).

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Growth parameters Most of the plant growth parameters analyzed, such as number of stems per plant, number of stolons per plant, number of commercial tubers per plant, steam dried weight were not affected by treatments. Also, no clear trend in harvest index and other variables was found. However, leaf dry weight was significantly higher for CI1, suggesting a trend to a higher aerial development in plants well supplied with water.

CONCLUSIONS As might be expected, tuber yield was related to the amount of water received by the crop. However, the ratio of relative yield attained (percentage of yield relative to maximum attained yield) to relative total water (percentage of total water applied relative to maximum total water) was higher for both PRD treatments compared to CI treatments. Since WUE was higher for PRD treatments during the experimental phase (when water supply was critically linked to tuberization), we think that the overall results have been negatively affected by the lengthy and liberal water application during the pre-experimental period. As to tuber quality, it was improved by PRD. In future field work, PRD will be applied much earlier and at lesser levels than in the present case. Also, water cost scenarios will be included in the analysis. Another issue that might be relevant to be considered is the potential effect of PRD on soil pathogens. More controlled indoors experiments already in course will assess physiological responses (ABA, leaf turgor, leaf expansion, reflectance, photosynthesis, root growth, tuberization dynamics) to PRD.

REFERENCES Ali, M., Jensen, C.R., Mogensen, V.O., Andersen, M.N. and Henson, I.E. 1999. Root signalling and osmotic adjustment during intermittent soil drying sustain grain yield of field grown wheat. Field Crops Research, 62(1):35-52. Bacon, M.A. 2003. Partial root-zone drying: A sustainable irrigation system for efficient water use without reducing fruit yield. The Lancaster Environmental Center, Lancaster University. Bahrun, A., Jensen, C.R., Asch, F. and Mogensen, V.O. 2002. Drought-induced changes in xylem pH, ionic composition, and ABA concentration act as early signals in field-grown maize (Zea mays L.). Journal of Experimental Botany, 53(367):251-263. Bailey, R.J. 2000. Practical use of soil water measurement in potato production. In: Haverkort, A.J. and MacKerron, D.K.L. Management of nitrogen and water in potato production. Wageningen, The Netherlands: Wageningen Pers., pp. 206-218.

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Beukema, H.P. and Van der Zaag, D. E. 1979. Potato improvement. Some factors and facts. International Agricultural Center. Wageningen, The Netherlands, 224 p. Davies, W.J. and Zhang, J. 1991. Roots signals and the regulation of growth and development of plant in drying soil. Annual Review of Plant Physiology and Plant Molecular Biology, 42:5-76. Davies, W.J., Bacon, M.A., Thompson, D.S., Sobeih, W. and Gonzales-Rodriguez, L. 2000. Regulation of leaf and fruit growth in plants growing in drying soil: exploitation of the plants' chemical signalling system and hydraulic architecture to increase the efficiency of water use in agriculture. Journal of Experimental Botany, 51(350):1617-1626. Davies, W.J. and Hartung, W. 2004. Has extrapolation from biochemistry to crop functioning worked to sustain plant production under water scarcity? In: Proceedings of the 4th International Crop Science Congress, Sep. 26-Oct. 1, 2004, Brisbane, Australia. Dry, P.R., Loveys, .B.R, Stoll, M., Stewart, D. and McCarthy, M. 2000. Partial root zone drying an update. The Australian Grapegrower and Winemaker 438a:35-39. Egúsquiza, B. R. 2000. La Papa: Producción, Transformación y Comercialización. Primera edición. Lima, Perú, 192 p. Fulton, J.M. 1970. Relationship of root extension to the soil moisture level required for maximum yield of potatoes, tomatoes and corn. Canadian Journal of Soil Science 50:92-94. Gu, S., Du, G., Zoldoske, D., Hakim, A., Cochran, R., Fugelsang, K., and Jorgensen, G. 2004. Effects of irrigation amount on water relations, vegetative growth, yield and fruit composition of Sauvignon blanc grapevines under partial rootzone drying and conventional irrigation in the San Joaquin Valley of California, USA. Journal of Horticultural Science and Biotechnology, 79(1):26-33. Hanson, A.D., and Hitz, W.D. 1982. Metabolic responses of mesophytes to plant water deficits. Annual Review of Plant Physiology, 33:163-203. Harris, P.M. (Ed.) 1978. The potato crop production. The Scientific Basis for Improvement. Ed. Chapman and Hall, London, 730 p. Harris, P.M. 1992. The influence of genotype and water stress on the nitrogen requirement of the potato crop. Conference: Meeting of the Section Physiology of the EAPR. Le Conquet (France). 2428 Jun 1991. Potato Research (Netherlands), 35(1):72. Haverkort, A.J. and. MacKerron, D.K.L 2000. Management of Nitrogen and Water in Potato Production. Wageningen, The Netherlands: Wageningen Pers., 353 p.

P A R T I A L

R O O T - Z O N E

D R Y I N G :

A N

A L T E R N A T I V E

C I P

â&#x20AC;˘

P R O D U C T I O N

S Y S T E M S

A N D

T H E

E N V I R O N M E N T

W O R K I N G

P A P E R

2 0 0 8 - 2

Horton, D. and Sawyer, R.L. 1985. The potato as a world food crop, with special reference to developing area. In: Li, P.H. (Ed.). Potato Physiology. Academic Press Inc. Ltd., (London), UK, pp.1-34. Hsiao, T.C. 1973. Plant responses to water stress. Annual Review of Plant Physiology, 24:519-570. Jackson, S.D. 1999. Multiple signaling pathways control tuber induction in potato. Plant Physiology, 119(1):1-8. Jensen, C.R., Jacobsen, S.-E., Andersen, M.N., NuĂąez, N., Andersen, S.D., Rasmussen, L. and Mogensen, V.O. 2000. Leaf gas exchange and water relation characteristics of field quinoa (Chenopodium quinoa Willd.) during soil drying. European Journal of Agronomy, 13(1):11-25. Jia, W., Zhang, J., and Zhang, D.P. 1996. Metabolism of xylem-delivered ABA in relation to ABA flux and concentration in leaves of maize and Commelina communis. Journal of Experimental Botany, 47(8):1085-1091. Jones, H.G. 1980. Interaction and integration of adaptive responses to water stress: the implications of an unpredictable environment. In: Turner, N.C., Kramer, P.J., (Eds.). Adaptation of plants to water and high temperature stress. New York, Wiley, pp. 353-365. Jones, H.G. 1992. Plants and microclimate: A quantitative approach to environmental plant physiology, 2nd ed., Cambridge University Press, Cambridge, U.K., 428 p. Kaiser, W. 1987. Effects of water deficit on photosynthetic capacity. Physiologia Plantarum, 71(1):142-149. Kang, S., Liang, Z., Hu, W. and Zhang, J. 1998. Water use efficiency of controlled alternate irrigation on root-divided maize plant. Agricultural Water Management, 38(1): 69-76. Kang, S., and Zhang, J. 2004. Controlled alternate partial root-zone irrigation: Its physiological consequences and impact on water use efficiency. Journal of Experimental Botany, 55(407): 24372446. Khalil, A.A.M. and Grace, J. 1993. Does xylem sap ABA control the stomatal behavior of water stressed sycamore (Acer pseudoplatanus L.) seedlings? Journal of Experimental Botany, 44(7):1127-1134. Kleinkopf, G.E. and Westermann, D.T. 1981. Predicting nitrogen requirements for optimum potato growth. Proc. Univ. Idaho Winter Commodity School, pp. 81-84.

P A R T I A L

R O O T - Z O N E

D R Y I N G :

A N

A L T E R N A T I V E

C I P

•

P R O D U C T I O N

S Y S T E M S

A N D

T H E

E N V I R O N M E N T

W O R K I N G

P A P E R

2 0 0 8 - 2

Kriedemann, P.E. and Goodwin, I. 2004. Regulated deficit irrigation and partial rootzone drying. An information package on two irrigation methods for high-input horticulture. Irrigation Insights No. 4. Land and Water Australia Pub., 102 p. Kumar, D., Minhas, J.S. and Singh, B. 2003. Abiotic stress and potato production. In: Khurana, S.M.P.; Minhas; J.S., Pandey, S.K. (Eds.). The Potato: Production and utilization in sub-tropics. Mehta Publishers, New Delhi (India), pp. 314-322. Lesczynski, D.B, and Tanner, C.B. 1976. Seasonal variation of root specific factors controlling the potato crop’s root growth distribution of irrigated, field-grown Russet Burbank potato. American Potato Journal, 53:69-78. Liu, F., Jensen, C.R., Shahnazari, A., Andersen, M.N. and Jacobsen, S.-E. 2005. ABA regulated stomatal control and photosynthetic water use efficiency of potato (Solanum tuberosum L.) during progressive soil drying. Plant Science, 168(3):831-836. Liu, F., Shahnazari, A., Andersen, M.N., Jacobsen, S.-E. and Jensen, C.R. 2006a. Effects of deficit irrigation (DI) and partial root-zone drying (PRD) on gas exchange, biomass partitioning, and water use efficiency in potato. Scientia Horticulturae, 109(2):113-117. Liu, F., Shahnazari, A., Andersen, M.N., Jacobsen, S.-E. and Jensen, C.R. 2006b. Physiological responses of potato (Solanum tuberosum L.) to partial root-zone drying: ABA signaling, leaf gas exchange, and water use efficiency. Journal of Experimental Botany, 57(14):3727-3735. Loveys, B., Grant, J., Dry, P. and McCarthy, M. 2001. Progress in the development of partial rootzone drying. CSIRO, Plant Industry, University of Adelaide, SARDI. Loveys, B.R., Stoll, M., Dry, P.R. and McCarthy, M.G. 2000. Using plant physiology to improve the water use efficiency of horticultural crops. Acta Horticulturae, 537:187-197. Loveys, B.R., Stoll, M., Dry, P.R. and McCarthy, M.G. 1998. Partial rootzone drying stimulates stress responses in grapevine to improve water use efficiency while maintaining crop yield and quality. Australian Grapegrower and Winemaker 414a:108-113. Martínez y Huamán, C.A. 1993. Expresiones Metabólicas de resistencia a la sequía en dos clones de papa sometidos a estrés hídrico. Tesis para optar el Grado de Magister Scientiae. Universidad Nacional Agraria La Molina (UNALM), Lima, Perú. Miller, D.E. and Martin, M.W. 1987. The effect of irrigation regime and subsoiling on yield and quality of three potato cultivars. American Potato Journal, 64(3):109-117.

P A R T I A L

R O O T - Z O N E

D R Y I N G :

A N

A L T E R N A T I V E

C I P

â&#x20AC;˘

P R O D U C T I O N

S Y S T E M S

A N D

T H E

E N V I R O N M E N T

W O R K I N G

P A P E R

2 0 0 8 - 2

Salter, P.J. and Goode, J.E. 1967. Crop responses to water at different stages of growth. Commonwealth Agricultural Bureaux, Farnham Royal, Bucks, England, 246 p. Shahnazari, A., Andersen, M.N., Liu, F., Jacobsen, S.-E. and Jensen, C.R. 2008. Partial root zone drying (PRD) sustains yield of potato (Solanum tuberosum L.) at reduced water supply. Acta Horticulturae, 792:581-586. Shock, C.C. and Feibert, E.B.G. 2002. Deficit irrigation of potato. In: Moutonnet, P. (Ed.). Deficit Irrigation Practices. Food and Agriculture Organization of the United Nations, Rome. Water Reports, 22:47-55. Sood, M.C. and Singh, N. 2003. Water Management. In: Khurana, S.M.P., Minhas; J.S., Pandey, S.K. (Eds.). The Potato: Production and utilization in sub-tropics. Mehta Publishers. New Delhi, India, pp. 111-120. Stikic, R., Popovic, S., Srdic, M., Savic, D., Jovanovic, Z., Prokic, L.J., and Zdravkovic, J. 2003. Partial root-zone drying (PRD): a new technique for growing plants that saves water and improves the quality of fruit. Bulgarian Journal of Plant Physiology, Special Issue:164-171. Stoll, M., Loveys, B., and Dry, P. 2000. Hormonal changes induced by partial rootzone drying of irrigated grapevine. Journal of Experimental Botany 51(350):1627-1634. Vayda, M.E. 1994. Environmental stress and its impact on potato yield. Wallingford (UK). Centre for Agriculture and Biosciences International (CABI), pp. 239-261. Wright, J.L. and Stark, J.C. 1990. Potato. American Society of Agronomy (ASA). Crop Science Society of America (ASA). Soil Science Society of America (SSSA). Wisconsin (USA), pp. 859-888. Xu, X., van Lammeren, A.A.M., Vermeer, E., and Vreugdenhil, D. 1998. The Role of Gibberellin, Abscisic Acid, and Sucrose in the Regulation of Potato Tuber Formation in Vitro. Plant Physiology. 117(2): 575-584. Zhang, S.Q. and Outlaw, W.H. Jr 2001. Abscisic acid introduced into the transpiration stream accumulates in the guard-cell apoplast and causes stomatal closure. Plant, Cell and Environment 24(10):1045-1054. Zhang, J. and Tardieu, F. 1996. Relative contribution of apices and mature tissues to ABA synthesis in droughted maize root system. Plant and Cell Physiology. 37(5):598-605. Zhang, J., Schurr, U. and Davies, W.J. 1987. Control of stomatal behavior by abscisic acid which apparently originates in the roots. Journal of Experimental Botany 38(7):1174-1181.

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CIP’s Mission The International Potato Center (CIP) seeks to reduce poverty and achieve food security on a sustained basis in developing countries through scientific research and related activities on potato, sweetpotato, and other root and tuber crops, and on the improved management of natural resources in potato and sweetpotato-based systems. The CIP Vision The International Potato Center (CIP) will contribute to reducing poverty and hunger; improving human health; developing resilient, sustainable rural and urban livelihood systems; and improving access to the benefits of new and appropriate knowledge and technologies. CIP will address these challenges by convening and conducting research and supporting partnerships on root and tuber crops and on natural resources management in mountain systems and other less-favored areas where CIP can contribute to the achievement of healthy and sustainable human development. www.cipotato.org CIP is supported by a group of governments, private foundations, and international and regional organizations known as the Consultative Group on International Agricultural Research (CGIAR). www.cgiar.org

International Potato Center Apartado 1558 Lima 12, Perú • Tel 51 1 349 6017 • Fax 51 1 349 5326 • email cip@cgiar.org