Key words: maize, mechanical stress, penetrometer, root growth, sandy loam
Abstract
Root penetration resistance and elongation of maize seedling roots were measured directly in
undisturbed cores of two sandy loam soils. Root elongation rate was negatively correlated with root
penetration resistance, and was reduced to about 50 to 60% of that of unimpeded controls by a
resistance of between 0.26 and 0.47 MPa. Resistance to a 30 semiangle, 1 mm diameter penetrometer
was between about 4.5 and 7.5 times greater than the measured root penetration resistance. However,
resistance to a 5 semiangle, 1 mm diameter probe was approximately the same as the resistnace to root
penetration after subtracting the frictional component of resistance. The diameter of roots grown in the
undisturbed cores was greater than that of roots grown in loose soil, probably as a direct result of the
larger mechanical impedance in the cores.
0.6
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Materials and methods
o %
0.4 0
Preparation of soil cores
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0.2 Undisturbed soil cores were collected from two
fields (Big Ground and Plum Orchard) at the
0.0 i i t i i !
Institute of Horticultural Research, Welles-
0.0 0.1 0.2 0.3 0.4 bourne.
Two shallow pits several metres apart were dug
Root resistance (MPa)
at each site to provide replicate samples, and soil
Fig. I. Axial root force/root cross sectional area versus cores (56 mm diameter x 40 mm) were extracted
elongation rate of pea seedling roots in remoulded cores of
from 0.12 to 0.20 m depth. The cores were sea-
sandy loam soil (calculated from Eavis, 1967).
led at field moisture content and stored at 10C
until required for use. Details of the physical and
The results reported by Eavis are of sufficient chemical properties of the soils are given in
importance to merit an independent study using Young (1987), and some of the soil properties
an improved technique for measuring root force are summarised in Table 1.
on a different species. The first aim of this paper The soil cores were saturated overnight with
is, therefore, to quantify the effect of mechanical distilled water (containing 'Phorate' systemic in-
impedance on maize root elongation rate in un- secticide to kill small numbers of springtail in-
disturbed soil cores by measuring root penetra- sects which had been observed on the cores) and
tion resistance directly and recording the average then equilibrated at - 1 0 kPa matric potential on
elongation rate of these roots. a tension table for 5 to 7 days. After equilibra-
Root penetration resistance and penetrometer tion, cores were wrapped in a layer of plastic film
resistance are defined as the force exerted by the containing a small (10 mm diameter) hole for the
root or probe divided by its cross sectional area root. The rate of water loss from the polythene
(Bengough and Mullins, 1990b). Greacen et al. covered core was less than about 0.3 g day -1. To
(1968) suggested that both roots and 'sharp' test whether the insecticide affected maize root
penetrometers (i.e. penetrometers with a small elongation rates, root elongation rates were mea-
cone angle) deform the soil cylindrically, which sured for 56 seedlings grown in moist horticultur-
requires less pressure than the spherical de- al vermiculite which was watered with either
formation caused by 'blunt' penetrometers. They distilled water or with insecticide solution. The
also suggested that roots experience virtually no average root elongation rate of the insecticide
frictional resistance, in contrast to penetrometers treated seedlings (averaged over 3 days) was
which often have a high component of frictional within 3% of that of the controls, and this differ-
resistance. Greacen and Oh (1972) used the ence was not statistically significant.
resistance to a 5 semi-angle probe minus the
frictional resistance as an estimate of the root Seed germination and pre-selection
penetration resistance, while Voorhees et al.
(1975) found better correlations between root To reduce variability between replicate seed-
elongation rate and penetrometer resistance (to lings, maize seeds (cv. Artus) were graded by
a 5 semi-angle probe) after the frictional compo- weight, and only seeds with masses within one
nent of resistance was subtracted off. Our second standard deviation of the mean were used. Seven
aim was, therefore, to test the hypothesis that seeds per core were sown on moist vermiculite,
and left in the dark to germinate (at about 20C).
Root resistance and elongation rate 61
4ram
Root lengths were measured 2.0 and 2.5 days (")
after planting, and root growth rates during this I
V- s 1
P i
microcomputer interfaced to the balance output trations were performed at a penetration rate of
and stored on cassette tape. There was negligible 4 m m min -1. In a separate experiment, soil re-
drift in the balance readings during each ex- sistance to a 5 1 mm diameter penetrometer was
perimental run. The root was allowed to elon- measured on a replicate set of equilibrated cores,
gate for a further 8 hours and then was exca- and the angle of soil-metal friction estimated
vated out of the soil core. Final root diameter using an inclined plane (Bengough, 1988).
was measured 2 to 5 mm behind the apex by
hand sectioning the root using pith and a fine
razor blade. Because the root circumference was
often quite distorted and irregular, several sec- Results
tions were cut and two perpendicular diameters
were measured for each section using a mi- Root force, diameter and elongation rate
croscope with graticule scale. The average of
these readings was taken as the final root diam- Seven out of eight maize roots penetrated cores
eter, and root length was also recorded. of both soils. Average root force was calculated
Because it was not possible to maintain accur- over the growth period when the root was be-
ate temperature control, temperature was con- tween 2 and 5 mm long (assuming a constant rate
tinuously monitored using a thermograph. After of elongation). The mean root penetration resist-
completing each experiment, part of the soil was ance was calculated by dividing the root force by
passed through a 2-mm sieve and repacked ve T either the initial or the final root cross sectional
loosely to a bulk density of about 0.7g c m - . area averaged between 2 and 5 mm behind the
Maize root elongation rates were measured for tip. Average values of root resistance, diameter
eight seedlings in sieved Big G r o u n d soil and and elongation rate for both soils are given in
seven seedlings in the sieved Plum Orchard soil Table 2.
to obtain a reference rate of elongation in soil Root elongation rates and diameters did not
with negligible root penetration resistance. differ significantly between the two soils, and
neither did the root resistances based on the
Penetrometer resistance initial root cross sectional area. The root resist-
ance based on the final root cross sectional area
Penetration resistance to 0.5 and 1 mm diameter was significantly greater for Plum Orchard than
30 semi-angle penetrometer probes was meas- for Big G r o u n d soil. Final root diameter was
ured in each intact soil core, both before and significantly greater than the initial diameter in
after each root resistance experiment. All pene- each soil.
Table 2. Mean maize root penetration resistances, elongation rates and diameters in undisturbed soil cores (standard errors of the
mean are shown in brackets)
Soil
Big Ground Plum Orchard
Root penetration resistance
Force/initial area (MPa) 0.38 (0.08) 0.47 (0.06)
Force/final area (MPa) 0.26 (0.03) 0.38 (0.04)
Root diameter: Initial (mm) 1.14 (0.02) 1.12 (0.02)
Final (mm) 1.35 (0.07) 1.36 (0.08)
Table 3. Penetrometer resistance in Big Ground and Plum Orchard soils, for different probes before and after the root resistance
experiment (standard errors of the mean are shown in brackets)
Penetrometer probe Penetration Penetrometer resistance (MPa)
Diameter (ram) Semiangle () performed Big Ground Plum Orchard
1 30 before 1.72 (0.24) 2.53 (0.34)
1 30 after 2.43 (0.22) 3.23 (0.28)
0.5 30 before 2.61 (0.43) 4.09 (0.64)
0.5 30 after 3.11 (0.37) 4.73 (0.99)
l 5 n.a. 2.60 (0.44) 2.88 (0.33)
n.a. = not applicable: root resistance not measured in these soil cores.
,,o
1
5 10 15 20
Depth (mm)
Fig. 3. Typical plot of depth versus penetration force for a 1 mm probe in undisturbed Plum Orchard soil.
soil being heterogenous on the size scale of the There was a significant linear correlation be-
fluctuations. The resistance to root growth may tween the elongation rate and the resistance
also increase during the experiment due to the experienced by individual roots ( r = - 0 . 7 7 6 ;
soil around the root drying out. This drying area P < 0.01, Fig. 4), when root resistance was based
gave rise to significantly higher penetrometer on the initial cross-sectional area of the root.
resistances (P < 0.01) at the end of the experi- The relation (not shown) between maize root
ment (Table 3). Thus, the resistance to root elongation rate and root resistance based on the
growth measured during the initial penetration final cross-sectional area was much more scat-
was not necessarily the same as the resistance tered (r = -0.343) showing no significant corre-
deeper in the core, although the initial root lation (P>0.05). It is not clear which of the
penetration resistance should provide some indi- initial or the final root cross sectional area mea-
cation of the root penetration resistance deeper surements is the more appropriate from which to
in the core. calculate the root penetration resistance. To re-
solve this question simultaneous measurements
of root force and diameter must be made, so that
1.8 I the root penetration resistance can be continu-
"%% ously monitored.
1.4 %
E
%%0 Penetrometer resistance and root penetration
resistance
1.1
%
%%
%,A Root and probe penetration resistances varied
C %
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._, 0.7 A~%%
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c
ent regions within each core, but there was a
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LU general trend for cores with higher penetrometer
0.4
,.,% resistances to offer higher resistance to root
%%
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penetration so that the two were significantly
0.0 i i ! i %1 correlated (r=0.741, P < 0 . 0 1 ; Fig. 5). Soil re-
0.0 0.2 0.4 0.6 O.S 1.0 sistance to the 1 mm diameter probe was be-
Root resistance (MPa)
tween about 4.5 and 7.5 times greater than the
measured penetration resistance to maize roots.
Fig. 4. Root force/initial cross-sectional area versus elonga-
tion rate of maize seedling roots, in Big Ground (O) and These ratios are similar to those found by direct
Plum Orchard (&) soil cores. (l'q) represents the estimated comparisons in other studies (Whiteley et al.,
value for the unimpeded root growth rate in loose soil. 1981).
Root resistance and elongation rate 65
1.0
,/
/ little frictional resistance and deform the soil
,s
/i cylindrically. A further implication is that a reli-
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/ able quantitative indication of the penetration
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resistance experienced by roots in structureless
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/ soils may be obtained from the resistance ex-
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A / perienced by penetrometers with a small cone
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angle and of diameter comparable to the roots
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once a correction has been made for the soil-
metal friction on the penetrometer. More work
Z
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cf J is needed to test this suggestion in a wider range
0.2 / J of soil types and soil conditions. Such work must
/
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/ also take into account the effects of penetration
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i i i i i rate which are probably most important in im-
O0 05 1.0 15 2.0 2.5 3.0 35 40 45 permeable soils at high moisture content.
Penetrometer resistance (MPa)
growth of seedling roots. Soil Sci. Soc. Am. Proc. 20, Stolzy L H and Barley K P 1968 Mechanical resistance
154-157. encountered by roots entering compact soils. Soil Sci. 105,
Goss M J 1977 Effects of mechanical impedance on root 297-301.
growth in barley (Hordeum vulgare L.). I. Effects on Taylor H M and Ratliff L F 1969 Root elongation rates of
elongation and branching of seminal roots. J. Exp. Bot. 28, cotton and peanuts as a function of soil strength and water
96-111. content. Soil Sci. 108, 113-119.
Greacen E L and Oh J S 1972 Physics of root growth. Nature Voorhees W B, Farrell D A and Larson W E 1975 Soil
(New Biol.) 235, 24-25. strength and aeration effects on root elongation. Soil Sci.
Greacen E L, Farrell D A and Cockroft B 1968 Soil resist- Soc. Am. Proc. 39, 948-953.
ance to metal probes and plant roots. Trans. 9th Congr. Whiteley G M, Utomo W H and Dexter A R 1981 A
Int. Soil Sci. Soc. 1,769-779. comparison of penetrometer pressures and the pressures
Misra R K, Dexter A R and Alston A M 1986 Penetration of exerted by roots. Plant and Soil 61, 351-364.
soil aggregates of finite size. II. Plant roots. Plant and Soil Young I M 1987 Soil Strength and Hard-Setting Behaviour of
94, 59-85. Some Structurally Unstable British Soils. Ph.D. thesis,
Richards B G and Greacen E L 1986 Mechanical stresses on University of Aberdeen, Aberdeen.
an expanding cylindrical root analogue in granular media.
Aust. J. Soil Res. 24, 393-404.