Forest Ecology and Management 188 (2004) 77–89

Spatial distribution of root length density and soil water of

linear agroforestry systems in sub-humid Kenya:
implications for agroforestry models
Simone Radersmaa,b,*, Chin K. Ongc
a
Department of Soil Quality, Agricultural University, P.O. Box 8005, 6700 EC Wageningen, The Netherlands
b
International Center for Research in Agroforestry (ICRAF), Maseno, Kenya
c
ICRAF, Nairobi, Kenya
Received 14 March 2002; received in revised form 6 May 2003; accepted 14 July 2003

Abstract

In simultaneous agroforestry systems trees can compete with crops for water, especially in semi-arid areas. However, in the
(sub)humid tropics, on P-fixing Oxisols/Ferralsols small decreases in soil water content caused a decrease in P-transport to roots
and therewith a soil-drying induced P-deficiency.
The aim of this study was to assess the spatial distribution of soil water content in crop fields bordering tree lines and its
relation with root length density distribution of the trees throughout the soil profile. To achieve this, soil water content and tree
root length densities throughout the soil profile were measured over a period of 2 years in an experiment with lines of four tree
species in the middle of maize fields in sub-humid western Kenya.
Soil water content was significantly reduced (2–7 vol.%) near two of the three fast-growing tree species, Eucalyptus grandis
and Grevillea robusta, but not near Cedrella serrata and the slower growing Markhamia lutea. These differences were related to
differences in water use. Eucalyptus and Grevillea showed high water use and Cedrella and Markhamia low water use. However,
soil water content distribution was not related to root length density distribution. Root length densities hardly decreased with
distance to Grevillea and clearly decreased with distance to Cedrella.
Most water-uptake models, including those of agroforestry models, assume that root length density distribution throughout the
profile is proportional to water extraction throughout the profile. The absence of a clear relation between root length density and
water extraction near Grevillea tree lines opposed this view. It can be explained by a decrease in water-potential gradient
between root and soil at increasing distance from the tree base. If the change in root length density is similar or smaller than the
change in water-potential gradient between root and soil, the decrease in water-potential gradient between root and soil is of
similar or larger importance for determining tree-water extraction distribution throughout the profile than root length density.
Thus, modeling of spatial agroforestry systems cannot assume a direct relation between tree-water extraction and root length
density, but needs to include decreasing water-potential gradient between root and soil along roots with increasing distance to the
stem base, especially over the horizontal dimension.
# 2003 Elsevier B.V. All rights reserved.

Keywords: Agroforestry; Water extraction; Root distribution; Simulation models

*
Corresponding author. Tel.: þ320-291-352; fax: þ320-230-479.
E-mail address: s.radersma@freeler.nl (S. Radersma).

0378-1127/$ – see front matter # 2003 Elsevier B.V. All rights reserved.
doi:10.1016/j.foreco.2003.07.021
78 S. Radersma, C.K. Ong / Forest Ecology and Management 188 (2004) 77–89
1. Introduction
Water deficiency can be an important limitation to
crop-growth, not only in semi-arid environments, but
also in environments where phosphorus (P) is the main
limitation to crop-growth, because transport of P in
soil is highly dependent on soil water content. On a P-
fixing Oxisol/Ferralsol in western Kenya we found
that a significant absolute soil water content reduction
of 2–3% (from 38 to 39 to 36 vol.%) at relatively high
pF, as measured close by tree lines, caused a decrease
in maize-production of 30–40%. Using model simula-
tions, the decrease in crop-growth near the tree lines
could only be explained by decreased P-transport
towards roots at the lower soil water contents, with
its cumulative effects on crop and crop-root growth. It
could not be explained by direct competition for
nutrients or water (unpublished data). In this way,
trees affected crop-growth by soil-drying induced
P-deficiency.
Trees can affect soil water contents in different ways.
They can increase soil water content by decreasing soil
evaporation (Wallace et al., 1999; Jackson and Wallace,
1999), by hydraulic lift (Caldwell and Richards, 1989;
Dawson, 1993) and by hampering growth of other
plants with a higher water use per unit surface area.
They decrease soil water contents by their own water
use, or by interception of rain (Broadhead, 2000; Wal-
lace et al., in press). In tropical climates with an annual
rainfall of 1000–1500 mm, interception losses decrease
soil water inputs under the trees. However, this decrease
in water input to the soil is only slightly larger than
evaporation reductions under canopies, which decrease
soil water losses under the trees. This leads to only a few
percent lower total water input under the tree canopies
(Wallace et al., 1999) compared to water inputs outside
the area shaded by tree canopies. Because of this
balance between canopy effects on interception and
evaporation in this climate, we considered tree water
uptake as the main determinant of soil water depletion
in the soil volume near tree lines.
Root density distribution of trees and crops is usually
seen as a main indicator of competition between trees
and crops (Schroth, 1995; Smith et al., 1999), assuming
that at equal supply of growth resources in the soil, the
resource uptake is related to the amount of root length
per unit soil volume. Similarly, most crop and forest
water-uptake models assume that water extraction rate
is closely related to root length density (Rowse et al.,
1978; Belmans et al., 1979; Lascano and van Bavel,
1984; Barataud et al., 1995). This relationship between
water extraction and root length is also used in agro-
forestry models like WaNuLCAS (Van Noordwijk
and Lusiana, 2000), Hypar (Mobbs et al., 1998) and
WIMISA (Mayus et al., 1999). Therefore, we consid-
ered root length density distribution as main deter-
minant of soil water content distribution in the soil
volume near trees.
In the research described in this paper we examined
the profiles of water extraction with distance from tree
lines and their relation with tree root distribution in a
field experiment on a deep P-fixing Ferralsol/Oxisol in
sub-humid western Kenya. Lines of four tree species
selected for pole-production were grown within maize
fields. During 2 years, soil water contents were mon-
itored on a monthly basis and roots were sampled
twice, over depth and with distance from the tree line,
in order to determine (i) the effect of the tree lines on
soil water contents over depth and with distance from
the tree line, (ii) differences in tree root distribution
among species and over time, and (iii) the relation
between root length density distribution and soil water
extraction.
2. Material and methods
2.1. Site description
An experiment was conducted on a degraded farm-
er’s field at Nyabeda, western Kenya (latitude 0.088N,
longitude 34.248E, altitude 1300 m). The site was sub-
humid with an annual rainfall of about 1500 mm,
distributed over two main cropping seasons; the long
rainy season from March to June/July (900 mm) and
the short rainy season from September to December/
January (600 mm). The experiment was established
on level land, which after maize cropping was left
fallow the last year (three blocks)/last 2–3 years (one
block).
The soil was an Oxisol/Ferralsol, which is a major
soil type in the area (Shepherd et al., 1992). Soil
texture was 58% clay, 16% sand and 26% silt in the
top 0–15 cm. Clay contents increased to 73% at depth
(3 m). Bulk density remained <1.3 kg dm3 over the
first 1.6 m depth, due to high sesquioxide content and a
 S. Radersma, C.K. Ong / Forest Ecology and Management 188 (2004) 77–89
good soil structure. Soil pH (1:2.5 soil/water suspen-
sion) was 5.6 over the entire depth, and Olsen-P was
1.2 mg kg1 in the top 15 cm, decreasing with depth
to 0.3 mg kg1. Soil organic carbon decreased from
1.7% in the top-soil to 0.3% at 2–3 m depth.
2.2. Experimental design and management
The experiment was a 5  2 factorial design, with
four replicates. The treatments were a complete fac-
torial combination of two phosphorus (P) levels: one
with P (þP) and one without P (P) and five crop/
agroforestry treatments: four treatments of a tree line
in the middle of maize plots and a sole maize treatment
without a tree line. The tree species were Cedrella
serrata, Grevillea robusta, Eucalyptus grandis and
Markhamia lutea. The þP treatments received an
addition of 250 kg P ha1 at the start of the experiment
and 50 kg P ha1 in April 2000. The P treatments
did not receive P-fertilizer. The plot size was 13:5 m
15 m with the tree lines in the middle of the plots
parallel to the 13.5 m length side. Trees were trans-
planted to the field experiment in April 1997 at an age
of 4–6 months. Tree spacing was 0.5 m in the first year
thinned to 1 m thereafter.
Nitrogen (70 kg ha1 per year) and potassium
(15 kg ha1 per year) were applied regularly in all
plots, to prevent these elements from becoming a
limitation to crop and tree growth.
Trees were pruned on 22 December 1997, 14 May
and 30 September 1998, 20 April and 27 October
1999, according to farmers’ practice to produce poles
(cutting all branches up to about 3/4 of the tree height),
to decrease shading of crops. The plots were weeded
manually by hoe, twice during a cropping season.
Growth of tree roots into neighboring plots was
avoided by trenching up to 1.5 m depth in February
1999 and February 2000.
2.3. Measurements
Soil water contents were measured by neutron
probe (Didcot Instruments, Wallingford, UK), at
monthly intervals between July 1998 and August
2000 in all treatments. Access tubes for these mea-
surements were installed at three distances from
the tree line, 0.94, 2.44 and 6.94 m. These distances
were chosen to be at one-quarter/three-quarter of the
79
distance between maize rows and at increasing dis-
tance to the tree lines. Measurements were done in
all four agroforestry treatments and all four replicates
at 0.20 m intervals to 1.60 m depth. Neutron-probe
measurements were calibrated in the same field. First,
10 neutron-probe measurements were taken in a tube
in a water-barrel (as usual), than each depth in an extra
tube in the field was measured seven times, and finally
again 10 water-barrel measurements were taken. This
resulted together in the usual soil-count:water-count
ratios. On the same day four fixed volume rings per
measuring depth were inserted horizontally in the soil,
dug-out, leveled to contain exactly 100 cm3, weighed
moist, oven-dried for 4 days at 108 8C, and weighed
dry. Seven access tubes all around the experiment were
used for calibration, three tubes in the driest periods,
two tubes in the wettest periods and two tubes in
periods with intermediate soil water contents.
Root densities were measured in three replicates
of maize þ Grevillea, maize þ Eucalyptus, maize þ
Cedrella and sole maize treatments at 15 and 27
months after transplanting the tree-seedlings. Mea-
surement dates coincided with maize tasseling in the
long rainy season of 1998 and 1999. A bucket-auger of
0.10 m diameter (Ø) was used to sample pairs of
points (one within a maize row and one in between
two maize rows) for the layers 0.10–0.30, 0.30–0.50,
0.50–1.00, 1.00–1.50 and 1.50–2.00 m (in 1998 the
1.00–2.00 m was sampled as one layer). In 1998 the
average distance of the pairs of points was 0, 0.94,
2.44 and 4.69 m, like the distances of soil water
content measurements. In 1998 when the trees were
still small and the spread of roots limited, samples
were taken at 4.69 m instead of 6.94 m and a sampling
at 0 m was included. In 1999 the 0 m distance was quit
because of the too large woody roots at the base of the
trees and the 6.94 m distance was added because of
the wider spread of tree roots at this age. The layer
0–0.10 m was not sampled, because weeding twice
during the cropping season cut most roots. If an auger
hole could not reach the specified depth due to occa-
sional stones, final depth of the hole was measured and
sample volume adjusted accordingly before calculat-
ing root length densities. Both sides of the tree lines
were sampled in this way. In 1998 the samples of the
two sides were taken separately, total root length
measured per sample and divided by the total volume
of each sample, after which the mean of both sides was
80 S. Radersma, C.K. Ong / Forest Ecology and Management 188 (2004) 77–89
used for further analysis. In 1999, samples from both
sides were bulked, total root length measured and
divided by the total volume of the two sides.
Samples of depth 1.00–2.00 m, which were too
large volumes (>20 l), were mixed thoroughly in the
field and a sub-sample of half or quarter of the total
weight was taken.
Buckets containing soil þ root samples were soaked
overnight. The next day the mixture was washed
through a double sieve with mesh size 0.5 mm for
the bottom sieve. Further measurement of root length
densities was done according to the procedure of hand
sorting, staining, spreading and scanning following the
method described by Smith et al. (1999).
2.4. Statistical analysis and calculations
Soil water content data of the four most contrasting
depths: 0.20, 0.60, 1.20 and 1.60 m, were analyzed
separately. Data for measurement times either within
the wet seasons or within the dry seasons showed
similar trends as far as differences in soil water con-
tents with distance from the tree lines were concerned.
Therefore the average of all measurement times in the
last three wet seasons (long rains of 1999, short rains
of 1999 and long rains of 2000) and the average of
all measurements times in the last two dry seasons
(dry season 1999 and dry season 2000) were used for
analysis of soil water content differences with distance
from the tree line (Dd), with Dd ¼ yfar  ynear , and yfar
the average volumetric soil water content of the mea-
surements at 2.44 and 6.94 m away from the tree line
and ynear the average volumetric soil water the content
at 0.94 m away from the tree line. These Dd values
were analyzed by ANOVA as 5  2 factorial design
with four replicates.
Because the trees were not all of the same size, the
root density data for each plot were adjusted to the
average of basal stem cross-sectional surfaces (S) of
all trees per treatment (average of three replicates). S is
related to total cross-sectional area of finer roots and
root length according to the fractal branching theory
(Van Noordwijk et al., 1994). Thus root densities in
each plot/replicate were multiplied by the ratio of
Smeasured:Saverage, where Smeasured is the sum of the
surface of the three tree-stem cross-sections closest to
the sampling transect in one plot/replicate, and Saverage
the average of the three replicates of Smeasured.
After adjustment of all root length densities to this
average S, the means of the three replicates of root
length density profiles with distance from the tree
line were analyzed by non-linear fitting to derive
the variables of the elliptical function as described
by (Van Noordwijk and Lusiana, 2000):
rldði; jÞ
¼ X0  DD expðDDðDepth2j þ ðDS  Disti Þ2 Þ1=2 Þ
(1)
where rld(i, j) is the root length density at distance i
and depth j in cm cm3, X0 the total root length per
unit area (cm cm2) at a distance of 0 m from the tree
stem, DD the parameter (cm1) for the distribution of
total root length per unit area with depth, and DS the
dimensionless parameter governing the shape of the
tree root system (when rld at 5 m distance is the same
as at 1 m depth than DS ¼ 0:2)
3. Results and discussion
3.1. Tree growth
Fig. 1 shows the growth in height and diameter of
the trees over time in the different treatments. Growth
rates of Eucalyptus þP (þP ¼ with P-application)
were clearly the highest, followed by Eucalyptus
P (P ¼ without P-application), Cedrella (þP and
P) and Grevillea (þP and P). Grevillea trees were
slightly taller than Cedrella trees, but Cedrella had a
larger diameter than Grevillea. Markhamia grew very
little compared to the other trees. The slow-down in
diameter-growth between the last two measurements
before 900tp (¼900 days after transplanting), is due to
a severe pruning event (April 1999). The first mea-
surement after 900tp marks another change: the height
of diameter measurements changed from 0.05 to
0.30 m above soil surface.
3.2. Soil water content
Average soil water contents over 2 years in sole
maize fields are shown in Fig. 2. The small time-lag
in drying and wetting cycles of the different soil
depths, shows that hydraulic conductivity of the soil
is quite high, in both wet and dry circumstances, as
 S. Radersma, C.K. Ong / Forest Ecology and Management 188 (2004) 77–89 81

Tree height over time Tree diameter over time

11,0 120
10,0 110
9,0 100
8,0 Mark.-P
90

basal diameter (mm)

Cedrel.-P
7,0
Grevil.-P 80
height (m)

6,0 Eucal.-P
70
5,0 Mar.+P
Cedrel.+P 60
4,0
Grevil.+P
50
3,0 Eucal.+P
2,0 40

1,0 30

0,0 20
450 900 1350 450 900 1350
20/7/98 15/10/99 8/1/00 20/7/98 15/10/99 8/1/00
tp (days) and date tp (days) and date

Fig. 1. Tree height and tree-stem diameter over time, with tp as time in days after transplanting to the field: Mark: ¼ Markhamia lutea;
Cedrel: ¼ Cedrella serrata; Grevil: ¼ Grevillea robusta; Eucal: ¼ Eucalyptus grandis; P ¼ without phosphorus application; þP ¼ with
phosphorus application. Standard errors for tree height and basal stem diameter were between 0.4 and 1% of the means plotted in the figures.

may be expected for an Oxisol/Ferralsol (El-Swaify,
1980).
The amplitude and average level of soil water
contents during the dry and wet season, differed for
the different layers. The amplitude in volumetric soil
water content (y) was the largest for the upper soil
layer (around 11%), with y at a mean value of 32%. At
0.60 and 1.60 m depth amplitudes were lower, 6 and
3.5 vol.%, respectively, with y at higher mean values
of 43 and 38%, respectively. In between these layers,
at 1.20 m depth, the amplitude was larger again
(10 vol.%) with y at a mean value of 38%.

50

45
vol.% soil water

40 depth 0.20 m
depth 0.60 m
35 depth 1.20 m
depth 1.60 m
30

25
490 690 890 1090
1/9/98 19/3/99 5/10/99 22/4/00
tp (days) and date

Fig. 2. Mean volumetric soil water content and standard errors in sole maize fields over time, with tp as time after transplanting.
82 S. Radersma, C.K. Ong / Forest Ecology and Management 188 (2004) 77–89

Because water contents at 2.44 and 6.94 m distance of soil water content at 2.44 and 6.94 m minus the soil
away from the tree lines did not yet differ significantly, water content at 0.94 m distance from the tree lines,
their means were used for further analyses to reduce was used to calculate Dd values, shown in Table 1a
spatial variability in soil water content. This average and b. Thus Dd ¼ y244694  y94 and Dd > 0 if
Table 1
Changes in soil water contents (Dd) in maize fields approaching the tree line (average over last three (a) wet seasons and (b) dry seasons),
calculated by Dd ¼ yfar  ynear, with yfar as the volumetric soil water content far from the tree lines (mean of measurements at 2.44 and 6.94 m
distance from the tree lines) and ynear as the volumetric soil water content at 0.94 m from the tree linesa

Species P-supply Dd (vol.%)

Depth 0.20 m Depth 0.60 m Depth 1.20 m Depth 1.60 m

(a) Average over last three wet seasons: March–July 1999, September–November 1999 and March–July 2000
Cedrella þP 0.13 0.34 1.39 0.58
Cedrella P 0.00 1.28 0.43 0.92
Eucalyptus þP 2.51 2.72 6.92 3.86
Eucalyptus P 1.68 1.22 2.45 1.07
Grevillea þP 0.57 1.22 3.17 1.28
Grevillea P 0.23 0.50 2.07 2.40
Markhamia þP 0.72 0.51 0.61 0.71
Markhamia P 0.57 0.93 0.22 0.56
Sole maize þP 0.60 0.34 0.56 0.9
Sole maize P 0.94 0.48 0.91 0.17
S.E.D. 1.423 1.108 1.309 1.207
Cedrella 0.07 0.47 0.48 0.17
Eucalyptus 2.09 1.97 4.69 2.46
Grevillea 0.17 0.86 2.62 1.84
Markhamia 0.64 0.72 0.19 0.63
Sole maize 0.17 0.07 0.74 0.54
S.E.D. 1.006 0.784 0.925 0.853
þP 0.85 0.75 2.53 1.46
P 0.22 0.88 0.96 0.66
S.E.D. 0.636 0.496 0.585 0.54
(b) Average over last three dry seasons: December 1998–February 1999 and December 1999–February 2000
Cedrella þP 0.58 0.14 1.12 0.60
Cedrella P 0.09 0.91 0.11 1.13
Eucalyptus þP 0.24 0.88 3.51 3.11
Eucalyptus P 1.31 0.61 0.66 0.99
Grevillea þP 0.97 0.34 2.16 1.01
Grevillea P 1.54 0.58 2.44 2.74
Markhamia þP 1.80 0.34 1.22 0.94
Markhamia P 0.35 0.53 0.18 0.38
Sole maize þP 1.17 0.57 0.27 0.22
Sole maize P 0.56 0.73 1.95 0.28
S.E.D. 1.535 1.255 1.677 1.316
Cedrella 0.24 0.39 0.50 0.27
Eucalyptus 0.54 0.13 2.08 2.05
Grevillea 1.25 0.46 2.30 1.87
Markhamia 0.72 0.10 0.52 0.66
Sole maize 0.31 0.08 1.11 0.25
S.E.D. 1.085 0.888 1.186 0.931
þP 0.33 0.03 1.17 1.18
P 0.74 0.43 1.03 0.65
S.E.D. 0.686 0.561 0.750 0.589
a
A significant positive Dd indicates a decrease in soil water content near the tree line.
 S. Radersma, C.K. Ong / Forest Ecology and Management 188 (2004) 77–89
y244694 > y94 or positive Dd values indicate decrea-
sed soil water content near the tree lines. Table 1a
shows the Dd values for the means of the three wet
seasons, (which showed similar trends and were there-
fore averaged). Positive Dd values were the clearest
(2.5–6.9%) in Eucalyptus þP, and significantly dif-
ferent from 0 (Dd > 2 S:E:D: at a ¼ 0:05) in all layers
except the first. Positive Dd values (1.2–2.5%) seemed
to occur in all layers of Eucalyptus P as well, but
these were not significant ða ¼ 0:05Þ. Positive Dd
values (1.2–3.2%) occur in the three lower layers of
Grevillea þP, but are only significant at 1.20 m depth.
In Grevillea P positive Dd values occurred in the
two deepest layers, but this Dd was only significant at
1.60 m depth.
The significance of Dd values could be partly
explained by tree-growth. Eucalyptus þP was the
largest and showed the largest Dd. Markhamia was
very small and did not show any significant Dd.
Another part of the differences in Dd among species
could be explained by daily water use of a species at a
particular size. Cedrella and Grevillea of about the
same size, showed different daily water use (unpub-
lished data). Decreased water use by maize, due to its
slow growth caused by competition near especially
Eucalyptus, would have counteracted the development
of large Dd values in the top layers.
If significant positive Dd values of 2–7 vol.% are
compared with the normal annual amplitude (Amp) in
a sole maize field (Fig. 2), by calculating Dd:Amp.
The Dd:Amp at depth 0.20 m is 2.51%:11% ¼ 23%,
at depth 0.60 m is 2.71%:6% ¼ 45%, at depth
1.20 m is 6.92%:10% ¼ 70% and at depth 1.60 m is
3.86%:3:5% > 100%. This shows that the soil near
Eucalyptus þP dried out to 25–100% of the normal
annual soil water content minimum in sole maize
fields. Near the other tree lines, drying was less severe,
but compared with seasonal fluctuations it was still
considerable.
A reason for the strong tree-caused drying of deeper
layers compared to seasonal drying at depth may have
been that the roots caused a ‘shortcut’ contact between
the deep layers and the atmosphere. This shortcut was
absent without roots (e.g. maize field in dry season),
and caused that the low soil conductivity determined
the drying rate.
The Dd values in the dry season are shown in
Table 1b. The Dd values for the first two layers
83
(0.20 m depth and 0.60 m depth) were generally not
significantly different from zero ðDd < 2 S:E:D:Þ, and
thus soil water contents did not change significantly
with distance from the tree lines. The Dd values in the
lower layers (1.20 and 1.60 m depth for Eucalyptus
þP and 1.60 m for Grevillea P) were still significant
in the dry season.
The decrease in Dd in the dry season, indicating
decreased differences in soil water content close by
and far from the trees can be explained as follows.
Over moist and wet soil water content ranges (like in
our wet season), root resistance usually exceeded soil
resistance (Belmans et al., 1979; Reid and Hutchinson,
1986) and suction by the root was larger than suction
by the soil. The situation in which soil resistance
exceeds root resistance occurs only in drier soil or
at high transpiration rates in coarse soils (Hillel et al.,
1976; Weatherley, 1979; Hainsworth and Aylmore,
1989). Soil suction becomes higher than root suction
when the soil dries out, causing plant water uptake to
cease. This happened likely earlier in drier soil near
the tree lines than in the wetter soil far from the tree
lines and caused the decrease in Dd values in the dry
seasons as compared to the wet season.
3.3. Tree roots
The elliptical-function parameters differed per spe-
cies but were not affected by P treatment. Therefore,
the root length density (rld) data were multiplied by a
factor Smeasured:Saverage for all three þP and P repli-
cates (where S is the cross-sectional surface of the
stem base and the calculation is similar to the one
described in Section 2) before taking the average of
þP and P and determining the ellipse function
parameters of the species as given in Table 2, or
plotting root distribution per species in Fig. 3. Because
of the exponential decrease in root length density over
depth, the depth taken to represent the layers in Fig. 3
is at 1/4 of the layer thickness below the upper
boundary of the layer, e.g. the average rld of layer
0.1–0.3 m depth is plotted at 0.15 m depth.
The elliptical parameter, X0, is the total root length
per unit soil surface (cm cm2) at distance 0 m from the
tree line. Thus high maximum and average rld values,
especially near the stem base are reflected in higher X0
values. The X0 values were partly dependent on tree-
size (average cross-sectional surface of stem base, Sav),
84 S. Radersma, C.K. Ong / Forest Ecology and Management 188 (2004) 77–89

Table 2
Ellipse function parameters of Eq. (1), rldði; jÞ ¼ X0  DD expðDDðDepth2j þ ðDS  Disti Þ2 Þ1=2 Þ, all calculated with depth and distance
in (cm)a

1998 1999 1998 1999 1998 1999

Sav X0 S.E. Sav X0 S.E. DD S.E. DD S.E. DS S.E. DS S.E.

(cm2) (cm cm2) (cm2) (cm cm2) (cm1) (cm1) (–) (–)

Cedrella 13.5 23.0 6.76 49.8 15.3 5.51 0.025 0.0059 0.027 0.0040 0.21 0.055 0.14 0.031
Eucalyptus 11.7 19.0 5.92 55.9 16.1 3.93 0.033 0.0057 0.020 0.0030 0.14 0.039 0.17 0.032
Grevillea 5.2 10.1 1.78 30.4 4.1 0.85 0.034 0.0034 0.022 0.0036 0.04 0.033 1E11 *

With rld(i, j) as root length density at distance i and depth j from the stem base, X0—total root length per unit area (cm cm2) at a
a

distance of 0 cm from the tree stem, DD—parameter (cm1) governing the decrease with depth of root length density (for X ¼ 0 at a depth of
0.699/DD the root length density has half of its value at the soil surface), DS—parameter governing the shape of the tree root system (when rld
at 5 m distance is the same as at 1 m depth, then DS ¼ 0:2) and Sav—mean basal stem surface of trees next to root-sampling transect.

and because Eucalyptus and Cedrella were much larger
than Grevillea, their X0 value was larger as well. How-
ever, when the two sampling times (1998 vs 1999) were
compared, there was no relation between Sav and X0. In
1999 the trees and the Sav were larger, but the X0 was
lower than in 1998. Fig. 3 also shows that absolute
root densities were lower in 1999 compared to 1998.
The decrease in absolute root length densities and in
X0 values of the elliptical function, was likely due to
tree-management and root-system growth dynamics
as also found by Smith et al. (1999) and Odhiambo
et al. (1999). Before the long rains of 1999, tree cano-
pies were severely pruned, which likely caused root-
mortality and thus reduced the X0:Sav ratio.
The DD variable describes the shape of the vertical
root profile. A high DD value in Table 2 is represented
by a less steep slope of the rld lines in Fig. 3. The
DD values were not significantly different for the tree
species. However, the DD value of 1999 root-data was
significantly smaller than that of 1998 for Eucalyptus
and Grevillea but did not change for Cedrella. This
implies that for Eucalyptus and Grevillea relatively
more roots had grown at depth than in the surface
layers. Smith et al. (1999) mentioned a relatively
high root length density of Grevillea at depth as well.
However, the exponential decrease in rld of Grevillea
over depth they mentioned, is much less than what we
found, likely due to the dryer environment of their
experiment.
The DS variable of the elliptical function describes
the shape of horizontal root distribution in relation to
the vertical distribution. If DS is 0, root densities do
not decrease over distance, and the rld lines in Fig. 3
should overlap. If DS is large, the root densities
decrease rapidly with increasing distance from the
tree line and rld lines in Fig. 3 should be far apart.
Thus, the very low DS of Grevillea, which was not
significantly different from 0, and the overlap of the
rld lines in Fig. 3, showed that Grevillea root length
density did hardly decrease with increasing distance to
the tree. DS values of Eucalyptus and Cedrella were
significantly higher than 0, hence in the figure the rld
lines at increasing distance to the trees became closer
to the Y-axes, and root length density decreased with
increasing distance to the trees. DS values did not
change significantly between 1998 and 1999.
Root distribution profiles with distance from a tree
line differed between tree species, and changed over
time. Over a period of 1 year the root systems generally
extended horizontally without similar vertical exten-
sion. High root density at the first root sampling in 1998
might indicate that if a tree grows fast over a short
period of time in a wet period, its increase in root length
increases root length density, thus intensifying extrac-
tion of resource from a small soil volume. On the other
hand after a dry season and a severe pruning event, fine
roots may have died and new roots grew mainly to
extend the root system in order to cover a larger soil
volume. Fitter et al. (1991) mentioned that a rooting
architecture with long links between branching points
and branching predominantly on the main axis,
although expensive in use of carbohydrates, would
mainly occur in nutrient poor soils. This indicates that
changes in root distribution over time may not only be
caused by management, but also by a root-system
distribution strategy adapted to depleted soils. Whether
caused by management or by resource availability, not
only total root length and extent changed over time, but
 S. Radersma, C.K. Ong / Forest Ecology and Management 188 (2004) 77–89 85

Eucalyptus (1998) Grevillea (1998) Cedrella (1998)

-3 -3 -3
rld (cm cm ) rld (cm cm ) rld (cm cm )

0,001 0,010 0,100 1,000 0,001 0,010 0,100 1,000 0,001 0,010 0,100 1,000
0 0 0

20 20 20

0m

0m

0m
40 40 40
0.94 m

0.94 m

0.94 m
depth depth depth
(cm) 60 (cm) 60 (cm) 60

80 80 80
2.44 m

2.44 m

2.44 m
100 100 100

120 120 120

4.69 m

4.69 m

4.69 m
(a) 140 140 140

Eucalyptus (1999) Grevillea (1999) Cedrella (1999)

-3 -3 -3
rld (cm cm ) rld (cm cm ) rld (cm cm )

0,001 0,010 0,100 1,000 0,001 0,010 0,100 1,000 0,001 0,010 0,100 1,000
0 0 0

20 20 20

0.94 m
0.94 m

0.94 m

40 40 40

60 60 60

depth depth depth

2.44 m
2.44 m

2.44 m

(cm) 80 (cm ) 80 (cm ) 80

100 100 100

4.69 m

4.69 m

4.69 m
120 120 120

140 140 140

160 160 160

6.94 m

6.94 m

6.94 m

(b) 180 180 180

Fig. 3. Tree root length density (rld) distribution and standard errors in (a) 1998 and (b) 1999.

also tree root distribution. This change of root distribu-
tion profiles over time may pose a problem to agrofor-
estry models, like WaNuLCAS, because the elliptical
parameters derived from measurements at one point
in time may not be valid during the whole growth of
the tree.
3.4. Root length densities and soil water content
Comparing the root profiles of Table 2 and Fig. 3
with tree-water extraction as indicated by Dd in the
wet season (Table 1a), tree-water extraction and root
density distribution show no clear relation. We see that
86 S. Radersma, C.K. Ong / Forest Ecology and Management 188 (2004) 77–89
the root density of Grevillea hardly changed with
distance from the tree line, but soil water contents
were significantly lower close by the Grevillea tree
line ðDd > 0Þ. On the other hand, the root density of
Cedrella decreased with distance to the tree line but
soil water contents were not significantly different
when measured close by or far from the tree line.
This absence of a relation between root density dis-
tribution and soil water contents is in contrast with the
relations used in most computer models (Rowse et al.,
1978; Belmans et al., 1979; Lascano and van Bavel,
1984; Barataud et al., 1995), the general descriptions
of how water extraction is modeled (Campbell, 1991;
Simmonds and Kuruppuarachchi, 1995), and with the
way it is employed in agroforestry models such as
WaNuLCAS (Van Noordwijk and Lusiana, 2000)
Hypar (Mobbs et al., 1998) and WIMISA (Mayus
et al., 1999). In all these models water uptake is
closely related to relative root length density.
We consider soil water content in the wet season to
be mainly determined by tree and crop water uptake
rather than by rainfall interception of the trees. This
assumption is supported by a water balance study of
G. robusta–maize agroforestry systems in semi-arid
Kenya (750 mm of annual rainfall), which showed
that the small reduction in rainfall input due to canopy
interception was partly offset by a reduction in soil
evaporation (Wallace et al., 1999). It is therefore
unlikely that interception explains the large reduction
in soil water content observed in our present study and
we ascribe water extraction from soil by the tree as the
main cause of decreased soil water contents near the
tree lines.
The absence of Dd in fields with a Cedrella tree line,
although the tree roots were concentrated near the
stem base, may be explained by Cedrella’s relatively
low water use (unpublished data) and low water
extraction from soil. This low water use is likely
easily balanced by precipitation in the wet season,
as shown by soil water content values, which are not
significantly different from those in sole maize fields.
However, the clear Dd in fields with a Grevillea tree
line, showed that water extraction near the stem base
was higher than far from the tree, although root length
densities did not decrease with distance to the stem
base.
The discrepancy between the general assumption
(and models), that water uptake is closely related with
root length density, and our observation near Grevillea
tree lines that root length density distribution and
distribution of water extraction are not closely related,
can be explained by Fig. 4a. In Fig. 4a water flow in a
branching root, is represented by a branched electrical
circuit. The principles used here are those of water
extraction by plants analogous to Ohm’s electrical
circuit, with a potential gradient as driving force
and resistances limiting the flow rate: q ¼ Dc=r
(Hillel et al., 1976; Scott Russell, 1977), with q as
flux/flow, Dc as potential gradient and r as resistance.
Fig. 4a represents a main root with branches, in which
each branch/resistance between main flow path and
earth represents a similar total root length (case 1).
Fig. 4a can also represent a tiny piece of root in which
each branch/resistance between main flow path and
earth represents a channel for radial water inflow
into the root (case 2). The þ side of the electric circuit
in Fig. 4a represents the stem base. The earth-sign
(potential c ¼ 0) represents sites where water uptake
from the soil into the root takes place. The resistances
(r) connecting earth to main flow path are all chosen
to be the same size ðr ¼ 1Þ. This same resistance size
shows that either the total root length after each
branching point is equal (case 1) like the rld of
Grevillea with distance from the tree line, or radial
resistance along the root is the same (case 2) whether
close by or further from the stem base. The resistances
along the main flow path are chosen to decrease
towards the þ, like can be expected from the axial
resistance of a main root towards the stem base. After
this choice of resistances and earth, the flux (q) and
the potential (c) along the different parts of the
branched electrical circuit were calculated by
q ¼ Dc=r. This shows that with increasing distance
away from the þ or stem base, the potential (c)
decreases resulting in a drop in influx (q) through
the resistances connecting earth to main flow path
ðr ¼ 1Þ. Thus, although the root length density (case
1) or radial resistance (case 2) is the same close by the
stem base (þ) as further away from the stem base, the
water inflow (q) decreases with increasing distance to
the stem base, due to a decrease in potential gradient
between root and soil.
If these small sections were multiplied to form a
whole root, the decline in potential gradient between
root and soil (or þ and earth) and water uptake with
distance from the stem base would show a logarithmic
 S. Radersma, C.K. Ong / Forest Ecology and Management 188 (2004) 77–89 87

Fig. 4. (a) Model of an electric circuit representing a branching root, in which the root-branches have similar hypothetical uptake-resistances
(chosen r ¼ 1) and the main root has smaller hypothetical axial resistances (chosen r ¼ 1=2; 1=3; 1=4). The earth-sign ðc ¼ 0Þ represents the
water-uptake site/soil–root interface. The flux (q) and potential (c) at different sites along the branched circuit/root were calculated by
q ¼ Dc=r. (b) Hypothetical course of decrease in hydraulic potential along a root with increasing distance to the stem base.

course like in Fig. 4b. This course can be described by
a function exy, in which x is the distance and y would
be a function of radial resistance, root-branching and
axial resistance.
The origin of the assumption that water uptake is
closely related to root length density can be found in
earlier soil water extraction models. These models
were made for crop fields or forest stands with a one-
dimensional profile, in which root density, water-
potential gradient between root and soil and water
extraction only varied vertically, with depth. Gener-
ally, root length densities decreased exponentially
over depth, hence water-uptake resistance of the root
system increased exponentially over depth, and was
the main determinant of soil water extraction. As a
result, in models with only a vertical dimension, water
extraction was modeled as proportional to (relative)
root length density. However, if the root length density
does not decrease, like horizontally away from the
Grevillea tree base, differences in other flux determin-
ing factors like water-potential gradient between root
and soil gain importance.
Taylor et al. (1992) and Doussan et al. (1999) show
that indeed there is a severe drop in water-potential
gradient between root and soil along roots (cotton,
peach) away from the stem base. Decreasing water
uptake per unit root length as a consequence of
increasing axial resistance and decreasing water-
potential gradient between root and soil has also been
mentioned by Hainsworth and Aylmore (1989). From
water profile figures of Green and Clothier (1999) it is
clear that the highest water extraction occurred around
the tree bases, although root densities did not show
horizontal variation with distance to the tree base in
their orchard.
Decreases in axial resistance and potential gradient
between root and soil occur in the vertical direction
as well. However, in the vertical direction the decrease
88 S. Radersma, C.K. Ong / Forest Ecology and Management 188 (2004) 77–89
in water-potential gradient between root and soil is
smaller than the exponential decrease in root length
density causing the exponential increase in radial resis-
tance. Therefore over the vertical direction, changes
in water potential between root and soil could be
neglected. Nevertheless, Belmans et al. (1979) mention
that the fit of modeled and experimental data would
probably be better if axial resistance and therewith
change in water-potential gradient between root and
soil was included.
Summarizing, if the root length density decreases
far more than the decrease in water-potential gradient
between root and soil, water extraction can be mod-
eled as inversely proportional to (relative) root length
densities. In the soil volume this occurs usually in the
vertical direction. However, if root length density does
not change far more than water-potential gradient
between root and soil, the decrease in water-potential
gradient between root and soil and/or axial resistance
needs to be included in attempts to model water
uptake, to avoid large errors in water-uptake distribu-
tion. This occurred near our Grevillea tree lines in the
horizontal direction.
Acknowledgements
Thanks are due to Sammy Kyalo and Japheth
Kyengo and all the casuals for field and root work,
to Meine van Noordwijk, Jan Goudriaan and Herman
van Keulen for their analysis suggestions, and to the
KEFRI/ICRAF EU-funded project and the Dutch
government covering the labor-costs of this research.
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