Aust. J. Soil Res.

, 1991, 29, 717-28

Soil Structure and Plant Growth

J. B. Passioura
Division of Plant Industry, CSIRO,
G.P.O. Box 1600, Canberra, A.C.T. 2601.

Abstract
Soil structure affects plant growth in many ways. Roots grow most rapidly in very friable
soil, but their uptake of water and nutrients may be limited by inadequate contact with the
solid and liquid phases of the soil. This contact is much more intimate in hard soil, but
then the growth of the roots is strongly inhibited, so that their foraging ability is poor, and
the plant may eventually become short of water or nutrients. However, many soils, even
if hard, contain continuous macropores that provide niches for the roots to grow in. The
presence of such macropores increases the extent of the root system, but because the roots
are clumped within them, the rate at which the roots can extract water and nutrients from
the soil between the macropores is considerably slowed. These macropores also provide
niches for microorganisms, both symbiotic and pathogenic, so that the response of roots to
different tillage treatments may differ markedly on this account alone. Soil structure not
only affects the ability of roots to grow and to supply the leaves with water and nutrients;
if adverse, it also induces them to send hormonal signals that slow the growth of the shoot,
even if they are currently able to take up adequate water and nutrients.

Keywords: soil structure, plant growth, root growth, root signals, macropores, water uptake.

Introduction
Soil structure affects the behaviour of plants in many ways. The most
obvious effect is on the appearance of the roots, which are generally smooth
and cylindrical in friable soil, but are stubby and gnarled in compacted soil
and are greatly restricted in their range, with potentially deleterious effects on
the supply of water and nutrients. In addition there are many less obvious but
often equally influential effects on the overall performance of the plant. The
soil may sometimes be too porous for the roots to make enough contact with
the solid and liquid phases to extract seemingly available nutrients or water.
The presence of macropores (continuous large pores that roots preferentially
occupy) may enable the roots to traverse otherwise impenetrable soil, and
thereby gain access to a larger reservoir of water and mobile nutrients.
Such macropores are not only different physically, but also chemically and
microbiologically, from the bulk soil, and may be enriched in pathogenic or
symbiotic microorganisms or depleted of nutrients, so that roots growing in
them may be affected in many ways. Finally, the structure of the soil may
not only influence the current ability of the roots to provide the shoot with
 J. B. Passioura

adequate water and nutrients; it may also induce roots in adverse soil to send
inhibitory signals to the leaves even though the roots are currently able to take
u p adequate water and nutrients. I will discuss these various aspects in turn,
emphasising especially effects on the overall growth of plants, but ignoring
the well-established indirect effects arising from the influence of structure on
the aeration of the soil.

Root Growth in Hard Soil

The elongation rate of roots is highly correlated with the penetrometer
resistance of uniform soil in which they are growing, despite the different
mechanisms by which roots and penetrometers deform the soil. The commonly-
used 60" cone penetrometers tend to compress the soil in front of them,
whereas roots compress the soil radially, thereby avoiding a pile-up of soil
in front of them (Greacen et a/. 1968), with the result that the axial pressure
needed by a root to penetrate the soil may be only 20% of that needed with the
penetrometer (Whiteley et al. 1981; Hettiaratchi 1990; Bengough and Mullins
1 9 9 0 ~ ) . The penetrometer resistance is strongly affected by both the bulk
density and the water content of the soil (Williams and Shaykewich 1970).
Taylor and Ratliff (1969), who worked with peanut and cotton, found that
the correlation between elongation rate and penetrometer resistance remained
strong no matter whether the resistance was varied by changing bulk density
or water content (Fig. 1). This was true, surprisingly, even when the water
content was so low that the soil water suction was of the order of 1 MPa.
On the other hand, Veen and Boone (1990), working with maize, found that
the suction and the penetrometer resistance were effectively additive in their
effects on elongation rate. This apparent disagreement may be a result of
the differing species used, which may adjust the osmotic pressure of their
expanding cells in different ways in response to a high suction. Materechera
et al. (1991) have shown marked variation among seedlings of 22 different
plant species in the ability of their roots to penetrate hard soil.
There has been controversy about the mechanism by which the elongation
rate of roots is reduced by hard soil. Greacen and Oh (1972) proposed a
simple physical model of root growth in which the elongation was controlled
by the difference between the turgor pressure within the expanding cells and
the penetration resistance of the soil. In contrast, Russell and Goss (1974)
and Goss (1977) have argued that the elongation rate of roots is so sensitive
to very low pressures applied to soil in triaxial cells in which the roots are
growing, that a simple physical model is not feasible, and that hormonal
control is involved. Richards and Greacen (1986), Hettiaratchi (1990), and
Bengough and Mullins ( 1 9 9 0 ~ )countered by pointing out that the resistance
actually experienced by such roots is much greater than that applied to the
cell, so that a physical model of root growth into a hard soil is not ruled out
by the apparent sensitivity of the roots to small confining pressures.
Many have suggested that the plant hormone, ethylene, may be responsible
for the changed shape of impeded roots (e.g. Kays et al. 1974; Veen 1982), for
it has a similar effect on the shape when applied to unimpeded roots. But Moss
et al. (1988) have shown that roots treated with an ethylene inhibitor develop
Soil Structure and Plant Growth

similar shapes to untreated roots in compacted soil, so that the physical

explanation remains plausible. No doubt, when we eventually understand
this system better, we will find that biochemical and physical effects are
both important. The processes controlling the orientation of the cellulose
microfibrils in the walls of the expanding cells probably hold the key to this
understanding (Hettiaratchi 1990; Williamson 1990). Bengough and Mullins
(1990b) have written an excellent review of this general topic which covers
the points I have touched on in considerable depth.
r 60

Penetrometer resistance (MPa)

Fig. 1. Elongation rates of peanut and cotton roots, and leaf area of wheat seedlings, as
functions of penetrometer resistance of the soil in which they were growing. The discrete
data are not shown for reasons of clarity: they deviate little from the lines. In all three
cases the penetrometer resistance was varied by changing either the bulk density or the
water content of the soil. The root data are adapted from Taylor and Ratliff (1969), and the
leaf data from Masle and Passioura (1987).

Much of the research on the effect of soil strength on root growth has been
done in controlled conditions with uniform soil. But in the field it is common
for roots to grow in macropores (Russell 1977; Wang et al. 1986; Tardieu
1988; Hasegawa and Sato 1987; Hatano et al. 1988). These macropores may
have been formed by worms or by the roots of previous plants (biopores),
or by gross movement of the soil to form slickensides and similar fissures
(Hamblin 1985). Roots that gain access to them are able to penetrate the soil
much more deeply, although their ability to extract water and nutrients is
restricted owing to their being clumped, as I elaborate on below.
Uptake of Water
Because hard soil inhibits root extension, the roots may eventually be unable
to supply enough water or nutrients to the leaves (Morris and Daynard 1978;
Barraclough and Weir 1988), and overall growth is reduced. Such problems
are particularly evident if a plough pan is present. Spectacular responses are
sometimes achieved by deep ripping, which enables roots to penetrate the
subsoil thereby greatly increasing their effective supply of water (Hamblin 1985).
 J. B. Passioura

Where roots are predominantly clumped together in macropores, the clumps

may be so widely spaced in the soil that nominally available water is poorly
accessible. The result may be that roots fail to extract such water, especially
from deep in the subsoil, even though a substantial length of root is present
at this depth (Jordan and Miller 1980). The general principles involved can be
explored by extending the well-established single-root model for the uptake of
water (e.g. Tinker 1976) so that it applies to clumps of roots. The following
treatment is adapted from Passioura (1985).
The central assumption of the single-root model is that water flows radially
to a typical root in response to gradients in pressure (or water content),
whence the cylindrical diffusion equation applies. Although this equation is
generally difficult to-solve, several good approximate solutions are available
of which one of the most useful, once the soil is dry enough for the flow of
water through it to be limiting uptake by the roots, is:

where de/dt is the (quasi-steady) rate of fall of mean soil water content, 8,
with time, t; D is the diffusivity of soil water, which is approximately constant,
with a value of about 2 cm2 day-l, during the extraction of about the last
third of the available water in the soil, when flow through the soil is likely
to be limiting the rate of uptake; 8, is the soil water content at the surface
of the root; and b is the radius of a putative cylinder of soil surrounding the
root, to yhich that root has effectively sole access, and can be calculated as
b = (nL)-7, where L, the rooting density, is the length of root per unit volume
of soil.
If we assume further that 8, is constant, as it would be if the root were
maintaining a constant water potential of say -1 - 5 MPa at its surface, then
equation (1) can be integrated to give:

where 8 d is the difference between 3 and 8,, 8 d o is 8 d when t = 0, and t*

(equal to 2b2/D) is the time constant for the system, that is, the time taken
for 8 to fall to l / e (i.e. 0.37) of its initial value. If D is 2 cm2 day-l then t*
reduces to the very simple form, b2 day. If the roots are evenly distributed in
the soil, then, even if the rooting density, L cm ~ m - is ~ ,very low, say 0.1, t*
(calculated from b2 = l/[nLl) is only about 3 days, and it is hard to imagine
why plants would fail to extract all the available water from the soil occupied
by their roots.
However, if the roots are clumped into sparse biopores, it is no longer
appropriate to calculate t* by assuming that the roots are evenly distributed.
The clumped system is analagous to the single-root model, with the distinction
being that instead of each root having exclusive access to a cylinder of soil of
radius b, we assume that each clump of roots has sole access to a cylinder of
soil of radius B, where B2 = 1/(nLX),and L* is the length of occupied pore per
unit volume of soil. Equations (1) and (2) remain applicable provided that we
Soil Structure and Plant Growth

replace b with B. Radius B may be many times larger than b so that t* could
become of the order of weeks; for example, one can calculate from data of
Wang et al. (1986) for the frequency of biopores in the soil under a soybean
crop that t* might range from 10 to 100 days at depths in the soil between
1 . 1 and 1 - 3 m .
-r

slabs h = l )

!/

I-
Structural s c a l e (crnl
Fig. 2. Time constant for the uptake of water by roots as a function of the structural scale
of the macropores in which the roots are constrained. For the biopores (e.g. wormholes) the
structural scale is half the average distance between adjacent pores. For roots constrained to
fissures delineating impenetrable blocks in the soil, the structural scale is half the distance
between the faces of the slabs, prisms, or cubes, into which the soil is assumed to be
divided. The value of n denotes the number of dimensions in which the water is assumed
to flow through the blocks (see text).

Equations (1) and (2) deal with centripetal flow to single roots or clumps of
roots in biopores. Where the roots are clumped into fissures, these equations
have to be changed. It is useful to idealise the arrangement of such fissures
into: a set of parallel planes with a spacing of 2W; two sets of intersecting
planes at right angles, which cut the soil into prisms, or columns, of width
2W: or three sets of intersecting planes at right angles, which cut the soil into
cubes with sides of length 2W. Estimates of the time scales for the extraction
of water by roots occupying the fissures that delineate these blocks of soil
can be got by assuming further that the columns can be approximated by
cylinders, and the cubes by spheres, of radius W. Inspection of Figs. 4.6, 5.7,
and 6 - 4 of Crank (1975), which show the loss of material by diffusion out
of a slab, a cylinder, and a sphere, respectively, shows approximate values
of t* of w 2 / 4 n ~ ,where n is 1, 2, or 3, for the slab, cylinder, and sphere,
respectively. Note that t* at a given length scale increases as the flow lines
of water change from divergent (as in flow out of a cubic ped), through
parallel, to convergent (as in flow towards a wormhole). Fig. 2 illustrates the
dependence of t* on B or W with different types of geometry. Hasegawa and
Sato (1987) worked with soil that was fractured vertically into prisms whose
 J. B. Passioura

diameter (2W) varied from 5 to 35 cm. Roots grew predominantly within the
fissures, so, in the context of Fig. 2, n is equal to two, so that the t* ranges
from about 2 to 20 days.
Where roots are clumped we cannot expect to understand the extraction of
water by measuring L and using the single root model, and we must devise
some ways of measuring to what extent the roots are clumped. In practice
we may find, with well-developed crops or pastures whose transpiration rate
is limited by the rate of water uptake rather than by evaporative demand,
that t* is a robust parameter which is more a function of the soil than of the
type of crop or pasture, although it will vary with depth in the soil.
These models assume that uptake is limited by the transport of water
through the soil, but there is some evidence that species may differ in the
inherent ability of their individual roots to extract water from the subsoil. For
example, Bremner et al. (1986)showed that sunflower was much more effective
than sorghum at extracting water from a heavy fractured subsoil, even though
both crops appeared to have similar overall distributions of roots. Possible
explanations for this difference are that sunflower roots are better able to
establish hydraulic continuity with peds when growing in the cracks between
them, that is, that the sorghum roots had a large interfacial resistance with
the soil, or that the sunflower managed to produce lateral roots that could
penetrate the peds, while sorghum could not. If such differences between
species are confirmed, the possibility arises of breeding plants that are better
able to extract water from difficult subsoils.
There is also some evidence that a substantial interfacial resistance to the
flow of water between root and soil may sometimes occur (Faiz and Weatherley
1977;Herkelrath et al. 1977a;Zur et al. 1982). Such a resistance seems to be
associated with roots in very sandy or friable soils (Passioura 1988),and may
depend on the extent to which the surface of the roots is wetted (Herkelrath
et al. 1977b). A freely-drained sand will typically have a volumetric water
content below 0.1,so that, as a first approximation, the epidermis of the root
may have a similar proportion of its surface, i.e. 0 -1, covered with water-filled
pores. If the hydraulic resistance of the epidermis is substantial, then the
channelling of the water flow to a small area may induce sufficiently large
gradients in water potential that the underlying cortical cells may shrink a
little (Huck et al. 1970;Passioura 1988),thereby exacerbating the problem of
hydraulic continuity between root and soil and inducing a large interfacial
resistance. !n very friable soils the same phenomenon may occur even with
a large volumetric water content, for the roots may be able to push soil
aggregates aside so easily that neither root nor aggregate is deformed and
the area of contact between the two is small.

Nutritional Implications
If the large interfacial hydraulic resistance that occurs in sands or very
friable soils does indeed arise because of inadequate wetting of the epidermis
of the root, then the uptake of nutrients in such soils is probably also
affected. Robinson et al. (1991)found that only about 10% of the root length
of nitrogen-deficient wheat seemed to be effective in taking u p nitrate. It is
Soil Structure and Plant Growth

feasible that the ineffectiveness of their roots may have been at least partly
due to inadequate wetting of the epidermis.
A root growing in hard soil is morphologically different from one in friable
soil. As well as being shorter and thicker, it is much more closely appressed
to the soil particles (Baligar et al. 1975; Atwell 1990). Furthermore, it typically
exudes copious mucilage that binds it to the adjacent soil particles. The
intimate connection between root and soil in these circumstances may especially
improve the ability of the roots to extract poorly soluble nutrients from the
soil. For example, Passioura and Leeper (1963) showed that the ability of oats
to take up manganese from deficient soil was so improved by compacting the
soil that the plants grew many times faster.
Cornish et al. (1984) argued that bulk density affected the ability of ryegrass
roots to extract phosphorus in two contrasting ways. A higher bulk density
brought more phosphorus into the root-hair zones and thereby increased the
availability, but it also reduced the total root length, owing to its effect on
elongation rate, and thereby decreased the availability. The preponderance of
one effect over the other depended on the water content.

Microbial Interactions
Little is known about the microbiological heterogeneity in the soil that
presumably accompanies the interaction between roots and biopores. Evans
and Miller (1988) showed that maize growing in soil low in available phosphorus
grew well if the soil was undisturbed, but suffered severe phosphorus deficiency
if the soil was ploughed. This startling effect was due to differing infections of
the roots by mycorrhizae. The difference may have arisen in part because roots
in the undisturbed soil preferentially occupied biopores that were rich in the
fungus so that the symbiosis developed rapidly. However, later work by Evans
and Miller (1990) showed that it was the presence of a well-developed hyphal
network in the undisturbed soil that accounted for much of the effectiveness
of the symbiosis.
If biopores are a haven for symbiotic fungi, the chances are that they are also
a haven for pathogens. The evidence of Chan et al. (1989) that direct-drilled
wheat is more prone to disease may reflect such an interaction. Rovira et
al. (1990) discuss the general issue of soil conditions and root diseases in
considerable detail.

Hormonal Effects
Plants grow more slowly in hard soil than in loose even when their roots
are able to supply adequate amounts of water and nutrients (Fig. 1). The
roots are apparently able to sense the physical conditions in the soil and
then send a signal to the shoot that lowers the expansion rate and stomata1
conductance of the leaves (Masle and Passioura 1987). Richards and Rowe
(1977), Peterson et al. (1984), Carmi (1986), and Ruff et al. (1987) found a
similar effect in peach, wheat, cotton, and tomato, respectively, when they
grew the plants in pots of different sizes to which they supplied adequate
water and nutrients; plants in large pots grew much more quickly than those
in small pots, perhaps because the roots perceived the walls of the pots to
 J. B. Passioura

be impenetrable 'soil', and because the proportion of roots that encountered

the wall would have been much higher in small pots than in large.
Similar inhibitory signals are sent to the shoot by roots growing in drying
soil (Passioura and Gardner 1990). Given that the penetrometer resistance of
most soils increases sharply as the soil dries, it is likely that these signals
are induced as much by the hardening of the soil as it dries, as by the fall
in water potential in itself. Fig. 3 shows the relative expansion rate of the
leaves of wheat plants growing in a drying clay loam, at two bulk densities,
a s a function of soil water deficit. One of the bulk densities was so low that
the penetrometer resistance remained low at all water contents. The other
was high enough to induce the penetrometer resistance to rise to substantial
levels, in the context of Fig. 1, as soon as the soil dried below its initial water
content. The leaf expansion rate of the plants in the soil of high bulk density
fell below that of well-watered controls soon after the soil started to dry,
whereas the plants in the drying loose soil kept up with their well-watered
counterparts until the soil water potential fell below about -400 kPa.
-
-
bP 120
1

Soil water content (g g-'1

Fig. 3. Expansion rate of leaves of young wheat plants (as a percentage of that in well-watered
controls) growing in drying soil of two bulk densities. The open symbols denote that the soil
had a low bulk density; the penetrometer resistance never exceeded 1 . 0 MPa at any water
content. The closed symbols denote that the soil had a high bulk density; the pentrometer
-
resistance w2s 2 0 MPa at a water content of 0 - 25, and rese !inearly with decreasing water
content until it reached 6 MPa at a water content of 0.15 (cf. the data on leaf area as a
function of penetrometer resistance in Fig. 1). The starred points denote that the watered
and unwatered plants differed significantly in leaf expansion rate. (Redrawn from Passioura
and Gardner 1990.)

It is likely that the signal sent by the roots to the shoot when the roots
are constrained in some way is a chemical transported in the transpiration
stream. The chemical may be abscisic acid (ABA) as Zhang and Davies (1990)
have argued in relation to plants growing in drying soil, but Munns and King
(1988), also working with plants in drying soil, showed that the concentration
of ABA was much too low, at least in wheat plants, to account for the inhibitory
effects on the leaves and that a potent unknown inhibitor was involved.
Soil Structure and Plant Growth

Another physical feature of soil that appears to affect the growth rate of
plants without necessarily affecting the uptake of water and nutrients is that
of aggregate size. Donald et al. (1987) sieved aggregates of different sizes
from a loam and found that maize plants growing in pots filled with the
smallest of the aggregates (4- 6 mm diameter) grew substantially better than
plants growing in the larger aggregates. All plants seemed to have adequate
supplies of water and nutrients, judged from analyses of the leaves, so we
are left with the conclusion that, as with plants growing in hard soil or small
pots, a signal from the roots was affecting the growth rate of the shoot.
Misra e t al. (1988) grew cotton and sunflower plants in pots containing
aggregates of different sizes (ranging from 1 . 5 mm to 16 mm in diameter)
and also found that the plants grew best in the smallest aggregates. They
interpreted this behaviour in terms of the probable greater ability of the
roots in the small aggregates to extract phosphorus from the soil, but they
presented no evidence that the plants were deficient in phosphorus, and their
data showed no decrease in phosphorus concentration in the plants with
increasing aggregate size; indeed, the sunflower plants had lower phosphorus
concentrations in the leaves when grown in the small aggregates than in
the large. Thus, as with the experiments of Donald e t al. (1987), those of
Misra e t al. (1988) point to growth being affected by a non-nutritional signal
from the roots when the roots are growing mainly in large pores, i.e in the
interaggregate spaces.
These observations have implications for the performance of plants whose
roots are growing preferentially in macropores. If the macropores are particularly
large, for example wormholes, which have diameters many times that of a
root, the plants may react in a similar way to those of Donald e t al. (1987)
that were growing in soil comprised of large aggregates, and slow their growth
even if the roots are able to supply the leaves with adequate amounts of
water and nutrients.
Conclusion
Soil structure affects plant growth in many, often surprising, ways. The
most obvious effects are on root growth, which is strongly inhibited by hard
soil, and which in turn influences the ability of the root system to extract
adequate water and nutrients from the soil. A more subtle effect occurs
in soil that is hard but which also contains biopores and fissures through
which roots can readily grew. The r m t s are c!umped tegether in such sci!s,
and while the overall rooting density may seem adequate for the uptake of
water and nutrients, the soil in the middle of large peds, or remote from a
biopore, may be essentially inaccessible to the roots. The time needed for
water and nutrients to flow to the roots in such circumstances may be such
a large proportion of the life of a crop that time runs out before they can be
extracted. Perennial pastures presumably do not have this problem.
Another effect occurs because of the spatial heterogeneity of the soil's
microflora. Roots growing in biopores may be exposed to much higher
population densities of both symbiotic organisms such as mycorrhizae, and
pathogens, than would be expected by analysing an average sample of soil.
With plants such as maize that depend strongly on mycorrhizae to extract
 J. B. Passioura

phosphorus from the soil, having access to biopores rich in mycorrhizae may
mean the difference between health and severe phosphorus deficiency.
The most startling effects of all occur when the roots respond to the
physical nature of the soil by sending hormonal signals to the leaves. Roots
send such signals even if they are currently able to extract adequate water
and nutrients from the soil; and they may do so when they are growing in
soil that is too hard, or too restricted in volume, or which has pores that are
too large. Although these effects are subtle, they can be large in terms of
growth, and it may often be important to consider them when analysing the
behaviour of crops and pastures.

Acknowledgments
I thank John Kirkegaard and Richard Stirzaker of CSIRO, and Glyn Bengough,
David Robinson, and Iain Young of the Scottish Crop Research Institute, for
many helpful discussions.

References
Atwell, B.J. (1990). The effect of soil compaction on wheat during early tillering. I. Growth,
development and root structure. New Phytologist. 115, 29-35.
Baligar, V.C., Nash, V.E., Hare, M.L., and Price, J.A. (1975). Soybean root anatomy as influenced
by soil bulk density. Agron. J. 67, 842-4.
Barraclough, P.B., and Weir, A.H. (1988). Effects of a compacted subsoil layer on root and
shoot growth, water use and nutrient uptake of winter wheat. J. Agric. Sci., Camb. 110,
207-16.
Bengough, A.G., and Mullins, C.E. ( 1 9 9 0 ~ )The
. resistance experienced by roots growing in a
pressurised cell: a reappraisal. Plant Soil. 123, 73-82.
Bengough, A.G., and Mullins, C.E. (1990b). Mechanical impedance to root growth, a review
of experimental technique and root growth responses. J. Soil Sci. 41, 341-58.
Bremner, P.M., Preston, G.K., and Fazekas de St. Groth, C. (1986). A field comparison of
sunflower (Helianthus annuus) and sorghum (Sorghum bicolor) in a long drying cycle. I.
Water extraction. Aust. J. .Agric. Res. 37, 483-93.
Carmi, A. (1986). Effects of root zone volume and plant density on the vegetative and
reproductive development of cotton. Field Crops Res. 13, 25-32.
Chan, K.Y., Mead, J.A., Roberts, W.P., and Wong, P.T.W. (1989). The effect of soil compaction
and fumigation on poor early growth of wheat under direct drilling. Aust. J. Agric. Res.
42, 221-8.
Cornish, P.S., So, H.B., and McWilliam, J.R. (1984). Effects of soil bulk density and water
regimen on root growth and uptake of phosphorus by ryegrass. Aust. J. Agric. Res. 35,
631-44.
Crank, J. (1975). The Mathematics of Diffusion. (Clarendon: Oxford.)
Donald, R.G., Kay, B.D., and Miller, M.H. (1987). The effect of soil aggregate size on early
shoot and root growth of maize (Zea mays L.) Plant Soil. 103, 251-9.
Evans, D.G., and Miller, M.H. (1988). Vesicular-arbuscular mycorrhizas and the soil-
disturbance-induced reduction of nutrient absorption in maize. New Phytologist. 110,
67-74.
Evans, D.G., and Miller, M.H. (1990). The role of the external mycelial network in the effect
of soil disturbance upon vesicular-arbuscular mycorrhizal colonization of maize. New
Phytologist. 110, 67-74.
Faiz, S.M.A., and Weatherley, P.E. (1977). The location of the resistance to water movement
in the soil supplying the roots of transpiring plants. New Phytologist. 78, 337-47.
Goss, M.J. (1977). Effects of mechanical impedance on root growth in barley (Hordeurn
vulgare L.). I . Effects on elongation and branching of seminal roots. J. Exp. Bot. 28,
96-111.
Soil Structure and Plant Growth

Greacen, E.L., Farrell, D.A., and Cockcroft, B. (1968). Soil resistance to metal probes and
plant roots. Transactions of the 9th Congress of the International Society of Soil Science
1, 769-79.
Greacen, E.L., and Oh, J.S. (1972). Physics of root growth. Nature (Lond.) New Biol. 235,
24-25.
Hamblin, A.P. (1985). The influence of soil structure on water movement, crop root growth,
and water uptake. Adv. Agron. 38, 95-158.
Hasegawa, S., and Sato, T. (1987). Water uptake by roots in cracks and water movement in
clayey subsoil. Soil Sci. 143, 381-6.
Hatano, R., Iwanaga, K., Okajima, H., and Sakuma, T. (1988). Relationship between the
distribution of soil macropores and root elongation. Soil Sci. PI. Nutr. 34, 535-46.
Herkelrath, W.N., Miller, E.E., and Gardner, W.R. (1977~).Water uptake by plants. I. Divided
root experiments. Soil Sci. Soc. Am. J. 41, 1033-38.
Herkelrath, W.N., Miller, E.E., and Gardner, W.R. (1977b). Water uptake by plants. 11. The root
contact model. Soil Sci. Soc. Am. J. 41, 1039-43.
Hettiaratchi, D.R.P. (1990). Soil compaction and plant root growth. Philosophical Trans. R.
Soc. London B 329, 309-20.
Huck, M.G., Klepper, B., and Taylor, H.M. (1970). Diurnal variations in root diameter. PI.
Physiol. 45, 529-30.
Jordan, W.R., and Miller, F.R. (1980). Genetic variability in sorghum root systems: implications
for drought tolerance. In 'Adaptation of Plants to Water and High Temperature Stresses'.
(Eds N.C. Turner, P.J. Kramer.) pp. 383-399. (Wiley: New York.)
Kays, S.J., Nicklow, C.W., and Simons, D.H. (1974). Ethylene in relation to the response of
roots to mechanical impedance. PI. Soil 40, 565-71.
Masle, J., and Passioura, J.B. (1987). The effect of soil strength on the growth of young
wheat plants. Aust. J. PI. Physiol. 14, 643-56.
Materechera, S.A., Dexter, A.R., and Alston, A.M. (1991). Penetration of very strong soils by
seedling roots of different plant species. PI. Soil. 135, 31-41.
Misra, R.K., Alston, A.M., and Dexter, A.R. (1988). Root growth and phosphorus uptake in
relation to the size and strength of soil aggregates. I. Experimental studies. Soil Tillage
Res. 11, 103-116.
Morris, D.T., and Daynard, T.B. (1978). Influence of soil density on leaf water potential of
corn. Can. J. Soil Sci. 58, 275-8.
Moss, G.I., Hall, K.C., and Jackson, M.B. (1988). Ethylene and the responses of roots of maize
(Zea mays L.) to physical impedance. New Phytologist. 109, 303-11.
Munns, R., and King, R.W. (1988). Abscisic acid is not the only stomata1 inhibitor in the
transpiration stream of wheat plants. PI. Physiol. 88, 703-8.
Passioura, J.B. (1985). Roots and water economy of wheat. In 'Wheat Growth and Modelling'.
(Eds W. Day, R.K. Atkin.) pp. 185-98. (Plenum: New York.)
Passioura, J.B. (1988). Water transport in and to roots. Ann. Rev. PI. Physiol. PI. Mol. Biol.
39, 245-65.
Passioura, J.B., and Gardner, P.A. (1990). Control of leaf expansion in wheat seedlings growing
in drying soil. Aust. J. PI. Physiol. 17, 149-57.
Passioura, J.B., and Leeper, G.W. (1963). Soil compaction and manganese deficiency. Nature.
Lond. 200, 29-30.
Peterson, C.M., Klepper, B., Pumphrey, F.V., and Rickman, R.W. (1984). Restricted rooting
decreases tiliering and growth of winter wheat. Agron. J. 76, 861-3.
Richards, B.G., and Greacen, E.L. (1986). Mechanical stress on an expanding cylindrical root
analogue in granular media. Aust. J. Soil Res. 24, 393-404.
Richards, D., and Rowe, R.N. (1977). Effects of root restriction, root pruning, and
6-benzylaminopurine on the growth of peach seedlings. Annals Bot. (London) 41, 729-40.
Robinson, D., Linehan, D.J., and Caul, S. (1991). What limits nitrate uptake from soil? PI.,
Cell Environ. 14,77-85.
Rovira, A.D., Elliot, L.F., and Cook, R.J. (1990). The impact of cropping systems on rhizosphere
organisms affecting plant health. In 'The Rhizosphere'. (Ed. J. Lynch.) (Wiley: Chichester.)
Ruff, M.S., Krizek, D.T., Mirecki, R.M., and Inouye, D.W. (1987). Restricted root zone volume:
influence on growth and development of tomato. J. Am. Soc. Hort. Sci. 112, 763-9.
 J. B. Passioura

Russell, R.S. (1977). Plant Root Systems. Their Function and Interaction with the Soil
(McGraw-Hill: London.)
Russell, R.S., and Goss, M.J. (1974).Physical aspects of soil fertility-the response of roots
to mechanical impedance. Neth. J. Agric. Sci. 2 2 , 305-18.
Tardieu, F. (1988). Analysis of spatial variability in maize root density. 111. Effect of wheel
compaction on water extraction. PI. Soil 109, 257-62.
Taylor, H.M., and Ratliff, L.F. (1969). Root elongation rates of cotton and peanuts as a
function of soil strength and soil water content. Soil Sci. 108, 113-19.
Tinker, P.B. (1976).Transport of water to plant roots in soil. Philosophical Trans. Royal Soc.
London B. 273, 445-61.
Veen, B.W. (1982).The influence of mechanical impedance on the growth of maize roots. PI.
Soil 66, 101-9.
Veen, B.W., and Boone, F.R. (1990). The influence of mechanical resistance and soil water on
the growth of seminal roots of maize. Soil 7illage Res. 16, 219-26.
Wang, J., Hesketh, J.D., and Woolley, J.T. (1986). Preexisting channels and soybean rooting
patterns. Soil Sci. 141, 432-37.
Whiteley, G.M., Utomo, W.H., and Dexter, A.R. (1981).A comparison of penetrometer pressure
and the pressure exerted by roots. PI. Soil 61, 351-64.
Williams, J., and Shaykewich, C.F. (1970).The influence of soil water matric potential on the
strength properties of unsaturated soil. Soil Sci. Soc. Am. Proc. 34, 835-40.
Williamson, R.E. (1990).Alignment of cortical microtubules by anisotropic wall stress. Aust.
J. PI. Physiol. 17, 601-613.
Zhang, J., and Davies, W.J. (1990).Changes in the concentration of ABA in xylem sap as a
function of changing soil water status can account for changes in leaf conductance and
growth. PI. Cell Environ. 13, 277-85.
Zur, B.J., Jones, K.W., Boote, K.J., and Hammond, L.C. (1982).Total resistance to water flow
in field soyobeans: 11. Limiting soil moisture. Agron. J. 74, 99-105.

Manuscript received 1 1 June 1991, accepted 1 1 September 1991