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Mechanical Impedance To Root Growth A Review of Experimental Techniques and Root Growth Responses
Mechanical Impedance To Root Growth A Review of Experimental Techniques and Root Growth Responses
SUMMARY
Mechanicalimpedancetorootgrowthisoneofthemostimportantfactorsdeterminingroot
elongation and proliferation within a soil profile. Penetrometers overestimate resistance to
root growth in soil by a factor of between two and eight and, although they remain the most
convenient method for predicting root resistance, careful interpretation of results and
choice of penetrometer design are essential if improved estimates of soil resistance to root
elongation are to be obtained. Resistance to root growth through pressurized cells contain-
ing ballotini considerably exceeds the confining pressure applied externally to these cells.
Results from this work are reappraised. Existing models of soil penetration by roots and
penetrometers are reviewed together with the factors influencing penetration resistance.
The interpretation of results from mechanical impedance experiments is examined in some
detail and root responses, including possible mechanisms of response, are discussed.
INTRODUCTION
The type of soil strength characteristic (i.e. the variation of soil strength with soil water content)
favourable to crop growth depends on both the amount and the distribution of the annual rainfall,
and on the nature of the crop. The soil must have sufficient mechanical strength to provide adequate
anchorage for the plant throughout its development, and to prevent the collapse of soil water and air
pathways by soil overburden pressure and the weight of vehicle and animal traffic. Dense regions of
high strength may limit root growth and crop yield (Jamieson et al., 1988; Oussible, 1988) by
creating a large mechanical resistance to root growth and/or restricting the rate of oxygen supply to
roots. These dense regions occur in naturally compact soil horizons and also arise from compaction
by heavy farm machinery and by the formation of plough pans.
Mechanical impedance is experienced to varying degrees by virtually all roots growing through
soil. If continuous pores of sufficiently large diameter do not already exist, a root tip must exert a
force to deform the soil. This process may considerably decrease root elongation rates, increase the
root diameter and change the pattern of lateral root initiation (Russell, 1977).
In this paper, the effects of mechanical impedance on root morphology are reviewed and some
direct comparisons between soil resistance to root growth and resistance to a penetrometer are
discussed. The physical process of root growth through soil and artificial media is considered, with
emphasis on the interpretation of results from different experimental techniques. Changes which
occur in root elongation rate under both constant, and temporally and spatially varying levels of
mechanical impedance are considered together with the complicating effects of soil aeration and
water status. Finally, possible physiological mechanisms for the root responses are discussed.
Terminology
Penetrometers provide the best estimates of resistance to root growth in soil, short of direct measure-
ment of root force. Most penetrometers consist of a metal probe with a conical tip fixed onto a
34 1
342 A . G . Bengough & C . E. Mullins
cylindrical shaft (Fig. 1) that is generally of smaller diameter than the cone (normally 80% of the
cone diameter; Gill, 1968; Barley & Greacen, 1967;Bengough, 1990).Penetrometer resistance, Q,, is
defined in Equation (l), where F, is the force required to push the penetrometer probe through the
soil, and A , is the cross-sectional area of the penetrometer cone:
Qp.r= Fp,rlAp,r (1)
The pressure that is exerted on the soil by a growing root cannot at present be measured at every
point on the root surface. In this review, root penetration resistance, Q,, is defined similarly to
penetrometer resistance, but where F, is the component of force directed along the root-axis that the
section of root that moves through the soil must exert on the soil in order to extend, and A , is the root
cross-sectional area, measured behind the elongation region. In common with the literature, the
terms mechanical impedance and root penetration resistance have been used interchangeably.
Elongating Xylem
- Shaft region
Meristemati(
j ,,/ f Muclgel
sheoth
region
7 l;
I
I
Fig. 1. (a) A penetrometer, where F,, A,, oN.and a are as defined in Equations (1) and (2). and (b) a root tip.
M O D E L L I N G M E C H A N I C A L I M P E D A N C E T O P L A N T R O O T S A N D TO
PENETROMETERS I N SOIL
Barley & Greacen (1967) comprehensively reviewed the mechanics of soil deformation and failure
which occur around penetrometer probes, roots and underground shoots. There have since been
several attempts to predict penetration resistances in soil and in ballotini beads from bulk mechan-
ical properties. All but one of these models estimate root resistance by predicting the theoretical
pressure required to expand a cavity in the soil or ballotini. Penetrometer resistance, Qp,is then given
by
Qp=a,(l +pcota)
(Greacen et al., 1968) where uN is the pressure required to expand a cavity in the soil (and is equal to
the normal stress on the surface of the penetrometer cone), p is the coefficient of soil-metal friction,
344 A . G . Bengough & C . E. Mullins
Table 1. Studies involving direct measurement of penetration resistance both to plant roots and to
penetrometer probes
*Root diameter was also measured in the air gap above the aggregate; it is not clear which figure was used.
and a is the cone semi-angle. On the assumption that plant roots experience very little frictional
resistance, Greacen et al. (1968) have shown that this equation can account for much of the large
difference between the resistance experienced by plant roots and by metal probes: Equation ( 2 )
predicts that sharp penetrometers (i.e. small a) will experience a much higher component of
frictional resistance than blunter penetrometers. However, with a semi-angle of more than 30", soil
bodies (that move with the probe) have been observed to form around the probe tip so that soil-
metal friction is no longer involved and Equation ( 2 ) ceases to be applicable (Mulqeen et al., 1977;
Bengough, 1988).
Farrell & Greacen (1966) and Greacen et al. (1968) calculated the pressure required to expand
cavities in the soil by spherical and cylindrical deformation respectively. The advancing probe or
root was accommodated by compression of the surrounding soil. This was assumed to occur in two
distinct regions: an inner zone of compression with plastic failure immediately surrounding the
probe, and a zone of elastic compression outside this. Sharp penetrometers (5" semi-angle) and plant
roots were assumed to deform the soil cylindrically, whereas blunt penetrometers caused spherical
deformation. In calculations for three sandy loam soils, the cavity pressure for cylindrical
deformation was only 25 to 40% of that required for spherical deformation.
The major disadvantage of the Greacen et al. (1968) model is that it requires many laborious
measurements of soil mechanical properties. A simpler approach was adopted by Romkens & Miller
(1971), who equated the pressure required for void-ratio changes occurring in a cylinder of soil
around a root with the pressure required for one-dimensional soil consolidation. The resulting
equation was used to predict the rooting densities at which further root radial expansion would be
inhibited by the expansion of neighbouring roots. Unfortunately, the Romkens & Miller (1971)
model is valid only for saturated cohesionless media and, therefore, is of very limited applicability to
many agricultural soils.
Further confirmation that less stress is required for radial (cylindrical) soil deformation than for
axial (spherical) deformation was provided by Abdalla et al. (1969) and Hettiaratchi & Ferguson
(1973). For any given (elastic) strain in a cylinder of soil ahead of the root tip, it was theoretically
predicted that less stress is required to deform the soil radially than axially (Abdalla et al., 1969).
Mechanical impedance to root growth 345
This theory was complemented by experiments using a large modified penetrometer to demonstrate
that radial expansion behind a penetrometer (or root) tip can reduce axial resistance to soil pen-
etration. Hettiaratchi & Ferguson (1973) predicted theoretically that the pressure required for
cylindrical soil deformation in a frictionless cohesive medium was always less than for spherical
deformation, the difference increasing with cohesion.
Collis-George & Yoganathan (1985) used the spherical cavity expansion model of VesiC (1972)
to define limiting mechanical conditions for seed germination and root growth. Although this model
may be suitable to describe germination conditions, use of spherical expansion theory will have
resulted in overestimates of the resistance to root growth. The VesiC model requires fewer inputs
than the Greacen model, and includes a volumetric strain term. Collis-George & Yoganathan
assumed the volumetric strain to be zero, so that fewer soil mechanical measurements were needed
to perform their calculation. However, their zero-strain assumption is questionable because even a
tiny volumetric strain may, under certain conditions, alter the cavity pressure considerably (VesiC,
1972).
P H Y S I C A L D I F F E R E N C E S B E T W E E N S O I L P E N E T R A T I O N BY P L A N T
ROOTS A N D PENETROMETER PROBES
Roots are flexible organs that follow tortuous paths through the soil, apparently seeking out the
path of least resistance. They extract water from the soil, excrete mucilage from around their tips,
and swell when physically impeded. In contrast, penetrometers are rigid metal probes constrained to
a linear path through the soil. Penetrometers vary from about 0.1 mm in diameter for a small
(needle) penetrometer (e.g. Groenevelt et al., 1984) to over lOmm for a large (field) penetrometer
(the standard ASAE penetrometer cone has a diameter of 20.27mm; ASAE, 1969), and often
penetrate the soil at rates up to two or more orders of magnitude greater than roots (Whiteley et al.,
1981).
The differences between penetrometers and roots have resulted in the expression of much doubt
as to the usefulness of penetrometers (e.g. Russell, 1977, p. 188), but despite their limitations they
remain the best available method of estimating resistance to root growth in soil. It is important,
therefore, to determine what are the most important physical differencesbetween the action of roots
and penetrometers.
Dexter & Tanner (1973) field soil 10,20,30,40 sphere smallest probe
(various textures)
Barley ef al. (1965) remoulded conical no difference
sandy loam (30")
Gooderham (1973, cited - smallest probe
by *below)
Bradford (1980) undisturbed 3.8,5.1 conical no difference
(307
Whiteley ef al. (1981)* undisturbed clods 1.00,1.25,1.50 conical no difference
and remoulded cores 1.75.2.00 (30")
of sandy loam
Whiteley & Dexter remoulded 1.00,1.25,1.50 conical smallest probe
(1981) (various textures) 1.75,2.00 (307
Bengough (1988) undisturbed cores of 0.5,l .O conical smallest probe
sandy loam (307
It is important to distinguish between the vertical component of frictional resistance on the tip of
a penetrometer, and the friction on the shaft, which can account for a greater proportion of the total
resistance to small penetrometers (Groenevelt et al., 1984; Barley et al., 1965; Greacen, 1986).
Probes with a (relieved) shaft of smaller diameter than the tip are used to decrease the component of
shaft friction. The success of this feature may be restricted if the trajectory of the probe tip causes the
shaft to bow and come into contact with the soil, or if soil falls or deforms inwards around the shaft
and rubs against it. Penetrometer resistance in soil cores that are laterally confined inside rigid
cylinders may also be greater if the ratio of core diameter to probe diameter is less than about 20
(Greacen et al., 1969). This effect of confinement is smaller in more compressible soils, where the
probe volume can be accommodated by the compression of a smaller cylinder of soil.
Studies on penetration rate by Eavis (1967), Voorhees et al. (1975), Gerard et al. (1972)
and Gooderham (1973) revealed decreases of less than 20% in penetrometer resistance for decreases
in penetration rate of between one and three orders of magnitude down to penetration rates of
Mechanical impedance to root growth 347
1 mm h- or slower. Although Waldron & Constantin (1970) found a large effect of penetration rate,
an intermittently rotated penetrometer was used, which would have resulted in larger decreases in
soil frictional resistance at slower rates of penetration. In very wet soil, penetrometer resistance is
more clearly linked to penetration rate because of its interaction with pore water pressure (Cockroft
et al., 1969).This effect will be greater in less permeable soils (especially remoulded soil) containing a
higher propertion of silt and clay, than in sands. Penetrometer resistance doubled in a sandy loam
soil remoulded at approximately field capacity for a 100-fold increase in penetration rate, whereas a
250-fold increase in penetration rate resulted in only a 25% increase in penetrometer resistance in
air-dry sand (Bengough, 1988). Similarly, Cockroft et al. (1969) found a doubling of penetration
resistance for a 350-fold increase in penetration rate in saturated remoulded clay. Thus, excluding
very wet and remoulded soils, penetrometer resistance is only weakly dependent on penetration rate
for speeds between those normally used in needle penetrometer measurements and typical rates of
root elongation.
Shape andfriction
Root or probe shape determines both the mode of soil deformation and the amount of frictional
resistance on the tip. Observations of soil movement and density patterns surrounding probes and
roots suggest that both narrowly tapered probes and plant roots deform the soil cylindrically
compared with the spherical deformation caused by blunt probes (Cockroft et al., 1969; Greacen
et al., 1968). By using both lubricated and rotated penetrometer probes it has been demonstrated
that a large component of penetrometer resistance is frictional (Tollner & Verma, 1984; Bengough &
Mullins, 1988). Greacen et al. (1968) suggested that root tips experience virtually no frictional
resistance because of the lubricating action of mucilage secretion and the sloughing off of root cap
cells. If this is so, the best estimate of root resistance may be obtained by measuring the resistance
to a narrowly-tapered probe, and then subtracting the component of frictional resistance using
Equation ( 2 )(Greacen &Oh, 1972; Voorhees et al., 1975).
Interactions between roots, water extraction by roots, root swelIing and root nutation
Because roots seldom grow through soil in complete isolation, it is important to consider inter-
actions between neighbouring roots. Greacen et al. (1969) measured resistance to penetration of a
narrowly-tapered probe, surrounded by six identical probes. Penetration resistance for the central
probe of a group was considerably lower than when the probe was used on its own. Tensile cracking
occurred between the probes, and similar cracks were also observed in a separate experiment
between neighbouring pea radicles growing into a loam. The drying action of roots is very important
in the formation of such cracks, which must facilitate the subsequent growth of lateral roots in a soil
of high resistance (Gerard et al., 1972). Although this cracking is generally likely to be advan-
tageous, there are soils of high tensile strength (e.g. Mullins et al., 1987, 1990) which may not crack
readily under the drying action of roots. In such soils, the increase in penetration resistance caused
by the soil drying may further impede root growth.
It has been suggested by Abdalla et al. (1969) that roots penetrate soil by an alternating series of
radial and axial enlargements. Graf & Cooke (1980) used a finite-element model and, assuming a
low coefficient of root-soil friction and that the soil behaved as a homogenous linear elastic medium,
predicted that the radial expansion of impeded root tips could reduce the axial stress on the root cap.
A zone of stress relief caused by radial enlargement was also predicted by Richards & Greacen
(1986), and by Hettiaratchi & OCallaghan (1974, 1978) who also analysed theoretically the mech-
anics of the changes in cell size and shape that can occur in mechanically impeded root tips. It is clear
that rigid penetrometers cannot mimic this radial expansion of roots, but the potential importance
of this as a mechanism for soil penetration by roots has not yet been fully investigated.
Finally, oae factor rarely commented on with respect to soil penetration is the tendency for roots
to nutate (Greacen et al., 1969;Ney & Pilet, 1981). The magnitude of this motion in highly resistant
soil is probably very small, but it could be a process by which roots locate low-resistance pathways
through heterogeneous soils, and may also reduce soil frictional resistance to root tip penetration.
Nutation has also been suggested as a mechanism that aids soil penetration by rhizomes (Fisher,
1964).
348 A . G . Bengough & C . E. Mullins
EFFECTS O F MECHANICAL IMPEDANCE ON ROOT GROWTH
Experimental techniques
Existing experimental techniques can be divided into several different catagories.
Soil. Experiments in soil are more realistic, but it is difficult to ensure that mechanical impedance
is the only soil factor limiting root growth. If a soil is compacted to increase resistance to root
growth, the resulting decrease in porosity may result in poor aeration. Similarly, increasing soil-
water tension to increase soil strength may result in water stress. However, the greatest difficulty is in
determining the penetration resistance experienced by roots in soil. This is ideally determined by
direct measurement (e.g. Stolzy & Barley, 1968; Eavis & Payne, 1969; Bengough & Mullins, 1988),
but practical difficulties have led most researchers to use penetrometer resistance measurements to
estimate root resistance (e.g. Greacen &Oh, 1972).
Pressurizedcelis. Root growth has been studied in artificial systems where it was intended that a
constant, uniform and known value of mechanical impedance could be imposed, while simul-
taneously maintaining a carefully controlled supply of aerated nutrient solution to the roots. Goss
(1977), Abdalla et al. (1969) and Barley (1963) grew roots in ballotini contained in flexible-sided cells
which could be subjected to an external confining pressure (Fig. 2).
(a) (b)
Root inlet
Root inlet
Gas inlet
Gas under
pressure, ud
Polythene
diaphragm
Nylon cloth
Pressure
inlet
u Lu
0 2 4 01 2
cm cm
Fig. 2. (a) A triaxial cell (after Barley, 1963), and (b) a pressurized diaphragm apparatus (after Barley, 1962).
Fig. 2(a) is reproduced from K.P. Barley, Influence of soil strength on growth of roots, Soil Science 1963,96(3),
175-180 ( 0 Williams & Wilkins, 1963).
Resistance to root growth in ballotini has been taken as either equal to (Russell & Goss, 1974;
Goss, 1977; Goss & Russell, 1980; Veen, 1982), or an order of magnitude greater than (Richards &
Greacen, 1986; Bengough & Mullins, 1990) the pressure applied externally to the boundary of the
growth medium (u3in Fig. 2). Goss (1977), Veen (1982), Richards & Greacen (1986) and Bengough
& Mullins (1990) all estimated resistance to root growth by measuring the pressure required to
inflate a small rubber tube within the pressurized ballotini cell. The tube inflation pressure depends
entirely on the detailed procedure followed (Bengough & Mullins, 1990).
Goss (1977), Veen (1982) and Bengough & Mullins (1990) inserted a tube inflated under
atmospheric pressure only and unsealed at one end into an unpressurized cell of ballotini. An
external confining pressure was then applied to the cell, causing the tube partly to deflate. The
Mechanical impedance to root growth 349
pressure required to reflate the tube to its original volume was then found to be equal to the pressure
applied externally to the ballotini cell. In contrast, when the tube is not allowed to deflate during
pressurizing of the cell, the pressure required to expand the tube beyond its initial volume is between
5 and 10 times higher than the pressure applied externally to the ballotini cell (Richards & Greacen,
1986; Bengough & Mullins, 1990). This is attributable to the frictional resistance to deformation of
the ballotini. Since roots penetrating the ballotini must exert pressure to expand a new cavity in
previously undisturbed ballotini, the latter experiment gives more accurate representation of the
resistance experienced by growing roots. Although the pressure required to inflate a tube in the
ballotini will depend on both the tube diameter and on the frictional properties of the tube walls, it
seems reasonable to conclude that the resistance to root growth in the cells considerably exceeded
the external confining pressure.
Richards & Greacen (1986) also used a finite-element model to predict the effect of tube inflation
pressure on tube diameter in ballotini and in sand. Tube diameter increased slowly until the internal
pressure reached a certain critical value, when the diameter increased much more rapidly (Fig. 3).
Predicted inflation pressures were much higher than the external confining pressure, and the
difference was greater for sand than for ballotini.
36~
I I
I
I
I
I
i
I
I
I
I
I
I
E I
E I I i
v I
I I I
3.0} I / I I
/
/
/
I 1 I 1 I I
0 0.05 0.10 0.15 0.20 0.25
Inflation pressure (MPa)
Fig. 3. Outside diameter of tube vs inflation pressure in ballotini at several constant external cell pressures
(indicated at the top of each curve), after Richards & Greacen (1986). Experimental (-); theoretical
(----)
Pressurized diaphragms. Barley (1962) and Gill & Miller (1956) adopted slightly different tech-
niques, using an externally-pressurized rubber diaphragm to exert pressure on roots growing down a
porous plate, separated from the diaphragm by nylon cloth or ballotini respectively (Fig. 2). In these
experiments, asymmetric stress was applied to the whole root length. These authors suggested that
resistance to root growth might be considerably greater than the diaphragm pressure (o,,in Fig. 2).
Barley (1962) estimated resistance to root growth by measuring the pressure required to inflate a
small tube placed under the diaphragm. He assumed a coefficient of soil-root friction of 0.25 and
estimated that resistance to root elongation varied between 2 and 7 times the diaphragm pressure.
Gill & Miller (1956) observed that if roots were well-covered with ballotini, elongation ceased at
lower diaphragm pressures. The authors suggested that arching of the ballotini caused small
displacements of single glass beads to require larger displacements of the diaphragm. Thus,
resistance to root elongation was considerably greater than the diaphragm pressure.
Pressurized airlwater. Chaudhary & Aggarwal(l984) proposed growing seedlings in moist sand
inside a pressure vessel as an experimental technique to measure the effect of mechanical resistance
on root elongation. However, although application of air pressure would cause a corresponding
350 A . G . Bengough & C . E. Mullins
increase in the absolute value of root cell turgor pressure, the decrease in osmotic potential or cell
wall tension required for root cell extension would be independent of the externally applied pressure
(assuming the cell permeability to water remained unchanged). Thus, the experiment does not truly
represent the situation of roots elongating against an external mechanical resistance. The reason for
the large decrease in root growth rates observed by the authors could be the 10-fold increase in
dissolved gas concentration in the water inside the pressure chamber, and ultimately in the plant,
resulting from the 10-fold increase in air pressure that they applied (Henry's law).
m
?
5; 60
m
t
W
3
-
0
c
.-0
5c 20
-
0
W
There have been only three studies in which both root elongation rate and root resistance (as
directly determined from the root force) have been measured (Eavis, 1967; Stolzy & Barley, 1968;
Bengough & Mullins, 1988). The results of Eavis (1967) have been replotted in Fig. 5 to show the
relationship between root elongation rate and root penetration resistance. This shows that root
elongation rate was reduced by about 50% by a resistance to root growth of approximately
0.3 MPa.
Bengough & Mullins (1988) found that maize root elongation rates in cores of two sandy loam
soils were reduced to between 50% and 90% of that of control plants grown in loose sieved soils, by
root penetration resistances of between 0.39 and 0.48 MPa (based on the initial root tip cross-
sectional area). Stolzy & Barley (1968) found that the elongation rates of two pea radicles were
reduced to 44% of their unimpeded rate by a root penetration resistance of 0.46 MPa.
Mechanical impedance to root growth 35 1
- 100
+
c
E
+
0 80
P
+
ul
P
60
c
0
._
20
c
-
0
W
Fig. 5. Root elongation vs stress, as specified below (see Table 3 for details) obtained in six studies:
A. Abdalla et al. (1969):external confining pressure
9. Barley (1962):estimated root penetration resistance
C. Barley (1963):external confining pressure
D. Eavis (1967): measured root penetration resistance
E. Gill & Miller (1956): diaphragm pressure
F. Goss (1977):external confining pressure
G . Greacen & Oh (1972): normal stress on penetrometer cone.
The results of these direct experiments represent the maximum effect of mechanical impedance
to be expected on root elongation rate for any particular root resistance. Reduced oxygen or
nutrient supply to the roots can further limit the observed 'root elongation rate, and similarly
anchorage of the tip by root hairs can result in the resistance being underestimated.
All the experimental relationships in Figs 4 and 5 show similar patterns in which elongation rate
decreases with increasing mechanical resistance. If the results of direct experimental measurements
of root resistance and elongation rate (Stolzy & Barley, 1968; Eavis, 1967) are compared with results
from penetrometer studies (e.g. Taylor & Ratliff, 1969), the factor of between 2 and 8 by which
penetrometer resistance exceeds root resistance is sufficient to remove any apparent discrepancy.
Similarly, the factor of between 5 and 10 by which resistance to root growth exceeds the confining
pressure externally applied to the boundary of the growth medium is sufficient to bring the results of
experiments performed in ballotini cells into broad agreement with the other experimental
approaches.
Spatial variation. The elongation rate of roots growing into a zone of greater mechanical impe-
dance has been observed to decrease with time (Barley, 1962, 1963). However, Barley notes that
352 A . G . Bengough & C . E. Mullins
Table 3. Techniques used in studies of the effect of mechanical impedance on root elongation
rate (Figs 4 and 5 )
the effects may have merely been due to a greater length of root being compressed (Barley, 1962),
and resistance to root growth increasing gradually with depth (Barley, 1963), respectively. The
elongation rate of roots encountering a very small mechanical resistance has been found to decrease
transiently by a surprisingly large amount (Goss & Russell, 1980). Maize root elongation rates
decreased to less than one-third of their unimpeded rate during the 10 min period after starting to
grow through loosely packed ballotini. However, during the subsequent 10 min the elongation rate
Mechanical impedance to root growth 353
returned to its original unimpeded value. Identical experiments performed using roots that first had
their root caps removed showed no significant changes in root growth rate. Thus, the root cap must
have a role in regulating root response to very low resistances, although it is uncertain whether the
same mechanism operates at the greater resistances more often found in soil.
CONCLUSIONS
Except where direct measurements of root force are available, penetrometer resistance is currently
the best method for estimating resistance to root growth in soil, although subject to many limi-
tations. The relationship between penetrometer resistance and root resistance needs to be more
clearly established by direct measurements of both root force and penetrometer resistance in a
variety of differing soils over a range of bulk densities and matric potentials. It is probable that
improved estimates of root resistance may be obtained by subtracting the frictional component of
penetrometer resistance (determined by independent measurement of the coefficient of soil-metal
friction) from the total resistance to a narrowly tapered probe. The effects on penetration resistance
of penetrometer or root diameter still require a fuller investigation.
Root elongation rate is progressively decreased by increasing mechanical resistance to growth,
and ceases at root penetration resistances of about 1 MPa. Resistance to root elongation within a
ballotini cell confined by an externally applied pressure may be up to one order of magnitude greater
than the externally applied pressure. The mistaken interpretation of results from such experiments
has caused the apparent disagreement with results from penetrometer studies in soil. Pressurized
ballotini cells still provide a valuable technique for studying the effects of mechanical impedence on
root morphology and nutrient uptake (Goss, 1977; Lindberg & Pettersson, 1985), and are poten-
tially of considerable use for investigating the largely unresearched ability of different species and
varieties to grow against mechanical impedance (Goss, 1974). The changes in root growth rate
which occur as a root enters or leaves a zone of high mechanical impedance are of considerable
relevance to real soils, but have received surprisingly little attention.
The mechanism of root response to mechanical impedance may involve osmotic adjustment,
although insufficient evidence has been presented to test such a model. The role of ethylene must be
re-assessed in the light of some recent experiments, and its type of involvement in the response of
roots to mechanical impedance must be more clearly established.
ACKNOWLEDGEMENTS
We thank Drs D. Linehan, B. Marshall, I. Young, D. Robinson and other colleagues at SCRI for
their helpful discussions and constructive criticism of this manuscript.
356 A . G . Bengough & C . E. Mullins
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