Hydraulic redistribution

Hydraulic redistribution refers to the mechanism by which some vascular plants redistribute soil water. It occurs in vascular plants that commonly have roots in both wet soil and extremely dry soil, especially plants with both taproots that grow vertically down to the water table, and lateral roots that sit close to the surface.

Process

During hot, dry periods, when the surface soil dries out to the extent that the lateral roots exude whatever water they contain, will result in the death of such lateral roots unless the water is replaced. Similarly, under extreme wet conditions when lateral roots are inundated by flood waters, oxygen deprivation will also lead to root peril. In plants that exhibit hydraulic redistribution, there are xylem pathways from the taproots to the laterals, such that the absence or abundance of water at the laterals creates a pressure potential analogous to that of transpirational pull. In drought conditions, ground water is drawn up through the taproot to the laterals and exuded into the surface soil, replenishing that which was lost. Under flooding conditions, plant roots perform a similar function in the opposite direction. For a visualization of this process, see "Hydraulic Redistribution Cartoon," Dawson Lab, UC Berkeley, CA

Though often referred to as hydraulic lift, movement of water by the plant roots has been shown to occur in any direction.[1][2][3] This phenomenon has been documented in over sixty plant species spanning a variety of plant types (from herb and grasses to shrubs and trees)[4][5][6] and over a range of environmental conditions (from the Kalahari Desert to the Amazon Rainforest).[4][5][7][8]

Causes

The movement of this water can be explained by the theory of water transport through a plant. This well established theory is called the cohesion-tension theory. In brief, it states that movement of water through the plant depends on having a continuous column of water, from the roots to the leaves. Water is then pulled from the roots to the leaves, through the plant system, by the difference in water potential between the boundary layers of the soil and the atmosphere. Therefore the driving force for moving water through a plant is the cohesive strength of water molecules and a pressure gradient from the roots to the leaves. This theory can still be applied when the boundary layer to the atmosphere is closed, e.g. when plant stomata are closed or in senesced plants.[9] The pressure gradient is between soil layers with different water potentials; water moves through the roots from wetter to drier soil layers in the same manner as it does when the plant is transpiring.

Significance

The ecological importance of hydraulically redistributed water is becoming better understood as this phenomenon is more carefully examined. Water redistribution by plant roots has been found influencing crop irrigation, where watering schemes leave a harsh heterogeneity in soil moisture. The plant roots have been shown to smooth or homogenize the soil moisture. This sort of smoothing out of soil moisture is important in maintaining plant root health. The redistribution of water from deep moist layers to shallow drier layers by large trees has shown to increase the moisture available in the daytime to meet the transpiration demand.

The implications of hydraulic redistribution seem to have an important influence on plant ecosystems. Whether or not plants redistribute water through the soil layers can affect plant population dynamics, such as the facilitation of neighboring species.[10] The increase in available daytime soil moisture can also offset low transpiration rates due to drought (see also drought rhizogenesis) or alleviate competition for water between competing plant species. Water redistributed to the near surface layers may also influence plant nutrient availability.[11]

Observations and Modeling

Due to the ecological significance of hydraulically redistributed water, there is an ongoing effort to continue the categorization of plants exhibiting this behaviour and adapting this physiological process into land-surface models to improve model predictions.

Traditional methods of observating hydraulic redistribution include Deuterium isotope traces,[8][5][3][12] sap flow,[4][7][13][14] and soil moisture.[5][2][15][16] In attempts to characterize the magnitude of the water redistributed, numerous models (both empirically and theoretically based) have been developed.[17]

See also

References

  1. ↑ S.S.O. Burgess, M.A. Adams, N.C. Turner, C.R. Beverly, C.K. Ong, A.A.H. Khan, and T.M. Bleby (2001), An improved heat pulse method to measure low and reverse rates of sap flow in woody plants, Tree Physiology, 21, 589-598
  2. ↑ 2.0 2.1 J.H. Richards and M.M. Caldwell (1987), Hydraulic lift: Substantial nocturnal water transport between soil layers by Artemisia tridentata roots, Oecologia, 73, 486-489
  3. ↑ 3.0 3.1 D.R. Smart, E. Carlisle, M. Goebel, and B.A. Nunez (2005), Transverse hydraulic redistribution by a grapevine, Plant, Cell and Environment, 28, 157-166
  4. ↑ 4.0 4.1 4.2 R.S. Oliveira, T.E. Dawson, S.S.O. Burgess, and D.C. Nepstad (2005), Hydraulic redistribution in three Amazonian trees, Oecologia
  5. ↑ 5.0 5.1 5.2 5.3 T.E. Dawson (1993), Hydraulic lift and water use by plants: implications for water balance, performance and plant-plant interactions, Oecologia, 95, 565-574
  6. ↑ M.M. Caldwell (1990), Water parasitism stemming from hydraulic lift: a quantitative test in the field, Israeli Journal of Botany, 39, 395-402
  7. ↑ 7.0 7.1 S.S.O. Burgess and T.M. Bleby (2006), Redistribution of soil water by lateral roots mediated by stem tissues, Journal of Experimental Botany, 57:12, 3283-3291
  8. ↑ 8.0 8.1 E.D. Schulze, M.M. Caldwell, J. Canadell, H.A. Mooney, R.B. Jackson, D. Parson, R. Scholes, O.E. Sala and P. Trimborn (1998), Downward flux of water through roots (i.e. inverse hydraulic lift) in dry Kalahari sands, Oecologia, 115, 460-462
  9. ↑ A.J. Leffler, M.S. Peek, R.J. Ryel, C.Y. Ivans, and M.M. Caldwell (2005), Hydraulic redistribution through the root systems of senesced plants, Ecology, 86, 633-642
  10. ↑ M.M. Caldwell, T.E. Dawson, and J.H. Richards (1998), Hydraulic lift: consequences of water efflux from the roots of plants, Oecologia, 113, 151-161
  11. ↑ R.J. Ryel, M.M. Caldwell, C.K. Yoder, D. Or, and A.J. Leffler (2002), Hydraulic redistribution in a stand of Artemisia tridentata: evaluation of benefits to transpiration assessed with a simulation model, Oecologia, 130, 173-184
  12. ↑ J.R. Brooks, F.C. Meinzer, R. Coulombe, and J. Gregg (2002), Hydraulic redistribution of soil water during summer drought in two contrasting Pacific Northwest coniferous forests, Tree Physiology, 20, 909-913
  13. ↑ S.S.O. Burgess, M.A. Adams, N.C. Turner, and K.C. Ong (1998), The redistribution of soil water by tree root systems, Oecologia, 115, 306-311
  14. ↑ F.C. Meinzer, S.A. James, and G. Goldstein (2004), Dynamics of transpiration, sap flow and use of stored water in tropical canopy trees, Tree Physiology, 24, 901-909
  15. ↑ H.R. da Rocha, M.L. Goulden, S.D. Miller, M.C. Menton, L.D.V.O. Pinto, H.C. de Freitas, and A.M.E.S. Figueira (2004), Seasonality of water and heat fluxes over a tropical forest in eastern Amazonia, Ecological Applications, 14(4), S22-S34
  16. ↑ T.W. Davis and X. Liang (2013), The potential use of soil moisture sensors for observing hydraulic redistribution characteristics, Journal of Water Resource and Hydraulic Engineering, 2(3), 84-91. Available online: http://www.academicpub.org/DownLoadPaper.aspx?PaperID=13821
  17. ↑ R.B. Neumann and Z.G. Cardon (2012) The magnitude of hydraulic redistribution by plant roots: a review and synthesis of empirical and modeling studies, New Phytologist, 194, 337-352. Available online: http://onlinelibrary.wiley.com/doi/10.1111/j.1469-8137.2012.04088.x/pdf

Further reading