Tải bản đầy đủ - 0 (trang)
VI. Growth in the Soil

VI. Growth in the Soil

Tải bản đầy đủ - 0trang



grains ( McMurdie and Day, 1958), the elastic strain is not negligible.

Nevertheless, root growth does not seem to occur in such media, other

than in continuous pores commensurate in width with the normal, unstressed root tip. Wiersum (1957) found, for example, that the roots of

tomatoes having tips of 0.3 mm. diameter grew through sintered glass

disks in which the diameter of the pores ranged from 0.20 to 0.50 mm.,

but they were unable to penetrate disks in which the range was 0.15

to 0.20 mm.

b. Deformable matrix. Little attempt has been made to utilize the

relatively simple properties of purely cohesive media in experiments

with growing plants. Taylor and Gardner (1960) and Gardner and

Danielson (1964) measured the penetration of roots into cohesive

waxes of different hardness; these workers used the waxes simply as convenient materials with which to rank the penetrating ability of plant

roots that had been subject to various treatments or grown in various

soils ( Section VI, B, 1 ) .

Although saturated clays are often treated as purely cohesive ($I =

0 ) media in the classical theory of consolidation and bearing capacity

(Terzaghi, 1943), it is not profitable to regard them as such when dealing with deformations by plants. The reasons for this have been described

in Section 111; here we need only remind the reader that effective normal

stresses are in general not negligible around growing plant organs even

in saturated, impermeable clays.

As described in Section 11, seedlings often emerge from cohesive

soils by rupturing and lifting slabs of the overlying soil. When this kind

of deformation occurs, it is appropriate to relate emergence to the breaking forces, obtained from the dimensions of the slab and the modulus of

rupture of the soil. Relations of this kind were measured by Richards

(1953) and by Allison (1956), but their data are of little value as the

strength tests were conducted on slabs reconstituted in the laboratory

from fine earth and dried at 50°C., rather than on the crusts through

which the seedlings actually emerged. Moduli of rupture pertaining to

the moist crusts through which seedlings emerged were measured by

Hanks and Thorp (1956, 1957). But it is doubtful whether this measure

was appropriate in their experiment; emergence was not reduced by

increasing the thickness of the crust, as would have been expected if

tensile failure were the means of emergence. Rather, the shoots may have

penetrated the crust by causing continuous, local failure. On the other

hand, Parker and Taylor (1965) related the emergence of guar, Cyamopsis tetragorwloba ( L . ) Taub., to indentation test data, even though the

seedlings emerged in their experiment by rupturing a crust.

Clearly, the kind of deformation involved should be ascertained



before mechanical criteria are chosen, Careful interpretation is needed,

particularly when one kind of deformation tends to pass into another

with time or distance. For example, radial thickening of the proximal

parts of the root sometimes ruptures peds or layers of soil. When this

happens the failure may be propagated ahead of the root apex, so that

the point stress, qp, falls to a small value. The initial penetration of the

soil by the root always involves shear failure, but, in the later stages,

penetration may result from tensile failure of the soil.

So far we have been dealing with growth in soils where compression

is unimportant, We now turn to the more general situation, where growth

is accompanied by local shear failure and compression of the soil.

Working with compressible moist clays and with gelatin, Pfeffer

(1893) related mechanical resistance, q, to the growth of radicles. He

measured q with steel probes having relieved tips. The tips were similar

in size and shape to those of the broad bean and corn radicles used in his

experiment. Initially each radicle was placed in a 2-cm. deep channel, SO

that the resistance encountered by the growing tip would be constant

during the experiment. In separate tests the channel was formed adjacent

to a glass plate so that the radicle could be observed at intervals during

elongation. When bean radicles were placed in a clay, for which the value

of q measured with the probe was 4 bar, the roots did not elongate for

the first 4 hours. After this delay elongation proceeded at a rate 20 percent below that attained in water or in a slurry of clay in water. Similar

results were obtained with corn radicles. When bean radicles were grown

in a firm gelatin ( q = 1.0 bar) elongation proceeded with little or no

delay at a rate equal to that attained in water or in a gelatin sol. Pfeffer

also reported that roots would not elongate in clays when q exceeded

12 bar, However he did not provide any experimental data for such

strong clays, and it seems likely that he merely inferred this value of q

from his knowledge of the maximum plant pressures measured in his

plaster-block experiments.

As the clays used in Pfeffer’s work were quite moist, gaseous diffusion

through the clay would have been very slow. In commenting on this,

Pfeffer suggested that sufficient oxygen would be obtained by diffusion

through the intercellular air space, providing the radicles were not more

than a few centimeters long. To support this argument he quoted his

observation that bean radicles grew equally well and at moderate rates

(0.8 mm.hr.-l) for at least 48 hours in either de-aired or aerated water.

Unfortunately Pfeffer did not measure the pore water pressure, u.,,,, in

his clays. In the stronger samples uw may have been large and negative,

leading to loss of turgor in the radicles (Section VI, B, 1).

Except at high void ratios a hard-grained soil is difficult to compress




with an isotropic pressure; nevertheless it can be compressed rather

easily when the applied stress has a large shear component. In such a soil

y is related directly to the applied pressure, p , or more generally, taking

account of pore water pressure, to the effective pressure, p’. Barley

(1963) made use of these properties to separate strength from variables

that depend upon the porosity of the medium. In his experiment a

moistened pack of 10 to 70 p Ballotini beads was housed within a modified triaxial test cell. The level of p’ in the medium could then be controlled by regulating the ambient pressure, p, applied to the pack. AS

isotropic compression was negligible ( c v = -0.002) over the range of

pressures applied ( 0 to 1 bar), volumetric air and water contents of the

bulk of the medium were not affected by the change of pressure. Corn

radicles were allowed to grow through inlets; they penetrated the medium readily, when strength permitted, by producing local shear failure.

The rate of elongation decreased from 1.6 to 1.1mm.hr.-l as p‘ increased

from 0.08 to 0.28 bar; the radicles were prevented from elongating at

p‘ = 0.58 bar. At this value of p’ the resistance offered to a cylindrical

steel probe corresponded to y = 22 bar. This is about twice as great a

pressure as the radicles are thought to be able to exert (see Table 11).

Among a number of reasons that could account for this large discrepancy

(Section 111, B, 2), differences in shape between the root tips and the

probe may be important. The problem of relating probe data to the

resistance offered to a root tip is increased, as shown in the above

experiment, by the dependence of the shape and size of the tip upon the

strength of the medium through which the root grows (see Fig. 7 ) .

Recently, three related sets of experiments have been conducted with

ordinary, unsaturated soils. Phillips and Kirkham ( 1962a) studied the

influence of mechanical resistance on the growth of corn radicles in an

unsaturated clay; Taylor and Gardner (1963) and Taylor et al., (1966)

studied both root growth and seedling emergence for a range of plant

species and soil types; Barley et al. (1965) studied the root growth of

two contrasting species in a loam. The experiments will be discussed further in Section VI, B, 2, where factor interactions are considered. Here

we are concerned with the methods used to characterize the resistance, q,

and with the values of y at which elongation growth ceased. In the last

of the investigations mentioned above point pressure, qp,and skin friction,

yf, were measured independently, and theoretical values of qp were

calculated after data had been obtained for the relevant soil parameters.

Table I11 lists the methods used in the experiments, together with the

limiting values of y. The latter have been recalculated from the original

data where necessary, and expressed uniformly as resistance per unit

cross section, y, to the penetration of a deep ( z > 3 d ) steel probe. The

The effect of mechanical resistance on the form of the elongating root tip of corn, Zea mays

L. The roots were



Values of Probe Pressure (9) a t Which Growth Ceased in Several Soil-Plant Systems


Pore water





( X -1)







Bulk densitya


Plant organ

Phillips and Kirkham (1962a)

Taylor and Gardner (1963)

Corn radicles

Cotton radicles

Barley et ul. (1965)

Pea and wheat


Grass shoots

Taylor et al. (1966)


Soil texture


Fine sandy



Sandy loams

Void ratio not calculated as absolute density is not reported.

Measure of resistance


p (bar)

Constant load probe

Constant depth probe



Strength parameters,

constant rate probe

Constant depth probe














wide variation found in the limiting values of q is not surprising. The low

values of q obtained by Phillips and Kirkham may arise for two reasons:

First, the samples of clay that roots failed to penetrate were nearly

saturated, and may have behaved as low 4 materials in tests with impermeable probes, Secondly, instead of allowing developed radicles to

penetrate the soil, as in the other investigations, Phillips and Kirkham

germinated the seeds within the body of cores of consolidated clay.

If pIant forces are osmotic in origin, then as x rarely exceeds 10 bar

in roots, the roots that grew through the stronger loams cannot have had

to overcome resistances as large as those opposed to the probes (see

Table 111).As we have seen in Section 111, there are a number of reasons

why root tips may meet less resistance than probes in unsaturated loams.

Briefly, the root may penetrate the soil by cylindrical rather than spherical compression; qf is likely to be less for root tips than for steel probes;

also, the propagation of cracks produced by thickening of the proximal

part of the root may reduce resistance to penetration.

It is even more difficult to relate the resistance encountered by shoots

to the limiting values of q measured by Taylor et al. As mentioned

previously, more relevant data could have been obtained if the authors

had used upward rather than downward acting probes. Apart from this,

qf may be underestimated in shallow tests or in deep tests where probes

with relieved tips are used, since the zone of maximum elongation often

lies some distance behind the apex of the shoot.

2. Heterogeneous Media

Lack of anchorage or support in loose layers of the soil may hinder

roots or shoots from penetrating stronger layers or crusts. If seed is

planted in loose soil, and the seedling shoot meets a superficial crust, the

shoot may push the seed deeper rather than emerge (Carnes, 1934).

Also, the shoot is more likely to be bent in a loose soil because of the

lack of radial support. Similarly, in a study of the ability of corn radicles

to penetrate a layer of hard wax, Taylor and Gardner (1960) found that

more radicles penetrated when grown in lightly compacted soil than

when grown in loose crumbs. In a subsequent paper Gardner and Danielson (1964) reported that compacting the soil above a wax layer failed

to improve the penetration of roots into the wax. This result appears to

be inconsistent with that obtained in 1960; but in the 1964 experiment

the hard wax was separated from the soil with a layer of soft wax, and

anchorage may have been obtained in the soft wax rather than in the soil.

Although mechanical resistance was not measured, morphological

evidence suggests that this may have been the controlling factor in an

experiment conducted by Schuurman ( 1965). His observations of the



growth of oat roots across the boundary between layers of a humic sand

showed that roots penetrated a highly compacted layer more readily

when this was overlain by a moderately compact rather than by loose

sand. The main roots grown in loose sand branched profusely just above

the boundary with a highly compacted sand. Laterals penetrating the

highly compacted layer were short, thickened, and distorted.

Little attention has yet been given to the possible role of mechanical

factors in controlling the entry of roots into discrete peds. A recent study

of the distribution of corn roots in a silty clay loam B horizon (Edwards

et al., 1964) revealed that, although the main roots were restricted to the

voids between the peds, laterals penetrated about one-half of the total

number of discrete peds. The peds entered by the laterals were, on the

average, less dense than those which the laterals failed to penetrate; but

the factors limiting entry were not determined,

The shape and orientation of a ped in the soil may have a large influence on root entry, insofar as these factors influence the angle at which

the geotropic root tip approaches the surface of the ped. The chance of

entry is known to be reduced when the angle at which the root tip approaches a slab of hard wax becomes more acute (Gardner and Danielson, 1964). Ped size may also be important, since small peds are more

easily ruptured by internal pressure.






1. The Nature of the Interaction

The growth of roots and underground shoots at a given temperature

is influenced strongly by the physical factors: mechanical resistance,

water supply, and aeration. These factors interact for two kinds of

reasons. First, as shown in Section 111, C, the factors themselves are

interdependent; secondly, the response of the plant to a change in one

factor may modify its response to another. Two examples will serve to

show the importance of this second kind of interaction.

As a soil dries uw decreases, and the plant exhibits a loss of turgor.

This is true even when transpiration is slow, as the plant and soil water

are then in or close to osmotic equilibrium. If the forces exerted by

plants arise almost entirely from osmotic turgor, as suggested in Section

IV, C, the ability of an organ to exert force should decline as uw decreases. Also, the rigidity of the root decreases as turgor is lost. Gardner

and Danielson (1964) show that the penetrating ability of plant roots

is indeed reduced for physiological reasons when u, decreases. In their

experiment cotton roots were grown through a loose soil to meet a hard



layer of wax. The percentage of roots that penetrated the wax decreased

continuously at u, < -0.5 bar. No penetration occurred with U, <

-11 bar, Over this range of pore water pressures the water content of

the root decreased from 1200 to 600 percent, implying a considerable loss

of turgor. The authors did not determine whether the roots failed to

penetrate because they exerted less force or because they were more

easily bent. If the roots had met a firm soil rather than a wax, the effect

of u, on the resistance offered by the soil would have operated also.

The interplay of mechanical stress and oxygen supply provides a

second example of soil factor interaction mediated by plant response. In

their pressure cell experiments (Section V, A, 3), Gill and Miller ( 1956)

and Barley (1962) found that a smaller confining pressure prevented the

elongation of corn radicles when the ambient concentration of oxygen

was reduced below 5 percent. Data on root weight increase, Aw, obtained

in the latter experiment show that neither a small increase in confining

pressure ( 0.0 to 0.5 bar) nor a reduction in oxygen concentration (20 to 5

percent) reduced A w when acting singly, but, when the variabIes acted

together, A w was halved. The simplest explanation of this interaction is

provided by the effect that mechanical stress had on the shape assumed

by the radicles, the volume per unit length being twice as great when the

radicles grew under compression. This increase in bulk, taken together

with compression of the intercellular spaces, would have increased the

ambient concentration of oxygen needed to maintain a diffusive supply

of oxygen to cells in the interior of the root.

Changes in shape similar to those described above occur when roots

grow in firm media (Fig. 7 ) ; moreover, high mechanical resistance is

frequently associated with compaction and poor aeration. In a study of

the penetration of compacted soils by cotton roots, Tackett and Pearson

(1964) found that at a bulk density of 1.3 g.cc.-l roots elongated at a

high and constant rate when the oxygen concentration remained greater

than 5 percent, but at a density of 1.5 g.cc.-l the threshold concentration

of oxygen rose to 10 percent. Morphological observations suggested that

mechanical resistance was likely to have been the other factor involved.

2. Correlation of Growth with Mechanical Resistance

Although the simple correlation of mechanical properties of the soil

and plant growth may be useful in diagnosing physically adverse soil

conditions ( Culpin, 1936), correlation does not identify the particular

factors controlling growth. Fortunately, it is often possible to detect

mechanical effects by visual observation, as when shoots are seen to be

bent beneath a superficial crust. Further examples of gross morphological

symptoms are given by TayIor and Burnett ( 1964).



Circumstantial evidence that mechanical resistance may be limiting is

sometimes presented by showing that other physical factors are unlikely

to be limiting. Having found a simple correlation ( r = 0.6 - 0.7) between probe resistance and corn yield in a field experiment on soil compaction, Phillips and Kirkham (1962b) proceeded to show that the soil

temperature and the oxygen content of bulk samples of the soil air were

similar at the various levels of compaction. Also, they noted that the soils

were kept moist by sprinkling during the growing period. But arguments

of this kind are often too tendentious to lead to any satisfying conclusion.

While it is possible to design experiments in which other than mechanical factors are nonlimiting, this may limit the range of values explored,

as the soil properties, e and uw,that govern air and water supply also

determine the level of mechanical resistance. A nonlimiting supply of

water can be assured by working at small pore water and osmotic pressures, and by keeping the shoots in a humid atmosphere so that transpiration is minimized. Paul (1965) found, for example, that when wheat

seedlings were covered with Mylar film to minimize transpiration, the

rate at which the roots elongated in a sand remained constant at volumetric water contents ranging from 4 to 17 percent, corresponding to

uw = -0.25 to -0.03 bar. It is more difficult to ensure a nonlimiting

supply of oxygen, particularly when mechanical properties are varied by

compaction. Various workers have employed either forced aeration

(Smith and Cook, 1946), a source of oxygen within the soil (Scott and

Erickson, 1964), or elevated ambient concentrations of oxygen (Rickman

et al., 1966). The effectiveness of such methods is best assessed by

sampling and analyzing the gas permeating the interior of the soil under

test (Tackett and Pearson, 1964). Although data on mechanical properties were not obtained in these several investigations, the results show

clearly that aeration is not the only factor involved in reducing root

growth in compact soils.

Generally we wish to examine soil-plant systems in states where each

of the physical factors capable of influencing growth may vary over a

wide range. If we aim to determine the relative importance of the different factors it becomes necessary to design experiGents to separate the

variables. To do this Phillips and Kirkham (1962a) took advantage of the

known relations between mechanical resistance, q, pore water pressure,

uw,void ratio, e, and air void ratio, e,. When uw is varied at a constant

e, q and e, change in the same sense; but when e is varied at a constant

uw, q and e, change in the opposite sense. It is true that the capillary

conductivity, k, changes in the opposite sense to q when either uw or e

is varied. However even wide changes in k may have little influence on

growth if the rate of transpiration is minimized (Paul, 1965). Also, by



employing a soil with a high friction angle, it is not necessary to use

large negative values of u, or to vary ufowidely to obtain a useful range

of q.

The system studied by Phillips and Kirkham is described in Table I11

together with systems studied by later investigators in experiments of

similar design. Although data obtained in experiments of the kind described in Table I11 lend themselves to covariance analysis, this was

utilized only in one of the investigations (Barley et al., 1965). The partial

correlation coefficients were significant for root length, L, and resistance,

q, but not for L and e, or for L and the gaseous diffusivity parameter

( e,/e)4 proposed by Currie (1961). Though given only in graphical form,

the data of Taylor and Gardner (1963) show clearly that most of the

variation in the number of roots penetrating a sandy loam over a range

of e and u, values was associated with the variation in q. Taken together

with morphological observations, the correlative data obtained in the

experiments described in Table I11 provide unmistakable evidence that

mechanical resistance can exert a considerable influence on root growth

and seedling emergence in finely structured soils at field densities and

water contents.



In the past the role of mechanical resistance has been relatively

neglected in agronomic studies, probably because of the academic separation of “soil mechanics” in schools of engineering from “soil physics”

in schools of agriculture. When applied to agronomy, engineering theories

of soil mechanics need to be modified to place more emphasis on the

compressibility of the soil, and to be combined with a knowledge of the

mechanics of plant growth.

The evidence presented in this review suggests that mechanical resistance should be regarded as having a widespread influence on the

growth of roots and underground shoots, rather than as a factor that

operates only in unusually strong soils.


Allison, L. E. 1956. Soil Sci. SOC. Am. Proc. 20, 147-151.

Amdt, W. 1965. Australian J . Soil Res. 3, 45-54.

Balla, A. 1961. Proc. 5th Intern. Conf. Soil Mech. Found. Eng. 2, 569-576.

Barley, K. P. 1954. Soil Sci. 78,205-210.

Barley, K. P. 1962. J . Exptl. Botany 13,95-110.

Barley, K. P. 1963. Soil Sci. 96, 175-180.

Barley, K. P. 1965. Australian J . Biol. Sci. 18, 499-503.

Barley, K. P., and Stolzy, L. M. 1966. Australian Soil Sci. Conf. Hbk. Paper 7-1.

Barley, K. P., Farrell, D. A., and Greacen, E. L. 1965. Australian J. Soil Res. 3, 6979.

Bell, G . G. E. 1961. J . Theoret. Biol. 1, 104-106.



Bennett, 0. L., Ashley, D. A., and Doss, B. D. 1964. Agron. J. !%,162-165.

Bennett-Clark, T. A. 1959. In “Plant Physiology” (F. C. Stewart, ed.), Vol. 11, pp.

105-191. Academic Press, New York.

Bishop, A. W. 1960. “Pore Pressure and Suction in Soils,” pp. 3 U 6 . Butterworth,


Bishop, A. W., and Henkel, D. J. 1962. “The Measurement of Soil Properties in the

Triaxial Test.” Arnold, London.

Bishop, R. F., Hill, R., and Mott, N. F. 1945. Proc. Phys. Soc. (London) 57, 147-159.

Bordner, J. S. 1909. Botan. Gaz. 48,251-274.

Brush, W. D. 1912. Botan. Gaz. 53,453-477.

Bunning, E. 1941. Ber. Deut. Botan. Ges. 41, 2-9.

Carlson, L. 1948. Proc. 2nd Intern. Conf. Soil Mech. Found. Eng. 1, 265-270.

Carnes, A. 1934. Agr. Eng. 15, 167-171.

Childs, E. C. 1955. Proc. Natl. Acad. Sci. (India) %A, 86-92.

Croney, D., and Coleman, J. D. 1954. J. Soil Sci. 5, 75-84.

Culpin, C. 1936. J . Agr. Sci. 26, 22-34.

Currie, J. A. 1961. Brit. J. Appl. Phys. 12, 275-281.

Dainty, J. 1963. Aduan. Bot. Res. 1, 279424.

de Jong, G., and Geertsma, J. 1953. Ingenieur 65,5pp.

de Vries, H. 1879. Botan. Zh. 37, 648-654.

de Vries, H. 1884. Jahrb. Wiss. Botan. 14, 427-601.

Edwards, W. M., Fehrenbacker, J. B., and Vavra, J. P. 1964. Soil Sci. SOC. Am. Proc.

28, 560-564.

Eide, O., Hutchinson, J. N., and Landva, A. 1961. Proc. 5th Intern. Conf. Soil Mech.

Found. Eng. 2 , 4 5 5 3 .

Evans, I., and Sherratt, G. G. 1948. J. Sci. Instr. 25, 411-414.

Farrell, D. A., and Greacen, E. L. 1966. Australian J. Soil Res. 4, 1-17.

Farrell, D. A., Greacen, E. L., and Larson, W. E. 1967. Soil Sci. SOC. Am. Proc. (in

press ) .

Fountaine, E. R., and Brown, N. J. 1959. J. Agr. Eng. Res. 4, 5359.

Frey-Wyssling, A. 1952. In “Deformation and Flow in Biological Systems” (A. FreyWyssling, ed. ), pp. 194-254. North-Holland Publ., Amsterdam.

Gardner, H. R., and Danielson, R. E. 1964. Soil Sci. SOC. Am. Proc. 28, 457-461.

Gardner, W. R. 1960. Soil Sci. 89, 63-73.

Gessner, F. 1961. In “Encyclopaedia of Plant Physiology” (W. Ruhland, ed.), Vol. 16,

pp. 66&-680.Springer, Berlin.

Gill, W. R., and Miller, R. D. 1956. Soil Sci. SOC. Am. Proc. 20, 154-157.

Greacen, E. L. 1960. J. Soil Sci. 11, 313-333.

Griffith, A. A. 1924. Proc. 1 s t Intern. Congr. Appl. Math. pp. 55-63.

Haberlandt, G. 1914. “Physiological Plant Anatomy” (Transl., M. Drummond ), Macmillan, London.

Hallbauer, W. 1909. Dissertation. Leipzig. (Reviewed by 0. Damm. 1911. Botan.

Cents. 116, 201.)

Hanks, R. J., and Thorp, F. C. 1956. Soil. Sci. Soc. Am. Proc. 20, 307410.

Hanks, R. J., and Thorp, F. C. 1957. Soil Sci. SOC.Am. Proc. 21, 357359.

Hermans, J. J. 1949. In “Colloid Science” (H. R. Kruyt, ed. ), Vol. 2, p. 86. Elsevier,


Heyn, A. N. J. 1931. Records Trav. Botan. N e e d 28, 113.

Hottes, C. F. 1929. Plant Physiol. 4, 1-29.

Hvorslev, M. J. 1937. Ingenioeruidenskab. Skrifter A&, 1-159.

Kirkham, D., De Boodt, M. F., and De Leenheer, L. 1959. Soil Sci. 87, 141-144.

Tài liệu bạn tìm kiếm đã sẵn sàng tải về

VI. Growth in the Soil

Tải bản đầy đủ ngay(0 tr)