Published online 20 November 2006
Published in Vadose Zone J 5:1264-1277 (2006)
DOI: 10.2136/vzj2006.0056
© 2006 Soil Science Society of America
677 S. Segoe Rd., Madison, WI 53711 USA
ORIGINAL RESEARCH
Root Water Extraction and Limiting Soil Hydraulic Conditions Estimated by Numerical Simulation
Quirijn de Jong van Liera,*,
Klaas Metselaarb and
Jos C. van Damb
a Exact Sciences Dep., Esalq-Univ. of São Paulo, 13418-900 Piracicaba (SP), Brazil, currently at Dep. of Environmental Sciences, Wageningen Univ., Wageningen, the Netherlands
b Dep. of Environmental Sciences, Wageningen Univ., Nieuwe Kanaal 11, 6709 PA Wageningen, the Netherlands
* Corresponding author (qdjvlier{at}esalq.usp.br)
Received 13 April 2006.
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ABSTRACT
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Root density, soil hydraulic functions, and hydraulic head gradients play an important role in the determination of transpiration-rate-limiting soil water contents. We developed an implicit numerical root water extraction model to solve the Richards equation for the modeling of radial root water extraction. The average soil water content at the moment root water potential dropped below a defined critical value was then estimated. The dependence of average water content at the onset of plant water stress on potential transpiration and root density was compared with an analytical solution for hydraulic conditions in the root sphere. The critical value was a function of potential transpiration rate, soil hydraulic properties, and root density. Matric flux potential appears to be a convenient hydraulic property to determine the onset of limiting hydraulic conditions, as numerical simulations showed that, at onset, matric flux potential vs. distance from the root surface is independent of soil type. This was also determined analytically under the constant-rate assumption. Mean water content occurs at about 0.53 times the half-distance between roots. This allows calculation of the mean limiting soil water content and pressure head from the matric flux potential at this distance, which is a function of transpiration rate and root density only. A nomogram was developed thatgiven the transpiration rate, the root density, and the soil hydraulic functionsallows determination of the values of mean water content and mean pressure head that occur at the onset of transpiration reduction.
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INTRODUCTION
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THE INTERACTION between crop transpiration rates and soil water availability is a complex issue. In an engineering approach used in irrigation scheduling and hydrologic modeling, a threshold value for water content (
) or pressure head (h) is often assumed below which transpiration rates decrease below maximum values. Theoretical considerations (Gardner, 1960; Cowan, 1965) and experiments (Carbon, 1973; Zur et al., 1982) have established that root density, soil hydraulic functions, and hydraulic head gradients play an important role in the determination of this threshold value, and that the permanent wilting point (based on soil bulk values) may also vary as a function of these variables. At present, these factors are not considered in management or hydrological models. To illustrate this, Fig. 1
shows the widely used FAO irrigation guidelines for estimating limiting water content as a function of potential transpiration rate (Doorenbos and Kassam, 1986). Although four crop groups are defined and limiting water content is a function of water content at field capacity and at the permanent wilting point, root characteristics and soil hydraulic properties are not considered in a direct way.
Modeling tools may provide insight as to how the interaction between soil and roots determines root water uptake. A great number of root water extraction models have been developed. Such models describe root water uptake based on the behavior of a single root (the microscopic approach) or describe uptake using properties of the overall root system (the macroscopic approach). Analytical solutions and numerical analyses for both types of models have been presented. Macroscopic models treat the root layer as a continuum of soil, water, and air with a sink (Gardner, 1964; Hillel et al., 1976; Hupet et al., 2002, 2003; Zuo and Zhang, 2002; Homaee et al., 2002; Dardanelli et al., 2004; Raats, 2006). Although they are often more easily applicable to specific situations than microscopic models, they lack the capacity of a process-based estimation of the limiting
or h value, which has to be estimated empirically. These models do not take into consideration the reduction of soil water content and, hence, the increase in hydraulic resistance close to water-extracting roots as confirmed experimentally by, e.g., Hainsworth and Aylmore (1986).
Microscopic models (e.g., Moldrup et al., 1992; Personne et al., 2003; Roose and Fowler, 2003; Raats, 2006) describe water extraction at an individual root level. The most common approach in these models is to consider the root as a straight cylindrical water sink, provoking a radial water flow toward it. This flow can be modeled with the Richards equation, which has been applied to solve numerous problems of unsaturated flow. A major constraint with respect to this equation is the highly nonlinear behavior of involved parameters, which limits analytical solutions to strict assumptions and boundary conditions. Analytical solutions for the microscopic approach have been presented by Gardner (1960) and Cowan (1965). A family of analytical solutions involves the quasilinear approximation (Pullan, 1990), but the advent of powerful computers now allows iterative numerical calculation techniques with less restrictive boundary conditions.
A mixed approach in which an analytical solution of the microscopic uptake profile has been combined with the overall behavior of the root system in a numerical three-dimensional model has been presented by Heinen (2001). At present, combined modeling of root architecture and flow patterns is envisaged (Darrah et al., 2006).
Explicit numerical models, in which flow processes are simulated in discrete steps in space and time, allow simulation of any kind of scenario but the resulting calculation procedures are usually very time consuming. Implicit schemes, in which a flow problem is described by a system of equations to be solved iteratively through time, may be adapted to many situations and require much less computer time to be processed. Van Dam and Feddes (2000) presented an implicit finite difference solution using the Richards equation for the simulation of infiltration, evaporation, and shallow groundwater levels. Their solution allows a flux-controlled or head-controlled surface boundary condition.
In this study, we developed an implicit numerical scheme that solves the Richards equation for radial root water extraction. The scheme uses an algorithm adapted from Van Dam and Feddes (2000). By defining a critical root water potential, the model was used to estimate the mean soil water content at the moment root water potential drops below the critical value. The dependence of this value on potential transpiration and root density was evaluated for three soils of the Dutch Staring series (Wösten et al., 2001) with different textures. An analytical solution for hydraulic conditions in the root sphere at first occurrence of water stress was used to support the conclusions. The overall objective was to derive critical parameter values for macroscopic root water extraction models from the microscopic (single-root) models.
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MATERIAL AND METHODS
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Soils and Hydraulic Properties
Root water extraction and limiting hydraulic conditions were analyzed using hydraulic data for three topsoils from the Dutch Staring series (Wösten et al., 2001) as listed in Table 1. For these soils, K
h relations can be described by the van Genuchten (1980) equation system:
 | [1] |
 | [2] |
in which
= (
r)/(
s
r);
,
r, and
s are water content, residual water content and saturated water content (m3 m3), respectively; h is pressure head (m); K and Ksat are hydraulic conductivity and saturated hydraulic conductivity, respectively (m d1); and
(m1), n, and
are empirical parameters.
A convenient soil hydraulic property that is often used in soil water movement studies (e.g., Raats, 1977; Warrick and Amoozegar-Fard, 1977; Pullan, 1990) and will be used in this study is the matric flux potential Mh0 (m2 d1), the integral of hydraulic conductivity over pressure head starting at the pressure head at permanent wilting point:
 | [3] |
Soil water content, hydraulic conductivity, and matric flux potential (integration started at h0 = 150 m) as a function of hydraulic head are shown for the three soils in Fig. 2
.

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Fig. 2. (a) Soil water content, (b) hydraulic conductivity, and (c) matric flux potential as a function of pressure head for sand, clay, and loam soils.
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Implicit Numerical Modeling
Darcy's equation combined with the mass conservation principle leads to the Richards equation, which for one-dimensional axisymmetric flow is written as
 | [4] |
where t is time (d), C is the differential water capacity (d
/dh, m1), q is the water flux density (m d1), r is the distance from the axial center (m), S is a sink or source term (d1), and H is the hydraulic head (m). To solve the Richards equation for the case of radial root water extraction, a numerical scheme based on the scheme presented by Van Dam and Feddes (2000) was used with the following modifications:- (i) the gravitational component is not considered in the root extraction model
- (ii) fluxes are adapted to the one-dimensional axisymmetric nature of root extraction, i.e., fluxes converge as they approach the root, proportional to the distance from the axial center
- (iii) there is no sink (the only water exit is the root surface located at the inner side of the first compartment)
- (iv) flux density at the outermost compartment is set to zero
- (v) flux density at the innermost compartment (the one immediately neighboring the root) is set equal to flux density entering the root, which is determined by transpiration rate and total root area
- (vi) if the pressure head in the innermost compartment drops below a critical value hlim (in this study hlim = 150 m), pressure head is fixed to this critical value and the flux density adjusted to that
Segment size (dr) was chosen smaller near the root and larger at greater distance, according to
 | [5] |
with dr,min = 108 m, dr,max = 5.104 m, S = 0.5, r0 (m) is the root radius and rm (m) is the radius of the root extraction zone, equal to the half-distance between roots (rm), which relates to the root density R (m m3) as
 | [6] |
It can also be shown that
 | [7] |
and
 | [8] |
in which L (m) is the root length, z (m) is the total rooted soil depth, Ap (m2) is the surface area and Ar (m2) is the root surface area. Combining these equations, it follows that
 | [9] |
Figure 3
shows a schematic representation of the root and axial segments considered for the simulations, including r, r0 and rm. The chosen segment size distribution resulted in 24, 73, and 228 segments for the high, medium, and low root density simulations, respectively. Further reduction of the segment size did not lead to significant differences in simulation results. In addition, intermediate K values were calculated using arithmetic or harmonic means, but no significant difference was found, confirming the chosen space step to be sufficiently small. Time steps were automatically adapted by the computer program to reach convergence of the iterative matrix solution in three to five iterations. In this way, observed time steps ranged from 1 s to 1 h.

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Fig. 3. Schematic representation of root and axial segments as used for the simulations; rm is the half-distance between roots, r0 is the root radius, dr,i is the segment size for segment i, and ri is the distance from the axial center for segment i.
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Assuming no sink or source (S = 0) and no gravitational or osmotic component (H = h), Eq. [4] reduces to
 | [10] |
Equation [10] can be discretized by explicit linearization of K and C:
 | [11] |
where i is the segment number (Segment 1 closest to the root) and j is the time step. In this equation, indexes refer to axial segments as illustrated in Fig. 3.
Analogous to Van Dam and Feddes (2000), Eq. [10] can be discretized using the finite difference scheme of Eq. [11], yielding the following equation to be solved iteratively:
 | [12] |
where p is the iteration level.
A solution to the problem can be found by applying Eq. [12] to each segment and solving the resulting equation system described by a tridiagonal matrix as follows:
 | [48] |
in which
i, ßi,
i, and fi are defined as follows:
1. Intermediate segments (i = 2 to i = n 1):
 | [13] |
 | [14] |
 | [15] |
 | [16] |
2A. Root segment (i = 1) with flux boundary condition (applied while h1 > hlim, i.e., when transpiration meets potential transpiration demand):
 | [17] |
 | [18] |
 | [19] |
in which qroot = TpAp/Ar, Ap being the surface area (m2) and Ar the root area (m2).
2B. Root segment (i = 1) with head boundary condition (applied while h1 = hlim, i.e., when transpiration is less than potential transpiration demand):
 | [20] |
 | [21] |
 | [22] |
3. Outer border segment (i = n):
 | [23] |
 | [24] |
 | [25] |
Intermediate K values (at positions i 0.5 or i + 0.5) were calculated from K values at positions i 1, i, and i + 1 obtained by Eq. [2] using the geometric mean.
Relations between Bulk Properties and Microscopic Properties
Mean water content (
mean) was calculated as the weighted average of all segment water contents:
 | [26] |
Mean pressure head (hmean), mean hydraulic conductivity (Kmean) and mean matric flux potential (Mmean) were calculated from the mean water content by Eq. [1]
to [3]:
 | [27] |
Transpiration demand, a function of the atmospheric conditions, together with soil hydraulic state variables, determines pressure head at the root surface. Its value, however, is limited to a minimum, hlim, here assumed to be equal to the commonly used pressure head at the permanent wilting point, 150 m. The instant of first occurrence of hlim at the root surface is very important, as from that moment on, water root uptake no longer meets transpiration demand. As depletion continues, relative transpiration will decrease and stop completely when the pressure head has dropped to the permanent wilting point everywhere in the soil cylinder surrounding the root. When the root potential reaches hlim for the first time, hmean as calculated by Eq. [26] and [27] is not equal to hlim but, instead, has a higher value that depends on the entire root extraction history. We will refer to the hydraulic conditions at this instant as limiting hydraulic conditions, which will be defined by the subscript mean,lim, as in hmean,lim. Thus, hmean,lim is hmean at the first occurrence of hlim at the root surface. The determination of hmean,lim and
mean,lim is a major challenge in soil physics. Simulation of flux phenomena in the root zone can help in doing so, as we will explain.
Simulation Scenarios
The main simulation scenario parameters are listed in Table 2. The variation in root densities (0.011 m m3) and transpiration rates (36 mm d1) was chosen in such a way that main crop types and climatic conditions are included (see, e.g., Stalham and Allen [2001], Pietola and Alakukku [2005], and Graveel et al. [2002] on root density, and Allen et al. [1998] on transpiration rates). Potential transpiration (Tp, mm d1) was supposed to vary daily according to a sine-wave function with its amplitude equal to its mean value Tp,mean. In this way, during a 24-h period, the transpiration rate varied between a minimum of zero and a maximum of 2Tp,mean. In the simulations, we assumed pressure head at the root surface to be variable, but not able to fall below 150 m. The value of 150 m is assumed to be independent of all other parameters.
The scenarios were set up to illustrate (i) how pressure head at the root surface, mean soil pressure head, and transpiration rate behave with time for different combinations of potential transpiration and root density; and (ii) how mean soil hydraulic conditions (pressure head, hydraulic conductivity, matric flux potential, and water content) vary according to soil type, potential transpiration rate, and root density at first occurrence of a limiting pressure head at the root surface.
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Results and Discussion
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Root Pressure Head, Mean Pressure Head, and Transpiration as a Function of Time
Simulation results are shown in Fig. 4
for the two potential transpiration rates and three root densities for the clay soil. The daily fluctuation of the pressure head in the root (hroot) to match varying transpiration can be observed in this figure, especially at low and medium R. For the low R, within a few days hroot reaches much more negative values than hmean, with a minimum every 24 h simultaneous with the maximum in the daily transpiration curve. In contrast, at high R, hroot remains almost equal to hmean from beginning to end of the simulations.

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Fig. 4. Simulated pressure head at the root surface (hroot), mean pressure head (hmean) and relative transpiration as a function of time for low and high potential transpiration rates and low, medium, and high root density in a clay soil.
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This difference between low and high R is caused by two things. In the first place, at high R, with total root area equal to 100 times the total root area at low R, flux densities toward the root need to be only 1/100 of their values at low R. Consequently, hydraulic gradients will be much smaller at high than at low R, and there will be less difference between values of pressure head at the root surface and in the surrounding soil. Second, at low R, distances between neighboring roots are on the order of 10 cm, while in a high R scenario this distance is only about 1 cm. This allows much greater differences to occur between pressure heads at the root surface and at the outer side of the root influence sphere. This effect is clearly observed when comparing Fig. 4a with 4c, but even more pronounced at higher Tp (Fig. 4d with 4f). In Fig. 5
, the same can be seen from another point of view, showing pressure head as a function of distance from the axial center at the moment of first occurrence of limiting root water potential. In this figure, it can be seen that pressure head gradients need to be much higher at the lower R. As a consequence, the difference between mean and root pressure head increases.

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Fig. 5. Pressure head as a function of distance from the axial center (hr) and mean pressure head (hmean) for low and high potential transpiration rates (Tp) for simulations with low, medium, and high root density, at the moment of first occurrence of limiting root water potential; rm is the radius of the root extraction zone or the half-distance between roots.
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At medium root density (Fig. 4b and 4e), during the first days when water content is still high, hydraulic conductivity is sufficiently high to allow low pressure head gradients, resulting in relatively small differences between hroot and hmean. At a certain moment, water content and hydraulic conductivity become small and higher gradients are necessary during the hours of the day with high transpiration. From that moment on, the daily fluctuation of hroot is similar to that at low R (Fig. 4a and 4d).
After the first occurrence of hlim, root water uptake no longer meets the potential transpiration demand, as shown by the simultaneous reduction in relative transpiration in all simulations (cf. Fig. 4). Comparing the three root densities at one transpiration level, it is clear that a higher R increases the time to first occurrence of limiting root water potential. For the case of low Tp (Fig. 4a4c) it takes 14, 31, and 34 d at low, medium, and high R, respectively; at high Tp (Fig. 4d4f) this changes to 3, 14, and 17 d, respectively. Therefore, in this soil and as far as water uptake is concerned, there is a great benefit for a crop in increasing R from low to medium, whereas the gain of an increase from medium to high R is small.
These results depend on the actual soil hydraulic functions, although the overall tendency is not expected to change: higher transpiration demand and lower root density cause greater discrepancy between hroot and hmean and, therefore, limiting hydraulic conditions are reached at higher values of hmean and
mean. Low hydraulic conductivity in the root zone water content range will cause greater discrepancy between hroot and hmean. Under these conditions, root density becomes more important for root water extraction than in soils with a higher conductivity.
Once the limiting conditions are reached, relative transpiration decreases. In the low-R scenarios, at this moment much water is still available farther away from the roots, as can also be seen in Fig. 5. During the night, transpiration decreases but established pressure head gradients maintain water movement toward the root, increasing the water content near the root. This results in a temporary relief of water stress: root pressure head becomes less negative and, during the first part of the next day, extraction can cope with potential transpiration. Due to this effect, relative transpiration decreases more slowly in the low-R scenarios. In contrast, in high-R scenarios, the exploited soil volume is almost completely depleted at the first occurrence of limiting pressure head at the root surface. As a consequence, the transpiration rate decreases much more abruptly from its maximum value to zero.
Dependence on Transpiration Rate: Diurnal Fluctation vs. Constant Rate
To analyze the dependence of hmean,lim and the corresponding
mean,lim on potential transpiration rates, a series of simulations were performed with the three soils and different root densities.
Results for the clay soil at transpiration rates from 2 to 8 mm d1 are shown by the sawtooth curves identified with A in Fig. 6
. The variable shape of these lines is caused by the fact that daily variations in transpiration are simulated: when root water potential approaches its limiting value at the beginning of a night period with low potential transpiration, the occurrence of the limiting value is postponed to the next day, consequently allowing an additional small reduction in water content and pressure head. On the other hand, if the limiting root potential is reached simultaneous with a high daytime transpiration, the first occurrence of water stress will tend to be sooner and at less negative hmean.

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Fig. 6. Mean pressure head at first occurrence of limiting hydraulic conditions (hmean,lim) as a function of potential transpiration rate (Tp) for low, medium, and high root density in a clay soil simulated with and without daily transpiration amplitude (for lines identified with "A", mean transpiration equals Tp and amplitude equals Tp; for lines identified with "B", mean transpiration equals Tp and amplitude is zero; for lines identified with "C", mean transpiration equals 2Tp and amplitude is zero).
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If daily amplitude is disregarded, simulations result in the smooth lines marked B in Fig. 6. The associated estimates of hmean,lim are less realistic for not considering the variation in daily transpiration. To avoid underestimation of hmean,lim, the most secure estimate of hmean,lim as a function of transpiration rate would be given by a line connecting the maxima in the sawtooth A lines. This can be achieved, in good approximation, using double the transpiration rate with zero amplitude, as shown by the lines marked C in Fig. 6 that refer to simulations with a constant transpiration rate twice as high as the axis labels indicate. This confirms that the local maxima in A correspond to the highest daily transpiration rates, equal to 2Tp. Subsequent simulations were performed at a constant level equal to 2Tp, corresponding to lines C. Hence, when referring to hmean,lim at transpiration rate Tp considering daily amplitude, the corresponding simulation was run at 2Tp without daily amplitude.
Selection of a Key Soil Physical Variable Controlling Transpiration Rate
Macroscopic limiting soil water conditions are often defined with respect to some spatial average of h (e.g., Feddes et al., 1988) or some spatial average of
(e.g., Doorenbos and Kassam, 1986). Equating these spatial averages with the mean values calculated using the numerical results, Fig. 7
and 8
show that these mean values are a function of transpiration rate, soil type, and root density. A function relating relative transpiration to matric head or soil water content should then include transpiration rate, soil type (soil hydraulic parameters), and root density as parameters. Comparing Fig. 8 with the FAO guidelines (Doorenbos and Kassam, 1986) for estimating limiting water content depicted in Fig. 1 shows similarity in order of magnitude between the FAO guidelines and the simulated low root density. The FAO approach, however, does not include soil physical factors.

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Fig. 7. Mean pressure head at first occurrence of limiting hydraulic conditions (hmean,lim) as a function of potential transpiration rate (Tp) for low, medium, and high root density in the sand, loam and clay soils (dotted lines refer to minimum root pressure head h = 150 m).
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Hydraulic conductivity is a key parameter in soil water movement and therefore it might be an alternative indicator of limiting hydraulic conditions. Figure 9
shows K values corresponding to hmean,lim as a function of transpiration rate for the three root densities and soils. Although values are relatively similar, especially for medium to high root densities and low transpiration rates, hydraulic conductivity does not seem to be a better indicator than h or
for limiting hydraulic conditions.

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Fig. 9. Hydraulic conductivity at first occurrence of limiting hydraulic conditions (Kmean,lim) as a function of potential transpiration rate (Tp) for low, medium, and high root density in the sand, loam and clay soils. (Kpwp is the hydraulic conductivity at the permanent wilting point for the respective soil).
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Given its definition (Eq. [3]), matric flux potential (M) integrates hydraulic conductivity and pressure head, providing another option for analysis. An analytical expression for M is not straightforward using the Mualemvan Genuchten equation system (Eq. [1] and [2]). Therefore M(h) was calculated by numerical integration of K, choosing h0 = hpwp (h at the permanent wilting point) = 150 m. For h less than 100 m, the interval was subdivided in 1.000 equal steps; for h between 100 and 30 m, this number was 10.000; between 30 and 10 m the number of steps was 30.000; and for h greater than 10 m, the number of steps was 100.000. Evaluating the value of M corresponding to hmean,lim, the result is a linear relation between M and Tp with intercept equal to zero, the slope of which is a function of root density, but independent of soil type (Fig. 10
). Based on this result, matric flux potential M is a convenient parameter to define limiting hydraulic conditions. Disregarding hysteresis, M is a strictly monotonic function of h, allowing conversion between M and h or
. Limiting hydraulic conditions can be characterized by a single M value, independent of soil type, due to the fact that M combines both factors that determine water movement, hydraulic conductivity K and pressure head difference dh. Water flux toward roots is then only defined by transpiration rate and root density.

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Fig. 10. Matric flux potential at first occurrence of limiting hydraulic conditions (Mmean,lim) as a function of potential transpiration rate (Tp) for low, medium, and high root density. Lines for all three soils are coincident.
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The ratio between Mmean,lim and Tp was determined for several half-distances between roots ranging from 5.65 to 56.5 mm, corresponding to the range between high and low root density in this study. A power function showed a good fit to the results (Fig. 11
), with (for M/Tp and rm expressed in meters):
 | [28] |
with p = 23.5 and q = 2.367 (n = 7, r2 = 0.9997).

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Fig. 11. Ratio between mean matric flux potential at first occurrence of limiting hydraulic conditions and potential transpiration rate (Mmean,lim/Tp) as a function of half-distance between roots (rm).
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A Nomogram to Estimate Soil Water State at First Occurrence of Limiting Hydraulic Conditions
For any given root density and transpiration rate, Eq. [28] can be used to calculate the corresponding Mmean,lim, which then can be converted to
mean,lim or hmean,lim by numeric means using the van GenuchtenMualem Eq. [1] and [2]. Using this result, a nomogram can be designed as shown in Fig. 12
, allowing determination of hmean,lim and
mean,lim if potential transpiration and root density are known. Dotted arrows illustrate nomogram use for a transpiration rate of 6 mm d1 and a medium root density (0.1 m m3) for the sand soil. In this case,
mean,lim corresponds to 0.17 and hmean,lim is shown to be close to 20 m, in agreement with the corresponding value from Fig. 7.

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Fig. 12. Nomogram for the determination of pressure head at first occurrence of limiting hydraulic conditions (hmean,lim) and relative water content at first occurrence of limiting hydraulic conditions ( mean,lim) for three soils from transpiration and root density data. Dotted arrow lines illustrate nomogram use for a high transpiration rate (6 mm d1) and a medium root density (0.1 m m3) for the sand soil.
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Alternatively, the nomogram can be used to determine which combination of root density and potential transpiration results in a hmean,lim of a certain magnitude. For example, choosing hmean,lim = 10 m and Tp = 6 mm d1, it can be read from the nomogram that the corresponding root density is between 0.02 and 0.05 m m3 (i.e., in the range between low and medium R) for the sand and clay soils, and between 0.002 and 0.005 m m3 (i.e., smaller than low R) for the loam soil. This is in agreement with Fig. 7 in which all represented values (except for clay and sand in combination with low R) are below 10 m. As water stress in field crops is commonly observed at values less negative than 10 m, while root densities are reported in the range of medium to high R, the values from Fig. 7 are lower than expected. Some explanations for this can be addressed. First, it is reported in literature (Tardieu et al., 1992; Amato and Ritchie, 2002) that the uneven distribution of roots can have significant influence on pressure head gradients in the neighborhood of roots and, hence, on water availability. Therefore, to transform experimentally observed root densities into mean distance between roots, information about the root system architecture should be considered. Second, partial rootsoil contact may decrease the roots' effectiveness in taking up water (Van Noordwijk et al., 1993). Third, the chosen value of hlim (150 m) may be too negative. Divergent values are reported in the literature: Adeoye and Rawlins (1981) measured 120 m in young maize (Zea mays L.) plants; Liu et al. (2005a) observed 250 m in soybean [Glycine max (L.) Merr.], while Liu et al. (2005b) found 80 m in potato (Solanum tuberosum L.). Sensitivity for this value may be an object of future research.
Comparison of Simulation Results to Analytical Solutions
The simulation results show that, at the first occurrence of limiting head at the root surface when transpiration rate is still equal to potential rate but about to drop below that, matric flux potential as a function of the space coordinate r is in good approximation independent of soil characteristics (Fig. 13
). The difference between the three soils can be expressed as the relative r-weighted root mean square difference. Comparing M for clay and loam to sand, calculated values are in the range of 0.8 to 3.0%. The weighted mean water content corresponds to the water content in the root zone at some location between 0.53rm and 0.57rm for most cases, slightly depending on soil type, transpiration rate, and root density (Table 3). Further analysis of these values shows a mean of 0.56, a standard deviation of 0.06, and a median of 0.53.

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Fig. 13. Matric flux potential (M) at first occurrence of limiting hydraulic conditions as a function of distance from axial center for low, medium, and high root density at low and high potential transpiration (lines for three soils coincide at this resolution); rm is the radius of the root extraction zone or the half-distance between roots.
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Table 3. Fraction of the half-distance between roots (rm) at which volumetric water content ( ) matches water content at first occurrence of limiting hydraulic conditions ( mean,lim) in numerical simulations.
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Analytical solutions for root water extraction for specific boundary conditions are reported in the literature, accompanied by simplifying assumptions about the boundary conditions and Kh or D
relations. For a soil with constant diffusivity D, infinite radial extent, and for large values of t, a solution to the differential Eq. [10] was presented by Philip (1957), analogous to the Carslaw and Jaeger (1946) solution for the process of thermal conduction:
 | [29] |
assuming that the average water content
mean is found at a distance rmean. In this equation,
i is the initial water content and
is the EulerMascheroni constant (
0.5772).
Defining t = tlim as the moment of first occurrence of hlim at the root surface, this equation can be rewritten to express rmean as a function of
mean at tlim:
 | [30] |
The average water content can be determined from the r-weighted integral of the
(r) profile over r according to Eq. [26] as follows:
 | [31] |
Substitution of Eq. [31] into [30] yields
 | [32] |
which, for rm >> r0 reduces to
 | [33] |
indicating that, under the assumptions made, the mean value of
corresponds to
at a distance of 0.6065rm from the axial center.
For each scenario of transpiration rate and rooting density, and while the transpiration rate matches potential transpiration rate, flux density at the root surface is independent of soil characteristics. Numerical simulation results show that this independency can be extended to the whole root sphere for the three evaluated soils (Fig. 13). These results also show that the weighted mean water content corresponds to the water content in the root zone at some location between 0.53rm and 0.57rm for most cases, slightly depending on soil type, transpiration rate, and root density (Table 3).
The simulation-derived values in Table 3 are in rough agreement with Eq. [33], except for some values for the high-root-density scenarios. The assumption of a constant diffusivity causes analytical
(r) profiles to be less depleted close to the root surface and, consequently,
mean,lim to occur at a greater distance from that surface than in the numerical simulations. For the high-root-density scenarios, it is probably not realistic to expect Eq. [33] to be valid, as one of the assumptions is an infinite extension of the root zone.
If an analytical expression for M(r) could be found, it would allow the calculation of M at the distance of mean
, providing a good estimation of
mean,lim or hmean,lim. The continuity equation for one-dimensional, axisymmetric flow (see also Eq. [4]) is
 | [34] |
From mass conservation, it follows that
 | [35] |
Ignoring the root volume (assuming rm >> r0), Eq. [35] reduces to
 | [36] |
To solve Eq. [34], an assumption has to be made about the behavior of d
/dt as a function of r. Figure 14
shows hr and
r profiles from numerical simulations with clay soil, medium root density, and a high transpiration rate. In this figure, hr profiles show a rather sharp increase of dh/dr near the root surface up to the moment of first occurrence of hlim, whereas
r profiles are much more parallel with time, suggesting that d
/dt is independent of r and constant during the period in which transpiration is equal to its potential rate. We assume that as long as and up to the moment the transpiration is (still) potential, the profiles of the soil hydraulic properties are in equilibrium with a constant uptake rate at the root surface. De Willigen and Van Noordwijk (1987) argued that, in the case of a roughly constant D, this implies that water content decreases at equal rates independent of the distance; corresponding matric flux potential profiles show a parallel downward shift with time (for studies using the same assumption, see, e.g., De Willigen and Van Noordwijk, 1987; Heinen, 2001; Raats, 2006). Therefore Eq. [36] can be generalized for any r:
 | [37] |

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Fig. 14. (a) Pressure head and (b) water content as a function of distance from axial center at several times after start of water extraction (clay soil, medium root density, high transpiration rate). Dotted lines indicate pressure head or water content profile at the moment of first occurrence of limiting hydraulic conditions at the root surface.
|
|
Equation [36] can be combined with Eq. [34] to yield the following second-order differential equation:
 | [38] |
for which the following general solution is found:
 | [39] |
where C1 and C2 are integration constants. This solution is known as the steady-rate solution. It was presented by Van Noordwijk and De Willigen (1987) and is also used in the model described by Heinen (2001).
Values for C1 and C2 can be found assuming the following boundary conditions to be valid at the moment of first occurrence of limiting root water potential:
 | [40] |
 | [41] |
Equation [40] combined with Eq. [9] yields
 | [42] |
Combining these boundary conditions with Eq. [39] yields
 | [43] |
 | [44] |
and
 | [45] |
Equation [45] shows that, under the assumption that d
/dt has the same value for all r, M(r) is indeed independent of soil characteristics, as shown in the numerical simulations (Fig. 14). This, combined with the fact that
mean,lim occurs at a fraction of the root mean distance that is not very sensitive to soil properties, allows estimation of Mmean,lim with Eq. [45] and determination of the corresponding hmean,lim and
mean,lim.
Figure 15
shows the values of hmean,lim calculated from values of M at first depletion estimated by the analytical model, compared with estimates by the numerical model for the three soils at low root density and high transpiration rate. Among the simulated combinations of root density and transpiration rate, and then especially for the sand soil, this was the only case in which a small discrepancy between numerical and analytical estimates could be observed. For all other cases, discrepancies were much smaller and lines approximately matched the 1:1 line. This confirms that the assumption resulting in Eq. [37] is reasonably well supported by the numerical simulation results.

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Fig. 15. Pressure head at the first occurrence of limiting root water potential (hmean,lim) calculated from analytical and numerical matric flux potential (Manalytical and Mnumerical, respectively) for the three soils at low root density and high transpiration rate.
|
|
Given the quality of this analytical result, it can be used to understand why the ratio M/Tp behaves like a power function of rm (Eq. [28]). The value of Mmean is found at the same distance as
mean (Eq. [27]), which we define here as a distance arm from the axial center. Evaluating Eq. [45] at arm, neglecting the r02 terms and multiplying by 2 to deal with the fact that critical hydraulic conditions correspond to double transpiration rates (as discussed above), it reduces to
 | [46] |
As a is between 0 and 1, and as generally arm >> r0, it is reasonable to neglect the a/2 term, further simplifying Eq. [46] to
 | [47] |
Evaluated for a = 0.53 and simulation parameters r0 = 0.5 mm and z = 0.5 m, values calculated by Eq. [47] for rm ranging from 15 to 75 mm fit to Eq. [28] with p = 18.1 and q = 2.277 (n = 61, r2 = 0.999) are similar to the values obtained with numerical simulation.
These results indicate that analytical and numerical methods are in good agreement. As a first approximation, the analytical solution may therefore be used. Results from the numerical method do not depend on assumptions about d
/dt, however, and are therefore preferable. The difference between the numerically estimated distance of mean water content, 0.53rm, and its analytical equivalent, 0.6065rm, is an indicator of the degree of dependency between d
/dt and r. Further investigation of this value for soils with different hydraulic properties, as well as detailed evaluation of the depletion phase after the first occurrence of limiting hydraulic conditions, will be important to provide a physically supported way to estimate soil water availability and actual transpiration rates.
CONCLUSIONS
- The limiting values (water content or pressure head) of macroscopic root water extraction models are a function of potential transpiration rate, soil hydraulic properties, and root density.
- In contrast, numerical simulations showed the matric flux potential at first occurrence of limiting hydraulic conditions (Mmean,lim) to be independent of soil type and to depend only on potential transpiration rate and root density. Analytically, it was shown that this can be explained if water depletion d
/dt has the same value within the entire root sphere. In this case, M(r) is a soil-independent function of transpiration rate and root density.
- Analytically, mean soil water content is shown to be expected to occur at a distance from the axial center 0.6065 times the half-distance between roots (rm). Numerical simulations showed that, in fact, this distance varies according to soil type, root density, and transpiration rate. For most simulations, however, the value was around 0.53.
- Combining the above two conclusions, we suggest that mean soil water content and pressure head at first occurrence of limiting hydraulic conditions (hmean,lim and
mean,lim) can be calculated from the matric flux potential at 0.53 rm, which is a function of transpiration rate and root density and which depends on a soil-specific K(h) function.
- Thus, either the described numerical model, nomograms as depicted in Fig. 12, or the analytical solutions can be used for a comprehensive determination of hmean,lim and
mean,lim.
 |
ACKNOWLEDGMENTS
|
|---|
Quirijn de Jong van Lier is funded by CAPES-MEC, Brazil; Klaas Metselaar is funded by the framework of the Dutch National Research Programme Climate changes Spatial Planning. We thank Pieter A.C. Raats and Sjoerd E.A.T.M. van der Zee for a valuable discussion of the analytical solutions.
 |
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