Forest Ecology and Management 171 (2002) 43–57

Changes with age in the spatial distribution of roots

of Eucalyptus clone in Congo
Impact on water and nutrient uptake
Jean-Pierre Bouilleta,*, Jean-Paul Laclaua, Michel Arnaudb,
Armel Thongo M’Bouc, Laurent Saint-Andréd,
Christophe Jourdane
a
CIRAD-Forêt/UR2PI, TA/10C, Campus International de Baillarguet, 34398 Montpellier Cedex 5, France
b
CIRAD-Tera, Ave. Agropolis, TA 60/15, 34398 Montpellier Cedex 5, France
c
IDR/UR2PI, BP 1291, Pointe-Noire, Congo
d
CIRAD-Forêt, TA/10C, Campus International de Baillarguet, 34398 Montpellier Cedex 5, France
e
CIRAD-CP, Ave. Agropolis, TA 80/01, 34398 Montpellier Cedex 5, France

Abstract

Clonal Eucalyptus plantations in the Pointe-Noire region have been established on sandy and nutrient poor soils, and carry
potential risk of depleting water and nutrient. To assess these risks, water and nutrient budgets should be calculated. However,
the accuracy of hydrological models is strongly dependent on a realistic description of root distribution in soil. The spatial
distribution of root systems in a chronosequence of clonal stands was then studied, using root intersect counting. The stands were
3 months, 6 months, 1 year, 2 years and 9 years old. For each stand we studied three vertical profiles perpendicular to the planting
row at different spacing from a representative tree. The profiles were divided into square grid cells of 25 cm2, and the number of
roots belonging to three diameter classes (between 0.1 and 1, 1 and 10 mm, and over 10 mm) were counted in each grid cell. The
profiles were 2.50 m wide (half the distance between the planting rows) and 3 m deep, except for the two youngest stands (1.5 m
deep). Spatial statistical analyses and analysis of variance were carried out to describe root distributions and their spatial and
temporal changes. We observed that:
(1) The trees quickly developed an extensive root system. One year after planting the root system extended to depth beyond
3 m deep and up to the middle of the inter-row.
(2) Root density decreased sharply with depth, with most fine roots in the surface layers. The number of fine root intersects in
the 0–25 cm surface soil layer represented 16–53% of the total throughout the profile depending on stand age and the type
of profile.
(3) The percentage of root intersects in surface layers increased with stand age: this may reflect the greater concentration of
nutrients in surface layers.
(4) High variability in root densities was observed in the surface soil layers but no gradient of decrease from the planting row
to the middle of the inter-row could be underscored. However, the highest root densities were generally observed under the
stump.

*
Corresponding author. Tel.: þ33-4-67-59-38-66; fax: þ33-4-67-59-37-33.
E-mail address: jean-pierre.bouillet@cirad.fr (J.-P. Bouillet).

0378-1127/02/$ – see front matter # 2002 Elsevier Science B.V. All rights reserved.
PII: S 0 3 7 8 - 1 1 2 7 ( 0 2 ) 0 0 4 6 0 - 7
44 J.-P. Bouillet et al. / Forest Ecology and Management 171 (2002) 43–57
These patterns of distribution allow rapid access to soil water and nutrients, reduce loss by deep drainage, and may partially
explain the fairly high productivity of these plantations.
# 2002 Elsevier Science B.V. All rights reserved.
Keywords: Eucalyptus; Root intersect counting; Root system; Root distribution; Sustainability; Hydrological models
1. Introduction
Since 1978, 42 000 ha of clonal Eucalyptus planta-
tions have been established around Pointe-Noire, mainly
for pulpwood production. These plantations, owned by
ECO SA (Eucalyptus du Congo Société Anonyme),
are based on two natural hybrids (Eucalyptus PF1;
E: teriticornis  E: grandis) and, increasingly, on the
artificial hybrid E: urophylla  E: grandis. These
plantations are established on sandy soils charac-
terised by low reserves of available nutrients and a
low water retention capacity (Laclau et al., 2001).
Short rotations are used, resulting in the export of high
amounts of biomass and nutrients every 7 years. This
leads to potential risks of water and nutrient deficien-
cies and non-sustainable production. In order to assess
these risks, studies have focused on the biogeochem-
ical cycle of nutrients, and on the quantification of
water and nutrient budgets in clonal stands during the
planted crop rotation (Bouillet et al., 1999). In such
studies, hydrological models are used to estimate the
water and nutrient status and flows in the soil, including
nutrient outputs by drainage. The ability of these models
to accurately estimate water and nutrient uptake is
strongly dependent on a realistic description of the
distribution of roots in the soil (Bruckler et al., 1991;
Habib et al., 1991; Lafolie et al., 1991; Molz, 1981;
Nouvellon et al., 2000). Moreover, detailed knowledge
of the architecture of the root system is essential
because of its role in improving the soil structure (Jama
et al., 1998) and in the tree anchoring (Coutts, 1983;
Ennos et al., 1993; Fitter, 1986), or the contribution of
roots to carbon and nitrogen cycles (Hendricks et al.,
1993; Ruess et al., 1996). This information may also be
used to assist management decisions such as weeding
or fertilisation to ensure the most efficient use of water
and nutrients (Nambiar, 1983).
Since 1997 research at UR2PI (Unité de Recherche
sur la Productivité des Plantations Industrielles) has
focused on the spatial and temporal variability of root
distribution in Eucalyptus stands. A study, using the
method of root intersect counting (Tardieu, 1988) was
conducted on a 6.5-year-old planted crop where soil
water, using TDR was monitored to 5 m depth (Laclau
et al., 2001). This study showed a strong vertical
anisotropy of the distribution of fine roots, the density
of roots decreasing sharply with depth, a marked
horizontal anisotropy, and a grouping of fine roots
at depth due to the possible influence of preferential
drainage paths. The objective of the present study
was to build upon this work, studying the dynamics
of root distributions at different stages of the whole
rotation. An accurate analysis of the spatial variabil-
ity of root distribution was carried out in stands of
a chronosequence, within the environment of one
representative tree. Relationships were investigated
among root distributions, changes in biological
cycles, and characteristics of water and nutrient
uptake.
2. Material and methods
2.1. Site description
Eucalyptus plantations cover a band about 30 km
wide on the Atlantic coast of Congo (48S, 128E). The
climate is characterised by a high atmospheric humid-
ity (85% on average) with low seasonal variations
(2%), a mean annual rainfall of about 1200 mm with a
dry season usually between May and September.
Mean annual temperature is high (25 8C) with seaso-
nal variations of around 5 8C. Within the plateau
where most plantations are located, altitude ranges
from 80 to 120 m and the topography is slightly
undulating. The geological bedrock is composed of
thick detritic layers of continental origin dated from
plio-pleistocene (Jamet, 1975). The soils are arenosols
(FAO classification), characterised by their depth
(down to 100 m), homogeneity in the landscape in
colour (from greyish on soil upper layers to ochre
in deep layers), texture (sand content over 85%),
 J.-P. Bouillet et al. / Forest Ecology and Management 171 (2002) 43–57
structure and poor chemical content (Laclau et al.,
2000a).
2.2. Plant material
This study concerns one of the most productive
clones of the hybrid Eucalyptus PF1 (clone 1-41). This
hybrid is a product of natural crosses between two or
three individual plants of Eucalyptus alba Reinw. ex
Blume (female tree) and a group of Eucalyptus hybrid
not well identified (male tree) coming from a Brazilian
arboretum. The latter might be a cross between
the E. grandis W. Hill ex Maiden, E. robusta Smith,
E. urophylla S.T. Blake, and E. botryoı̈des Smith
(Delwaulle, 1988).
2.2.1. Particularities of the clone 1-41
About 7000 ha have been planted with this clone
since 1978. It is hardy and grows well with appropriate
weed control and application of fertiliser (20 m3 ha1
per year). Inventories of the plantations has given
detailed knowledge of its patterns of growth through-
out successive rotations at a variety of sites. It is
therefore used as a reference clone in breeding
and silviculture trials, and for the development of tree
growth models and for the quantification of nutrient
budgets (Bouillet et al., 1997; Laclau et al., 2000a,b).
Fig. 1. Mean stand heights of Eucalyptus stands of different ages.
45
2.2.2. Stand characteristics
Studies based on a chronosequence approach assume
that the stands sampled represent the main stages of
development of the same plantation (Dyck and Cole,
1994; Ranger et al., 1996). The clonal plantations
studied here constituted an appropriate situation in
which to use this approach. Indeed, the stands of the
chronosequence represented the first year of the rotation
(3 months, 6 months, 1 year) when weeding is carried
out, the second application of fertiliser (2 years) and the
logging age (9 years) (Table 1). They were located
within a 1 km radius area in a plateau with homoge-
neous soils. The native savannah was ploughed with
a disc-harrow and the stands were planted at 5 m  3 m
spacing (666 trees ha1). A starter fertiliser (150 g
N–P–K 13–13–21 per cutting) was applied and chemi-
cal sprays used to control weeds.
In order to check the validity of this chronose-
quence, a height growth curve was established for
the 9-year-old stand, from measurements made when
this plantation was 10, 63, 88 and 108 months old. The
mean height of the different stands of the sequence
was very close to the curve and the hypothesis that age
is the main variable among stands can be accepted
(Fig. 1). The mean height of the 6.5-year-old stand
studied by Laclau et al. (2000a) is also close to this
curve (Fig. 1).
46 J.-P. Bouillet et al. / Forest Ecology and Management 171 (2002) 43–57

Table 1
Main characteristics of the chronosequence stands

Stand age 3 months 6 months 1 year 2 years 9 years

Mean height (m) 1.4 1.8 5.5 11.8 25.1

Mean CBH (cm) – – 26 34 60
Date of measurements June 1999 September 1999 June 2000 April 1999 March 1999
Silvicultural treatments Savannah burned and ploughed. Starter fertilisation (150 g per tree cutting of N–P–K 13–13–21).
Chemical weeding

2.3. Methodology plane under study. The number of root intercepts

The spatial distribution of roots was assessed using
the methodology described by Laclau et al. (2000a).
One tree with a basal area corresponding to the mean
basal area of the stand was selected. Excavation was
performed and three vertical planes perpendicular
to the planting row at different spacing from this
representative tree were studied: under the stump
(P0), 0.75 m from the stump (P1), and 1.50 m from
the stump (P2), corresponding to half of the tree
spacing in the planting row (Fig. 2). Vertical profiles
were divided into 5 cm  5 cm grid cells and in each
grid cells roots were exposed using a small knife to
remove surrounding soil. We used the root intersect
method in this study (Tardieu, 1988). By root intersect
we mean any intersection of a root with the vertical
within three diameter classes were counted in each
grid cell of 25 cm2 (fine roots between 0.1 and 1 mm,
medium-sized roots between 1 and 10 mm, and coarse
roots over 10 mm). The soil profiles were 2.50 m wide
(half the distance between the planting rows) and 3 m
deep, except for the two youngest stands (1.5 m deep).
The 6-month-old stand was measured at the end of the
dry season. The other observations were carried out at
the end of the rainy season or the beginning of the dry
season (Table 1).
The root measures were made on a single tree in
each age category, because of the labour intensive
methodology adopted. So, within age stand variability
in root distribution was not estimated. But a previous
study showed that the mean length of shallow roots
was around 5 m in a 1-year-old stand and that some

Fig. 2. Location within the stand of the three profiles P0, P1 and P2, according to the location of the representative tree, the tree spacing in the
planting row and the distance between the planting rows.
 J.-P. Bouillet et al. / Forest Ecology and Management 171 (2002) 43–57
roots could reach 12 m at 6 years old (Hamel, 1992).
Thus, we consider that the spatial distribution of roots
observed represented that of the stand and not just of
the nearest tree.
2.4. Statistical analyses
The number of intercepts of fine and medium-sized
roots per area of 25 cm2 of soil are presented as fine
root density (FRD) and medium-sized root density
(MRD), respectively. Coarse roots were not taken into
account because they were only found in the profile P0
below the stump. In each profile, FRD and MRD
distribution were studied, according to the spacing
from planting row and depth. Two statistical tools
were used to map FRD and MRD:
(i) Automatic classification was performed in each
profile (procedure TREE using SPLUS software).
This technique enables an explicative model of
root density to be built according to the spatial
co-ordinates of the values observed. The algo-
rithm divides the profile into areas of low
variability for the variable by successive dichot-
omy on both co-ordinates, detecting the threshold
at which the criterion of deviance is maximised
(Breiman et al., 1984).
(ii) Kriging was performed in each profile (ISATIS
software). This method is based on the experi-
mental variogram expressing the variance of the
values measured, according to the spacing and
direction between measurements (Goovaerts,
1997). After modelling the variogram, kriging
allows an assessment of the values which were
not measured and smoothes the data, thus
preserving the main spatial features of the
variable (independence, anisotropy, etc.).
Additionally, the GLM procedure of the SAS soft-
ware (SAS Institute, 1988) was used to analyse the
variance of FRD. The statistical model used to
describe the FRD was
FRDijk ¼ m þ Li þ Pj þ Ak þ yij þ yik þ yjk þ eijk ;
where FRDijk is the mean value of measured at layer i
in profile j at age k; m the overall mean; L, P, A account
for the effects of depth (layer), distance from the
stump (profile), and age, respectively; yij the interac-
tion between layer i and profile j; yik the interaction
47
between layer i and age k; yjk the interaction between
profile j and age k; and eijk the residual effect.
In order to predict the change in FRD distribution
with depth for each age, a model was adjusted to FRD
measurements, using the NLIN procedure of the SAS
software. The model used was FRD ¼ a0  a1 depth þ
b expðc depthÞ þ e, where a0 þ b is the FRD at
depth ¼ 0; a0  a1 depth tend to 0 when depth
increases; c controls the curve’s shape, and e is the
residual error.
3. Results
Although MRD was 10–30 times lower than FRD,
its spatial distribution was similar. Therefore, results
are presented only for FRD (fine roots being the
most important in regard to the water and nutrient
absorption), except for the 3- and 6-month-old stands
for which FRD was not measured because it was not
possible to separate the fine roots of the savannah from
those of the Eucalyptus stand.
3.1. Spatial distribution of the medium-sized roots
in the 3- and 6-month-old stands
In the 3-month-old stand all root intersects (11/11)
were confined to the surface soil layer (0–25 cm depth).
This proportion decreased to 11/15 in the 6-month-old
stand; the four other intersects were observed between
0.9 and 1.5 m deep.
The profile close to the tree stump (P0) had the most
intercepts. In this profile there were eight root inter-
sects at 3 months and 10 root intersects at 6 months.
Most root intersects were within 30 cm of the plant-
ing row (7/11 and 12/15 for the 3- and 6-month-old
stands, respectively). However, the rapid lateral exten-
sion of the root system should be emphasised because
four intersects were recorded between 1.25 and 2.50 m
from the planting row in the 3-month-old stand.
3.2. Spatial distribution of fine roots in the
1-year-old stand
In the 1-year-old stand, root intersects were
observed down to 3 m (Fig. 3). For P0, the surface
soil layer had a high density of roots (32% of the total
root intersects between 0 and 25 cm), and FRD
 48
J.-P. Bouillet et al. / Forest Ecology and Management 171 (2002) 43–57
Fig. 3. FRD automatic classification in profiles P0, P1 and P2 for the 1-, 2- and 9-year-old stands (FRD in root intersects per 25 cm2).
 J.-P. Bouillet et al. / Forest Ecology and Management 171 (2002) 43–57
Fig. 4. Kriged map of fine root distribution in profiles P0, P1 and P2 for the 1-, 2- and 9-year-old stands.

49
50 J.-P. Bouillet et al. / Forest Ecology and Management 171 (2002) 43–57
decreased with depth. This trend was less pronounced
in the two other profiles: the proportion of root inter-
sects in 0–25 cm layer was 19 and 16% in P1 and P2,
respectively. Regardless of the profile, soil >20 cm
depth had few roots with large volumes of soil without
any roots (Fig. 4).
There was large horizontal heterogeneity in FRD in
the 0–20 cm layer in P0 and P1. Automatic classifica-
tion (Fig. 3) showed that FRD varied from 0.3 to 2.0
root intersects per 25 cm2 in P0 and from 0.4 to 1.1
root intersects per 25 cm2 in P1. The profile P0 was
also characterised by higher FRD values under the
stump.
3.3. Spatial distribution of fine roots in
the 2-year-old stand
Although roots were observed down to 3 m, FRD
decreased sharply with depth in the three profiles
(Fig. 3). Regardless of the profile, the surface layer
had the highest density of roots. The number of root
intersects counted in the 0–25 cm layer was 32, 32 and
36% of the total in P0, P1 and P2, respectively. A high
spatial heterogeneity in root distribution was observed
under this layer (Fig. 4).
Fig. 5. Change with age in vertical distribution of fine roots: data averaged for profiles P0, P1 and P2.
FRD distribution in the P0 profile differed from that
in P1 and P2 profiles. Automatic classification for P0
(Fig. 3) showed that the highest FRD were close to the
stump, in the surface soil layer. A layer with inter-
mediate values extending to a depth of 60 cm was also
observed. By contrast, in P1 and P2, the highest values
in the surface layer were not observed close to the
planting row (shortest distance to the stump) and FRD
was constant under this layer.
There was a marked horizontal heterogeneity in
FRD in the 0–5 cm layer: automatic classification
showed FRD varied from 2.9 to 21.7 root intersects
per 25 cm2 in P0, from 3.7 to 10.4 root intersects per
25 cm2 in P1, and from 3.8 to 10.6 root intersects per
25 cm2 in P2. FRD decreased from the planting row to
the middle of the inter-row for this layer in P0, but not
in the two other profiles. There was no effect of
spacing from the planting row in the deeper layers.
3.4. Spatial distribution of fine roots in
the 9-year-old stand
Similar to the 2-year-old stand, FRD decreased
sharply with depth in the three profiles (Fig. 3). The
number of root intersects in the 0–25 cm layer was 42,
 J.-P. Bouillet et al. / Forest Ecology and Management 171 (2002) 43–57 51

45 and 53% of the total in P0, P1 and P2, respectively. Table 3

Change in FRD (root intersects per 25 cm2) with age and depth in
By contrast, the soil below a depth of 2 m had few fine
the three profilesa
roots (Fig. 4).
The main differences among the profiles was a Stand age Depth (cm) Profile, P0 Profile, P1 Profile, P2
much higher FRD in P0 under the stump (Fig. 3) 1 year 0–25 1.21 (1.44) 0.54 (0.79) 0.34 (0.67)
down to 1.15 m depth (1.2 vs. 0.2 root intersects per 25–100 0.33 (0.75) 0.24 (0.53) 0.20 (0.54)
25 cm2 for P1 and P2). Although less pronounced than 100–300 0.20 (0.42) 0.19 (0.48) 0.15 (0.64)
in the 2-year-old stand, there was also horizontal 2 years 0–25 2.49 (4.66) 2.62 (3.53) 2.69 (3.45)
heterogeneity in FRD in the 0–10 cm layer of soil 25–100 0.61 (0.87) 0.54 (0.83) 0.37 (0.67)
and no effect of spacing from the planting row as 100–300 0.44 (0.71) 0.50 (1.05) 0.45 (0.51)
determined by the automatic classification. In con- 9 years 0–25 2.60 (2.68) 2.05 (2.72) 2.83 (3.14)
trast, below this layer, FRD was higher under the 25–100 0.48 (0.82) 0.28 (0.57) 0.32 (0.78)
stump, in P0. 100–300 0.27 (1.07) 0.21 (0.53) 0.19 (0.78)
a
Standard errors are given in parentheses.
3.5. Effects of soil layer, profile placing and
stand age on FRD
The analysis showed significant effects of age and
Using the previous results, the analysis of variance depth (Table 2) and a significant age  depth interac-
took into account: tion ðP < 0:001Þ. FRD increased considerably
between 1 and 2 years. In contrast, FRD was not
 Three layers corresponding to the surface soil
significantly different at 2 and 9 years, but there
(between 0 and 25 cm depth) which is highly
was a trend of more fine roots in the 2-year-old stand
prospected by fine roots, an intermediate layer
(Fig. 5). We also observed that mean FRD decreased
(between 0.25 and 1 m depth) and the deeper layer
substantially with depth, from 1.93 root intersects per
(over 1 m in depth).
25 cm2 in the top soil layer to 0.38 root intersects per
 The three profiles P0, P1 and P2.
25 cm2 between 25 cm and 1 m in depth, and 0.29 root
 The 1-, 2- and 9-year-old stands, where FRD were
intersects per 25 cm2 beyond this depth.
measured.
The significant age  depth interaction illustrated
the differences in root densities of soil layers, accord-
ing to stand age (Table 3). There were proportionally
Table 2
more fine roots in surface soil as stands aged. The
Effect of age (year), profile (type) and depth (cm) on FRD (in root number of root intersects in the 0–25 cm of surface
intersects per 25 cm2)a was 24% of the total down to 3 m in the 1-year-old
stand, 33% in the 2-year-old stand and 47% in the
FRD
9-year-old stand. Two profiles of the latter stand also
Age had the highest FRD values in surface soil. In contrast,
1 0.38 a
the marked decrease in FRD in the deeper layers
2 1.19 b
9 1.03 b observed in the 9-year-old stand explained that the
total root intersects was lower than in the 2-year-old
Profile
P0 0.96 a
stand.
P1 0.80 a
P2 0.84 a
Depth 4. Discussion
0–25 1.93 a
25–100 0.38 a Results obtained from this study in several stands
100–300 0.29 b of a chronosequence were consistent with those
a
Letters indicate significant differences (threshold: 0.001) previously obtained in a 6.5-year-old stand. These
between modalities of factors using Bonferonni test. observations have revealed several major trends that
52 J.-P. Bouillet et al. / Forest Ecology and Management 171 (2002) 43–57
characterise the establishment and the extension of the
root system of Eucalyptus clonal stands, planted under
the conditions of the Pointe-Noire region.
4.1. Root system dynamics
The clone quickly developed its root system, med-
ium-sized roots being observed at 1.5 m from the
stump 3 months after planting, and roots explored
at least down to 3 m in the 1-year-old stand. These
results show that the growth rate of the medium-sized
roots in the upper soil layers was at least 1.6 cm per
day (average over a 3-month period) and approxi-
mately 0.8 cm per day (average over a 1-year period)
in the deeper horizons. Field rhizotrons have been
established to confirm these results. It was also shown
that in the stands older than 6 months FRD was not
significantly different among profiles. This indicates
that after a few months root development was not
limited to areas close to the stump, but to the whole
stand. These results are consistent with previous
observations made in Congo on the same clone
(Hamel, 1992), or on E: urophylla  E: grandis clones
(unpublished data).
Similarly, Jama et al. (1998) confirmed the fast-
growing characteristics of the root system of Euca-
lyptus stands in Kenya. They found that 11 months
after planting roots developed to 4 m depth. Roots of
Eucalyptus were observed in Brazil down to 3.9 m, 8
months after planting and down to 6.2 m on 2-year-old
stands (Pacheco and Louzada, 1991).
4.2. Root distribution with depth
The models adjusted to FRD measurements for the
each stand age showed the change in root distribution
during the stand rotation (Table 4 and Fig. 6).
Table 4
Change with stand age in the parameter estimates, coefficients of determination and RMSE of the model:
FRD ¼ a0  a1 depth þ b expðc depthÞ, with FRD in root intersects per 25 cm2, and depth in m
Stand age (year) R2 RMSE
1 0.935 0.080
2 0.994 0.101
6.5 0.972 0.195
9 0.970 0.173
a
All parameters were significantly different from 0.
Surface soil had high root density with the number
of fine root intersects in the 0–25 cm surface soil layer
representing 16–53% of the total root intersects (from
0 to 3 m) which depended on stand age and profile.
Consistent results were observed in the 6.5-year-old
stand studied by Laclau et al. (2000a), where this
percentage was 35% for the profile P1.
High densities of roots in the upper soil layers are
common in pure and mixed forest stands (Santantonio
et al., 1977; Nambiar, 1983; Fabiao et al., 1991, 1995;
Heilman et al., 1993; Fredericksen and Zedaker, 1995;
Mello et al., 1997; Jama et al., 1998; Misra et al.,
1998). This allows for efficient uptake of water and
nutrients from these surface layers which are char-
acterised by accumulation of organic matter, and by
large amounts of available nutrients (Tiarks et al.,
1998). This root pattern is particularly adapted to
the nutrient poor soils of Pointe-Noire where tree
growth is highly dependent on the soil organic matter
content and the production of mineral N (Bouillet
et al., 1999). The ability of Eucalyptus stands to
rapidly withdraw water and nutrients inputs has been
demonstrated by Laclau et al. (2000a) by comparing
the change in calcium concentrations between
throughfall and the soil solution. During the rainy
season the concentration in the solution collected at
a depth of 15 cm was around 30% of that measured
under the forest floor, and decreased up to 10% during
the dry season when there was little rain (less than
1 mm) and the calcium concentration in rainfall was
high.
Mean FRD values decreased sharply with depth and
were lower than 0.5 root intersect per 25 cm2 below
2 m, regardless of stand age. However, these scarce
roots play a crucial role in water uptake during the dry
season, when the soil moisture of the upper layers is
close to the wilting point (Laclau et al., 2001). Sana
a0a a1a ba ca
0.346 0.00091 0.641 0.078
0.629 0.00091 8.059 0.382
1.019 0.00316 5.140 0.161
0.401 0.00099 5.495 0.132
 J.-P. Bouillet et al. / Forest Ecology and Management 171 (2002) 43–57 53

Fig. 6. Change with age in vertical distribution of fine roots: models FRD ¼ a0  a1 depth þ b expðc depthÞ, fitted on data averaged for
profiles P0, P1 and P2. Data for the 6.5-year-old stand obtained from Laclau et al. (2001).

(1997) showed that water uptake in a 5.5-year-old
stand of the same clone, occurred down to a depth of
5 m during the dry season. Similar results were
obtained in Ivory Coast on deep Tertiary sandy soil
under adult oil-palm stands where roots exceeded 6 m
deep (Jourdan and Rey, 1997). The major role of deep
roots in supplying water to trees were also widely
reported in forest stands (Nambiar and Sands, 1992;
Dye, 1996; Roupsard et al., 1999).
The development of a deep root system could
explain that phosphorus is not a limiting factor for
the growth of the Eucalyptus plantations in the Pointe-
Noire region (Safou-Matondo and Bouillet, 1999), and
the low efficiency of use of this nutrient (Laclau et al.,
2000a). The concentration of available phosphorus in
the soil is not high (0.20 g kg1) but constant down to
6 m. Thus, the amount of this nutrient available for the
trees is large due to the considerable volume of soil
explored by roots. Moreover, the proportion of phos-
phorus taken up from the deep horizons could con-
siderably increase during the dry season when soil
moisture is higher in depth than in the surface soil
(Comerford et al., 1997).
4.3. Changes in root density with age
In a chronosequence of the same clone, Laclau et al.
(2000b) observed an increment in stand biomass
(aerial parts plus big roots) of 8.3 Mg ha1 in the first
year and 16.9 Mg ha1 in the second year. This greater
current annual increment coincided with an increase in
root density, which could be related to the higher
demand in water and nutrients. Between the first year
and the second year, the mean value of FRD increased
from 0.24 to 0.65 root intersect per 25 cm2. Other
authors reported, in the early growth stages, a close
relationship between the current annual increment in
aerial biomass and in fine root biomass (Misra et al.,
1998), or in medium-sized root biomass (Fabiao et al.,
1995; Misra et al., 1998). In our study, such relation-
ship was also observed in older stages when decrease
in root density in deeper layers between 6.5 and 9
years of age corresponded to a marked decrease in
growth (Mabounou, 2000).
The increase with age in the percentage of roots in
the surface layer was consistent with the changes in
nutrient cycling in this ecosystem. Litterfall starts at
54 J.-P. Bouillet et al. / Forest Ecology and Management 171 (2002) 43–57

the end of the first year after planting (Nzila et al.,
2002) and leads to a continuous accumulation in the
forest floor, and a slight enrichment of organic carbon
in 0–10 cm of soil (Trouvé, 1995; Bernhard-Reversat
and Loumeto, 1999). The increase in root density in
surface soil may result from an increase in the con-
centration of nutrients in the solution of these top soil
layers due to the decomposition of the forest floor
(Laclau et al., 2000b). Then the biological cycle
becomes more and more active allowing the trees to
feed mainly from the upper layers, while, in the young
stages, the deep layers might be proportionally more
prospected by roots to supply the nutrient requirement
of the stand.
4.4. Effect of the spacing from the planting row
A high variability in FRD was observed in surface
soil but no gradient of decrease from the planting row
to the middle of the inter-row. However the highest
FRD and MRD values were generally observed under
the stump. This trend is common for tree plantations
(Fabiao et al., 1995; Pagès et al., 2000), but Nambiar
(1983) obtained the opposite result in a radiata pine
plantation, where the rooting density tended to
increase towards the mid-point of the rows. In our
Eucalyptus stand, stemflow averaged only 1.5% of
throughfall, but the nutrient concentrations were much
higher in stemflow than in throughfall (þ300% for K
or þ500% for P and Mn) (Laclau et al., 2000b). So,
under the stump, roots proliferated in an area of
preferential flow of water carrying high concentration
of nutrients. A similar root pattern was observed by
Laclau et al. (2000a) in a 6.5-year-old stand, where the
root development under the stump and the planting
row was even higher than in our study. It could be due
to the soil decompaction resulting from the ripping of
the planting line performed in that stand. This assump-
tion is supported by the larger vertical ramification of
big roots close to the stump and in the ripping row
observed by Bouillet et al. (1997) in a 4-year-old stand
of the same clone.
Under surface soil there was also a strong variability
in root density. High MRD and FRD values were
measured in areas where the roots were grouped
together, whereas there were areas without any root
intersects. This pattern of distribution could be
explained by the existence of preferential pathways
for water infiltration, resulting from the severe hydro-
phobicity of the surface soil (Laclau et al., 2001).
These eucalypts might be able to concentrate their fine
roots under these preferential drainage channels,

Fig. 7. Time-course of soil water content in the 15–50 cm soil layer, simulated with a daily water balance model as compared to TDR
measurements (replicates of five probes). Soil water measured from 225 to 547 Julian days.
 J.-P. Bouillet et al. / Forest Ecology and Management 171 (2002) 43–57
where high amounts of nutrient are transferred in
gravitational solutions.
4.5. Hydrological models
The results on FRD distribution have been used to
parameterise a hydrological bucket model (Granier
et al., 1999) applied for the 6.5-year-old stand. The
predicted water content compared well with the mea-
surements obtained with time domain reflectometry
probes (Laclau et al., 2001) (Fig. 7). The changes in
FRD distribution with age will be integrated into an
age-dependent hydrological models in order to estab-
lish water budgets for the whole rotation.
4.6. Silvicultural practices
This study suggests that weeding should not be
performed by ploughing because large amounts of
nutrients are likely to be lost by deep drainage as a
consequence of the destruction of the superficial root
mat. Furthermore, when medium-sized roots are cut,
they regenerate only one small root without ramifica-
tion (Bouillet et al., 1997). Fertiliser should be placed
close to the stump in the earlier growth stages. Mid-
rotation fertiliser can be broadcast, but location close
to the stump or along the planting row might be better.
5. Conclusion
In Congo the clonal Eucalyptus plantations are
established on soils characterised by low nutrient
content and low water retention capacity. Moreover,
rainfall is low and irregularly distributed with a dry
season of about 5 months. Despite these ecological
conditions, several factors may explain their fairly
high productivity. The trees quickly develop an exten-
sive root system which enables a rapid access to water
and nutrients. One year after planting the root system
extended to depth beyond 3 m, although the higher
densities of fine and medium-sized roots were
observed close to the surface. The change with age
in the percentage of roots in the top layers might be
related to the increasing role of biological cycles in the
nutrition of the stand. Some of the nutrients taken up
by the trees in the deep layers might be recycled
through mineralisation of the forest floor. The trees
55
then would absorb more and more nutrients from the
upper horizons as the stand ages. A marked horizontal
heterogeneity was observed with more root prospect-
ing under the stump and under areas of preferential
drainage. This root distribution allows rapid nutrient
uptake from soil solution, and reduces nutrient loss by
deep drainage.
Further research is needed to confirm and extend
these results. In particular, works including more trees
and intermediate ages will focus on the influence of
the season, vegetal material or stand location on the
development of the root system.
Acknowledgements
The founders of UR2PI (Republic of Congo, CIRAD-
Forêt and ECO SA) are gratefully acknowledged for
providing the financial support for this research project.
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