Plant Water Relations, Plant Stress and Plant Production

  • Abraham Blum


Plant water deficit is initiated as the crop demand for water exceeds the supply. The capacity of plants to meet the demand and thus avoid water deficit depends on their “hydraulic machinery.” This machinery involves firstly the reduction of net radiation by canopy albedo, thus reflecting part of the energy load on the plant. Secondly, it determines the ability to transport sufficient amount of water from the soil to the atmosphere via the stomata (which take in CO2) in order to provide for transpiration, transpirational cooling and carbon assimilation. Water is transported by way the SPAC (soil-plant-atmosphere continuum). SPAC is largely controlled by the resistances in the continuum as determined by root, stem, leaf, stomata and cuticular hydraulic resistances. Resistances are generally a function of the plant basic anatomy, development and metabolism. Some resistance such as those of stomata is also variable depending on plant responses and environment effects.

The primary force driving water against plant resistances is the soil-to leaf gradient of water potential which is expressed in reduced leaf water potential. Reduced leaf water potential may induce osmotic adjustment which helps maintain leaf hydration at low leaf water potential. As plants enter a state of water deficit, hormones, mainly ABA are produced in the root and the shoot, causing an array of responses, most of which cannot be defined as productive in the agronomic sense. Thus, the combination of hydraulic stress and hormonal metabolism carry various impacts on plant adaptation to stress on one hand and reductions in growth and productivity on the other. The most susceptible growth stage to water deficit is flowering and reproduction, which in many crop species cannot be recovered upon rehydration. Some (not all) of the heritable plant traits and adaptive responses to water deficit can be counterproductive in term of allowing high yield potential.


Drought Stress Arbuscular Mycorrhizal Water Deficit Guard Cell Leaf Water Potential 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.


  1. Ahmadi A, Baker DA (2001) The effect of water stress on grain filling processes in wheat. J Agric Sci 136:257–269CrossRefGoogle Scholar
  2. Alexandersson E, Danielson JÅH, Råde J et al (2010) Transcriptional regulation of aquaporins in accessions of Arabidopsis in response to drought stress. Plant J 61:650–660PubMedCrossRefGoogle Scholar
  3. Aneja M, Gianfagna T, Ng E (2004) The roles of abscisic acid and ethylene in the abscission and senescence of cocoa flowers. J Plant Growth Regul 27:149–155Google Scholar
  4. Araus JL, Febrero A, Vendrell P (1991) Epidermal conductance in different parts of durum wheat grown under Mediterranean conditions – the role of epicuticular waxes and stomata. Plant Cell Environ 14:545–558CrossRefGoogle Scholar
  5. Araus JL, Slafer GA, Reynolds MP et al (2002) Plant breeding and drought in C3 cereals: what should we breed for? Ann Bot 89:925–940PubMedCrossRefGoogle Scholar
  6. Aroca R, Porcel R, Ruiz-Lozano JM (2007) How does arbuscular mycorrhizal symbiosis regulate root hydraulic properties and plasma membrane aquaporins in Phaseolus vulgaris under drought, cold or salinity stresses? New Phytol 173:808–816PubMedCrossRefGoogle Scholar
  7. Aroca R, Vernieri P, Ruiz-Lozano JM (2008) Mycorrhizal and non-mycorrhizal Lactuca sativa plants exhibit contrasting responses to exogenous ABA during drought stress and recovery. J Exp Bot 59:2029–2041PubMedCrossRefGoogle Scholar
  8. Asseng S, Ritchie JT, Smucker AJM et al (1998) Root growth and water uptake during water deficit and recovering in wheat. Plant Soil 201:265–273CrossRefGoogle Scholar
  9. Auge RM, Kubikova E, Moore JL (2001) Foliar dehydration tolerance of mycorrhizal cowpea, soybean and bush bean. New Phytol 151:535–541CrossRefGoogle Scholar
  10. Austin RB, Bingham J, Blackwell RD et al (1980) Genetic improvement in winter wheat yields since 1900 and associated physiological changes. J Agric Sci 94:675–689CrossRefGoogle Scholar
  11. Barker DJ, Sullivan CY, Moser LE (1993) Water deficit effects on osmotic potential, cell wall elasticity, and proline in five forage grasses. Agron J 85:270–275CrossRefGoogle Scholar
  12. Bauerle TL, Richards JH, Smart DR et al (2008) Importance of internal hydraulic redistribution for prolonging the lifespan of roots in dry soil. Plant Cell Environ 31:177–186PubMedGoogle Scholar
  13. Beis A, Patakas A (2010) Differences in stomatal responses and root to shoot signalling between two grapevine varieties subjected to drought. Funct Plant Biol 37:139–146CrossRefGoogle Scholar
  14. Blum A (1970) Nature of heterosis in grain production by the sorghum panicle. Crop Sci 10:28–31CrossRefGoogle Scholar
  15. Blum A (1975) Effect of the BM gene on epicuticular wax deposition and the spectral characteristics of sorghum leaves. SABRAO J 7:45–52Google Scholar
  16. Blum A (1977) The basis of heterosis in the differentiating sorghum panicle. Crop Sci 17:880–882CrossRefGoogle Scholar
  17. Blum A (2004) Sorghum physiology. In: Nguyen HT, Blum A (eds) Physiology and biotechnology integration for plant breeding. CRC Press, Boca RatonGoogle Scholar
  18. Blum A (2009) Effective use of water (EUW) and not water-use efficiency (WUE) is the target of crop yield improvement under drought stress. Field Crops Res 112:119–123CrossRefGoogle Scholar
  19. Blum A, Arkin GF (1984) Sorghum root growth and water-use as affected by water supply and growth duration. Field Crops Res 9:131–142CrossRefGoogle Scholar
  20. Blum A, Ritchie JT (1984) Effect of soil surface water content on sorghum root distribution in the soil. Field Crops Res 8:169–176CrossRefGoogle Scholar
  21. Blum A, Sinmena B (1995) Isolation and characterization of variant wheat cultivars for ABA sensitivity. Plant Cell Environ 18:77–83CrossRefGoogle Scholar
  22. Blum A, Mayer J, Golan G (1988) The effect of grain number (sink size) on source activity and its water-relations in wheat. J Exp Bot 39:106–114CrossRefGoogle Scholar
  23. Blum A, Munns R, Passioura JB et al (1996) Genetically engineered plants resistant to soil drying and salt stress: how to interpret osmotic relations? Plant Physiol 110:1051PubMedGoogle Scholar
  24. Boomsma CR, Vyn TJ (2008) Maize drought tolerance: potential improvements through arbuscular mycorrhizal symbiosis? Field Crops Res 108:14–31CrossRefGoogle Scholar
  25. Borras L, Westgate M, Otegui ME (2003) Control of grain weight and grain water relations by post-flowering source-sink ratio in maize. Ann Bot 91:857–867PubMedCrossRefGoogle Scholar
  26. Boyer JS, Westgate ME (2004) Grain yields with limited water. J Exp Bot 55:2385–2394PubMedCrossRefGoogle Scholar
  27. Bramley H, Turner DW, Tyerman SD et al (2007) Water flow in the roots of crop species: the influence of root structure, aquaporin activity, and waterlogging. Adv Agron 96:33–196Google Scholar
  28. Brodrib TJ, Feild TS, Sack L (2010) Viewing leaf structure and evolution from a hydraulic perspective. Funct Plant Biol 37:488–498CrossRefGoogle Scholar
  29. Burow GB, Franks CD, Xin Z (2008) Genetic and physiological analysis of an irradiated bloomless mutant (epicuticular wax mutant) of sorghum. Crop Sci 48:41–48CrossRefGoogle Scholar
  30. Caird MA, Richards JH, Hsiao TC (2007) Significant transpirational water loss occurs throughout the night in field-grown tomato. Funct Plant Biol 34:172–177CrossRefGoogle Scholar
  31. Cameron KD, Teece MA, Smart LB (2006) Increased accumulation of cuticular wax and expression of lipid transfer protein in response to periodic drying events in leaves of tree tobacco. Plant Physiol 140:176–183PubMedCrossRefGoogle Scholar
  32. Chazen O, Neumann PM (1994) Hydraulic signals from the roots and rapid cell-wall hardening in growing maize (Zea mays l) leaves are primary responses to polyethylene glycol-induced water deficits. Plant Physiol 104:1385–1392PubMedGoogle Scholar
  33. Cheikh N, Jones RJ (1994) Disruption of maize kernel growth and development by heat stress – role of cytokinin abscisic acid balance. Plant Physiol 106:45–51PubMedGoogle Scholar
  34. Chimenti CA, Marcantonio M, Hall AJ (2006) Divergent selection for osmotic adjustment results in improved drought tolerance in maize (Zea mays L) in both early growth and flowering phases. Field Crops Res 95:305–315CrossRefGoogle Scholar
  35. Christmann A, Grill E (2009) Are GTGs ABA’s biggest fans? Cell 136:21–23PubMedCrossRefGoogle Scholar
  36. Christmann A, Weiler EW, Steudle E et al (2007) A hydraulic signal in root-to-shoot signalling of water shortage. Plant J 52:167–174PubMedCrossRefGoogle Scholar
  37. Cochard H, Casella E, Mencuccini M (2007) Xylem vulnerability to cavitation varies among poplar and willow clones and correlates with yield. Tree Physiol 27:1761–1767PubMedGoogle Scholar
  38. Cochard H, Holtta T, Herbette S et al (2009) New insights into the mechanisms of water-stress-induced cavitation in conifers. Plant Physiol 151:949–954PubMedCrossRefGoogle Scholar
  39. Cosgrove DJ (2000) Loosening of plant cell walls by expansions. Nature 407:321–326PubMedCrossRefGoogle Scholar
  40. Cutler JM, Rains DW, Loomis RS (1977) The importance of cell size in the water relations of plants. Physiol Plant 40:255–260CrossRefGoogle Scholar
  41. Davies FT, Olalde-Portugal V, Aguilera-Gomez L et al (2002a) Alleviation of drought stress of Chile ancho pepper (Capsicum annuum L cv San Luis) with arbuscular mycorrhiza indigenous to Mexico. Sci Hort 92:347–359CrossRefGoogle Scholar
  42. Davies WJ, Wilkinson S, Loveys B (2002b) Stomatal control by chemical signalling and the exploitation of this mechanism to increase water use efficiency in agriculture. New Phytol 153:449–460CrossRefGoogle Scholar
  43. Davies WJ, Kudoyarova G, Hartung W (2005) Long-distance ABA signaling and its relation to other signaling pathways in the detection of soil drying and the mediation of the plant’s response to drought. J Plant Growth Regul 24:285–295CrossRefGoogle Scholar
  44. de Wit CT (1958) Transpiration and crop yields. Versl Landabouwk Onderz 64:1–88Google Scholar
  45. Dodd IC (2009) Rhizosphere manipulations to maximize ‘crop per drop’ during deficit irrigation. J Exp Bot 60:2454–2459PubMedCrossRefGoogle Scholar
  46. Fan L, Neumann PM (2004) The spatially variable inhibition by water deficit of maize root growth correlates with altered profiles of proton flux and cell wall pH. Plant Physiol 135: 2291–2300PubMedCrossRefGoogle Scholar
  47. Fan L, Linker R, Gepstein S et al (2006) Progressive inhibition by water deficit of cell wall extensibility and growth along the elongation zone of maize roots is related to increased lignin metabolism and progressive stelar accumulation of wall phenolics. Plant Physiol 140:603–612PubMedCrossRefGoogle Scholar
  48. Fang X, Turner NC, Yan G et al (2010) Flower numbers, pod production, pollen viability, and pistil function are reduced and flower and pod abortion increased in chickpea (Cicer arietinum L) under terminal drought. J Exp Bot 61:335–345PubMedCrossRefGoogle Scholar
  49. Fletcher AL, Sinclair TR, Allen LH Jr (2007) Transpiration responses to vapor pressure deficit in well watered ‘slow-wilting’ and commercial soybean. Environ Exp Bot 61:145–151CrossRefGoogle Scholar
  50. Frascaroli E, Tuberosa R (1993) Effect of abscisic acid on pollen germination and tube growth of maize genotypes. Plant Breed 110:250–254CrossRefGoogle Scholar
  51. Fulai L, Christian RJ, Mathias NA (2004) Drought stress effect on carbohydrate concentration in soybean leaves and pods during early reproductive development: its implication in altering pod set. Field Crops Res 86:1–13CrossRefGoogle Scholar
  52. Galmés J, Pou A, Alsina MM et al (2007) Aquaporin expression in response to different water stress intensities and recovery in Richter-110 (Vitis sp): relationship with ecophysiological status. Planta 226:671–681PubMedCrossRefGoogle Scholar
  53. Garrity DP, Vidal ET, O’Toole JC (1986) Manipulating panicle transpiration resistance to increase spikelet fertility during flowering stage water stress. Crop Sci 26:789–795CrossRefGoogle Scholar
  54. Granier C, Tardieu F (1999) Water deficit and spatial pattern of leaf development Variability in responses can be simulated using a simple model of leaf development. Plant Physiol 119:609–620PubMedCrossRefGoogle Scholar
  55. Gutschick VP, Simonneau T (2002) Modelling stomatal conductance of field-grown sunflower under varying soil water content and leaf environment: comparison of three models of stomatal response to leaf environment and coupling with an abscisic acid-based model of stomatal response to soil drying. Plant Cell Environ 25:1423–1434CrossRefGoogle Scholar
  56. Harrison MA, Kaufman PB (1980) Hormonal regulation of lateral bud (tiller) release in oats (Avena sativa L). Plant Physiol 66:1123–1127PubMedCrossRefGoogle Scholar
  57. Havlová M, Dobrev PI, Motyka V et al (2008) The role of cytokinins in responses to water deficit in tobacco plants over-expressing trans-zeatin O-glucosyltransferase gene under 35S or SAG12 promoters. Plant Cell Environ 31:341–353PubMedCrossRefGoogle Scholar
  58. Henson IE, Mahalakshmi V (1985) Evidence for panicle control of stomatal behaviour in water-stressed plants of pearl millet. Field Crops Res 11:281–290CrossRefGoogle Scholar
  59. Holmes MG, Keiller DR (2002) Effects of pubescence and waxes on the reflectance of leaves in the ultraviolet and photosynthetic wavebands: a comparison of a range of species. Plant Cell Environ 25:85–93CrossRefGoogle Scholar
  60. Hooker TS, Millar AA, Kunst L (2002) Significance of the expression of the CER6 condensing enzyme for cuticular wax production in Arabidopsis. Plant Physiol 129:1568–1580PubMedCrossRefGoogle Scholar
  61. Hose E, Steudle E, Hartung W (2000) Abscisic acid and hydraulic conductivity of maize roots: a study using cell- and root-pressure probes. Planta 211:874–882PubMedCrossRefGoogle Scholar
  62. Hsiao TC, Xu LK (2000) Sensitivity of growth of roots versus leaves to water stress: biophysical analysis and relation to water transport. J Exp Bot 51:1595–1616PubMedCrossRefGoogle Scholar
  63. Huang D, Wu W, Abrams SR et al (2008) The relationship of drought-related gene expression in Arabidopsis thaliana to hormonal and environmental factors. J Exp Bot 59:2991–3007PubMedCrossRefGoogle Scholar
  64. Jacqmard A, Houssa C, Bernier G (1995) Abscisic acid antagonizes the effect of cytokinin on DNA-replication origins. J Exp Bot 46:663–666CrossRefGoogle Scholar
  65. Javot H, Maurel C (2002) The role of aquaporins in root water uptake. Ann Bot 90:301–313PubMedCrossRefGoogle Scholar
  66. Jeschke WD, Hartung W (2000) Root-shoot interactions in mineral nutrition. Plant Soil 226:57–69CrossRefGoogle Scholar
  67. Ji X, Shiran B, Wan J et al (2010) Importance of pre-anthesis anther sink strength for maintenance of grain number during reproductive stage water stress in wheat. Plant Cell Environ 33:926–942PubMedCrossRefGoogle Scholar
  68. Kaldenhoff R, Ribas-Carbo M, Flexas J et al (2008) Aquaporins and plant water balance. Plant Cell Environ 31:658–666PubMedCrossRefGoogle Scholar
  69. Khanna-Chopra R, Sinha SK (1988) Enhancement of drought-induced senescence by the reproductive sink in fertile lines of wheat and sorghum. Ann Bot 61:649–653Google Scholar
  70. Kholova J, Hash CT, Lava Kumar P et al (2010) Terminal drought-tolerant pearl millet [Pennisetum glaucum (L.) R. Br.] have high leaf ABA and limit transpiration at high vapour pressure deficit. J Exp Bot 61:1431–1440PubMedCrossRefGoogle Scholar
  71. Koch K (1996) Carbohydrate-modulated gene expression in plants. Annu Rev Plant Physiol Plant Mol Biol 47:509–540PubMedCrossRefGoogle Scholar
  72. Kosma DK, Bourdenx B, Bernard A et al (2009) The impact of water deficiency on leaf cuticle lipids of Arabidopsis. Plant Physiol 151:1918–1929PubMedCrossRefGoogle Scholar
  73. Landi P, Sanguineti MC, Conti S et al (2001) Direct and correlated responses to divergent selection for leaf abscisic acid concentration in two maize populations. Crop Sci 41:335–344CrossRefGoogle Scholar
  74. Landi P, Sanguineti MC, Liu C et al (2007) Root-ABA1 QTL affects root lodging, grain yield, and other agronomic traits in maize grown under well-watered and water-stressed conditions. J Exp Bot 58:319–326PubMedCrossRefGoogle Scholar
  75. Laplaze L, Benkova E, Casimiro I et al (2008) Cytokinins act directly on lateral root founder cells to inhibit root initiation. Plant Cell 19:3889–3900CrossRefGoogle Scholar
  76. Le Bris M, Michaux-Ferrière N, Jacob Y et al (1999) Regulation of bud dormancy by manipulation of ABA in isolated buds of Rosa hybrida cultured in vitro. Aust J Plant Physiol 26:273–281CrossRefGoogle Scholar
  77. Li Y, Wang G-X, Xin M et al (2004) The parameters of guard cell calcium oscillation encodes stomatal oscillation and closure in Vicia faba. Plant Sci 166:415–421CrossRefGoogle Scholar
  78. Li Y, Sperry JS, Shao M (2009) Hydraulic conductance and vulnerability to cavitation in corn (Zea mays L) hybrids of differing drought resistance. Environ Exp Bot 66:341–346CrossRefGoogle Scholar
  79. Liu F, Andersen MN, Jensen CR (2003) Loss of pod set caused by drought stress is associated with water status and ABA content of reproductive structures in soybean. Funct Plant Biol 30:271–280CrossRefGoogle Scholar
  80. Liu F, Jensen CR, Andersen MN (2005) A review of drought adaptation in crop plants: changes in vegetative and reproductive physiology induced by ABA-based chemical signals. Aust J Agr Res 56:1245–1252CrossRefGoogle Scholar
  81. Lockhart JA (1965) An analysis of irreversible plant cell elongation. J Theor Biol 8:264–276PubMedCrossRefGoogle Scholar
  82. Lopez G, Behboudian MH, Vallverdu X et al (2010) Mitigation of severe water stress by fruit thinning in ‘O’Henry’ peach: implications for fruit quality. Sci Hort 125:294–300CrossRefGoogle Scholar
  83. Lu ZJ, Neumann PM (1998) Water-stressed maize, barley and rice seedlings show species diversity in mechanisms of leaf growth inhibition. J Exp Bot 49:1945–1952CrossRefGoogle Scholar
  84. Luan S (2002) Signalling drought in guard cells. Plant Cell Environ 25:229–237PubMedCrossRefGoogle Scholar
  85. Mambelli S, Setter TL (1998) Inhibition of maize endosperm cell division and endoreduplication by exogenously applied abscisic acid. Physiol Plant 104:266–272CrossRefGoogle Scholar
  86. Marshall JG, Dumbroff EB (1999) Turgor regulation via cell wall adjustment in white spruce. Plant Physiol 119:313–320PubMedCrossRefGoogle Scholar
  87. McMichael BL, Lascano RJ (2010) Evaluation of hydraulic lift in cotton (Gossypium hirsutum L) germplasm. Environ Exp Bot 68:26–30CrossRefGoogle Scholar
  88. Mills D, Genfa Z, Benzioni A (2001) Effect of different salts and of ABA on growth and mineral uptake in jojoba shoots grown in vitro. J Plant Physiol 158:1031–1039CrossRefGoogle Scholar
  89. Miyamoto N, Steudle E, Hirasawa T et al (2001) Hydraulic conductivity of rice roots. J Exp Bot 52:1835–1846PubMedCrossRefGoogle Scholar
  90. Miyazawa S-I, Yoshimura S, Shinzaki Y et al (2008) Deactivation of aquaporins decreases internal conductance to CO2 diffusion in tobacco leaves grown under long-term drought. Funct Plant Biol 35:553–564CrossRefGoogle Scholar
  91. Moore JP, Nguema-Ona E, Chevalier L et al (2006) Response of the leaf cell wall to desiccation in the resurrection plant Myrothamnus flabellifolius. Plant Physiol 141:651–662PubMedCrossRefGoogle Scholar
  92. Morgan JM (1980) Possible role of abscisic acid in reducing seed set in water-stressed wheat plants. Nature 285:655–657CrossRefGoogle Scholar
  93. Morgan JM (1992) Osmotic components and properties associated with genotypic differences in osmoregulation in wheat. Aust J Plant Physiol 19:67–76CrossRefGoogle Scholar
  94. Munne-Bosch S, Alegre L (2004) Die and let live: leaf senescence contributes to plant survival under drought stress. Funct Plant Biol 31:203–216CrossRefGoogle Scholar
  95. Munns R, Richards RA (2007) Recent advances in breeding wheat for drought and salt stresses. In: Jenks MA, Hasegawa PM, Mohan Jain S (eds) Advances in molecular breeding toward drought and salt tolerant crops. Springer, DordrechtGoogle Scholar
  96. Munns R, Sharp RE (1993) Involvement of abscisic acid in controlling plant growth in soils of low water potential. Aust J Plant Physiol 20:425–437CrossRefGoogle Scholar
  97. Nakashima K, Ito Y, Yamaguchi-Shinozaki K (2009) Transcriptional regulatory networks in response to abiotic stresses in Arabidopsis and grasses. Plant Physiol 149:88–95PubMedCrossRefGoogle Scholar
  98. Nar H, Saglam A, Terzi R et al (2009) Leaf rolling and photosystem II. Efficiency in Ctenanthe setosa exposed to drought stress. Photosynthetica 47:429–436CrossRefGoogle Scholar
  99. Neumann PM (1995) The role of cell wall adjustment in plant resistance to water deficits. Crop Sci 35:1258–1266CrossRefGoogle Scholar
  100. Ober ES, Sharp RE (2007) Regulation of root growth responses to water deficit. In: Jenks MA, Hasegawa PM, Jain S (eds) Advances in molecular breeding towards drought and salt tolerant crops. Springer, DordrechtGoogle Scholar
  101. Ofir M, Kigel J (1998) Abscisic acid involvement in the induction of summer-dormancy in Poa bulbosa, a grass geophytes. Physiol Plant 102:163–170CrossRefGoogle Scholar
  102. Ohkuma K, Lyon JL, Addicott FT et al (1963) Abscisin II, an abscission-accelerating substance from young cotton fruit. Science 142:1592–1593PubMedCrossRefGoogle Scholar
  103. Oliver SN, Dennis ES, Dolferus R (2007) ABA regulates apoplastic sugar transport and is a potential signal for cold-induced pollen sterility in rice. Plant Cell Physiol 48:1319–1330PubMedCrossRefGoogle Scholar
  104. Or E, Belausov E, Popilevsky I et al (2000) Changes in endogenous ABA level in relation to the dormancy cycle in grapevines grown in a hot climate. J Hort Sci Biotechnol 75:190–194Google Scholar
  105. Ortega U, Duñabeitia M, Menendez S et al (2004) Effectiveness of mycorrhizal inoculation in the nursery on growth and water relations of Pinus radiata in different water regimes. Tree Physiol 24:65–73PubMedGoogle Scholar
  106. O’Toole JC (1982) Adaptation of rice to drought prone environments. In: Drought resistance in crops with emphasis on rice. International Rice Research Institute, Los BanosGoogle Scholar
  107. Palta JA, Turner NC, French RJ et al (2007) Physiological responses of lupin genotypes to terminal drought in a Mediterranean-type environment. Ann Appl Biol 150:269–279CrossRefGoogle Scholar
  108. Parent B, Hachez C, Redondo et al (2009) Drought and abscisic acid effects on aquaporin content translate into changes in hydraulic conductivity and leaf growth rate: a trans-scale approach. Plant Physiol 149:2000–2012PubMedCrossRefGoogle Scholar
  109. Passioura JB, Fry SC (1992) Turgor and cell expansion: beyond the Lockhart equation. Aust J Plant Phys 19:565–576CrossRefGoogle Scholar
  110. Porcel R, Ruiz-Lozano JM (2004) Arbuscular mycorrhizal influence on leaf water potential, solute accumulation, and oxidative stress in soybean plants subjected to drought stress. J Exp Bot 55:1743–1750PubMedCrossRefGoogle Scholar
  111. Quarrie SA (1991) Implications of genetic differences in ABA accumulation for crop production. In: Davies WJ, Jones HG (eds) Abscisic acid: physiology and biochemistry. Bios Scientific Publishers, LondonGoogle Scholar
  112. Quintero JM, Fournier JM, Benlloch M (1999) Water transport in sunflower root systems: effects of ABA, Ca2+ status and HgCl2. J Exp Bot 50:1607–1612CrossRefGoogle Scholar
  113. Rasmussen RD, Hole D, Hess JR et al (1997) Wheat kernel dormancy and plus abscisic acid level following exposure to fluridone. J Plant Physiol 150:440–445Google Scholar
  114. Reymond M, Muller B, Leonardi A et al (2003) Combining quantitative trait loci analysis and an ecophysiological model to analyze the genetic variability of the responses of maize leaf growth to temperature and water deficit. Plant Physiol 131:664–675PubMedCrossRefGoogle Scholar
  115. Reynolds MP, Balota M, Delgado MIB et al (1994) Physiological and morphological traits associated with spring wheat yield under hot, irrigated conditions. Aust J Plant Physiol 21:717–730CrossRefGoogle Scholar
  116. Riederer M, Schreiber L (2001) Protecting against water loss: analysis of the barrier properties of plant cuticles. J Exp Bot 52:2023–2032PubMedCrossRefGoogle Scholar
  117. Rose TJ, Rengel Z, Ma Q et al (2008) Hydraulic lift by canola plants aids P and K uptake from dry topsoil. Aust J Agric Res 59:38–45CrossRefGoogle Scholar
  118. Ruiz-Lozano JM, Collados C, Barea JM et al (2001) Arbuscular mycorrhizal symbiosis can alleviate drought-induced nodule senescence in soybean plants. New Phytol 151:493–502CrossRefGoogle Scholar
  119. Sack L, Holbrook NM (2006) Leaf hydraulics. Annu Rev Plant Biol 57:361–381PubMedCrossRefGoogle Scholar
  120. Sade N, Vinocur BJ, Diber A et al (2009) Improving plant stress tolerance and yield production: is the tonoplast aquaporin SlTIP2;2 a key to isohydric to anisohydric conversion? New Phytol 181:651–661PubMedCrossRefGoogle Scholar
  121. Sanguineti MC, Conti S, Landi P et al (1996) Abscisic acid concentration in maize leaves – genetic control and response to divergent selection in two populations. Maydica 41:193–203Google Scholar
  122. Schachtman DP, Goodger JQD (2008) Chemical root to shoot signaling under drought. Trends Plant Sci 13:281–287PubMedCrossRefGoogle Scholar
  123. Schussler JR, Westgate ME (1995) Assimilate flux determines kernel set at low water potential in maize. Crop Sci 35:1074–1080CrossRefGoogle Scholar
  124. Serpe MD, Matthews MA (2000) Turgor and cell wall yielding in dicot leaf growth in response to changes in relative humidity. Aust J Plant Physiol 27:1131–1140Google Scholar
  125. Sharp RE (2002) Interaction with ethylene: changing views on the role of abscisic acid in root and shoot growth responses to water stress. Plant Cell Environ 25:211–222PubMedCrossRefGoogle Scholar
  126. Sharp RG, Davies WJ (2009) Variability among species in the apoplastic pH signalling response to drying soils. J Exp Bot 60:4363–4370PubMedCrossRefGoogle Scholar
  127. Sharp RE, LeNoble ME (2001) ABA, ethylene and the control of shoot and root growth under water stress. J Exp Bot 53:33–37CrossRefGoogle Scholar
  128. Sharp RE, Poroyko V, Hejlek LG et al (2004) Root growth maintenance during water deficits: physiology to functional genomics. J Exp Bot 55:2343–2351PubMedCrossRefGoogle Scholar
  129. Shaterian J, Georges F, Hussain A et al (2005) Root to shoot communication and abscisic acid in calreticulin (CR) gene expression and salt-stress tolerance in grafted diploid potato clones. Environ Exp Bot 53:323–332CrossRefGoogle Scholar
  130. Shearman VJ, Sylvester-Bradley R, Scott RK et al (2005) Physiological processes associated with wheat yield progress in the UK. Crop Sci 45:175–185Google Scholar
  131. Shepherd T, Wynne GD (2006) The effects of stress on plant cuticular waxes. New Phytol 171:469–499PubMedCrossRefGoogle Scholar
  132. Sherson SM, Alford HL, Forbes SM et al (2003) Roles of cell-wall invertases and monosaccharide transporters in the growth and development of Arabidopsis. J Exp Bot 54:525–531PubMedCrossRefGoogle Scholar
  133. Shinozaki K, Yamaguchi-Shinozaki K (2007) Gene networks involved in drought stress response and tolerance. J Exp Bot 58:221–227PubMedCrossRefGoogle Scholar
  134. Smeekens S (1998) Sugar regulation of gene expression in plants. Curr Opin Plant Biol 1:230–234PubMedCrossRefGoogle Scholar
  135. Sperry JS, Hacke UG, Oren R et al (2002) Water deficits and hydraulic limits to leaf water supply. Plant Cell Environ 25:251–263PubMedCrossRefGoogle Scholar
  136. Sperry JS, Stiller V, Hacke UG (2003) Xylem hydraulics and the soil plant-atmosphere continuum: opportunities and unresolved issues. Agron J 95:1362–1370CrossRefGoogle Scholar
  137. Steinbach HS, Benech-Arnold RL, Sanchez RA (1997) Hormonal regulation of dormancy in developing sorghum seeds. Plant Physiol 113:149–154PubMedGoogle Scholar
  138. Steudle E (2000) Water uptake by plant roots: an integration of views. Plant Soil 226:45–56CrossRefGoogle Scholar
  139. Steudle E, Peterson CA (1998) How does water get through roots? J Exp Bot 49:775–788CrossRefGoogle Scholar
  140. Stirzaker RJ, Passioura JB (1996) The water relations of the root-soil interface. Plant Cell Environ 19:201–208CrossRefGoogle Scholar
  141. Stratton L, Goldstein G, Meinzer FC (2000) Stem water storage capacity and efficiency of water transport: their functional significance in a Hawaiian dry forest. Plant Cell Environ 23:99–106CrossRefGoogle Scholar
  142. Tardieu F, Granier C (2000) Quantitative analysis of cell division in leaves: methods, developmental patterns and effects of environmental conditions. Plant Mol Biol 43:555–567PubMedCrossRefGoogle Scholar
  143. Thompson AJ, Andrews J, Mulholland BJ et al (2007) Overproduction of abscisic acid in tomato increases transpiration efficiency and root hydraulic conductivity and influences leaf expansion. Plant Physiol 143:1905–1917PubMedCrossRefGoogle Scholar
  144. Troughton A (1980) Production of root axes and leaf elongation in perennial ryegrass in relation to dryness of the upper soil layer. J Agric Sci Camb 95:533–538CrossRefGoogle Scholar
  145. Tsuchihira A, Hanba YT, Kato N (2010) Effect of overexpression of radish plasma membrane aquaporins on water-use efficiency, photosynthesis and growth of Eucalyptus trees. Tree Physiol 30:417–430PubMedCrossRefGoogle Scholar
  146. Vandeleur RK, Mayo G, Shelden MC et al (2009) The role of plasma membrane intrinsic protein aquaporins in water transport through roots: diurnal and drought stress responses reveal different strategies between isohydric and anisohydric cultivars of grapevine. Plant Physiol 149:445–460PubMedCrossRefGoogle Scholar
  147. Wall GW, Garcia RL, Kimball BA et al (2006) interactive effects of elevated carbon dioxide and drought on wheat. Agron J 98:354–381CrossRefGoogle Scholar
  148. Wan CG, Xu WW, Sosebee RE et al (2000) Hydraulic lift in drought-tolerant and -susceptible maize hybrids. Plant Soil 219:117–126CrossRefGoogle Scholar
  149. Wang Z, Cao W, Dai T et al (2001) Effects of exogenous hormones on floret development and grain set in wheat. Plant Growth Regul 35:225–231CrossRefGoogle Scholar
  150. Welcker C, Boussuge B, Bencivenni C et al (2007) Are source and sink strengths genetically linked in maize plants subjected to water deficit? A QTL study of the responses of leaf growth and of anthesis-silking interval to water deficit. J Exp Bot 58:339–349PubMedCrossRefGoogle Scholar
  151. Westgate ME, Passioura JB, Munns R (1996) Water status and aba content of floral organs in drought-stressed wheat. Aust J Plant Physiol 23:763–772CrossRefGoogle Scholar
  152. Whalley WR, Clark LJ, Gowing DJG et al (2006) Does soil strength play a role in wheat yield losses caused by soil drying? Plant Soil 280:279–290CrossRefGoogle Scholar
  153. White RG, Kirkegaard JA (2010) The distribution and abundance of wheat roots in a dense, structured subsoil – implications for water uptake. Plant Cell Environ 33:133–148PubMedCrossRefGoogle Scholar
  154. Wilkinson S, Davies WJ (2002) ABA-based chemical signalling: the co-ordination of responses to stress in plants. Plant Cell Environ 25:195–210PubMedCrossRefGoogle Scholar
  155. Wu YJ, Spollen WG, Sharp RE et al (1994) Root growth maintenance at low water potentials – increased activity of xyloglucan endotransglycosylase and its possible regulation by abscisic acid. Plant Physiol 106:607–615PubMedCrossRefGoogle Scholar
  156. Wu YJ, Sharp RE, Durachko DM et al (1996) Growth maintenance of the maize primary root at low water potentials involves increases in cell-wall extension properties, expansin activity, and wall susceptibility to expansins. Plant Physiol 111, 765–772PubMedGoogle Scholar
  157. Xiong Y-C, Li F-M, Zhang T et al (2007) Evolution mechanism of non-hydraulic root-to-shoot signal during the anti-drought genetic breeding of spring wheat. Environ Exp Bot 59:193–205CrossRefGoogle Scholar
  158. Xu XD, Bland WL (1993) Reverse water flow in sorghum roots. Agron J 85:384–388CrossRefGoogle Scholar
  159. Yang JC, Zhang JH, Wang ZQ et al (2001) Activities of starch hydrolytic enzymes and sucrose-phosphate synthase in the stems of rice subjected to water stress during grain filling. J Exp Bot 52:2169–2179PubMedGoogle Scholar
  160. Yang JC, Zhang JH, Wang ZQ et al (2003) Involvement of abscisic acid and cytokinins in the senescence and remobilization of carbon reserves in wheat subjected to water stress during grain filling. Plant Cell Environ 26:1621–1631CrossRefGoogle Scholar
  161. Yang JC, Zhang JH, Ye YX et al (2004) Involvement of abscisic acid and ethylene in the responses of rice grains to water stress during filling. Plant Cell Environ 27:1055–1064CrossRefGoogle Scholar
  162. Yang JC, Zhang J, Liu K et al (2006) Abscisic acid and ethylene interact in wheat grains in response to soil drying during grain filling. New Phytol 171:293–303PubMedCrossRefGoogle Scholar
  163. Zhang J-Y, Broeckling CD, Blancaflor EB et al (2005) Overexpression of WXP1, a putative Medicago truncatula AP2 domain-containing transcription factor gene, increases cuticular wax accumulation and enhances drought tolerance in transgenic alfalfa (Medicago sativa). Plant J 42:689–707PubMedCrossRefGoogle Scholar
  164. Zhang J, Jia W, Yang J et al (2006) Role of ABA in integrating plant responses to drought and salt stresses. Field Crops Res 96:111–119CrossRefGoogle Scholar
  165. Zhu J, Brownjonathan KM, Lynch P (2010) Root cortical aerenchyma improves the drought tolerance of maize (Zea mays L.). Plant Cell Environ 33:740–749PubMedGoogle Scholar
  166. Zimmermann U, Schneider H, Wegner LH et al (2004) Water ascent in tall trees: does evolution of land plants rely on a highly metastable state? New Phytol 162:575–615CrossRefGoogle Scholar

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© Springer Science+Business Media, LLC 2011

Authors and Affiliations

  1. 1.Tel AvivIsrael

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