Plant Soil (2009) 324:1–30

DOI 10.1007/s11104-009-0159-y

MARSCHNER REVIEW

Desirable plant root traits for protecting natural

and engineered slopes against landslides
Alexia Stokes & Claire Atger &
Anthony Glyn Bengough & Thierry Fourcaud &
Roy C. Sidle

Received: 26 February 2009 / Accepted: 26 August 2009 / Published online: 23 September 2009
# Springer Science + Business Media B.V. 2009

Abstract Slope stability models traditionally use
simple indicators of root system structure and strength
when vegetation is included as a factor. However,
additional root system traits should be considered when
managing vegetated slopes to avoid shallow substrate
mass movement. Traits including root distribution,
length, orientation and diameter are recognized as
influencing soil fixation, but do not consider the spatial
and temporal dimensions of roots within a system.
Thick roots act like soil nails on slopes and the spatial
Responsible Editor: Yongguan Zhu.
A. Stokes (*)
INRA, UMR AMAP,
34398 Montpellier Cedex 5, France
e-mail: alexia.stokes@cirad.fr
C. Atger
Pousse Conseil, Domaine de Fitzgerald, Le Mas Rouge,
Chemin du Mas Rouge,
34970 Lattes, France
position of these thick roots determines the arrange-
ment of the associated thin roots. Thin roots act in
tension during failure on slopes and if they traverse the
potential shear zone, provide a major contribution in
protecting against landslides. We discuss how root
traits change depending on ontogeny and climate, how
traits are affected by the local soil environment and the
types of plastic responses expressed by the plant. How
a landslide engineer can use this information when
considering slope stability and management strategies
is discussed, along with perspectives for future
research. This review encompasses many ideas, data
and concepts presented at the Second International
Conference ‘Ground Bio- and Eco-engineering: The
Use of Vegetation to Improve Slope Stability—
ICGBE2’ held at Beijing, China, 14–18 July 2008.
Several papers from this conference are published in
this edition of Plant and Soil.
Keywords Soil cohesion . Root architecture .
Slope stability . Soil mass wasting . Suction

A. G. Bengough
Scottish Crop Research Institute, What is a trait?
Dundee DD2 5DA, UK

T. Fourcaud A trait is defined as a distinct, quantitative property of

CIRAD, UMR AMAP,
34398 Montpellier Cedex 5, France
R. C. Sidle
Appalachian State University, Department of Geology,
P.O. Box 32067, Boone, NC 28608, USA
organisms, usually measured at the individual level
and used comparatively across species. A functional
trait is one that strongly influences organismal
performance (McGill et al. 2006). Plant quantitative
traits are extremely important for understanding the
2 Plant Soil (2009) 324:1–30
local ecology of any site. Plant height, architecture,
root depth, wood density, leaf size and leaf nitrogen
concentration control ecosystem processes and define
habitat for other taxa (Westoby and Wright 2006). An
engineer conjecturing as to how plant traits may
directly influence physical processes occurring on
sloping land needs to consider how, for example,
canopy architecture and litter properties influence the
partitioning of rainfall among interception loss,
infiltration and runoff. Plant traits not only influence
abiotic processes occurring at a site, but also the
habitat for animals and invertebrates. Depending on
the goal of a landslide engineer or restoration
ecologist, the immediate and long-term effects of
plant traits in an environment must be considered if a
site is to remain viable and ecologically successful. To
stabilise a slope against shallow landslides, vegetation
can be used, as plant root systems fix soil against
slippage (Schiechtl 1980; Coppin and Richards 1990;
Norris et al. 2008). Although root systems are
designed to anchor the plant to the soil and provide
access to nutrients and water, an understanding of the
interactions between root traits and soil physical
processes would enable the landslide engineer to
better manage an unstable slope.
What types of mass wasting processes
can be stabilized by plant roots?
Woody vegetation, particularly trees, can help prevent
shallow landslides in two ways: (1) modifying the soil
moisture regime via evapotranspiration; and (2)
providing root reinforcement within the soil mantle
(Fig. 1). The first factor is generally not very
important for shallow landslides and debris flows that
occur during an extended rainy season, except
possibly in the tropics and sub-tropics where evapo-
transpiration is high throughout the year; deeper-
seated landslides, such as earthflows, may experience
slightly prolonged activity due to higher moisture
contents after vegetation is removed (Sidle and Ochiai
2006). In most temperate regions, soils are nearly
saturated and evapotranspiration is low during autumn
and winter rainstorms when shallow slope failures
typically occur, thus, soil water content is only
minimally affected by the small water losses attribut-
ed to evapotranspiration (Sidle et al. 1985). In
Canada, reduced evapotranspiration following log-
ging may increase pore water pressures during
moderate-sized winter storms, but for the large storms
in which landslides generally occur, differences in
pore water pressure due to logging were difficult to
detect (Dhakal and Sidle 2004a). However, evapo-
transpiration facilitated by trees could extend the
“window of susceptibility” for shallow landslides and
debris flows if a large storm occurred near the
beginning or end of the rainy season (Megahan
1983; Sidle et al. 1985). Also, when large and high
intensity storms occur during drier conditions, evapo-
transpiration may reduce the potential of shallow
landslides (Sidle and Ochiai 2006).
A much more significant contribution of woody
vegetation to the stability of shallow soils on steep
slopes is the additional soil strength or cohesion
attributed to root systems. In shallow soils, tree roots
may penetrate the entire soil mantle and anchor the
soil into more stable substrate (e.g., Wu et al. 1979)
(Fig. 1a). Dense lateral root systems in the upper soil
horizons form a membrane that stabilizes the soil
(e.g., Schmidt et al. 2001) (Fig. 1a), and larger tree
roots can provide reinforcement across planes of
weakness along the flanks of potential slope failures
(Sidle et al. 1985). Past research suggests that while
woody roots significantly reduce shallow (<1 to 2 m
deep) landslide potential on steep slopes, deeper soil
mantles (>5 m) benefit little from such reinforcement,
as root density decreases dramatically with depth and
few large roots are able to anchor across the basal
failure plane. The only benefit of root strength to the
stabilization of deep-seated landslides, e.g., earth-
flows and slumps, is when very large lateral woody
roots cross planes of weakness along the flanks of
potential failures (Sidle and Ochiai 2006). However,
such reinforcement would only benefit small deep-
seated landslides whereas lateral roots also offer
protection against most shallow landslides (Swanston
and Swanson 1976) (Fig. 1a). Basal woody vegetation
cover, organic matter, and dense, shallow root mats
also prevent surface mass wasting, e.g. dry gravel on
steep slopes (Mersereau and Dyrness 1972).
What information is used by engineers
when modelling slope stability?
Investigating the stability of vegetated slopes implies
taking into consideration the interaction among three
Plant Soil (2009) 324:1–30
Fig. 1 a Conceptual illustration of a potential slip plane at the
soil—bedrock interface of a hillslope, dynamic shallow
groundwater table (landslide triggering mechanism), and
stabilizing components of root systems (lateral reinforcement
within the upper soil and anchoring of shallow soil mantle to
bedrock). b A herringbone root system with one main axis on
which are borne first order lateral roots. c A dichotomous root
system with two external root tips borne on every lateral. 2nd
and 3rd order lateral roots are present. Magnitude is defined as
the number of root tips (n=16 in both cases), and altitude the
longest path from root base to a root tip. Depending on the
3
branching pattern, the same volume of soil is not exploited
equally, even when root volume and magnitude is the same in
both cases. Dashed lines indicate the potential shear zone, the
thickness of which can be influenced by the number of roots
crossing through it, root length and branching angle. The
circled area illustrates how root proliferation in e.g. a nutrient
rich patch can influence root system architecture near a slip
surface. In b), initiation and emergence of a lateral root on the
upper side of the mother root can occur if the latter encounters
an obstacle
4 Plant Soil (2009) 324:1–30
physical or biological systems, i.e. soil, water and
plants, through a multi-disciplinary approach (Coppin
and Richards 1990). However, there are very few
models that implement all these aspects as a whole,
and even fewer which consider the time evolution of
these interactions.
Modelling the mechanical reinforcement of soil
by roots
The effect of roots on soil fixation has been reported
by several authors, but quantifying the gain in soil
shear strength is difficult to achieve. Pioneering
modelling work by Wu (1976) and Waldron (1977)
have introduced the root mechanical contribution as
additive soil cohesion in the Coulomb’s failure criteria
using a simple mechanistic model. The additional
cohesion at the slip surface was defined by two
variables: the average root tensile strength and root
area ratio (RAR, or the fraction of a plane of soil
occupied by roots). It was assumed that roots are
initially perpendicular to the slip surface and bend
according to the relative displacement of soil on both
sides of the shear zone (see Fan and Su 2009). The
tangential component of root tensile force thus
directly contributes to the increase in soil shear
strength, whilst the normal component augments the
confining pressure. These first models of soil rein-
forcement have been shown to overestimate the
additional cohesion due to roots in tension, as all
roots are assumed to break at the same time (Bischetti
et al. 2009a; Loades et al. 2009). A significant
improvement of this approach has been proposed by
Pollen and Simon (2005) and Pollen-Bankhead and
Simon (2009) who applied a Fibre Bundle Model to
rooted soils. This model considers that the root
network breaks progressively from the weakest to
the strongest roots, and that stresses of broken roots
are redistributed to the remaining elements. More
recent papers also pointed out the limitation of
assuming that roots are initially oriented perpendicu-
lar to the slip surface, and propose using root
architectural models as an improvement in slope
stability analyses (Reubens et al. 2007; Danjon et al.
2008).
In most models, roots are considered as very
flexible elements, thus limiting their application to
the finest roots. However, from a mechanical point of
view, a distinction must be made between fine and
thin roots, which behave like cable elements, i.e. with
a very low bending stiffness, and structural roots that
are similar to beams, developing longitudinal shear
stresses (Fournier et al. 2006; Reubens et al. 2007).
The orientation of structural roots near the slip surface
cannot be estimated using a simple geometrical rule,
as performed by Wu et al. (1979), due to flexural
rigidity that limits their deformations. Woody roots
play an important role in tree anchorage and their
mechanical interactions with the soil medium have
been previously investigated using finite element
models (FEM). These models take into account root
architecture and structural roots have been considered
as individual beam elements (Dupuy et al. 2005a, b;
Dupuy et al. 2007; Fourcaud et al. 2008). Few
numerical analyses have been carried out at a local
scale to quantify the effect of such root element
inclusions on soil shear strength (Wu 2007). Nor has
stiffness of structural roots been introduced in slope
stability models so far, except for the notable
exceptions of models developed by Nakamura et al.
(2007) and Tosi (2007).
In addition to root tensile strength, the pull-out
resistance of roots (Table 1) has been incorporated as
a criterion in slope stability models (Waldron 1977;
van Beek et al. 2005; Pollen-Bankhead and Simon
2009). The maximum force required to pull-out a root
is usually correlated with root diameter at the clamp
position, root length and/or total root biomass
including lateral ramification. However, pull-out tests
do not allow the root-soil interface behaviour to be
rigorously calibrated due to the difficulty in separat-
ing the effect of lateral root branching. Bond
resistance of a single root is defined by the product
between the root-soil interface shear strength and the
total root surface (Ennos 1990). The interface shear
strength can be approximated theoretically consider-
ing that shear resistance of the root-soil interface is
proportional to soil shear strength. The influence of
branching patterns on pull-out resistance in different
soil types has been investigated using numerical
(Dupuy et al. 2005a) and experimental (Stokes et al.
1996; Mickovski et al. 2007) approaches. These
studies have shown that root topology, branching
angle and branching density can significantly change
the distribution of stresses and plastic strains within
the soil medium, thus modifying resistance to pull-
out, but these aspects are not yet directly considered
in soil reinforcement models.
Table 1 Ranges (or means) of pull-out resistance for roots of different diameters and lengths

Species Soil type Range (mean) of root Range (mean) of Pull-out force range Range (mean) of equivalent References
diameter at clamp root length (mm) (mean) (N) failure stress at clamp
position or at breaking position or at position of
point (*) (mm) root failure (*) (MPa)

Picea sitchensis Bong. Carr. Brown earth (8.8) (351) (465) Anderson et al. 1989
Plant Soil (2009) 324:1–30

Picea sitchensis Bong. Carr. Peat (8.4) (359) (517) Anderson et al. 1989
Casuarina glauca Sieb. Brown loams and 0.56–17.23 (4.49) 150–200 9–2,148 3.34–81.68 (22.43) Docker and Hubble 2008
sandy loams
Eucalyptus amplifolia Naud. Brown loams and 0.45–16.43 (5.00) 150–200 9–2,482 8.88–130.65 (27.33) Docker and Hubble 2008
sandy loams
Eucalyptus elata Dehn. Brown loams and 0.21–15.93 (4.85) 150–200 2–2,357 8.57–198.06 (31.49) Docker and Hubble 2008
sandy loams
Acacia floribunda (Vent.) Willd. Brown loams and 0.31–13.33 (4.09) 150–200 10–2,969 11.19–217.89 (58.09) Docker and Hubble 2008
sandy loams
Triticum aestivum L. Sandy clay loam (1.38) (4.8) Easson et al. 1995
Triticum aestivum L. Sandy loam (1.38) (3.9) Easson et al. 1995
Helianthus annuus L. Sandy loam (1.8) 25–160 0.5–1.7 Ennos 1989
Allium porrum L. Dry sandy loam 0.35–0.45 3–40 0.02–0.28 Ennos 1990
Allium porrum L. Wet sandy loam 0.35–0.45 5–40 0.02–0.22 Ennos 1990
Alnus incana L. Sand and gravel 2.42–3.19 (2.76) * 250–357 (299) 23.5–39.2 (30.4)* Karrenberg et al. 2003
Populus nigra L. Sand and gravel 3.25–4.07 (3.64) * 436–517 (475) 26.4–37.3 (31.4)* Karrenberg et al. 2003
Salix elaeagnos Scop. Sand and gravel 3.33–3.97 (3.62)* 578–705 (638) 43.3–57.8 (50.1)* Karrenberg et al. 2003
Vetiveria zizanioides L. Silt 0.30–1.45 (1.02) 110–275 (219)a 190–620 (470) Mickovski et al. 2005
Crataegus monogyna Jacq. Brown sandy clay 7–62 (21.6) (1,290) 300–12,000 (2,880) 3–15 (8) Norris 2005
Quercus robur L. Brown sandy clay 2–9 (5.4) (360) 30–440 (150) 2–14 (7) Norris 2005
Pinus halepensis Silt 1–11 – – 3–24 (9) Van Beek et al. 2005
a
Rooting depth, as the whole plant was pulled-out
5
6 Plant Soil (2009) 324:1–30
Integration of root reinforcement models in slope
stability models
Root reinforcement models are used to determine the
contribution of vegetation to the factor of safety
(FOS) of a particular slope. The FOS is defined as the
ratio of the actual soil strength to the minimum shear
strength required for equilibrium. To calculate the
FOS of a vegetated slope, 2D or 3D numerical
techniques exist which couple soil mechanics, soil
hydrology and plant growth (Stokes et al. 2008), e.g.
limit equilibrium (Greenwood 2006), finite difference
(Wilkinson et al. 2002; van Beek et al. 2005) and finite
element (Kokutse et al. 2006) models. Vegetation must
be taken into account as reinforcement elements and
elements that modify the hydrological regime of the
slope (Simon and Collison 2002; Wilkinson et al.
2002; Dhakal and Sidle 2004b).
What are the traits useful for reinforcing soil
on slopes?
Although the plant root traits most commonly used by
engineers when modelling slope stability are tensile
strength and RAR, several other important traits
(Burylo et al. 2009) should also be considered
(Table 2). First however, the type of root must be
defined. Therefore, to better understand the different
processes governing soil fixation by roots, three classes
of roots will be referred to in this paper, depending on
their diameter (although these classes do not necessar-
ily reflect the developmental stage of a root, as the
diameter of roots at different ages of maturity will
depend on species):
– Fine roots (>0.0–2.0 mm). A major function of
these roots is to take up water and nutrients. If
roots are young and not yet woody, root hairs
may be present which aid uptake. The turnover of
fine roots is relatively rapid.
– Thin roots (>2.0–10.0 mm). In woody species
most roots in this class will be lignified. Turnover
is not known because some roots will become
thicker, depending on a certain number of
external and internal processes.
– Thick roots (>10.0 mm). These roots are
important for anchoring the plant to the soil
and preventing uprooting. The spatial position of
these roots will determine the position of the
fine and thin roots in the soil, and thus influence
indirectly nutrient and water uptake.
The first root (radical) which emerges from a seed
grows downwards and in some species, becomes the
taproot (Figs. 1b, c, 2) along which are borne lateral
roots of different orders (Fig. 1b, c). Even if this
taproot is removed through, e.g. injury, a new vertical
root will emerge or an existing lateral root will grow
downwards to replace the taproot, thus ensuring that
the functional role of the taproot is maintained
(Champagnat et al. 1974; Amin et al. 1987; Khuder
et al. 2007). In other species, the radical dies shortly
after germination and is never replaced. Roots are
borne on the root collar, or stem base (Fig. 2). In this
paper we shall refer to the first lateral roots emerging
from the taproot or root collar as first order roots,
lateral roots emerging from the first order roots as
second order roots and so on (Fig. 1b, c).
With regard to soil fixation by plant roots, the most
important trait to consider initially is rooting depth.
To stabilize a slope against a shallow landslide, roots
must cross the shear surface (Fig. 1) which can be up
to 2 m deep in the middle of the slope (Cammeraat et
al. 2005; van Beek et al. 2005; Norris et al. 2008).
Rooting depth is species dependent when soil
conditions are not limiting and root development
changes significantly with depth, although in general,
>80% of biomass is in the top 0.4–0.5 m of soil. The
number and orientation of roots that the shear surface
intersects will change significantly with rooting depth
for the same plant, even for distances of only several
cm (Fig. 1b, c). Roots vertically aligned to the
potential slip surface will be mobilised throughout
their length whereas those with a larger angle will be
more likely to slip before failure in tension occurs. A
greater number of roots and diversity of branching
angles will increase the thickness of the shear zone
(Abe and Ziemer 1991a; Mickovski et al. 2007).
Although the true angle of roots in the shear zone is
often ignored, and assumed to be between 40° and
70°, Greenway (1987) and Danjon et al. (2008)
showed that such assumptions lead to an overesti-
mation of the shear resistance contributed by roots.
Even for the same root, the angle at which it grows
with regard to the vertical can change several times
along its length. Due to the physiology of roots, a
Plant Soil (2009) 324:1–30
Table 2 Definitions and desirability of root traits useful to a landslide engineer
Trait Description
Root area ratio (RAR) The fraction of a plane of soil occupied by
roots
Tensile strength Resistance to breaking in tension
Root thickness Diameter of root
Shallow/deep rooting Inherent rooting depth
Root length density (RLD) Defined as either root length per unit volume
of soil or root length per unit surface of a
soil
Specific root length (SRL) Root length per unit root dry mass
Root angle Angle between mother and daughter root in
vertical or horizontal plane
Production of adventitious roots Roots initiated from the base of the stem, with
later roots emerging higher up the stem
Topology Physical organisation of root branching
Forking Forks appear when the growth potential of
one lateral root changes as a shift occurs in
the mother root’s functioning
Root clustering Proliferation of roots within a given volume
of soil
7
How can the trait be desirable for landslide
engineers?
The greater the RAR, the more soil shear
strength will be increased (Fig. 5b)
A higher resistance will enable a root to
mobilise its full strength during pull-out and
increase soil shear strength. Therefore, for a
given RAR, many small roots are more
desirable than a few thick roots
Thin roots are relatively stronger in tension.
Thickness is an indicator of root longevity
(thicker roots live longer), bending stiffness
and the ability to penetrate soil
If deeper growing roots cross the potential
soil shear surface, slope stability is
enhanced. Anchorage of roots to bedrock
will improve substrate fixation, but root
growth into cracks may cause cracks to
enlarge and result in substrate failure
Increasing root length augments the pull-out
resistance up to a critical length, from which
roots will break in tension instead of slipping
out of the soil. The water uptake rate from a
given horizon will increase with RLD. Along
with evapotranspiration rate, slope stability is
affected by soil water content, particularly in
tropical regions, therefore higher RLD is
desirable, especially in deeper horizons
High SRL implies more numerous thinner
roots and low SRL means less but thicker
roots (see ‘Tensile strength’ above). Is a
useful descriptor for herbs and perennials
where a high SRL is desirable
Soils containing roots with a range of
orientations develop wider shear zones and
can slowly mobilise reinforcement from
roots via their tensile strength even at large
shear displacements
Adventitious roots can grow into soil and
debris deposited on the upslope side of a
plant, thus fixing shallow soil layers
Root topology influences resource uptake and
efficiency and can significantly change the
distribution of stresses and plastic strains
within the soil medium, thus modifying root
resistance to pull-out. Dichotomous systems
are better anchored than herringbone sys-
tems
Production of forks will anchor more
superficial roots to the soil, especially if
forks grow vertically downwards
If roots are clumped within cracks and
biopores in the soil, the time required for
total water extraction increases because of
the greater distance that water must flow
8 Plant Soil (2009) 324:1–30

Table 2 (continued)

Trait Description How can the trait be desirable for landslide

engineers?

through the soil structural unit to the roots.

Root response to soil stresses
Root decay rate
Resprouting ability
Mycorrhizal interaction
Effect of adverse physical, chemical, or
biological conditions on root growth
Rate of dry mass loss per unit dry mass
initially present
The ability of a plant to produce a new shoot
from a root or stem if part of the plant’s
biomass has been removed
Symbiotic association between a fungus and
plant roots. Arbuscular mycorrhizas hyphae
enter into the root cells and ectomycorrhizas
consist of a hyphal sheath covering the root
tip and a hartig net of hyphae surrounding
cells within the root cortex
But if clusters of roots occur in resource-
rich patches, water uptake is faster around
the cluster compared to a zone without
roots, thus preferential water extraction
occurs from specific parts of the soil. A
homogenous distribution of roots within the
soil is more desirable
Physical—large variation in plant responses
to soil compaction, drought and
waterlogging
Chemical—certain genotypes may overcome
nutrient deficiencies or grow in presence of
toxic compounds
Biological—species dependent susceptibility
and resistance to disease
Roots which decay slowly fix soil for a longer
period. Root decay may create preferential
flow pathways in soil. Root decay rates vary
greatly according to root diameter, chemical
composition and species
Usually in perennials only. If a plant can
resprout, from either the shoot or root
system, it will be able to survive in a newly
disturbed environment e.g. in a landslide
deposition zone, even if above-ground
biomass is destroyed
Mycorrhizal infection usually enhances root
branching density. Tree species
predominately forming ectomycorrhiza can
have a higher branching intensity than
those forming arbuscular mycorrhiza.
Ectomycorrhizal infected roots have been
found to have a higher tensile strength.
Mycorrhizal inoculation may be desirable
on degraded soil

root branch can be initiated at any point along a
parent root, but not necessarily emerge fully from the
parent root. Emergence is largely controlled by auxin
fluxes and the bending of a root around an obstacle
or as it grows through the soil can trigger the
initiation and emergence of a lateral root (Fig. 1b,
Lucas et al. 2008).
Thick roots act like soil nails on slopes, reinforcing
soil in the same way that concrete is reinforced with
steel rods (Fig. 1a). The spatial position of these thick
roots also has an indirect effect on soil fixation in that
the location of thin and fine roots will depend on the
arrangement of thick roots. Thin and fine roots act in
tension during failure on slopes and if they cross the
slip surface, provide a major contribution to slope
stability (Fig. 1, Waldron and Dakessian 1982). Root
thickness is thus an important trait to consider, and is
also an indicator of root longevity (thicker roots live
longer), bending stiffness and the ability to penetrate
soil (Clark et al. 2008) as well as to store and
transport water (Roumet et al. 2006). For the landslide
engineer, one of the first considerations with regard to
root thickness will be how tensile strength changes
with diameter. Although thicker roots require more
force to be pulled out of the soil, when root strength is
calculated (force/root cross-sectional area), thinner
Plant Soil (2009) 324:1–30
a) b)
Fig. 2 Depending on the strategy type, root systems develop
differently for given species. In a short-lived and perennial
lateral roots are borne on the taproot (black) and in b directly
on the stem (shaded). Arrows show the direction in which the
root system develops
roots are significantly stronger than thick roots, due to
changes in cellulose content (cellulose is highly
resistant in tension) (Genet et al. 2005). Tensile
strength values can vary from 20 to 730 MPa for
roots in a diameter range of 0.15–4.5 mm (Bischetti et
al. 2005), this surprisingly high latter value for Fagus
sylvatica L. being stronger than that for steel (400–
700 MPa). A list of tensile strength values for 67
species is available in Stokes et al. (2008), but in a
model of slope stability, Danjon et al. (2008) used a
generic power equation of root tensile strength versus
diameter for several broadleaf and conifer species
(equation from Genet et al. 2005):
Tensile strength ¼ 28:97x
0:52
Genet et al. (2009) suggested that differences in
tensile strength among tree species have little influ-
ence on the calculation of a slope’s FOS and that root
distribution within the soil will affect slope stability
much more.
Root length is one of the most studied traits in root
systems and in herbaceous species is usually studied in
conjunction with biomass, resulting in a parameter
called specific root length (SRL). SRL is thus calculated
as the root length per unit root dry mass. This ratio will
provide information about the proportion of thin and
thick roots within a root system (high SRL implies more
numerous thinner roots and low SRL means fewer but
9
thicker roots). A higher SRL would therefore be
desirable, particularly at greater soil depths. Annual
species usually have a higher SRL than perennials,
possibly associated to a faster relative growth rate (RGR)
and metabolic activity (Roumet et al. 2006). The RGR
of a plant organ is calculated as the instantaneous rate
of dry mass increase per unit dry mass already present
(Wright and Westoby 1999). For a landslide engineer,
how fast a plant grows will be of particular importance
when a gap has occurred on a site after, e.g. tree fall,
thinning of a plantation or clear-felling (Sidle 1992).
The roots of plants remaining around the newly-created
gap, or those of naturally generated or planted nursery
seedlings, need to grow quickly to colonize the new
patch of soil if soil slippage is to be prevented on a
fragile slope. The inherent RGR of a species depends
largely on its life form and ecological niche: in
optimum conditions, plants from fertile habitats grow
faster than plants from nutrient poor or stressful habitats
and annuals grow faster than perennials. Although not a
trait in itself, RGR can be correlated with various traits,
e.g. SRL, although results from studies are conflicting.
Wright and Westoby (1999) found that in seedlings of
33 species growing down a rainfall gradient in
Australia, SRL decreased due mainly to an increase in
root diameter (a larger proportion of thicker roots was
present in lower rainfall areas) and was highly
correlated with RGR. Such a strategy could indicate
selection for increased efficiency of water uptake
during seedling establishment in low rainfall habitat
or possibly a greater diameter might confer an
enhanced ability to penetrate dry soil (Wright and
Westoby 1999). However, Poorter and Remkes (1990)
found no relationship between SRL and RGR in 24
species. If the investment of resources in fast and slow-
growing roots does depend on inherent RGR, one can
suppose that these differences may be reflected in root
tensile strength.
Although traits such as root diameter and SRL are
useful indicators of plant structure, they are only simple
descriptors of how a root system is configured. These
traits will also change depending on the root order
(Fig. 1), development stage and position within the
branched hierarchy of the root system (Eissenstat et al.
2000). To better understand how root systems occupy
soil over time and space, a landslide engineer needs to
consider the 3D root architecture of any given species.
This task is not easy, nevertheless, certain general-
izations can be made, based on developmental obser-
10
vations and a knowledge of plant ontogenetic processes
(the history of an organism from birth to death).
Root system architecture
The architecture of a plant depends on the nature and
the relative arrangement of each of its parts (see
Barthélémy and Caraglio 2007). From germination to
death, various well-ordered and definite develop-
mental stages can be characterized taking into
account the developmental and functional features
and the relative arrangement of the different parts.
Until fairly recently, quantitative data on root
architecture has been scanty not only due to the
difficulty in accessing roots, but also because the
morphological analysis of the root system is not as
efficient as that of its aerial counterpart. Root
systems do not bear morphological markers of
growth rhythms e.g. leaf nodes, bud-scales or even
well-defined annual growth rings, therefore temporal
changes cannot be dated with hindsight.
The main differences between plant root architec-
tural types are those which can be highlighted by
comparing woody with non woody species. By
assessing the initial rooting development, which is
largely under genetic control in the first stages
(Hermann 1977; Zobel 1996), two main rooting
strategies can be seen depending on the development
of the radical (Fig. 2):
– In the first strategy type (Fig. 2) the whole root
system is derived by the growth and lateral
branching of the seedling radical borne opposite
to the stem in the embryo. This primary root
grows downwards and branches in a given
orderly sequence, building up all the lateral
woody root system of the plant (Kahn 1983; Bell
1991). During the branching process, the tap-
root is responsible for the gravitropic response
and for the morphological and functional
differentiation of lateral roots borne on the
taproot (Dyanat Nejad and Neville 1972;
Champagnat et al. 1974). The best illustration
of such a rooting habit is shown in woody
seedlings which possess a taproot, regardless of
the taproot’s growth in the ensuing stages of
plant development. Many dicotyledonous herbs
and shrubs also exhibit such a rooting strategy
Plant Soil (2009) 324:1–30
(Kutschera and Lichtenegger 1997). In the
juvenile developmental stages of plants exhibiting
this strategy type, even if the taproot is
removed through e.g. an injury, a new vertical
root will emerge or an existing lateral root will
grow downwards to replace the taproot, thus
ensuring that the functional role of the taproot
is maintained (Khuder et al. 2007).
– In the second strategy type (Fig. 2), the radical
stops growing or aborts soon after the seed
germinates (Jourdan et al. 1995; Charles-
Dominique et al. 2009). The adult root system is
entirely adventitious1 and made up of many
homologous roots initiated in an upward order
at the base of the stem (Fig. 2). The best
illustration of such a rooting habit is that present
in cereals or bulbs, but tropical arborescent palms
or ferns are also concerned (Hallé and Oldeman
1970; Jenik 1978; Bell 1991).
These two main strategies define very early on the
way in which the root system will colonize the soil
and thus reinforce unstable soil, because they greatly
influence root angle, relative horizontal versus verti-
cal root spread and root density. In particular, in the
first rooting strategy type, the spreading of horizontal
roots away from the stem can result in large gaps in
the soil with no roots present (Figs. 3, 4), whereas, in
the second strategy type, many roots are continuously
renewed from the base of the stem over a long period
during plant development. On a slope where loose soil
and debris can collect upslope of a plant, adventitious
roots can therefore grow into this material, stabilising it
better than deeper growing roots could.
Roots of herbaceous plants
Root systems of herbaceous plants are usually more
diffuse, or fibrous than those of woody plants,
although bulbs, tuberous and storage roots are
common. Fibrous root systems generally possess
1
Adventitious root development can be encountered in the first
rooting type. Many tropical trees exhibit a very large cone of
adventitious roots at the base of the trunk which reinforces the
original primary root system. This type can be seen as a mixed
type. In temperate species the adventitious development of root
is often less spectacular because only the few first centimetres
of the trunk base are concerned.
Plant Soil (2009) 324:1–30 11

Fig. 3 a–d Development of

a root system at different a) b) c)
stages of ontogeny. Large
central roots build up the
framework of the root sys-
tem (1, 2). The taproot (1)
bears the woody laterals (2)
which spread the thinner
roots (3) away from the base
of the trunk in order to
explore the soil. These col- Taproot (1) Taproot (1), Taproot (1), thin (3),
onisation roots branch to and fine (5) exploitation (4) exploitation (4) and fine
develop thin woody ephem- roots and fine roots (5) roots (5)
eral exploitation roots (4).
All the woody roots branch 4
into non woody laterals
which are usually the fine 5
absorbing roots (5) 3
d)

4
5

All root types present

more fine and thin roots than those found in woody
species. Tensile root strength can be similar to that of
roots from woody species, whereas RAR can be
higher (Mattia et al. 2005; De Baets et al. 2008;
Loades et al. 2009), therefore, if shallow reinforce-
ment of soil is required, fast-growing herbs can be a
Fig. 4 Forks occur in the lateral root branches of woody
species. This branching pattern allows the root system to radiate
widely along the soil surface
useful alternative to slower growing shrubs and trees.
However, shallow mats of homogenously distributed
fibrous roots can easily tear away from the subsoil in
humid alpine conditions (Tasser et al. 2003). While
such shallow root systems may provide protection
against shallow surface erosion, they exert little to no
protection against shallow or deep-seated landslides
(Rice and Foggin 1971; Gabet and Dunne 2002).
At a more detailed level, topological indices have
been used frequently to describe root systems of
herbaceous species (Fitter et al. 1991) as well as
young tree roots and the root systems of juvenile trees
(Stokes et al. 1995; Tsakaldimi et al. 2009). The
topology of a root system largely determines how
roots occupy a given unit of soil and plays an
important role in resource uptake and anchorage
efficiency. Fitter (1985) classed herbaceous root
branching patterns into three types, depending on
magnitude (the number of branch tips), altitude (the
number of links, or distance between two branches, in
12
the longest path from the base to the tip) and the
relationships between these two parameters. Herring-
bone root systems are composed of a single main root
and one order of lateral roots (Fig. 1b) whereas in
dichotomous systems, a mother root always possesses
two daughter branches (Fig. 1c). Most branching
patterns lie between these two extreme types and are
called ‘random’ patterns. All three different types of
topology can be found within the same root system,
depending on root age and nutrient heterogeneity. In
trees, shrubs and herbs, very young roots may need to
explore nutrient rich patches, therefore send out
‘foraging’ herringbone roots which proliferate into
random or dichotomous systems once a pocket of
nutrients has been found. The branching pattern can
affect the resistance to uprooting of a plant and for an
equal root volume, dichotomous systems are signifi-
cantly better anchored than herringbone systems if
roots are rigid. This change in anchorage is because
during uprooting, the total amount of soil mobilised
during uprooting increases as it is carried upwards on
the forks of lateral root branches (Stokes et al. 1996;
Dupuy et al. 2005a, b). Similarly, in sand, the pullout
resistance of model roots depends on the number and
depth of lateral roots. The deeper the laterals, the greater
the resistance to pullout due to the increase in stresses
acting in the sand at depth, due to the overburden of
sand above (Mickovski et al. 2007). The Coulomb
friction law also predicts that if soil moves in a
horizontal plane around the root, above a critical depth,
the soil is lifted upwards. Below this critical depth, less
energy is required to compress the soil and to move it
horizontally around the soil (McKyes 1985).
Roots of woody plants
Woody plants generally develop two main classes of
roots, long roots (called woody, coarse, skeleton,
anchoring roots or macrorhizae) and short roots
(called non woody, fine, feeding, absorbing roots or
brachyrhizae). The long roots can become a perma-
nent part of the root system acting as a framework and
for anchorage. More ephemeral short roots emerge as
lateral branches that do not undergo secondary thicken-
ing, and so are thinner than long roots. Discrimination
between these two classes of root is usually on the basis
of root diameter within the range 1 to 10 mm (Hermann,
1977), as defined previously in this paper.
Plant Soil (2009) 324:1–30
Generally, trees develop large conical-shaped
woody central roots which are perennial and build
up the framework of the root system (Fig. 3). The
central and vertical taproot ensures the anchorage of
the plant and bears all the woody laterals (Atger and
Edelin 1994a). Near the collar, the biggest woody
laterals spread the thinner roots away from the base of
the trunk in order to explore the soil (Figs. 3, 4). This
framework develops laterally into two types of woody
roots that are usually shorter and thinner (cylindrically
shaped) and rarely live for a long time. These
colonisation roots occupy briefly the soil around the
skeleton roots as they are spread away from the tree
stem, before branching to develop thin woody
ephemeral exploitation roots (Fig. 3). Woody roots
branch into non woody laterals which are usually the
fine absorbing roots (Fig. 3) and which are more or
less branched, depending on species. For a landslide
engineer, it is important to remember that a slope is
stabilised by the whole community of plants, and not
individual root systems in isolation. The concept of
root occupancy is useful here, representing the sum of
the reinforcement due to new root material, plus the
remaining root reinforcement of any residual decay-
ing root systems (Watson et al. 1999). Watson et al.
(1999) viewed the root system of a tree as an
expanding circle, finding a rate of lateral root
expansion of 0.44 m year−1 for radiata pine (Pinus
radiata D. Don), as compared with 0.25 m year−1 for
kanuka (Kunzea ericoides A. Rich), on clear-felled
slopes in New Zealand. However, as the natural stand
density of trees for kanuka (16,000 stems ha−1) is
much greater than for radiata pine (typically 400 to
1,250 stems ha−1), the kanuka site reached its target
root occupancy in only half the time taken by the
densest stand of radiata pine. (In such studies
however, it is always necessary to consider the plant’s
developmental stage, as root extension rates of young
trees are greater than those of mature trees).
Tree root systems have traditionally been classed
into three types, depending on their overall shape:
plate, heart and tap (Köstler et al. 1968). Plate root
systems have large lateral roots and vertical sinker
roots, heart systems possess many horizontal, oblique
and vertical roots and tap systems one large central
root and smaller lateral roots (species list available in
Stokes et al. 2008). This classification is over-
simplified however, as many species have a mixture
of root system types. In particular, a ‘sinker’ root
Plant Soil (2009) 324:1–30 13

system type is frequently encountered and is com- soil and so increasing the effective stress on potential
posed of lateral roots from which descend vertical shear surfaces (Fig. 5a). Root systems extract water
sinker roots (Danjon et al. 2005). With regard to tree from deeper in the soil than would occur for
overturning during storms, trees possessing heart and evaporation from the bare soil surface, and calcu-
taproot systems have been classified as being the most lations based on the Penman-Monteith Equation for
resistant to uprooting and plate systems the least evapotranspiration suggest that this phenomenon may
resistant (Dupuy et al. 2005b; Norris et al. 2008). For significantly strengthen slopes against shallow fail-
stabilizing a slope, the type of root system could play a ures (Tarantino et al. 2002; Sidle and Ochiai 2006).
role in species choice, with deeper taprooted systems However, evapotranspiration is often small during
planted in the middle of a slope and plate rooted extended rainy seasons, especially in temperate
systems at the top of the slope (Danjon et al. 2008). regions and, as mentioned previously, slopes often
fail when saturated by heavy rainfall. In Malaysia,
where evapotranspiration is high throughout the year,
Root traits and their interaction with soil stable slopes were associated with large root length
properties densities (RLD) and relatively dry water contents,
whereas unstable slopes tended to be relatively wet
Roots reinforce slopes by the direct mechanical and unrooted (Osman and Barakbah 2006). RLD is a
reinforcement of their roots, but also by drying the trait defined as either root length per unit volume of
soil (i.e. root length distribution with depth) or root
250 length per unit surface of a soil (i.e. the amount of
(a) Fladbury
Alluvial gley roots per unit ground surface or crack surface)
Wicken
200 (Miyazaki 2006). However, simple water balance
Shear stress (kPa)

simulations in Peninsular Malaysia suggest that

150 tropical forest removal would only moderately in-
crease the period of movement of deep-seated failures
100 and would have little effect on landslides in shallow
soils (Sidle and Ochiai 2006).
50

0 Root traits and the soil water dynamics of slopes

0 100 200 300 400 500 600
Soil matric potential (-kPa)
Root extraction of soil water
50
(b)
Shear strength increase (kPa)

Willow (sandy loam)

Alfalfa (clay loam)
Alfalfa (chalk)
The shear strength of a soil depends on the balance of
40 Pistacia (chalk)
Rosemary (chalk)
forces operating on a potential shear plane. The matric
suction associated with unsaturated soil tends to
30
increase the resistance to shear as soil dries, by
increasing the effective stress between particles in the
20
soil matrix (Fig. 5). Plant roots act as small suction
pumps, drawing water from the soil at matric
10
potentials as negative as −1.5 MPa. Individual root
tips may continue to grow at matric potentials as dry
0
0.000 0.001 0.002 0.003 0.004 0.005 0.006
as −2.3 MPa, and even to −4 MPa if the rest of the
Root area ratio plant is adequately supplied with water (Portas and
Taylor 1976; Sharp et al. 2004).
Fig. 5 a Shear strength as a function of soil matric potential for The rate of water extraction achieved from a soil
three clay soils (data from Spoor and Godwin 1979) and b
increase in shear strength as a function of root area ratio (RAR)
layer depends on the RLD in a particular zone of soil
for clay loam and chalk (data from Operstein and Frydman and the uptake rate per root length. If the root system
2000), and from sandy loam (data from Mickovski et al. 2009) is uniformly distributed in a volume of soil, a RLD as
14 Plant Soil (2009) 324:1–30

small as 0.1 cm cm−3 should be adequate to extract

the water from that soil volume in a few days
(Passioura 1991). However, if roots are clumped
within cracks and biopores in the soil, the time
required for water extraction may increase by an
order of magnitude or more (Passioura 1991). This
phenomenon is because of the greater distance that
water must flow through the soil structural unit to the
roots—several roots clustered within a single biopore
may only be marginally more effective than an
individual root in the same pore. Such clustering
occurs particularly within the subsoil where the
strength of the soil may be relatively large and restrict
growth to continuous channels and planes of weak-
ness or, e.g. if new roots grow preferentially along
decomposing root channels. Little is known about the
relative ability of roots to exploit biopores, though
Fig. Illustration of
Fig. 66 Illustration of how
how aa decayedly root
alongchannel
the channel formeda
can initiate
methods are being developed that might allow decayed root
preferential flowchannel can in soil.byThe
path (PFP) thedark
decomposing
coloured area ligni-
in
screening for this trait in crop plants (McKenzie et initiate a preferential flow fied tissue of a
the Bt horizon indicates where dye has flowed preferentiallyroot, stained
al. 2009). There are at least indications that species path (PFP)
along in soil. formed
the channel The dark black. (Reprinted
by the decomposing fromtissue
lignified Per-
may differ substantially in their ability to extract coloured area in the Bt illo et al. (1999),
of a root, stained black. (Reprinted from Perillo et al. (1999),with per-
horizon
with indicates
permission from where
Elsevier) mission from Elsevier)
water from the subsoil, even if the distribution of dye has flowed preferential-
RLD is similar (Bremner et al. 1986; Passioura 1988).
There are large species and varietal differences in of these preferential flow paths increases with rainfall
the distribution of root length with depth (Jackson et intensity and soil water content (Tsuboyama et al.
al. 1996), and so it should be possible to choose 1994; Perillo et al. 1999), and may result in relatively
vegetation for slopes that extract water from greater rapid transport of water through the surface horizons
depths. In terms of the total amount of water extracted to less permeable horizons or to bedrock (Fig. 6). In
from a slope, generally the larger the biomass the gently sloping terrain preferential flow paths generally
larger the potential evapotranspiration rate. Water serve to facilitate vertical transport of water and
extraction by the root system must be viewed in the solutes; however, in steep topography, especially
wider context of the water balance of the whole slope. during high rainfall, these preferred flow networks
can rapidly transport water downslope potentially
Effects of roots on preferential flow pathways increasing pore water pressure at the soil-bedrock
in the soil interface or within the soil when hydrologic disconti-
nuities are encountered (Sidle and Ochiai 2006).
Preferential flow paths are regions of the soil where It is likely that differences exist in the preferential
relatively rapid transport of water and solutes may flow pathways associated with root systems of
occur under heavy rain or flooding (Perillo et al. different species, although this is not well docu-
1999). Rapid macropore flow paths may be difficult mented. Root systems with relatively rapid root
to locate or predict using core-sampling techniques, turnover rates will presumably result in more decay-
and may only become apparent at large lysimeter or ing root channels than plants with slow turnover.
hillslope scales (Deeks et al. 2008). Macropores Whilst the thick structural roots of trees may persist
formed by plant root channels comprised at least for many years, finer roots belonging to the same root
55% of all macropores in a Japanese forest soil system e.g. <2 mm tree roots, may be born and die
(Noguchi et al. 1997), whilst 60% of all preferential within a period of weeks or months (Eissenstat et al.
flow paths under recently tilled alfalfa (Medicago 2000; Block et al. 2006). Even within a single root
sativa L.) plots were associated with roots or decom- system, there can be very large differences in root
posed root channels (Perillo et al. 1999). The length lifespan related to root diameter—for example, fine
Plant Soil (2009) 324:1–30
tree roots turnover relatively rapidly (e.g. 30 to 300 d
for fine poplar roots; Block et al. 2006), as compared
with much thicker structural tree roots that can persist
for many years. This temporal nature of fine root
dynamics needs to be reflected in models (Abe and
Ziemer 1991b).
Roots play a major role in the formation of soil
structure by providing a source of carbon, via root
turnover and exudation, and by subjecting the
rhizosphere to alternating wet-dry cycles that bind
rhizosphere particles together with mucilage (Watt et
al. 1993). The interaction of the root system with soil
type influences the nature of structure formation. For
example, grass root systems often have RLD of
20 cm cm−3 or more in the surface 0.1 m of soil,
with relatively rapid turnover rates. This gives rise to
the typical crumb-like soil structure found under grass
swards. The presence of the grass roots and their
associated fungal and bacterial communities rapidly
binds the soil together and decreases erosion from
establishing newly engineered slopes. The average
depth of most grass root systems is usually <1.0 m
and so the effect is confined to the surface horizons
and has little affect on landslides as previously noted.
Soil constraints on root system morphology
Root system form varies not only between species but
also within a species. The first stages of root
development are genetically driven and shape is not
highly variable. As a plant matures, the morphology
of its root system and the distribution of roots in the
soil are greatly affected by the immediate soil
environment as well as aboveground influences.
However, for a given soil constraint, e.g. mineral
deficiency, plant responses can be significantly
different depending on species and the part of the
root system considered. Even the response of a single
root to any one stress will vary according to the root
type and its branching order (Drew 1975; Thaler and
Pagès 2000).
When analysing the consequences of environmen-
tal constraints on root development, roots which
undergo many of the common stresses often respond
by first reducing and then by arresting lateral root
growth. Therefore, a mother root inhibits lateral root
emergence before restricting its own growth (May et
al. 1965, 1967; Hackett 1972; Drew et al. 1973). In
15
heterogeneous soil, when a stressed root reaches a
zone with conditions ideal for growth, any inhibition
of branching can be followed by a proportional
stimulation of lateral root development. Therefore,
when one part of a root or a root system has been
suppressed, the remainder can react and exhibit what
has been called compensatory growth (Drew et al.
1973). Studying the competitive effects of neighbours
on lateral root spread in a Creosote bush (Larrea
tridentata Sesse and Moc. ex DC.) population,
Brisson and Reynolds (1994) demonstrated that the
occurrence of such compensatory growth induced a
sharply asymmetrical distribution of horizontal roots.
As the asymmetrical formation of root systems is
frequently encountered on mountain slopes (see
below), studies of plant growth on slopes must be
analyzed carefully, taking into account this potentially
compensatory root growth. Nevertheless, not all
species can exhibit compensatory growth to avoid
neighbours e.g. Platanus sp. and the west African
species Aucoumea klaineana Pierre. Such species
have roots which can fuse and graft with each other
(both within the same tree and between individuals),
thus allowing sap exchange between individuals
(Norris et al. 2008). The fused root systems can act
as a network, holding soil and stones in place and
interlocking with rocks and coarse fragments in the
soil, enhancing the lateral membrane strength of the
soil-root complex (Fig. 1). Nevertheless, many tree
species demonstrate compensatory growth which
results in the whole form of the root system changing
so that, e.g. a taprooted or heart rooted system can be
transformed into a plate root system when influenced
by local soil conditions, such as the presence of a hard
pan or a seasonal water table (Fig. 1a, Danjon et al.
2005; Norris et al. 2008). The expression of these
different forms is known as plasticity.
Shallow and deep root endemics
Schenk and Jackson (2002) collated a database of
rooting depth for 1,300 plants from water-limited
(arid to sub-humid) environments. These authors
found that plant rooting depths in water-limited
environments are highly correlated with infiltration
depth. Perennial herbs and forbs had average rooting
depths of approximately 0.75 m whereas the rooting
depth of annuals was usually <0.5 m, although these
16
depths could reach several metres. Shrubs and trees
on deep soils have a mean rooting depth of 2.2 m
whereas shrubs and trees growing on shallow soils
over bedrock have a mean rooting depth of 7.9 m.
What is happening in this latter case is that roots of
woody plants on shallow soils tend to grow along
fractures deep into the underlying bedrock (Fig. 1a).
Schenk (2008) therefore asks the question—how do
roots at the boundary between soil and bedrock locate
rock fractures that are large enough for them to grow
in? Poot and Lambers (2008) studied six species of
Hakea, two of which are endemic to shallow soils
over ironstone and four are commonly found on
deeper soils in the same region. When grown in
containers, the shallow soil endemics explored the
bottom of the containers more efficiently than the
deep soil endemics. This superior foraging capacity
will allow roots to locate cracks in the underlying
bedrock, leading in turn to possible supplies of
nutrient and water rich pockets. Therefore, the
shallow soil endemic will be able to better tolerate
water and nutrient stress, but the disadvantages of
occupying this particular ecological niche include
higher energy costs for constructing deeper roots, the
possibility of oxygen deficiency and lower water and
nutrient availability (Schenk 2008). For the landslide
engineer, the ability of vertical roots to grow into
cracks in the bedrock is of extreme but conflicting
interest. If plants on shallow soils are able to grow
through the soil and anchor the plant to the underlying
bedrock, soil on a slope would be better fixed
(Fig. 1a). However, the roots of many species, e.g.
Pinus pinaster Ait., are not able to penetrate the
bedrock and grow along the surface of the rock,
resulting in an unstable shallow root plate (Fig. 1a;
Danjon et al. 2005). In some species, e.g. eucalyptus
(see Stone and Kalisz 1991), roots can grow as deep as
60 m, often being found in subterranean or aquatic
caves (Hubble et al. 2009). However, in plants where
roots are anchored to the bedrock, root growth through
cracks could enlarge them over time, thus destabilising
the rock and causing it to fracture, leading in turn to
slope failure (Sati and Sundriyal 2007).
Plasticity in root traits
The ability of a genotype to change its phenotype
was originally thought to be an unstable process.
Plant Soil (2009) 324:1–30
However, it is now recognized that this ability to
change, termed phenotypic plasticity, is central to
plant ecological development (Bradshaw 2006).
Under genetic control, plasticity enables sessile plants
to adjust to spatial and temporal heterogeneity, thus
minimizing stress (Fourcaud et al. 2008). Plasticity
has been found in relation to nutrient supply (Drew
1975), water availability, and soil strength (Bingham
and Bengough 2003).
Heterogeneity in resource supply
When plant fine roots encounter a nutrient rich patch,
they usually proliferate, or cluster, within it (Fig. 1b),
where proliferation is the increase in initiation of new
laterals (Hodge 2004). Lateral root length can also
increase (Drew 1975), but by producing more
branches, a given unit volume of soil will be
exploited more efficiently. Total root system biomass
does not necessarily change because an increase in
root mass within a nutrient patch can occur to the
detriment of the root system growth outside the patch
(Drew 1975). Within a patch, roots can be longer and
thinner (Farley and Fitter 1999) and the trait root
tissue density (RTD) may differ in the proliferated
roots (Eissenstat 1991). Lignified roots are not
sensitive to changes in nutrient supply whereas in
fine roots, changes in physiological plasticity can also
occur so that ion uptake is increased, which might in
turn act as a feedback mechanism, triggering the
production of new lateral roots (Hodge 2004). A trait
describing lateral root spread may be useful for
estimates of the area over which plants anchor the
surface soil (Fig. 1). In shrubs and trees, lateral spread
was found to be typically between 2 and 16 m in
radius (Stone and Kalisz 1991; Schenk and Jackson
2002) but only 0.10–0.60 m for herbaceous plants,
with semi-shrubs and stem succulents intermediate.
Most (95%) herbaceous perennials have lateral root
spreads of <1.5 m, and 95% of all shrubs have been
found to possess lateral root spreads of <15 m
(Schenk and Jackson 2002).
Added to the complexity of understanding root
plastic responses to soil nutrient status is the
consideration of microbial communities. Mycorrhizal
fungi colonise roots of a host plant, and benefit from
this mutual symbiosis by having direct access to
carbohydrates produced by the plant. In return, the
plant can access a larger surface area of soil via the
Plant Soil (2009) 324:1–30
mycelial network of the mycorrhiza and take advan-
tage of the mycorrhiza’s ability to ‘scavenge’ or
‘mine’ phosphate in the soil (Lambers et al. 2008).
Mycorrhizas can also proliferate in nutrient rich
patches, and increase root production and alter RLD
(Cui and Caldwell 1996; Hodge et al. 2000). In 25
North American tree species, species predominately
forming ectomycorrhiza were found to have a higher
branching intensity than those forming arbuscular
mycorrhiza (Comas and Eissenstat 2009). Root tensile
strength increases in ectomycorrhizal-infected roots
(Ba 2008) but this response appears to be species and
site dependent. In a growing literature on the subject,
mycorrhizas are seen as useful tools in the restoration
of degraded soil, but again, species and site are
important factors to consider before undertaking any
restoration project using mycorrhizas (e.g. Palenzuela
et al. 2002; Estaun et al. 2007). In certain conditions,
the introduction of mycorrhizas at a site may even
be detrimental to plant establishment (see Walker
et al. 2009).
Root response to zones of increased water avail-
ability are less clear. Lateral soil moisture heteroge-
neity can result from variations in topography, water
infiltration, differential root competition and agricul-
tural irrigation whereas vertical heterogeneity, e.g.
waterlogging, is often seasonal and predictable
(Bauerle et al. 2008). Root growth is usually reduced
in dry soils and within dry patches; resource alloca-
tion is also reduced so that outside the dry patch,
preferential root growth occurs (Coutts 1982). Differ-
ent species have different strategies to water supply;
cacti may produce low-cost, short-lived fine roots in
response to periods of precipitation (Schenk and
Jackson 2002; Snyman 2006), whereas Eissenstat
(1991) showed that Citrus sp maintain roots in dry
soil and increase root growth only slightly in wetter
soil. Bauerle et al. (2008) demonstrated that morpho-
logical plasticity in root systems of fast growing vines
(Vitis vinifera cv. Merlot) was greater than that in
slow-growing vines, but that the tolerance to moisture
stress was similar in both types. Plastic changes will
influence how the plant extracts water from the soil,
with some root morphologies being more efficient
than others (Tsutsumi et al. 2003). When clusters of
roots occur in resource-rich patches, water uptake is
faster around the cluster compared to non-rooted
zones, thus preferential water extraction occurs from
specific regions of soil (Kazda and Schmid 2008).
17
Soil texture and compaction
Soil texture and compaction have major implications
for the physical stresses that roots will experience (da
Silva et al. 1994; Bengough et al. 2006). Sandy soils
will store less plant-available water (water held
between field capacity and wilting point) than clays,
and so will be more susceptible to drought. Clay soils
hold more available water, but they may have
insufficient air-filled porosity, limiting root growth
by hypoxia under wet conditions.
Mechanical impedance is often the major limitation
associated with soil compaction. The elongation rates
of both main axes and lateral roots may be decreased,
unless small pores in the soil allow fine lateral roots to
proliferate (Goss 1977). Due to the slower elongation
rate of the main root axes, the number of lateral roots
per unit length of the main axis will tend to increase in
compacted soils. Roots thicken and follow a more
tortuous path in more compacted soils (Konopka et al.
2009). Root depth is often restricted on former-mining
sites, and operations such as deep ripping may be
required to enable tree roots to penetrate deeper into
dry profiles to obtain sufficient water (Szota et al.
2007). Species differences exist in the extent to which
trees can root into compacted soils, with Corsican Pine
(Pinus nigra Arnold.) showing superior rooting than
Italian alder (Alnus cordata Loisel.), Japanese larch
(Larix kaempferi Lindl.) and birch (Betula pendula
Roth.; Sinnett et al. 2008). Plants may exhibit
compensatory growth as they respond plastically to
localised areas of compaction, this response varying
between species (Bingham and Bengough 2003).
Physical limitations to root growth can be thought of
in terms of the Least Limiting Water Range (Bengough
et al. 2006; da Silva et al. 1994), which defines critical
values whereby root growth will be significantly
limited if exceeded. For matric potential, soils drier
than −1.5 MPa will severely limit root growth and in
such dry soils mechanical impedance will also limit
root growth. The limit chosen for mechanical imped-
ance is a penetrometer resistance of 2 MPa whilst
aeration is likely to be limiting if there is less than
0.1 cm3 cm−3 air filled porosity in a region of soil.
Root growth on hillslopes
Landslide engineers are particularly interested in how
plants grow on slopes. Several studies have been
18
carried out on the direction of root growth of plants
growing on sloping terrain. Chiatante et al. (2003)
argue that in Quercus cerris L., preferential root
growth occurs either up- or downslope, thus enhanc-
ing anchorage along the axis of static mechanical
loading. Mullen et al. (2005) also showed that in
Arabidopsis seedlings inclined at different angles,
lateral root growth was strongly controlled by gravity;
although roots emerged on the upper side of the
taproot, their direction of growth was quickly altered
and they curved downwards, towards the lower side
of the taproot, resulting in a significantly higher
number of lateral roots downslope. However, in other
species, it has been found that preferential growth of
lateral roots occurred upslope or even perpendicular
to the slope direction (Nicoll et al. 2006; Khuder et al.
2006; Khuder 2007). Nevertheless, although lateral
roots of seedlings might grow preferentially in a given
direction, as they elongate they become less sensitive
to gravity (Kiss et al. 2002). At the tissue level,
differences have also been observed. In 8 month old
Spartium junceum L., a significantly higher lignin
content was found in root systems growing on slopes
compared to those growing on flat ground (Scippa et
al. 2006). As lignin and cellulose content are usually
inversely proportional, more lignin means less cellu-
lose. Lignin is resistant in compression, and it is not
known why it should be present in higher quantities
in roots growing on slopes. Differences in root tissue
structure will influence tensile strength (Genet et al.
2005) and in trees growing on slopes, tensile strength
was found to be greater in upslope roots compared
with downslope and horizontal lateral roots (Schiechtl
1980), although similar studies have shown no
differences between roots from young Robinia
pseudoacacia L. grown on a slope in controlled
glasshouse conditions (Khuder 2007).
The diversity of root responses to slope angle
observed thus probably depends on several factors
including species, experimental conditions and plant
age. For example, Di Iorio et al. (2008) found that in
S. junceum, soil type had a greater influence than
slope angle on the distribution of surface roots.
Surface roots were more numerous in clay than in
loam soil, independent of the slope angle, and grew
upslope in clay soil only. Stone and Kalisz (1991)
suggest that upslope laterals grow obliquely or
Plant Soil (2009) 324:1–30
horizontally into weathered joints of bedrock and
can attain great lengths. Nutrient and water availabil-
ity also play a major role: Pierret et al. (2007) found
that in fields on sloping terrain in Laos, both crop
(rice—Oryza sativa L. and Job’s tear—Coix lacryma-
jobi L.) and fallow vegetation (15 native herbaceous
species) developed more shallow root length per unit
soil volume when growing in gentle than steep slopes.
In conclusion, it is not possible to say that roots grow
preferentially up- or downslope; the combined effects
of all interacting factors must be taken into account.
Few studies have been carried out on how root
traits change with altitude, and this topic is of much
interest to landslide engineers. In the few studies
existing, high altitude grasses in New Zealand were
found to have thicker roots and smaller SRL than
plants at low altitude (Craine and Lee 2003).
However, Körner and Renhardt (1987) found that
SRL was greater in herbaceous dicots at higher
elevations. An investigation into the spatial arrange-
ment of roots within the root system, and how their
distribution affects resource uptake, may help explain
these differences in results. Soethe et al. (2006)
examined root architecture of several tree species
growing at two different altitudes in Ecuador. At the
higher altitude, soil depth was restricted and root
extension significantly greater, resulting in a wide-
spreading but shallow root-soil plate. The increase in
root plate width would help compensate for the
shallowness of the root system, thereby maintaining
anchorage ability. Genet et al. (2006) found that root
tensile strength decreased at very high altitudes in
Abies georgii var. smithii., although the functional
significance of this decrease is not yet clear. It could
be that there is less investment in roots at high
altitudes, which have been found to have a faster
turnover rate (Graefe et al. 2008).
Plasticity in root traits can therefore be highly
diverse, and to add to their complexity, responses may
be quite different depending on whether a plant is
grown individually, as a monoculture, or with a
mixture of species (Huber-Sannwald et al. 1996,
1997). Not only do traits change over time because
of their inherent genetic expression (Barthélémy and
Caraglio 2007), but depending on the frequency and
intensity of temporal signals, e.g. nutrient availability
and rainfall events, morphological and physiological
Plant Soil (2009) 324:1–30
responses will differ within a plant root system
(Hodge 2004).
How do plant root traits change with time?
At the beginning of this paper, we said that a trait is
defined as a distinct, quantitative property of organisms.
However, it must not be forgotten that traits are not static
properties and that they evolve over time as the plant
matures (Barthélémy and Caraglio 2007). The nature of
a trait will thus depend on endogenous growth
processes and be influenced by exogenous constraints.
Developmental stages in woody root systems
From germination to death, successive developmental
stages can be defined according to the nature and
topology of the different root categories developed in
the branched system (Fig. 3). Tree species have
complex rooting forms and can undergo reiterative
branching processes which influence root distribution
and interactions with the local environment. One such
process leads to the formation of forks (Fig. 4) which
pin more superficial roots to the underlying soil
substrate. Forks appear when the growth potential of
one lateral root changes as a shift occurs in the mother
root’s functioning. Horizontal perennial roots can
build up two kinds of forks; those near the taproot
are often made up of one or two homologous
horizontal, perennial roots and one vertical tap root
which branches from the lower surface of the fork,
away from the trunk. Forks give rise only to
horizontal roots, no vertical members can develop.
This branching pattern allows the root system to
radiate widely along the soil surface (Fig. 4, Atger
and Edelin 1994a, b; Edelin and Atger 1994). A
second type of reiterative branching pattern can occur,
which involves the outgrowth of a new lateral root
(neoformation) or the activation of a dormant lateral
root on an old part of the root system, e.g. after
wounding or pruning.
Fine root dynamics
Although many data are available on the subject,
engineers rarely consider the dynamics of fine root
19
production in models of landslide risk. Fine root
production reflects the annual changes that can occur
belowground and how a system responds to distur-
bance (Vogt et al. 1996). Usually, only roots <3 mm
in diameter are considered, which are those having
the highest tensile strength (Bischetti et al. 2005,
2009a; Genet et al. 2005), and thus of great interest to
landslide engineers. The biomass of thick roots is less
affected by environmental change and so is fairly
static over the short-term.
Hendricks et al. (2006) showed that the method by
which fine root production is estimated is of upmost
importance. Fine root biomass was measured using
nitrogen and carbon budget techniques, standard soil
cores, ingrowth cores (soil cores are removed and the
soil replaced with root-free soil into which new roots
can grow) and minirhizotrons (a transparent cylindri-
cal tube inserted into the soil and down which a
camera is inserted). By comparing the different
methods in three forest types, Hendricks et al.
(2006) demonstrated that annual fine root biomass
values ranged from 0 to 4,618 kg ha−1 year−1 when
production was measured from soil cores and by
using minirhizotrons, respectively. The minirhizotron
technique provided the most reliable data and this
study underlines the shortcomings in the various root
sampling techniques. As discussed earlier, root archi-
tecture is highly variable and plastic responses are
many. Soil cores therefore need to be large and
numerous enough to capture the heterogeneity in fine
root production across a slope (Genet et al. 2008).
Nevertheless, temporal changes in dynamics through-
out a year will only be achieved by regular sampling
or through using minirhizotrons or large scale
rhizotrons. The latter consist of panes of glass pressed
against the soil surface in a trench dug into the slope
(e.g. Thongo M’bou et al. 2008). Root initiation,
elongation rates and mortality can thus be observed as
frequently as desired. In a study of Eucalyptus clones
in Congo, seasonal dynamics were observed in such a
rhizotron and it was found that fine roots grew
continuously throughout the year, but that a signifi-
cant decrease in elongation rates occurred at the end
of the dry season (Thongo M’bou et al. 2008). These
authors could also follow root growth dynamics with
regard to depth and correlate elongation rates with
water variation in the soil profile.
20
Integrating information for the management
of mountain slopes
Natural regeneration or planting mixtures?
Surprisingly, the distribution of trees, and hence their
root systems, in forests prone to landslides has been
studied little (Schmidt et al. 2001; Sakals and Sidle
2004; Kokutse et al. 2006; Danjon et al. 2008; Genet
et al. 2008, 2009) and yet is of major importance as
landslides tend to occur in areas of reduced root
distribution (Roering et al. 2003), where gaps occur
between trees. Genet et al. (2008) studied root
reinforcement in plantations of Cryptomeria japonica
D. Don. of three different ages and spatial densities. It
was found that in older plantations, where trees had
been thinned and large gaps occurred between trees,
the greater the number of trees in given transects and
the smaller the average distance between groups of
trees, the higher the FOS. No relationships between
FOS and stand structure were observed in the
youngest plantation, due to the high stand density
and low spacing between trees, resulting in a more
homogeneous structure. Sakals and Sidle (2004)
proposed that model simulations of root cohesion
over time and space would allow foresters to
determine how gap formation between trees would
affect slope stability. Similarly, Danjon et al. (2008)
suggested using maps of root cohesion in forest
stands to establish appropriate types of planting, e.g.
rows or staggered lines of trees, depending on the
FOS of the slope. Simulating changes in root
architecture using plant growth models (Jourdan et
al. 1995; Dupuy et al. 2005c; Drouet and Pagès 2007)
could in the future be combined with slope stability
models (Kokutse et al. 2006) and is an area where
much promising research could be carried out.
Once e.g. a clear-felling or a fire has occurred on a
slope, the site will be relatively fragile until new
vegetation colonizes the soil. Although the roots
remaining in the soil provide a certain amount of
strength for the early years (e.g. Watson et al. 1999;
Sidle et al. 2006; Jackson and Roering 2009), a
forester has to decide how best to manage the site
depending on the long-term goal. If not under the
obligation to plant fast-growing monocultures of
timber species, natural regeneration of the local
vegetation may be a solution. Very little information
exists on the slope failure risk associated with natural
Plant Soil (2009) 324:1–30
regeneration, although higher species diversity has
been shown to improve soil aggregate stability (Pohl
et al. 2009). Genet et al. (2009) showed that in the
first phase of succession in naturally regenerated
slopes in the Sichuan, China, when big node bamboo
(Phyllostachys nidularia Munro.) was the dominant
species, there was no contribution of vegetation to the
FOS due to the shallow rhizomatous rooting nature of
big node bamboo. The FOS did increase over time, as
deeper rooting deciduous tree species colonised the
sites studied. Similarly, Roering et al. (2003) observed
that where gaps occurred in a naturally regenerated,
temperate forest, sword fern (Polystichum munitum
Kaulf.) was present and always found in the vicinity
of the landslide scarp. Schmidt et al. (2001) discuss
the possibility of planting conifer seedlings immedi-
ately after clear-felling, to reduce the window of
landslide hazard which occurs due to subsequent root
decay and hydrologic response. The duration of this
window varies from 15 to 25 years and is highly site
and species specific (Ziemer 1981; Watson et al.
1999). For example, Cammeraat et al. (2005) showed
that in abandoned bench terraced slopes at different
phases of vegetation succession, mass wasting pro-
cesses increase after abandonment, due to the limited
contribution of anchorage by roots at potential slip
planes and the fast transfer of rainfall to the potential
slip plane. It took up to 40 years for mass wasting
process to decline. If natural regeneration is the
choice of the landslide engineer, rather than restora-
tion or prevention of landslides through plantations,
then promotion of the recovery of self-sustaining
communities is feasible by stabilization with native
ground cover, applications of nutrient amendments
and facilitation of dispersal to overcome establish-
ment bottlenecks (Walker et al. 2009).
Regeneration strategies
The ability of a plant to resprout after a disturbance is
an important consideration to be made in the
management or restoration process. Resprouting is
the ability of a plant to produce a new shoot from a
root or stem if part of the plant’s biomass has been
removed through a disturbance event such as grazing,
fire, windstorms, avalanche etc (Bond and Midgeley
2001). Not all species possess this ability. Annual
species usually cannot resprout as resources need to
be allocated and stocked in storage organs. As
Plant Soil (2009) 324:1–30
construction costs are higher for such organs, these
costs need to be weighed against the benefit that a
plant will gain after a disturbance. If a plant can
resprout, it will be able to survive in the newly
disturbed environment, even when above-ground bio-
mass is destroyed (Stokes et al. 2007). Very few
studies have been carried out on the influence of
edaphic factors on resprouting ability (Sakai et al.
1997). Guerrero-Campo et al. (2006) carried out a
study of 123 commonly occurring plant species found
on eroded land in north-east Spain. These authors
found that species with coarse and deep tap-roots
tended to be root-sprouting and those with fine,
fasciculate and long main roots (which generally
spread laterally), tended to be shoot-rooting. Landslide
engineers can find information about such traits useful
when deciding on management strategies with regard
to coppicing. However, Sati and Sundriyal (2007)
suggested that Alnus nepalensis D. Don can promote
landslide activity after resprouting. If the roots of this
species are exposed through soil loss, resprouting
occurs which initially reduces soil erosion. The roots
of several individuals then form an interlocking matrix
which acts as a single unit. As the plants mature, the
weight of this root-soil matrix increases and can result
in a downward movement of the integrated soil mass.
A similar phenomenon has also been observed in the
rhizomatous big node bamboo, which resprouts from
the running rhizomes located just underneath the soil
surface (Stokes et al. 2007). Vogt et al. (2006) showed
that abandoned stands of coppiced Sweet chestnut
(Castanea sativa Mill.) became more likely to uproot
as trees became taller and more mechanically unstable,
particularly when growing in pits or steep depressions.
On steep slopes, uprooted trees are likely to fall
downslope, transporting sediment attached to the root
plate and even resulting in soil mass movement
(Norman et al. 1995). After clear-felling of a coppiced
stand, it is not known if root decay occurs, thus
reducing soil reinforcement, and should be further
investigated. A careful management of coppiced stands
is therefore required to meet the balance between the
protection of surface soil through erosion and a deeper
reinforcement of the slope.
Choice of planting stock
If seeding or natural regeneration of vegetation at a
site is not an option and a site is to be planted with
21
container seedlings, the landslide engineer should use
the best quality planting stock available, to avoid
seedling mortality and poor growth when plants are
planted out. To promote lateral root growth in
container seedlings, different types of pots exist, e.g.
slitwall containers have open slits down the sides of
the container so that roots can grow through the slits
(Rune 2003). Ribwall containers have solid ribbed
walls along which roots grow, thus avoiding root
spiralling, but root area is less in these pots and
spiralling more frequent than in slitwall containers
(Rune 2003). Root spiralling occurs when lateral root
growth is restricted by the pot size and roots grow
around the pot walls, forming a root ball. A
symmetrical root system shape with unrestricted
lateral and vertical roots would fix soil earlier and
more efficiently than seedlings with root balls.
Container-grown trees can also have a significantly
different root architecture to naturally regenerated
seedlings, due largely to the availability of nutrient
and water resources in nursery conditions, e.g.
naturally regenerated Quercus ilex were found to
possess herringbone-like root systems whereas con-
tainer seedlings had more randomly branched root
systems (Tsakaldimi et al. 2009). Many different
techniques exist to manipulate root systems during
the juvenile stage of growth, e.g. by root pruning
before transplanting, thus changing the shape of the
existing root system and affecting the subsequent
branching pattern when roots regenerate from the
pruned areas (Girouard 1995). Cuttings can also have
a significantly different root architecture to seedlings.
Seedlings generally possess their original taproot
whereas cuttings do not necessarily, and only adven-
titious roots can be present, depending on the parent
material of the cutting (Charrier 1969: Sasse and
Sands 1997). Live pole cuttings are often used to
stabilize slopes and river banks (Norris et al. 2008)
and also develop an adventitious root system with
the mechanical function of the pole analogous to a
large taproot. It was found that not only does root
system development depend on soil type, but the
anchorage of adult Populus sp grown from live poles
depends on whether soil is frictional (sandy) or
cohesive (clayey), with anchorage improved in the
latter (Dupuy et al. 2007).
The engineer must therefore carefully select plant-
ing stock, as seedlings grown in ideal nursery
conditions and a particular soil type may not be
22
adapted to stressful field conditions. In such cases,
root growth of healthy seedlings can be arrested when
substrate conditions change drastically. Before plant-
ing out therefore, it would thus be wise to determine
how the most limiting factors to growth at the given
site affect root traits. A list of desirable root traits for
the landslide engineer is given in Table 2.
Perspectives for future research
One of the current areas where more information is
needed is the development of a simple yet elegant
slope stability model which takes into account the
impact of vegetation. User-friendly softwares, e.g.
Slip4ex (Greenwood 2006), exist which calculate the
FOS of a slope, but rely on data from Wu’s (1976)
and Waldron’s (1977) models of additional cohesion,
which overestimate the cohesion provided by roots.
The FBM (Pollen and Simon 2005; Pollen 2007;
Pollen-Bankhead and Simon 2009) is highly promis-
ing but requires further development with regard to
the spatial arrangement of roots in soil along with
their mechanical properties. It is unlikely that a simple
model can consider 3D root architecture, therefore
parallel studies using techniques like FEM need to be
carried out to determine how the position of roots in a
block of soil influence its shear strength, taking into
account root size, mechanical properties and soil
physical properties.
Most of the existing slope stability models which
consider vegetation are of a static nature (but see
Pollen 2007), i.e. they just consider the distribution of
roots at a given time, nevertheless, such models are
still useful for estimating landslide risk for a given
slope configuration. Exploring the effect of reforesta-
tion scenarios or more generally the impact of slope
management can be performed making assumptions
on the expected rooting patterns and using the
previously mentioned static models (see Dhakal and
Sidle 2003 as an example of considering root decay
and regrowth after harvesting). However, coupling
growth models, in particular based on structural-
functional notions (Fourcaud et al. 2008), with soil,
climate and hydrological models is an exciting
challenge to answer the question of the consequences
of climate change on substrate mass movement. Such
models can also take into account the temporal nature
of root dynamics and trait expression.
Plant Soil (2009) 324:1–30
How root systems interlock with rocks and coarse
fragments in the soil is not quantified, but is a
character which could be manipulated by the land-
slide engineer to enhance the lateral membrane
strength of the soil matrix component (Fig. 1a). We
also need more information on the properties of the
interface zone between roots and soil and how this
changes friction along the root surface. Roots have
rough surfaces due to the presence of bark, roots hairs
and exudates (Mickovski et al. 2007). The latter can
influence soil aggregate and rhizosheath formation
which is also an important aspect for the landslide
engineer to consider (Pohl et al. 2009). How root
tensile strength changes along a root, within a root
system, with plant and root age and how it is
influenced by the soil environment, needs to be
elucidated along with its interaction with other traits
and plant form and history.
It is particularly important that we develop better
ways of measuring root distribution with depth. If
root traits could be predicted from aboveground
measurements or characteristics, it would be possible
to map the contribution of vegetation to slope stability
and thus highlight zones of potential failure (Roering
et al. 2003; Preti et al. 2009). This type of mapping
could also consider species diversity and the effect of
different species mixtures on slope stability. How root
systems grow when planted individually or in
competition with other species differs, and soil
exploration and exploitation can be highly influenced
by a diversity of root forms (Fig. 7). More work needs
carrying out therefore on plant diversity with regard
to slope stability, as well as how to plant and manage
along a slope, taking into account local variations in
topography and ecology (Genet et al. 2009). To this
end, the development of expert systems and species
databases would be very useful, especially open
source systems where data can be added freely by
end-users and researchers (Norris et al. 2008).
With regard to slope stability in a changing
climate, the impacts of potential climate change
scenarios outlined in the recent Fourth Assessment
Report by the Intergovernmental Panel on Climate
Change (Cruz et al. 2007) on landslide and severe
surface erosion are mixed. For slow, deep-seated
landslides, the likelihood that future climate change
will decrease susceptibility or movement rates is
actually greater than cases where rates would increase
(Sidle and Burt 2009). Shallow landslides are likely to
Plant Soil (2009) 324:1–30
Fig. 7 Illustration of rooting types and depths for seven
herbaceous species growing at 470 m altitude, Klagenfurt,
Austria. From left to right, Dactylis glomerata L., Knautia
arvensis L., Arrhenatherum elatius L., Pastinaca sativa L.,
Bromus hordeaceus L., Carum carvi L., Holcus lanatus L. and
Crepis biennis L.. Where species’ mixtures are diverse, plant
root systems occupy space differently than if grown individu-
increase in a warmer climate if the local frequency of
intense storms increases. Since such landslides are
more influenced by rooting strength, modification of
vegetation cover and harvesting methods in steep,
managed forests may be required to ensure stable
slopes in a changing climate. However, before such
expensive measures are implemented on a widespread
basis, better predictions of the spatial changes in
rainfall response are needed than are currently
available from climate change models. For landslides
and severe surface erosion, more focus needs to be
placed on sustainable anthropogenic practices, which
are much more manageable in the context of current
knowledge and practical policy-making decisions,
than the uncertain impacts of global warming.
Climate change can also affect the expression of
root traits; the increased carbon gain under elevated
CO2 may augment RLD and mycorrhizal coloniza-
tion, whilst at the same time decreasing tissue
nitrogen concentrations (Eissenstat et al. 2000). Tree
wood strength may increase but genetic variation is
great (Beismann et al. 2002). Increases in soil
23
ally or as a monoculture, e.g. the root system of K. arvensis
exploits soil horizontally beneath the root systems of its
neighbours. By increasing the diversity of root forms, the
occupation of soil will be increased, and hence the overall
fixation of soil along a slope (reprinted with permission from
the publisher; see Kutschera and Lichtenegger 1997)
temperature and changes in precipitation patterns will
affect root growth dynamics. Therefore, we need
more understanding of how the expression of traits
and their interactions are affected by the local
environment, in particular soil temperature and
precipitation. At the moment it is not possible to say
with any certainty how traits will be affected because
responses of soil to changes in climate will be
localized. One thing is certain though, once a
landslide has occurred through either land use
mismanagement or the consequences of climate
change, the replacement of soil on the denuded slope
can take up to thousands of years through natural
processes. In a world where the population is
expected to reach 9 billion by 2040 (http://www.
census.gov/ipc/www/idb/worldpopgraph.html), agri-
cultural soil is precious and hillslope stability should
be a priority for governments needing to feed rapidly
increasing populations.
In conclusion, it is clear that biologists and
landslide engineers need to work together to develop
long-term strategies for the ecological management of
24
steep slopes (Norris et al. 2008). We have included
many ideas, data and concepts presented at the
Second International Conference ‘Ground Bio- and
Eco-engineering: The Use of Vegetation to Improve
Slope Stability—ICGBE2’ held at Beijing, China,
14–18 July 2008. Several papers from this conference
are published in this edition of Plant and Soil
(Bischetti et al. 2009a; Fan and Su 2009; Mickovski
and van Beek 2009; Pohl et al. 2009; Reubens et al.
2009; Stangl et al. 2009; Tsakaldimi et al. 2009;
Walker et al. 2009) and several more are published in
a special edition of Ecological Engineering (Bathurst
et al. 2009; Bischetti et al. 2009b; Genet et al. 2009;
Hubble et al. 2009; Loades et al. 2009; Preti et al.
2009; Schwarz et al. 2009; Stokes et al. 2009
submitted; Wang et al. 2009; Zhang and Dong
2009). The third conference in this series will be held
in Vancouver, Canada, 2012 and the first author of
this paper can be contacted for further details.
Acknowledgements Funding for AS, CA and TF was
received from INRA (Jeune Equipe), CNRS (EcoPente project)
and Agropolis Fondation, Montpellier. AMAP (Botany and
Computational Plant Architecture) is a joint research unit which
associates CIRAD (UMR51), CNRS (UMR5120), INRA
(UMR931), IRD (2M123), and Montpellier 2 University
(UM27). The Scottish Crop Research Institute receives grant-
in-aid support from the Scottish Government Rural and
Environment Research and Analysis Directorate. Thanks are
due to the Chinese Academy of Sciences and LIAMA, in
particular B. Hong and X. Zhang, for their help in the
organisation of the ICGBE2 conference.
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