European Journal of Agronomy 13 (2000) 111 124

www.elsevier.com/locate/eja

Modelling vertical and lateral seed bank movements during

mouldboard ploughing
Nathalie Colbach a,*, Jean Roger-Estrade b, Bruno Chauvel a, Jacques Caneill c
a

Unite dAgrononomie-Malherbologie, INRA, 17 Rue Sully, BV 1540, 21034 Dijon Cedex, France
b
Unite dAgronomie, INRA-INA PG, 78850 Thi6er6al-Grignon, France
c
Departement des Sciences et Techniques Agronomiques, ENESAD, 26 Bd du Dr Petitjean, 21036 Dijon Cedex, France
Received 19 February 1999; received in revised form 12 August 1999; accepted 27 October 1999

Abstract
The vertical distribution of weed seeds in the soil is of fundamental importance because seedling emergence depends
on seed depth. The lateral displacement of the earth during mouldboard ploughing contributes to the dispersal of the
weeds inside the tilled field. In order to model vertical and lateral seed displacements during ploughing, an existing
model describing soil particle movements for different ploughing characteristics (depth and width) and soil structures
was tested on a multilocal field trial. The trials were carried out in 1996 and 1997 and comprised two soil textures
and three soil structures; tillage was performed with a mouldboard plough at varying ploughing widths and depths.
Seeds were simulated by beads that were introduced immediately before ploughing with an auger at different depths
and lateral positions (relative to the future passage of the coulter) within and just below the ploughed horizon. Lateral
displacement and the final vertical position of the beads were measured and compared to the simulations obtained
with the model. The model correctly simulated the final vertical seed co-ordinate and lateral seed displacement as a
function of soil structure, ploughing width and depth and initial seed position, if ploughing depth is lower than
ploughing width. If, however, the former exceeds the latter and/or if the furrows are not properly rotated, the model
does not simulate the seed movements correctly. The model was then used to calculate seed transfer matrices
describing vertical seed movements between seed bank layers for different conditions and plough modes and to
determine the optimal ploughing mode for a given seed bank distribution. For instance, if most seeds are located in
the top layer ploughing should be as deep as possible, with a low depth to width ratio to maximise soil inversion and
seed burial. If, however, the seeds are concentrated in the bottom layer, the model can be used to decide how
shallowly to plough in order to avoid disturbing the deeper seeds and what ploughing width to associate to this depth
in order to minimise soil inversion and leave as many seeds as possible undisturbed. Ways of improving the model
are suggested, particularly the necessity to simulate the effect of a skim coulter. 2000 Elsevier Science B.V. All
rights reserved.
Keywords: Mouldboard ploughing; Soil structure; Weed management; Seed bank; Seed dispersal

* Corresponding author. Tel.: + 33-3-80693033; fax: + 33-3-80693222.

E-mail address: colbach@dijon.inra.fr (N. Colbach).
1161-0301/00/$ - see front matter 2000 Elsevier Science B.V. All rights reserved.
PII: S 1 1 6 1 - 0 3 0 1 ( 0 0 ) 0 0 0 6 9 - 1

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N. Colbach et al. / Europ. J. Agronomy 13 (2000) 111124

1. Introduction
One of the most important reasons for soil
tillage is weed management (Moss and Clarke,
1994). Among the various possible soil tillage
operations, mouldboard ploughing is widely used
in most European cropping systems. In weed
management, mouldboard ploughing is of special
interest because of its important effect on the
vertical distribution of the seeds in the soil. The
vertical seed bank distribution is of fundamental
importance because seedling emergence either decreases continuously with seed depth (FroudWilliams et al., 1983; Dyer, 1995) or increases
with slight burial and then decreases at greater
depth (Mohler and Galford, 1997). Simultaneously, the lateral displacement of the earth during ploughing contributes to the dispersal of the
weeds inside the tilled field.
It is, therefore, essential to improve our understanding of the effect of mouldboard ploughing
on seed bank movements in order to define efficient soil management rules for weed control,
leading to a decrease in chemical herbicides. This
is the reason why many weed demography models
include sub-models illustrating the effect of soil
tillage on seed bank dynamics (Aarts, 1986; Doyle
et al., 1986; Jordan et al., 1995). Many of these
sub-models are either based directly on the work
of Cousens and Moss (1990) or developed by
similar methods and include a quantification of
the vertical seed bank movement during ploughing. Cousens and Moss divided the seed bank of
the tilled horizon into four horizontal sub-layers
and estimated the proportion of seeds moved
between layers during mouldboard ploughing.
This model was deduced from statistical relationships observed in one experimental situation.
Parameters well known to have a great effect on
soil displacement during ploughing such as
ploughing depth or width (Henin et al., 1969;
Kouwenhoven and Terpsta, 1972) or pre-tillage
soil structure (Coulomb et al., 1993) were not
taken into account. It is thus difficult to extrapolate Cousens and Moss model to other soil textures and structures or to variations in tillage
depth or width.

No mechanistic model has yet been developed

specifically for weed seed movements, but RogerEstrade and coworkers (Roger-Estrade, 1995;
Roger-Estrade and Manichon, 1998) proposed a
model for vertical and lateral movements of soil
particles, depending on their initial vertical and
lateral position, on ploughing depth and width as
well as soil structure. Consequently, the objectives
of this paper were: (1) To evaluate the suitability
of this model to predict weed seed movements in
the soil and, therefore, the distribution of seeds in
the seed bank, a multilocal field trial was set up to
observe seed movements under various conditions
and to compare these observations to the simulations obtained with Roger-Estrades model. As
this model is not limited to the most relevant
variable for weed seed position, i.e. vertical seed
displacement, but integrates both vertical and lateral movements, observations and subsequent
evaluations must, of course, take into account
both dimensions. (2) To calculate vertical seed
transfer matrices, like those established by
Cousens and Moss (1990), for different conditions
and plough modes and to determine the optimal
ploughing mode for a given seed bank distribution. This second objective is only feasible if the
model is deemed acceptable for weed seed movements.

2. Material and methods

2.1. Modelling seed displacement during ploughing

(Roger-Estrade, 1995; Roger-Estrade and
Manichon, 1998)
To model the seed displacement during ploughing, the representation of the furrow movement
during ploughing shown on Fig. 1 was used. The
principle of the furrow rotation first appeared in
the literature in Bousfield (1880) and has since
then been adopted by numerous authors (Ashby,
1934; Sohne, 1959; Henin et al., 1969). RogerEstrade improved this model and introduced it
into a larger model describing the changes in soil
structure under the influence of cropping systems
(Roger-Estrade, 1995).

N. Colbach et al. / Europ. J. Agronomy 13 (2000) 111124

In the plane perpendicular to the direction of

the plough, the furrow of soil cut by the mouldboard plough follows the movement described in
Fig. 1. This movement comprises two successive
rotations of the furrow and ceases when the furrow is settled on the previously rotated furrow
(Fig. 1A). The inclination angle between the furrow and the plough pan only depends on ploughing width and depth, i.e. the sine of this angle
equates to the ratio of tillage depth to width.
Actually, the furrow breaks up during this movement and partially falls on the plough pan. This
phenomenon is modelled by Roger-Estrade by
separating the furrow into slices which slide
downwards until they reach the plough pan (Fig.
1B). The number of slices depends on the mechanical soil behaviour: it is low in the case of poor
fragmentation when the ploughed soil is dry or
compacted; and it increases with the fragmenta-

113

tion of the soil, when ploughing occurs in good

moisture conditions and/or when the ploughed
soil is uncompacted. Using this relationship it is
possible to calculate the final vertical and lateral
co-ordinates of any point of the furrow as a
function of its co-ordinates before ploughing and
of ploughing depth and width as well as soil
structure.

2.2. The field trials

To evaluate the above described model field
trials were set up in two situations: (a) The field
chosen at the INRA experimental station in the
Dijon area in 1997 (52% E, 4720% N) was on an
eutric cambisol (FAO). The texture of the
ploughed horizon (030 cm) was: clay 39%, silt
55% and sand 6%. The field had been cropped for
several years with small grain cereals that were

Fig. 1. Soil movement during ploughing according to Roger-Estrade (1995) explained as a succession of a rotation of the whole
furrow (A), followed by a breakup into slides and their translation, with the number of slides decreasing with soil compaction (B).

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N. Colbach et al. / Europ. J. Agronomy 13 (2000) 111124

Fig. 2. Profile view of the beads introduced with an auger into the ploughed layer and their relative position to the future passage
of the coulter. The beads marked
are located beneath the ploughed layer and are not moved by the plough.

always sown and harvested in dry conditions,

inducing a low risk of compaction. (b) The second
field, used in 1996, was located at the INRA
experimental station in Grignon (158% E, 4851%
N). The soil was an orthic luvisol (FAO) and the
texture was: clay 26%, silt 58% and sand 16%. For
the last two decades the crop rotation had been
maize/winter wheat. Therefore, one harvest in two
took place in autumn when conditions are frequently wet, thus inducing a high risk of soil
compaction. The aim of this choice was to obtain
two contrasted types of soil behaviour during
ploughing, with an uncompacted, fragmentary
soil structure in Dijon and a compacted structure
in Grignon. In order to extend this range of soil
structures and mechanical soil behaviours, an extreme, severely compacted situation was created
on one part of the Dijon field (later on called
Dijon II as opposed to Dijon I, i.e. the uncompacted part of the field) by rolling the whole area
in wet conditions with a heavy tractor, just before
ploughing.
The initial soil structure was assessed just before ploughing. Three-metre-wide and 50-cm-deep
observation pits were dug perpendicular to the
tillage direction, and the soil structure of the
ploughed layer was described using the method
proposed by Manichon (Manichon, 1982, 1987).
This method is based on the description of the
morphology of the clods created by the action of
tillage tools. The clod size, their distribution and
internal structural porosity are evaluated in situ.
Mean bulk density of the ploughed layer was also

measured with a rubber balloon type density apparatus with a piston.

Seeds were simulated by cubic plastic beads (of
about 1 mm3) that are more easily observed and
recovered than weed seeds while being similarly
dispersed by ploughing (Rottele and Koch, 1981;
Moss, 1988). Immediately before ploughing, these
beads were mixed with soil and introduced with
an auger (diameter 5 cm) within and just below
the ploughed layer (Fig. 2). Every 5 cm down to a
depth of 30 cm in Grignon and 40 cm in Dijon, a
different bead colour was used, with a total of six
to eight colours depending on the location. Furthermore, beads of yet another colour were dispersed on the soil surface to simplify the
recognition of the limits between adjacent furrows
after ploughing. Each vertical hole resulting from
the auger was considered as a replication and six
(Grignon) to eight (Dijon) replications were made
(for each structure location), introducing the
beads every w+ 5 cm (w= plough width) in the
direction perpendicular to the future tillage direction, thus resulting in different lateral positions
relative to the future passage of the plough, with
only one replication per future furrow. The deepest beads were not moved by tillage, and marked
the initial lateral position of the beads.
Soil tillage was performed with a three-bottom
mouldboard plough (Huart 370E with a universal-type mouldboard) without a skim-coulter,
with two adjoining passages at Grignon and four
per soil structure at Dijon. The ploughing width
was 40 cm (16 in.) in Grignon and 35 cm (14 in.)

N. Colbach et al. / Europ. J. Agronomy 13 (2000) 111124

in Dijon. Soil moisture at ploughing was measured by randomly choosing a dozen soil samples
from the freshly ploughed furrows and calculating
the ratio of their dry weight to their fresh weight;
mean soil moisture was 30% (S.D. 1.8%) at Dijon
and 24% (S.D. 1.5%) at Grignon.
After ploughing, a 50-cm-deep pit was dug
perpendicular to the tillage direction, immediately
in front of the original position of the beads. The
pit covered the complete width of the tilled field.
The form and location of the displaced furrows
were drawn following the procedure described by
Coulomb et al. (1993). The actual ploughing
depth was measured for each furrow. The soil was
then removed in the direction of the tillage to
locate the initial position of the beads, marked by
the unmoved beads located below ploughing
depth. The actual initial lateral position (relative
to the passage of the coulter) was measured for
each vertical hole. The removal of the soil was
continued to discover the new position of the
beads. Lateral displacement and the final vertical
position were then measured as shown on Fig. 1
and compared to the simulations obtained with
the model. The situation was slightly different in
Grignon where all lateral co-ordinates were measured relatively to a common origin and lateral
displacements were then deduced.

2.3. Statistics
The model was evaluated, using the formula
given by Mayer and Butler (1993) for the coefficient of determination or modelling efficiency:
%(zi zi )2
2

r = 1
%(zi zi )2
where zi are observed values (with mean zi ) and zi
simulated values. Another quality indicator which
is often used in statistical literature is the meansquared error of prediction (MSEP); as the data
used to evaluate the model were independent of
the data used to develop the model, MSEP was
estimated as simply the average squared deviation
between the model prediction and observations
(Wallach and Goffinet, 1987, 1989). To obtain an

115

error measure of the same unit as both observations and simulations, the square root of MSEP
was used.

3. Results and discussion

3.1. E6aluation of ploughing model

3.1.1. Description of furrows after ploughing and
choice of model input 6ariables
Observations and measurements of the initial
soil structure verified that the produced experimental situations indeed ranked as wished, i.e.
with the most compacted structure at Dijon II,
the less compacted one in Dijon I and Grignon
being intermediate. In Dijon II the soil structure
appeared homogeneous, massive, without any apparent structural porosity; mean bulk density was
1.49 Mg m 3 with an S.D. of 0.03. Because of
this compacted soil structure the furrows were
nearly unfragmented. Therefore, only two translation slides were used in the simulations. In Dijon
I the soil structure was fragmentary, characterised
by the dominance of fine earth with some clods of
which the diameter did not exceed 5 cm; mean
bulk density was significantly lower than in Dijon
II (1.29 Mg m 3, with an S.D. of 0.10). Furrow
fragmentation was high enough to obtain a
smooth soil surface and little void between adjacent furrows; therefore, five translation slides were
used for the simulations. The Grignon profile
showed a spatially variable soil structure: fragmentary zones alternated with compacted soil volumes. The degree of fragmentation of the furrow
and soil surface roughness were intermediate between Dijon I and II. Consequently, simulations
were performed with three translation slides.
At Grignon the tillage depth was a uniform 25
cm and the furrows had been properly rotated so
that the inclination angle of the furrows (angle A
in Fig. 1) was about 40 as foreseen by the model.
At Dijon the measurements of the actual ploughing depth performed for every furrow showed that
the ratios of ploughing width to depth varied
between furrows. Consequently, furrow inclination also varied considerably. At Dijon II the
tillage depth ranged from 29 to 33 cm. Because of

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N. Colbach et al. / Europ. J. Agronomy 13 (2000) 111124

this large ploughing depth (relative to the ploughing width) the average inclination angle was close
to 90 (sin A close to 1). At Dijon I the ploughing
depth, ranging from 33 to 37 cm, exceeded in
some furrows the ploughing width. In this treatment the furrow inclination angle was close to 90
in most of the furrows. However, some furrows
did not complete their first rotation, either because the depth was too great to accomplish that
movement or because their rotation movement
was blocked by their too-deeply ploughed neighbour furrow. This situation probably resulted
from the fact that the plough worked in a very
compacted clayey soil, so that the actual working
depth of the plough was very difficult to control.
The input value for ploughing width was 40 cm
for the Grignon data and 35 cm at Dijon. Whatever the location, the input value for tillage depth
in the model was the measured ploughing depth
of each furrow when this value was lower than the
ploughing width. When this was not the case the
depth to width ratio exceeded 1 and it was impossible to calculate the sine; the inclination angle in
the model was then set at 90.
By removing the soil in the direction of the soil
tillage the furrows could be observed at various
longitudinal positions. No supplementary fissures
or variations in the inclination angle or in the
degree of fragmentation of the furrows at the level

of auger penetration were observed. Therefore,

the use of an auger to introduce the beads did not
seem to influence furrow rotation and distortion.

3.1.2. Analysis of model performances

At a first look, the final vertical co-ordinate
(the distance to the plough pan) and the lateral
displacement were not excessively well simulated
by the model (Table 1): modelling efficiency (r 2)
was only slightly higher than 0.6, mean error
(
MSEP) was rather large, even compared to the
observed range of variations. No systematic overor under-estimation (mean of residuals close to
zero) was found. Fig. 3, comparing observed and
simulated values for the final vertical co-ordinate
and the lateral displacement respectively, reinforces this first impression, showing large discrepancies between simulated and observed
values, even though the points were generally
distributed around the equation representing
equality of simulated and observed values.
If however, the replications located in those
furrows identified by the above described analysis
of furrow characteristics (i.e. too deeply ploughed
furrows or furrows that had been rotated by less
than 90) were eliminated, the model performance
increased dramatically (Table 1): in that case
modelling efficiency was high, mean error considerably decreased and again, there still was neither

Table 1
Evaluation of Roger-Estrades model by analysing prediction accuracy of lateral displacement and final vertical coordinate of
displaced beads. Synthesis of three situations: Dijon I, Dijon II (1997) and Grignon (1996)a
Case

Number of
points

Evaluated output
variable

Mean of
residuals (cm)

r2

MSEP
(cm)

All points

155

Lateral
displacement
Final vertical
coordinate

1.1

0.69

12.0

0.5

0.63

8.2

Lateral
displacement
Final vertical
coordinate

1.4

0.85

8.7

1.1

0.85

4.4

Elimination of furrows with ploughing depth

\width and without completed 1st rotation

73

a
Residual= zizi, where zi are observed values (with mean zi ) and zi simulated values; modelling efficiency r2 = 1((zizi )2/
(zizi )2) (Mayer and Butler, 1993); MSEP = (zizi )2/n with n =number of observations (Wallach and Goffinet, 1987, 1989); ",
mean significantly different from zero at a=5%.

N. Colbach et al. / Europ. J. Agronomy 13 (2000) 111124

117

Fig. 3. Comparison of final vertical seed coordinates (A) and of lateral seed movements (B) simulated by the mouldboard plough
model (Roger-Estrade, 1995) and observed on three field trials. Each point represents beads of a given colour and replication.

systematic over- nor under-estimation. If the

residuals were analysed separately for each location it appeared that the errors for lateral displacement were significantly higher at Grignon
(mean of absolute residual values =7.9 cm) than
at Dijon (4.5 cm). However, this particularity was
probably related to the different measurement
system used at Grignon where, whatever the

beads, all lateral co-ordinates were established

relative to one common origin, with an error risk
increasing with the distance from this origin.
These results show that Roger-Estrades model
correctly simulates the final vertical seed co-ordinate as well as lateral seed displacement as a
function of soil structure, ploughing width and
depth and, of course, initial seed position, if

N. Colbach et al. / Europ. J. Agronomy 13 (2000) 111124

118

ploughing depth is lower than ploughing width.

If, however, the former exceeds the latter and/or if
the furrows are not properly rotated, the model
cannot be used. This restricts the possible use of
the proposed model, but admittedly, under field
conditions, ploughing is usually performed with a
ploughing depth lower than the ploughing width.
The model gives the position of the seeds immediately after ploughing. At least in compacted
structures the soil settles considerably later on,
either because of superficial tillage for soil bed
preparation or because of climatic interference
such as alternation of dry and humid or cold and
warm conditions. The seed displacement model
for ploughing must, therefore, be completed by a
further model describing the degradation and
compression of the furrows after tillage.

3.2. Using the model for simulation of the 6ertical

weed seed distribution
3.2.1. Determination of 6ertical seed transfer
matrixes
As shown by the model evaluation, RogerEstrades model can be used to simulate seed
movements and final seed positions immediately
after tillage. Most existing weed demography
Table 2
Proportion of seeds moved from layer i to layer j by a
mouldboard plough (depth 20 cm; width: 30.5 cm) in case of
a seedbank divided into four 5-cm-thick horizontal layers
Final

Initial layer I

Layer j

Aa
1 (top)
2
3
4 (bottom)

0.02
0.11
0.40
0.46

0.21
0.27
0.30
0.21

0.37
0.26
0.20
0.18

0.29
0.10
0.12
0.48

Bb
1 (top)
2
3
4 (bottom)

0.24
0.26
0.26
0.24

0.24
0.26
0.26
0.24

0.24
0.26
0.26
0.24

0.24
0.26
0.26
0.24

According to Cousens and Moss (1990).

According to the plough model (Roger-Estrade, 1995) in
the case of a fragmented soil structure (ten translation slides).
b

models are only dealing with vertical seed transfers and positions as they do not simulate horizontal movements (Colbach and Debaeke, 1998).
Among the few authors who attempted to quantify the effects of soil tillage on vertical seed
movements, Cousens and Moss (1990) proposed a
compartmental model. In this model the tilled
horizon is divided into four 5-cm-thick horizontal
layers that are considered as compartments. The
seed content of one compartment j of the posttillage seed bank can be predicted from the seed
content of the four layers of the initial seed bank
and a vertical seed transfer matrix. Each coefficient of this matrix represents the proportion of
seeds of layer i moved to layer j during soil tillage.
Roger-Estrades model can be used to determine such vertical seed transfer matrices. In order
to compare the result with the model of Cousens
and Moss, similar tillage conditions, i.e. settled
soil (resulting from a high number of translation
slides), a plough depth and width of 20 and 30.5
cm, respectively, are used for the simulation.
Table 2 gives the proportions of seeds moved
between layers during soil tillage for the matrices
presented by Cousens and Moss and calculated
with Roger-Estrades model. It appears that this
model predicts a homogeneous distribution of the
seeds of each layer among the four tilled layers
whereas Cousens and Moss model foresees that a
large proportion of the initially superficial seeds is
buried in the two deepest layers. This is not
surprising as these authors added a skim-coulter
to their mouldboard plough, thus ensuring that
superficially located seeds, residues and soil clods
are buried close to the plough pan, whereas simply ploughing tends to distribute seeds more or
less homogeneously among the layers (Fig. 4).
This appears to be an interesting strategy in the
case of a field with a superficial soil layer heavily
infested by weed seeds where the aim is to limit
immediate seedling emergence. Therefore, the description of the soil and seed movements due to a
skim-coulter should necessarily be added to
Roger-Estrades model.
However, despite this deficiency, RogerEstrades model has several advantages over
Cousens and Moss model: in contrast to the
latter, the first uses ploughing depth and width as

N. Colbach et al. / Europ. J. Agronomy 13 (2000) 111124

119

Fig. 4. Seed distribution after ploughing in the case of a soil with a superficial weed seed infestation. Simulations were performed
with the vertical transfer matrixes proposed by Cousens and Moss (1990) or calculated with Roger-Estrades model (Roger-Estrade,
1995).

well as soil structure as input variables and is not

restricted to a 20-cm-deep four-layer seed bank.
Indeed, not only can different ploughing modes
such as deeper or wider ploughing be simulated,
but, much more importantly, the seed bank can
be divided into more numerous, thinner layers.
This is essential if the plough model is to be
introduced into models describing the demography of species such as blackgrass (Alopecurus
myosuroides Huds.) for that only seeds located
close to soil surface can successfully emerge and
give rise to seedlings and seed-producing adults
(Barralis, 1968; Naylor, 1972) whereas seed germination and mortality rates vary considerably with
seed depth (Barralis, 1970; Horng and Leu, 1978;
Ballare et al., 1988; Cussans et al., 1996).
The separation of the seed bank into horizontal
layers is easy in the case of highly fragmented soil
where the post-tillage soil surface is smooth. But
this separation is considerably more complicated
if the soil structure is compacted and the
ploughed soil surface rough, i.e. when furrows are
poorly fragmented. In this case (as on the
Grignon and Dijon II trials), the layers are

defined as shown on Fig. 5, i.e. depending on the

distance of the seeds to soil surface. The layers are
thus almost horizontal in the case of highly fragmented soil structure (Fig. 5A), they appear to be
more zigzagged when the fragmentation is limited (Fig. 5B). This procedure of subdividing the
seed bank appeared more relevant as most physical conditions that are important for weed seed
evolution depend on the distance to the surface.
For instance, for many weed species (Barralis,
1970; Bouwmeester and Karssen, 1989; Bai et al.,
1995; Benvenuti, 1995; Jensen, 1995) and even
some cropped species occurring as volunteers
(Pekrun et al., 1997a,b; Pekrun and Lutman,
1998), the amount and quality of light is essential
for the onset of germination and these factors
were shown to decrease with depth (Benvenuti,
1995).
Roger-Estrades model was then used to calculate vertical transfer matrices for 30-cm-deep seed
banks divided into 1-cm-thick layers. These matrixes were calculated for different soil structures
as well as ploughing depths and widths and then
applied to various initial seed distributions.

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N. Colbach et al. / Europ. J. Agronomy 13 (2000) 111124

3.2.2. Simulations
In the case of an initial superficial seed infestation, if the aim is to bury as many seeds as
possible, then ploughing is, of course, advised
instead of superficial tillage or direct drilling, but
soil structure influences the efficiency of this operation via its effects on the final vertical seed
distribution: in the case of a fragmented structure
(Fig. 6A) ploughing distributes the seeds homogeneously among the ploughed layers, regardless of
ploughing width. Hence, the proportion of seeds
found at a given depth only depends on tillage
depth; the deeper the ploughing, the less seeds are
found at a given depth (and, therefore, close to
the soil surface). The situation is not as simple in
the case of a compacted structure (Fig. 6B) where
ploughing width also influences seed distribution
and both tillage depth and width must be reasoned together. Indeed, Fig. 6B shows that the
deepest ploughing does not necessarily result in
the lowest superficial seed content and that, in
fact, a high ratio of ploughing width to depth
(with a high inclination of the furrow) is necessary
to bury superficial seeds. In contrast, in the case
of a low width to depth ratio (with a low inclination of the furrow) the superficial seed concentration after deep tillage can be as high as that after
a more shallow tillage with a high width to depth
ratio.

If most seeds are, however, located in the

deeper soil layers (Fig. 6C and D), then shallow
ploughing (or even superficial tillage) is advised to
limit superficial seed content, whatever the soil
structure. Again, seeds are distributed homogeneously among the ploughed layers in the case of
the fragmented structure whereas the seed profile
is highly irregular for the compacted structure
with, moreover, an influence on the ploughing
width. However, in contrast to the above described situation with an initially superficial seed
concentration, ploughing depth remains the most
important factor, even for compacted structures.
Indeed, if the layers containing the weed seeds are
not disturbed it is unlikely that these seeds are
carried back to the soil surface, except that some
movement can take place as a result of soil fauna
activity for instance, albeit on a small scale. If
though shallow tillage is not a possible option,
then at least ploughing with a low width to depth
ratio should be attempted to decrease the proportion of exhumed seeds.
In this discussion the aim of ploughing was to
minimise seed content close to the soil surface to
limit weed seedling emergence immediately after
tillage (Yenish et al., 1992). This is, however, not
always the objective of tillage, even when its
ultimate aim is weed control. If ploughing precedes the seeding of a crop by several months (as
in the case of clay soils to be sown with spring

Fig. 5. Subdivision of seed bank into layers before and after ploughing, depending on soil structure. Each layer comprises all the
points located at equal distance from the soil surface which is defined as the nearest part of the contour of the furrow. Thus, each
layer has the shape of the surface of the freshly ploughed field: horizontal in case of a highly fragmented soil or rough (zigzagged)
when fragmentation due to the plough is poor.

N. Colbach et al. / Europ. J. Agronomy 13 (2000) 111124

crops) in some cases it can be more advantageous

to maximise superficial seed content in order to
make as many seeds as possible emerge before
crop seeding and thus deplete the seed bank, i.e.
the stale seedbed technique (Leblanc and Cloutier,

121

1996). Such a strategy would, of course, only

work with relatively non-dormant seeds (such as
Poa annua L., Orlando et al. (1995)) that respond
to tillage by emerging immediately. This strategy
would, on the other hand, be disastrous in the

Fig. 6. Seed distributions before and after ploughing based on simulations performed with the vertical transfer matrixes calculated
with Roger-Estrades model (Roger-Estrade, 1995). (A) Case of a highly fragmented soil structure with a superficial weed seed
infestation. (B) Case of a highly compacted soil structure with a superficial weed seed infestation. (C) Case of a highly fragmented
soil structure with a deeply located weed seed infestation. (D) Case of a highly compacted soil structure with a deeply located weed
seed infestation.

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N. Colbach et al. / Europ. J. Agronomy 13 (2000) 111124

Fig. 6. (Continued)

case of species such as Polygonum persicaria L.

(Orlando et al., 1995) that emerge predominantly
in spring; the seed bank dormancy would have
decreased between tillage and seeding and the
seeds concentrated in the top layers would just be
ready for emergence at crop seeding.

4. Conclusion
The evaluation of Roger-Estrades model
showed that it is not only appropriate for soil
clod displacement, but is also adequate to predict
vertical and lateral weed seed bank movements

N. Colbach et al. / Europ. J. Agronomy 13 (2000) 111124

and to quantify soil tillage effects for a variety of

soil structures and ploughing modes. However,
this model does not foresee the use of a skimcoulter, a tool that would considerably improve
the burial of initially superficial weed seeds. Further studies are, therefore, currently being undertaken by the authors to model the effects of this
additional implement to the mouldboard plough.
Another necessary addition to this ploughing
model describing seed positions immediately after
tillage concerns the long-term seed movements
under the influence of superficial soil tillage and
climate. This especially concerns compacted soil
structures where important soil and, therefore,
weed seed movements occur when the formerly
compacted furrows break up and the soil settles
on the plough pan.
Despite these considerations the model can already be used to optimise soil tillage for weed
management by indicating the optimal ploughing
depth and width, depending on the soil structure
and the initial vertical seed distribution in the soil,
as shown by the various simulations. Furthermore, due to the quantification of the soil tillage
effects on vertical seed positions and to the model
ability to distinguish layers of varying numbers
and thicknesses, Roger-Estrades model can be
used to manage by soil tillage weed species with
contrasting germination and emergence requirements. To optimise this weed management the soil
tillage model should be combined with further
models describing biological processes such as
seed mortality, dormancy and seedling emergence.
These processes not only depend on vertical seed
position, but also on soil properties such as temperature, humidity, light penetration, oxygen content, etc., which are also influenced by soil tillage
(Mohler and Galford, 1997).
The range of possible soil tillage solutions
could, of course, also be increased by proposing
other techniques such chisel ploughing or various
superficial interventions. To evaluate the relative
performances and advantages of these different
tillage options, models similar to the ploughing
model would then be necessary for these other
tillage implements.
Roger-Estrades model also constitutes a first
step on the way to weed demography models

123

integrating intra-field variability as, besides vertical seed movements, lateral displacements are described. Therefore, it is now already possible to
build models describing lateral (perpendicular to
the direction of the soil tillage) intra-field variability and thus describe lateral weed dispersal and
variations in weed densities. To accomplish complete horizontal variability, both in the direction
and perpendicular to the direction of tillage and
other agricultural interventions, it is, however,
necessary to tackle longitudinal seed movements
during tillage.

Acknowledgements
The authors thank Jacques Troizier, Head of
the Experimental Centre at Grignon, and his
team, and Luc Biju-Duval and his colleagues of
the Experimental Station of INRA-Dijon, for
conducting the field trials.

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