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From: N. K. Fageria and A. Moreira, The Role of Mineral Nutrition on Root Growth
of Crop Plants. In Donald L. Sparks, editor: Advances in Agronomy, Vol. 110,
Burlington: Academic Press, 2011, pp. 251-331.
ISBN: 978-0-12-385531-2
Copyright 2011 Elsevier Inc.
Academic Press.
F O U R
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
Introduction
Root-Induced Changes in the Rhizosphere
Root Systems of Cereals and Legumes
Contribution of Root Systems to Total Plant Weight
Rooting Depth and Root Distribution
Root Growth as a Function of Plant Age
RootShoot Ratio
Root Growth Versus Crop Yield
Genotypic Variation in Root Growth
Root Oxidation Activity in Oxygen-Deficient Soils
Root Growth in Conservation Tillage Systems
Mineral Nutrition Versus Root Growth
12.1. Nitrogen
12.2. Phosphorus
12.3. Potassium
12.4. Calcium
12.5. Magnesium
12.6. Sulfur
12.7. Micronutrients
13. Management Strategies for Maximizing Root Systems
13.1. Soil management
13.2. Plant management
14. Conclusions
Acknowledgment
References
Abstract
Agriculture is going through a profound revolution worldwide due to increasing
world demand for food, higher costs of energy and other inputs, environmental
pollution problems, and instability of cropping systems. In this context,
* Rice and Bean Research Center of Embrapa, Santo Antonio de Goias, GO, Brazil
Western Amazon Research Center of Embrapa, Manaus, AM, Brazil
Advances in Agronomy, Volume 110
ISSN 0065-2113, DOI: 10.1016/B978-0-12-385531-2.00004-9
{
251
1. Introduction
Roots are important plant organs. They absorb water and nutrients
from the soil and translocate them to plant tops (Merrill et al., 1996, 2002;
Sainju et al., 2005a; Stone et al., 2001). Roots also give mechanical support
to plants and supply hormones that affect many physiological and biochemical processes associated with growth and development. Roots exert control
253
255
Ectorhizosphere
Root hair
Mucigel
(plant and bacteria)
Endorhizosphere
Endodermis
Root cap
Figure 1 Root cross section showing ecto- and endorhizosphere (Fageria and Stone,
2006).
Physical changes
Temperature
Water holding capacity
Structure
257
Biological changes
Nitrogen fixation
PGPR bacterias
Mycorrhizal fungi
Harmful microorganisms
Chemical changes
pH
Redox potential
Release of organic compounds by roots
Nutrient concentration and availability
AI detoxification
Allelopathy
Figure 2 Major physical, chemical, and biological changes in the rhizosphere (Fageria
and Stone, 2006).
begins before the leaves have unfolded, with the result that the plant
establishes early contact with moist soil (Hoad et al., 2001). Generally,
roots are classified into four groups. These groups are the taproot, basal
roots, lateral roots, and shootborn or adventitious roots (Zobel, 2005a).
When plants produce secondary shoots (tillers) or shoot branches which
develop roots, these roots are commonly called adventitious roots. To
indicate the true origin of these adventitious roots, the term shootborn is
sometimes used (Zobel, 2005a). The primary function of the taproot, basal
roots, and adventitious roots is to establish the most optimum framework
from which to initiate small lateral roots to effect water and nutrient uptake
(Zobel, 2005b). The taproot penetrates relatively deeply to ensure an
adequate supply of soil water, the basal roots spread out laterally to ensure a
structure for lateral roots that take up P and other nutrients that are less
abundant in the lower levels of the soil profile (Zobel, 2005b), and to
provide a degree of lodging resistance to the plant as it matures and produces
seed (Barlow, 1986; Stoffella et al., 1979). For many grasses and other species
in which root secondary thickening is not important, the shootborn roots
take over the role of the basal roots. The shootborn roots continue to build
the framework with larger and larger conducting roots as the plant increases
in size (Zobel, 2005b). The basal and shootborn roots probably provide little
direct uptake of nutrients and water (St. Aubin et al., 1986).
Besides length and weight, surface area is an important parameter of the
root system in crop plants. The form of root systems and their development
conditions greatly affect the surface area of roots. The surface area of roots
has a high positive correlation with the amount of nutrient absorption
(Takenaga, 1995). Various studies show that 9095% or more of the root
length of an intact plant is made up of roots <0.6 mm in diameter (Zobel,
2003, 2005b).
Monocots and dicots typically have different root system structures.
Root systems of monocots are fibrous, whereas dicots often have taproots.
The fibrous root systems of monocots consist of seminal, nodal, and lateral
roots. Seminal roots develop from primordia within seeds and nodal roots
develop adventitiously from lower stem nodes. All adventitious roots of
stem origin are called nodal roots to distinguish them from other adventitious roots that emerge from the mesocotyl or elsewhere on the plant.
Nodal roots are identified by the node number from which they originate.
Nodal roots may be functional or nonfunctional (Thomas and Kaspar,
1997). Functional nodal roots are defined as roots that have emerged from
stem nodes, entered the soil, and developed lateral roots and/or root hairs.
Nonfunctional nodal roots are defined as roots that have emerged from
aboveground stem nodes and have not entered the soil or produced lateral
roots (Thomas and Kaspar, 1997).
Initial seminal or nodal roots develop laterals that are classed as roots of
the first order, roots that develop from first-order roots are classed as
second-order roots, and additional roots that develop from these laterals
are classed as third-order roots, fourth-order roots, etc. (Yamauchi et al.,
1987a,b). Nodal roots are also known as adventitious, coronal, and/or
crown roots. Roots of cereals such as rice include mesocotyl, radical
(seminal), and nodal or adventitious roots (Yoshida, 1981). Mesocotyl
roots emerge from the axis between the coleoptile node and the base of
the radical, and they typically develop only when seeds are planted very
deep or are treated with chemicals (Yoshida, 1981). Until adventitious roots
develop, seedlings must rely on roots which initiate on the subcoleoptile
internodes above the seed or seminal roots below the seed. Adventitious
roots are important to seedling establishment because they can conduct
259
3rd leaf
4th leaf
2nd leaf
1st tiller
1st leaf
Adventitious root
Radicle root
First trifoliate
Gem
Primary leaf
Epicotyl
Hypocotyl
Adventitious roots
Basal roots
Lateral roots
Figure 4
Tap root
such as rice, barley, and corn is typically lower than the CEC of legumes like
dry bean and broad bean. Roots with high CEC absorb more divalent
cations like Ca2 and Mg2 than monovalent cations such as K and
NH4. On the other hand, roots with lower CEC absorb more monovalent
than divalent cations. Hence, in grasslegume mixtures, legumes generally
suffer with K deficiency due to large uptake of this element by grasses.
Essau (1977), Fageria et al. (2006), Klepper (1992), Leskovarant and Stofella
(1995), OToole and Bland (1987), and Zobel (1991, 2005a,b) have dealt
extensively with various types of monocotyledonous and dicotyledonous
roots and root hairs, and their growth and morphology.
261
are under abiotic or biotic stress. The traditional view is that a large vigorous
root system, through avoidance of plant water deficits, is required for high
yields in water-limited environments (Ludlow and Muchow, 1990). In
addition, large root systems add more organic matter to the soil, creating
better environmental conditions for the growth and development of crops.
The distribution of photosynthetic products in the root, shoot, and grain is
determined genetically, but it also varies with environmental conditions
(Fageria, 1992, 2009; Sainju et al., 2005a). Because roots are so difficult to
separate from soils, little is known about the effects of soil and crop management practices on their growth and development (Sainju et al., 2005b).
In most annual plant species, only 1020% of mature plant biomass
consists of root tissue (Zobel, 1986). Similarly, Fageria (1989a) also reported
that the contribution of roots of food crops to the total plant weight varies
from 1020%, depending on species and cultivars within species. However,
Sainju et al. (2005b) reported that C accumulation in cotton and sorghum
roots ranged from 1% to 14%. Data in Table 1 show the contribution of the
root system of 20 upland rice genotypes grown at two N rates in a Brazilian
Oxisol. In this study, the nitrogen genotype interaction was significant
because some genotypes were highly responsive to the N application while
others were not. Thus, genotype selection is an important strategy for
upland rice production in Brazilian Oxisols. In the control treatment, the
contribution of the root system to total plant weight varied from 12% to
30%, with an average value of 22%. At the 300 mg N kg 1 soil treatment,
the contribution of root weight to total plant weight varied from 3% to
21%, with an average value of 14%. The proportionally lower root dry
weight at the higher N rate was associated with a significant increase in the
grain and shoot weight of rice genotypes with the addition of N fertilizer
(Fageria and Baligar, 2005).
The contribution of root systems to total plant weights of tropical
legume cover crops under three P levels is shown in Table 2. There was a
significant influence of P rate, cover crop species and P species interaction was significant, indicating different responses of cover crops at different
P rates. At 0 mg P kg 1 level, the contribution of roots to total plant weight
varied from 4.91% to 21.50%, with an average value of 12.98%. With
100 mg P kg 1, the root contribution to the total plant weight varied
from 11.30% to 25.16%, with an average value of 18.93%. At the
200 mg P kg 1 P rate, the contribution of roots to the total plant weight
varied from 6.36% to 25.91%, with an average value of 14.79%. Overall,
increase in root contribution to the total plant weight with the increase in P
rate may be associated with the response of legumes to P fertilization.
Significant differences have been reported among the crop species and
genotypes of the same species in the absorption and utilization of P
(Epstein and Bloom, 2005; Fageria, 2009; Marschner, 1995). Tian et al.
(1998) reported significant responses of legume cover crops grown on
Table 1 Contribution of upland rice roots in the total plant dry weight (%) as
influenced by N rates
Genotype
0 mg N kg 1 soil
BRA01506
BRA01596
BRA01600
BRA02535
BRA02601
BRA032033
BRA032039
BRA032048
BRA032051
BRA042094
BRA042156
BRA042160
BRA052015
BRA052023
BRA052033
BRA052034
BRA052045
BRA052053
BRS Primavera
BRS Sertaneja
Average
F-test
N rate (N)
Genotype (G)
NG
CV(%)
18abcd
18abcd
23abcd
25abcd
23abcd
20abcd
14cd
12d
23abcd
25abcd
24abcd
28abc
17abcd
16bcd
22abcd
30a
27abc
29ab
28abc
26abc
22
3g
3g
6fg
14cde
19abcd
15bcde
21ab
19abc
14cde
12e
15cde
15cde
13de
21a
15bcde
12e
17abcde
11ef
12e
14de
14
*
**
**
17
Table 2 Contribution of tropical legume cover crops root dry weight (%) in the total
plant weight as influenced by phosphorus rates
Cover crop
Crotalaria breviflora
Crotalaria juncea L.
Crotalaria mucronata
Crotalaria spectabilis Roth
Crotalaria ochroleuca G. Don
Calopogonium mucunoides
Pueraria phaseoloides Roxb.
Cajanus cajan L. Millspaugh
Cajanus cajan L. Millspaugh
Dolichos lablab L.
Mucuna deeringiana (Bort) Merr.
Mucuna aterrima (Piper & Tracy)
Holland
Mucuna cinereum L.
Canavalia ensiformis L. DC.
Average
F-test
P rate (P)
Cover crops (C)
PC
20.77a
12.28abc
4.91c
11.66abc
5.18c
11.66abc
7.22bc
21.50a
16.90abc
12.81abc
19.70ab
14.17abc
11.39ef
19.77abcdef
19.29abcdef
24.72ab
22.76abc
12.16def
17.17abcdef
21.12abcde
21.80abcd
22.22abc
15.08bcdef
25.16a
21.22ab
12.49bcd
21.96ab
9.26cd
17.38abc
25.91a
8.15cd
6.36d
13.77bcd
14.71bcd
13.36bcd
16.43abc
11.09abc
11.83abc
12.98
13.44cdef
11.30f
18.93
16.94abc
9.12cd
14.79
**
**
**
265
maturity. Kaspar et al. (1984) noted that the rate of soybean (Glycine max
L. Merr.) root depth penetration reached a maximum during early flowering and declined during seed fill. However, some root growth was observed
throughout the reproductive stage until physiological maturity (Klepper and
Kaspar, 1994).
Slaton et al. (1990) studied root growth dynamics of lowland rice and
found that maximum root growth rates were reached between active
tillering and panicle initiation, and maximum root length was reached by
early booting. Beyrouty et al. (1987) noted that the most rapid rate of root
and shoot growth in flooded rice occurred before panicle initiation, which
corresponds to the plant transition between vegetative and reproductive
growth. Approximately 77% and 81% of total shoot and root biomass,
respectively, was achieved before panicle initiation. Following panicle
initiation, the length of roots and shoots increased only slightly until harvest
(physiological maturity). Beyrouty et al. (1988) also reported that lowland
rice root growth was most rapid during vegetative growth, with maximum
root length occurring at panicle initiation. Root length either plateaued or
declined during reproductive growth.
Fageria and Santos (2011) studied the root and shoot growth of lowland
rice during its growth cycle (Fig. 5). Root dry weight increased in a
quadratic fashion with the advancement of plant age from 19 to 120 days,
but shoot dry weight increased linearly during the growth cycle. Development of the root system was slow during the first 40 days after sowing and
then it increased almost linearly until physiological maturity. The slow
40
30
20
10
Shoot dry
weight (g plant1)
0
60
Y = 12.5474 + 0.5287X
R 2 = 0.9702**
50
40
30
20
10
0
20
40
60
80
100
120
140
Figure 5 Root and shoot dry weight of lowland rice as a function of plant age (Fageria
and Santos, 2011).
Maximum root
length (cm)
increase in root dry weight early in the growth cycle may be associated with
low translocation of photosynthetic materials due to low leaf area (Fageria,
2007). When there is a low amount of photosynthetic product, a major part
goes to the shoot, and very little is translocated to the roots (Fageria, 1992).
Root growth generally parallels shoot growth in crop plants. When a
large amount of nutrients, especially N, is supplied to leaves from roots,
photosynthesis remains high during maturation, which secures the supply of
carbohydrates to roots. Hence, the activities of roots and shoots are mutually
dependent (Osaki et al., 1997). Figure 6 shows maximum root length and
root dry weight of dry bean during the growth cycle of a Brazilian cultivar
BRS Valente under greenhouse conditions. Maximum root length was
40
30
20
10
0
4
Number of
trifoliates (plant1)
0
20
Y = 0.4461EXP(0.0787X 0.00041X 2)
R 2 = 0.9216**
10
0
20
Y = 0.0297EXP(0.1161X 0.00054X 2)
R 2 = 0.9430**
10
20
40
60
80
100
Figure 6 Relationship between plant age and dry bean growth parameters (Fageria
and Santos, 2008).
7. RootShoot Ratio
The partitioning of photoassimilate between roots and shoots has
frequently been analyzed as a balance between root and shoot activity
(Brouwer, 1966; Davidson, 1969a; Werf, 1996). Different plant species
may have different patterns for photosynthate transportation and allocation
to shoot and root (Dyer et al., 1991; Freckman et al., 1991). There is an
interdependence of shoot and root for growth and development. The shoot
relies on the root for water and nutrients, while the roots depend on the
shoot for carbohydrates (Hoad et al., 2001). The terms shoot and root
are used here in a botanical sense and refer, respectively, to the entire aerial
and subterranean portions of higher seed plants (Aung, 1974). In the early
part of the twentieth century, shootroot ratios were used rather extensively
to characterize plant response to imposed nutritional changes. Root growth
is closely related to the whole plant growth. This relationship is called
allometry or relative growth. Root dry weight is related to the total dry
weight of a plant using the following equation (Yoshida, 1981):
WR HWTh ;
where WR is the root dry weight, WT is the total dry weight (shoot
dry weight root dry weight), and H and h are constants. The above
relationship has been tested for different rice cultivars grown under various
environmental conditions, and can be expressed by the following equation
(Yoshida, 1981):
WR 0:212WT0:936 :
When plants are small (substitute 1 for WT), WR/WT is 0.2; WR/WT
values approach 0.1 as plants grow larger (substitute 105 for WT). In other
words, ratio of root dry weight to total dry weight ranges from 0.2 at the
seedling stage to 0.1 at the reproductive stage (heading) for rice (Yoshida,
1981).
The above relationship between root and total dry weights gives an
estimate of root mass that remains in soil if shoot weight is known. For
example, when plants produce shoot dry weights of 3 Mg ha 1 at heading,
269
Grain yield
(g plant1)
Y = 3.6178EXP(0.9326X 0.0109X
20
R = 0.5101**
10
20
30
40
Figure 7 Relationship between maximum root length and grain yield of upland rice.
Not the root dry weight.
Grain yield
(g plant1)
20
10
Y = 1.0257EXP(0.9397X 0.0718X
R 2 = 0.8656**
0
Figure 8
Relationship between root dry weight and grain yield of upland rice.
found that yield stability was correlated with root system length. The first
author studied the relationship between maximum root length and root dry
weight and grain yield of upland rice (Figs. 7 and 8). Grain yield increased in
a quadratic fashion with increasing root length or root dry weight, and root
dry weight was a better predictor than root length of yield. Similarly, the
author studied the relationships between root length and root dry weight
and shoot dry weight of tropical legume cover crops (Figs. 9 and 10). There
was a significant increase in shoot dry weight of legume cover crops with
increasing root length and dry weight, and as with upland rice, root dry
weight was a better predictor than root length of shoot dry weight.
R 2 = 0.8783**
7
6
5
4
3
2
1
0
10
20
30
40
Maximum root length (cm)
50
Figure 9 Relationship between maximum root length and shoot dry weight of
tropical legume cover crops. Values are averages of 14 tropical legume cover crops.
Y = 0.3422EXP(5.5187X 2.7205X
8
7
R 2 = 0.9712**
6
5
4
3
2
1
0
0.2
0.4
0.6
0.8
1.0
Root dry weight (g plant1)
1.2
Figure 10 Relationship between root dry weight and shoot dry weight of tropical
legume cover crops. Values are averages of 14 tropical legume cover crops.
273
respectively. Similarly, Brown et al. (1987) and Pan et al. (1985) also
reported differences in rooting depths of corn and barley (Hordeum vulgare
L.), respectively. Similar variability exists in dicotyledonous species where
most work has been undertaken on legumes, including dry bean (Fageria,
2002a; White and Castillo, 1989), soybean (Taylor et al., 1978; Zobel, 2005a),
chickpea (Cicer arietinum L.; Vincent and Gregory, 1986), peanut (Mathews
et al., 1988), and white clover (Trifolium repens L.; Caradus, 1990). Hamblin and
Tennant (1987) compared root growth and water uptake of wheat and lupin
(Lupinus albus L.) and Gregory and Brown (1989) did similar work with barley
and chick pea. These authors concluded that there were differences between
the species in the root length necessary to extract water, and the rate of water
extraction was greater in legumes than in cereals. Fageria (1991) studied root
dry weight of the Brazilian upland rice cultivars IAC 47 and the International
Rice Research Institute (IRRI) cultivar IR 43 and concluded that root dry
weight of Brazilian cultivars was almost double that of the IRRI cultivar
during the entire growth cycle. Xiaoe et al. (1997) reported that hybrid rice
has a more vigorous root system, larger panicle, and more grains per panicle
than traditional rice cultivars.
The genotypic variability in the root growth of annual crops has been
used to identify superior genotypes for drought-prone environments
(Gregory, 1994; Hurd et al., 1972). Gregory and Brown (1989) reviewed
the role of root characters in moderating the effects of drought and concluded that roots may have a direct effect, by increasing the supply of water
available to the crop, or an indirect effect by changing the rate at which the
supply becomes available. Where crops are grown on deep soils and water is
stored throughout the whole soil profile, the depth of rooting has a major
influence on the potential supply of water (Gregory, 1994). Rain may
replenish the upper soil during the season, but later growth and grain filling
in many crops are accomplished during periods of low rainfall when soil
moisture stored deep in the profile must be utilized. Sponchiado et al. (1989)
reported that in dry bean, drought avoidance results from root growth and
soil water extraction deep in the profile.
Atkinson (1990) reported significant variation in the speed of root
penetration, specific root length, branching pattern, root density, total
root mass, and root hair development of 25 spring barley cultivars. Information reported in the literature on old and new cereal cultivars indicates
that more modern cultivars are more responsive than older cultivars to high
nutrient availability (Haberle, 1993; Haberle et al., 1995), although they
tend to have a lower root fraction (Wahbi and Gregory, 1995). Root
physiological characteristics also differ among cultivars and can affect processes like nutrient acquisition (Hoad et al., 2001; Marschner, 1998). The
rate of uptake of nutrient per unit root length depends on the nutrient
availability but also varies considerably among cultivars (Hoad et al., 2001;
Romer, 1985).
rice (O. sativa L.), and other species and crops growing in wet soils, allows
the roots to grow into saturated soils and still extract nutrients (Zobel,
2005b). In corn, the leaf bases also form aerenchyma when submerged,
while the enclosed stem bearing aerenchymatous nodal (adventitious) roots
may elongate more rapidly ( Jackson, 1994). Aerenchyma formation in corn
roots appears to be triggered by the loss of tonoplast integrity (Campbell and
Drew, 1983), although cell wall disintegration is also involved at an early
stage, at least in rice ( Jackson, 1994; Webb and Jackson, 1986). Aerenchymatous roots are formed either by some cell wall separation and cell
collapse (lysigeny) or by cell separation without collapse (schizogeny).
Figure 11 shows cross sections of rice and wheat roots, and lysigenous and
schizogenous intercellular space. Both forms result in large longitudinal
channels in root cortices, and such structures enhance diffusion of atmospheric or photosynthetic oxygen from shoots to roots so that aerobic
respiration and growth can be maintained (Armstrong, 1979). Changes in
root morphology occur after flooding for both wetland and nonwetland
plant species. Flooding may also increase branching of roots, development
of new adventitious roots, and superficial rooting (Laan et al., 1989).
Cellular spaces that exist in roots facilitate oxygen diffusion; however,
the amount of aeration varies greatly among plant species. In the case of
certain marsh plants like rice, root cortical cells are arranged in columns, and
when channels form, the spaces become large and continuous, facilitating
diffusion. However, in the case of certain terrestrial plants, the cellular
arrangement is oblique and the spaces formed are small (Horiguchi, 1995).
There are marsh plants, however, that do not display columnar forms, while
some terrestrial plants do (Horiguchi, 1995). In the case of rice, large
cortical aerenchyma spaces develop schizogenously and lysigenously even
in well-aerated soils. Horiguchi (1995) reported that in rice and other
graminaceous plants, aerenchymous cells are well developed in both the
nodes and the internodes. The oxidizing capacity of roots of crop plants also
depends on the soil fertility level. Higher soil fertility improves the oxidation activity of rice roots (Horiguchi, 1995). Application of silicon to rice
A
Rice
B
Lysigenous
intercellular
space
Wheat
Schizogenous
intercellular
space
Figure 11 Cross section of a rice (A) and wheat (B) root showing lysigenous and
schizogenous intercellular spaces (from Horiguchi, 1995).
crops increases the diffusion of oxygen from the tops to the roots, strengthening oxidation activity and the deposition of ferrous iron on root surfaces,
and restricting the transport of excess iron to the top (Horiguchi, 1995).
Development of adventitious roots in some crops is related to plant
tolerance of soil waterlogging. Such roots developed by corn after one
day of flooding were short and porous enough to allow significant O2
movement from the shoots to the root meristems (Miller, 1986; Wenkert
et al., 1981). Despite this adventitious root formation and increased root
porosity, corn and most other crop plants soon perish under such conditions
(Miller, 1986).
277
quality. However, conservation tillage may compact surface soil horizons and
may lead to poor root growth. Adverse effects of soil compaction on crop
production have been recognized for many years. Cato the Elder (234149 B.C.)
wrote that the first principle of good crop husbandry is to plow well and the
second principle is to plow again, presumably to provide a mellow (wellaerated) seedbed (Unger and Kaspar, 1994). Conservation tillage increases
soil bulk density (Martino and Shaykewich, 1994) which may inhibit root
growth in the upper part of the soil profile (Cannell, 1985; Lampurlanes
et al., 2001), reducing nutrient uptake and plant growth (Peterson et al.,
1984). Qin et al. (2004) reported that wheat root length density, mean root
diameter, and percentage of small-diameter roots were lower in no-till than
conventional tillage. Root diameter may be indicative of the effects of soil
strength on root growth and affects the utilization of nutrients in the soil.
Sidiras et al. (2001) reported thicker barley roots under conventional tillage
than under no-till. In general, bulk densities that impede root growth are
1.55 Mg m 3 for clay loams, 1.65 Mg m 3 for silt loams, 1.80 Mg m 3 for
sandy loams, and 1.85 Mg m 3 for loamy fine sands (Miller, 1986).
Tillage-induced differences in the soil nutrient status may also have a
significant impact on root growth (Qin et al., 2004). Conservation tillage
often results in the stratification of soil nutrients, especially of immobile
nutrients like P (Crozier et al., 1999; Holanda et al., 1998; Logan et al.,
1991). This produces greater soil fertility near the soil surface which, in
contrast to the effects of compaction described above, causes an increase in
root length density near the soil surface under conservation tillage (Cannell
and Hawes, 1994; Gregory, 1994). Frequently, root growth is greater from
0 to 5 cm in conservation and no-tillage systems than in conventional tillage
systems (Chan and Mead, 1992; Rasmussen, 1991; Wulfsohn et al., 1996).
Radial root swelling has been reported for lupins (Lupinus angustifolius
L.) grown in compacted soil (Atwell, 1989), for barley (H. vulgare L.) under
mechanical impedance (Wilson et al., 1977), and for mustard (Brassica sp.) in
drying soil (Vartanian, 1981). Studies on root elongation of cotton as a
function of soil strength and soil water content showed that root elongation
is more sensitive to soil strength than water content (Taylor and Ratliff,
1969). Root volumes were less affected than length, indicating an increase
in root diameter (Ball et al., 1994). Chassot et al. (2001) reported that
conservation tillage decreases soil temperature, and this may be the main
reason for the poor growth of the roots and shoots of corn seedlings
compared with conventional tillage under temperate humid conditions.
Considering the many advantages of conservation tillage compared to
conventional tillage, the effects of soil compaction produced by conservation tillage can be minimized. Unger and Kaspar (1994) reported that
growing deep-rooted crops in rotation will help avoid or alleviate compaction, improving root distribution and increasing rooting depth. These
authors also reported that weather conditions and soil moisture can enhance
120
100
80
60
40
20
u
C
F
e
g
M
a
C
AF
Figure 12 Relative shoot dry weight of dry bean as influenced by adequate fertility
level (AFL) and other nutrients were not applied or omitted from the Oxisol.
120
Shoot
Root
100
80
60
40
20
0
N+P+K
Figure 13 Relative dry weight of shoot and root of dry bean as influenced by N, P, and
K fertilization.
exists, both within and among crop species, in nutrient acquisition and use.
This variability reflects differences in root morphology and mechanisms that
either aid or prevent ion movement into the root (Gabelman et al., 1986).
12.1. Nitrogen
Nitrogen is one of the most yield-limiting nutrients in crop production in
most agroecosystems. Nitrogen plays numerous key roles in plant biochemistry, including being an essential constituent of enzymes, chlorophyll,
N+P+K
Wheat
Upland
rice
Figure 14 Root growth of wheat at the top and upland rice at the bottom grown at
N P K and N, P, and K levels.
N+P+K
Corn
nucleic acids, storage proteins, cell walls, and a vast array of other cellular
components (Harper, 1994). Consequently, a deficiency of N in crop plants
profoundly influences plant growth, development, and yield. The recovery
281
of applied N with chemical fertilizers is lower than 50% for most annual
crops. The low recovery of N is associated with loss of this element
by leaching, denitrification, volatilization, incorporation into soil microorganisms, and soil erosion (Fageria and Baligar, 2005). To improve
the efficiency of N uptake and use by crop plants, root systems play an
important role.
Root morphology is influenced by the amount of N fertilizer applied
(Eghball et al., 1993) and factors such as temperature (Feil et al., 1991) and
soil mechanical impedance (Bengough and Mullins, 1990). Eghball et al.
(1993) showed that N stress in corn reduced root branching. Similarly,
Maizlish et al. (1980) showed greater root branching in corn with increasing
levels of applied fertilizer N. Costa et al. (2002) reported that greater root
length and root surface area were obtained at an N fertilizer rate of
128 kg N ha 1 compared with either the absence of fertilizer N or the
higher rate of 255 kg N ha 1. Nitrogen fertilizer improves root growth in
soils having low-OM content (Gregory, 1994; Robinson et al., 1994).
Nitrogen fertilization may increase crop root growth by increasing soil N
availability (Garton and Widders, 1990; Weston and Zandstra, 1989). Sainju
et al. (2001) observed that tomato (Lycopersicon esculentum Mill.) root growth
was greater with hairy vetch and crimson clover cover crops and
90 kg N ha 1 than with no cover crops or N fertilization. Nitrogen also
improves production of lateral roots and root hairs, as well as increasing
rooting depth and root length density deep in the profile (Hansson and
Andren, 1987). Hoad et al. (2001) reported that surface application of
nitrogen fertilizer increases root densities in the surface layers of the soil.
Nitrogen fertilization can increase root length and root surface area and
decrease root mass per unit area of corn (Anderson, 1987; Costa et al., 2002).
It is well known that roots tend to proliferate in nutrient-enriched soil zones
(Drew et al., 1973; Qin et al., 2005). Russell (1977) refers to this as a
compensatory response. The results of pot experiments showed that corn
roots were longer and thinner in zones that were rich in N (Durieux et al.,
1994; Zhang and Barber, 1992, 1993).
Root mass was less affected by N than root length, but the effect may
depend on the stage of maturity of the crop (Baligar et al., 1998). Higher
rates of application of N reduced root growth and depth of rooting in wheat
(Comfort et al., 1988) and reduced root:shoot ratio in rye (Brouwer, 1966).
In corn, the primary root system was 16% thicker when NH4N was
applied rather than NO3N. The NH4N treatment also increased the
diameters of lateral and first- and second-order nodal roots (Anderson et al.,
1991). Baligar et al. (1998) reported that relative dry weights of roots of rice,
dry bean, corn, and soybean were reduced by 38%, 56%, 35%, and 11%,
respectively, when N was omitted from a complete fertilizer. Nitrogen
deficiency also reduces branching and root hairs in cereals and legumes
(Baligar et al., 1998).
2
Y =1.6407 + 0.0053X
R 2 = 0.5944**
50
100
150
200
250
300
350
400
Figure 16 Influence of nitrogen on root dry weight of upland rice. Values are averages
of 20 upland rice genotypes.
Table 3 Root length and root dry weight of 20 upland rice genotypes as influenced
by nitrogen fertilization
Root length (cm)
Genotype
BRA01506
BRA01596
BRA01600
BRA02535
BRA02601
BRA032033
BRA032039
BRA032048
BRA032051
BRA042094
BRA042156
BRA042160
BRA052015
BRA052023
BRA052033
BRA052034
BRA052045
BRA052053
BRS Primavera
BRS Sertaneja
Average
F-test
N rate (N)
Genotype (G)
NG
CV(%)
34.67ab
30.00b
27.00ab
31.67ab
28.00b
30.00a
43.00a
28.50b
30.67ab
30.33b
29.33b
29.67b
31.00ab
29.67b
30.33b
29.00b
28.67b
29.00b
29.33b
30.00b
30.49
NS
**
**
14.25
21.00cd
15.67d
25.33bcd
30.00abcd
32.67abc
28.00abcd
34.67abc
32.00abc
35.67ab
38.00ab
33.00abc
32.50abc
35.00abc
26.50abcd
31.33abc
37.67ab
37.67ab
36.50ab
36.33ab
40.33a
31.99
0.92a
0.87a
1.14a
1.33a
1.11a
1.12a
1.24a
1.05a
1.31a
1.78a
1.19a
1.51a
1.67a
1.48a
1.56a
1.78a
1.75a
1.65a
1.68a
1.45a
1.38
0.40f
0.45f
1.03ef
3.25abcd
3.73ab
2.41cd
3.77ab
3.62abc
2.82bcd
2.31de
2.84bcd
3.33abcd
2.83bcd
4.14a
3.38abcd
2.58bcd
3.66abc
2.72bcd
2.49bcd
2.68bcd
2.72
*
**
**
15.86
(2009) also reported that rice growth was better in 10 mg L 1 Al than with
0 mg Al L 1 in nutrient solution. Brazilian rice cultivars (lowland as well as
upland) are highly tolerant to soil acidity (Fageria et al., 2004). Another
possible explanation is that ammonium sulfate has about 24% S, which may
improve root growth if the extractable soil S level is lower than 10 mg kg 1.
The timing of nitrogen application can also influence the root growth of
crop plants (Table 4). The treatment T3, which produced maximum grain
yield, also produced minimum root length; treatment T2 which produced
minimum grain yield produced maximum root length. There was a
N300
Figure 17 Root growth of cultivar BRS Sertaneja at 0 and 300 mg N kg 1 soil
(Fageria, 2011).
BRS Primavera
N0
N300
Figure 18 Root growth of cultivar BRS Primavera at 0 and 300 mg N kg 1 soil
(Fageria, 2011).
285
BRA052053
N0
N300
BRA052045
N0
N300
5
(NH4)2SO4 (Y ) = 1.9132EXP(0.00093X + 0.000031X
2)
R 2 = 0.5296**
3
2
CO(NH2)2 (Y ) = 1.1194+0.0135X 0.000024X 2
R 2 = 0.4122**
50
100
150
200
250
300
350
400
CO(NH2)2
0N
50 N
300 N
Figure 22 Root growth of upland rice at 0 (left), 50 (center), and 300 (right)
mg N kg 1 soil supplied with urea.
The effects of two N rates on root length and root dry weight of six
tropical legume cover crops are showed in Table 5. Root length as well as
root dry weight were significantly increased with the addition of N fertilizer. Similarly, root length as well as root dry weight were significantly
different among cover crop species. Root growth of four legume cover
crops is shown in Figs. 2528. In all cases, root growth was better at the
higher N rate compared to low N rate.
12.2. Phosphorus
Phosphorus is one of the most yield-limiting nutrients in tropical highly
weathered soils (Fageria, 2009; Fageria and Baligar, 2003, 2008). The
deficiency of P in these soils may be related to the low natural level of this
element in these soils as well as to the immobilization of P in these soils
(Fageria and Baligar, 2008). Phosphate plays many roles in the physiology
and biochemistry of pants. It is a component of important compounds like
(NH4)2SO4
0N
300 N
400 N
Figure 23 Root growth of upland rice at 0 (left), 300 (center), and 400 (right)
mg N kg 1 soil with ammonium sulfate.
Table 4 Root length and root dry weight of upland rice as influenced by nitrogen
timing treatments
Nitrogen timing treatmenta
T1
T2
T3
T4
T5
Average
F-test
CV(%)
28.75b
35.00a
25.75b
27.00b
28.25b
28.95
**
8.7
4.21bc
2.56c
5.74ab
7.70a
4.98b
5.04
**
20.9
T1
T3
T2
T5
T4
Cover crops
Showy
crotalaria
Calopogonio
Pueraria
Pigeon pea
Lablab
Gray mucuna
bean
Average
F-test
N rate (N)
Cover crops
(C)
NC
CV(%)
22
29
0.48
0.77
28
23
30
27
28
34
24
34
32
31
0.30
0.16
0.73
0.74
2.01
0.46
0.15
0.48
1.72
1.84
26b
31a
0.74b
0.90a
**
**
**
**
NS
6.47
**
16.27
**, NS Significant at the 1% probability level and nonsignificant, respectively. Means in the same line for
each growth parameter, followed by the same letter are not significantly different by Tukeys test
at the 5% probability level.
0N
Figure 25 Root growth of black mucuna bean tropical legume cover crop at 0 (left)
and 100 (right) mg N kg 1 soil.
0N
Figure 26 Root growth of gray mucuna bean tropical legume cover crop at 0 (left)
and 100 (right) mg N kg 1 soil.
291
Lablab
0N
100 N
Figure 27 Root growth of Lablab tropical cover crops at two N levels: 0 (left) and 100
(right) mg N kg 1 soil.
phospholipids, phosphorylated sugars and proteins, DNA (deoxyribonucleic acid), and RNA (ribonucleic acid). It is also a component of ATP
(adenosine 5-triphosphate), PEP (phosphoenolpyruvate), NADPH (nicotinamide adenine dinucleotide phosphate, reduced), and other biochemicals
that use the phosphate bond in energy utilization and storage (Blevins,
1994).
Phosphorus is a key nutrient essential for root development in highly
weathered tropical soils. Baligar et al. (1998) reported that P increased the
root weight of wheat, dry bean, and cowpea in a quadratic fashion with
increasing P rate from 0 to 200 mg kg 1 of soil. The regression equations
related to P rates versus root dry weight were Y 0.4019 0.094X
0.00031X2, R2 0.74* for wheat, Y 0.4813Exp. (0.019X 0.000071
X2), R2 0.63* for dry bean, and Y 0.7351 0.0232X 0.000073
X2, R2 0.80** for cowpea. Based on these regression equations, maximum root dry weight for wheat was achieved at 152 mg P kg 1, whereas
maximum root dry weight for common bean and cowpea was achieved at
134 and 159 mg P kg 1 soil, respectively. These results indicate that
increasing P levels increased root growth, but root growth was reduced at
Crotalaria spectabilis
0N
100 N
Figure 28 Root growth of Crotalaria spectabilis tropical legume cover crops at two N
levels: 0 (left) and 100 (right) mg N kg 1 of soil.
Table 6 Root dry weight and root length of 20 upland rice genotypes as influenced
by P levels
Genotype
BRA01506
BRA01596
BRA01600
BRA02535
BRA02601
BRA032033
BRA032039
BRA032048
BRA032051
BRA042094
BRA042156
BRA042160
BRA052015
BRA052023
BRA052033
BRA052034
BRA052045
BRA052053
BRS
Primavera
BRS
Sertaneja
Average
F-test
P level (P)
Genotype
(G)
PG
Low P
(25 mg kg 1)
High P
(200 mg kg 1)
Low P
(25 mg kg 1)
High P
(200 mg kg 1)
3.92ab
2.78ab
2.81ab
3.12ab
4.42ab
3.70ab
2.91ab
3.96ab
2.00b
2.82ab
2.50b
5.68a
3.91ab
4.69ab
2.23b
3.18ab
3.07ab
2.57ab
3.56ab
3.22c
2.73c
3.03c
4.30c
3.20c
3.62c
4.36c
3.91c
2.58c
3.92c
2.91c
8.32ab
2.98c
8.55a
2.43c
3.99c
3.08c
3.87c
5.21bc
26.00ab
35.67ab
36.00ab
28.67ab
31.33ab
23.00b
27.67ab
37.00a
36.00ab
30.00ab
29.00ab
32.67ab
27.00ab
29.67ab
27.00ab
31.00ab
38.33a
28.33ab
29.67ab
26.67a
28.00a
29.00a
33.33a
27.00a
29.67a
27.00a
33.33a
30.67a
27.33a
27.00a
33.00a
27.00a
34.33a
29.00a
24.67a
24.67a
23.67a
25.00a
4.36ab
3.92c
34.00ab
23.67a
3.41
4.01
30.9
28.20
NS
**
*
**
**
of P level on root system dry weight was not significant. Root length varied
from 23.00 to 38.33 cm with an average value of 30.9 cm at low P level.
At high P level, root length varied from 23.67 to 34.33 cm, with an average
value of 28.20 cm. There was a significant 10% decrease in root length at the
high P level compared to the low P level. However, higher P level roots had
more fine hairs compared to lower P level (personal observations).
Figures 2931 show root growth of upland rice under different P rates.
Unlike the data cited above, root growth of all the cultivars/genotypes
increased with increasing P levels.
The first author studied the influence of P levels on the root growth of
14 tropical legume cover crops grown on a Brazilian Oxisol. Root dry
weight was significantly influenced by P; crop species and P crop species
interactions were significant (Fig. 32). The significant P crop species
interaction indicates significant variation in shoot dry weight with the
variation in P levels. At the lowest P level (0 mg kg 1), maximum root
dry weight of 0.77 g plant 1 was produced by white jack bean (Canavalia
ensiformis) and minimum root dry weight of 0.01 g plant 1 was produced by
crotalaria (Crotalaria mucronata) and pueraria (Pueraria phaseoloides). At the
medium P level (100 mg P kg 1), maximum root dry weight of
1.91 g plant 1 was produced by black mucuna bean (Mucuna cinereum)
and minimum root dry weight of 0.07 was produced by crotalaria (Crotalaria
0P
50 P
175 P
Figure 29 Root growth of upland rice under 0 (left), 50 (center), and 175 (right)
mg P kg 1 soil (Fageria and Stone, 1999).
0P
25 P
50 P
75 P 100 P 200 P
IAC 164
Figure 30 Root growth of upland rice cultivar IAC 164 at different P levels.
295
BRA 01596
25 P
200 P
Figure 31
1.6
a
200 mg P kg1
1.4
1.2
b
1.0
0.8
0.6
0.4
0.2
b
b
de
de
d
de
de
0
2.0
a
100 mg P kg1
1.8
1.6
1.4
1.2
1.0
bc
bc
bcd
0.8
cde
0.6
ef
0.4
0.2
ef
ef
df
0
0.8
a
0 mg P
0.6
kg1
0.4
c
cd
Figure 32
de
Canavalia ensiformis
Mucuna cinereum
Mucuna aterrima
e
Mucuna deeringiana
Crotalaria ochroleuca
Calopognium
mucunoides
de
cde
Dolichos lablab
de
Pueraria phaseoloides
Crotalaria spectablis
Crotalaria juncea
Crotalaria breviflora
de
cd
Crotalaria mucronata
0.2
The influence of P fertilization on dry bean root dry weight was studied
by Fageria (1989b). Root dry weight of three bean genotypes increased
significantly in a quadratic fashion but differed from genotype to genotype
(Table 7). The variability in root dry weight due to P fertilization was about
70
200 mg P kg1
60
a
ab
ab
ab
50
40
30
c
cd
20
cd
d
c
cd
cd
cd
10
0
60
100 mg P kg1
50
bc
40
cde cde
30
20
def def
f
def
cde
ef
bcd
def
10
0
40
0 mg P kg1
ab
ab
de
de cde
cd
cd
Cajanus cajan
(mixed color)
cde
20
cd
30
cd
bc
de
Canavalia ensiformis
Mucuna cinereum
Mucuna aterrima
Mucuna deeringiana
Dolichos lablab
Pueraria phaseoloides
Calopogonium
mucunoides
Crotalaria ochroleuca
Crotalaria spectablis
Crotalaria mucronata
Crotalaria juncea
Crotalaria breviflora
10
Figure 33 Maximum root length of 14 tropical legume cover crops at three P levels.
200 P
Figure 34 Root growth of tropical legume cover crop Crotalaria mucronata at 0 and
200 mg P kg 1 of soil.
200 P
Figure 35 Root growth of tropical legume cover crop gray mucuna bean at 0 (left) and
200 (right) mg P kg 1 of soil.
299
Crotalaria breviflora
0P
200 P
Figure 36 Root growth of tropical legume cover crop Crotalaria breviflora at 0 and
200 mg P kg 1 of soil.
12.3. Potassium
Potassium plays an important role in the growth and development of plants,
including the root system. Many enzymes are activated in plants by potassium, and it is also required for photosynthesis, transport of photosynthate,
and protein synthesis (Blevins, 1994). Potassium plays a role in cell growth
following cell division by serving as a major component in cell turgor. It also
maintains ionic balance and electrical neutrality in plants. Crops that produce large quantities of protein per unit area of land require more K than
those that produce less protein (Blevins, 1994). Potassium also plays an
important role in opening and closing the stomata.
Tennant (1976) reported that potassium deficiency stops root growth
completely within 1012 days of planting in wheat. Inadequate K reduces
root growth and consequently, crop yields (Baligar et al., 1998). A deficiency of K in an Inceptisol reduced root growth by 23% in lowland rice, by
30% in dry bean, by 12% in corn, and by 11% in soybean (Baligar et al.,
Table 7 Influence of phosphorus fertilization on the root growth of three dry bean
genotypes at harvest
Root dry weight (g per 2 plants)
1
P rate (mg kg
Carioca
CNF10
CNF4856
0
0.29
0.45
0.23
25
1.13
1.79
1.38
50
1.66
1.28
1.83
75
1.31
1.29
1.52
100
1.34
1.60
1.59
125
1.62
1.33
1.78
150
1.31
1.75
1.82
175
1.18
1.29
1.48
200
1.79
1.50
1.67
Regression analysis
P rate versus Carioca (Y) 0.6459 0.0128X 0.000044X2,
R2 0.5179**
P rate versus CNF10 (Y) 0.8704 0.0109X 0.000042X2,
R2 0.3509*
P rate versus CNF4856 (Y) 0.6145 0.0193X 0.000075X2,
R2 0.6953**
Source: Adapted from Fageria (1989b).
*,** Significant at the 5% and 1% probability levels, respectively.
1998). On an Oxisol, a 35% lower root dry mass was observed in 13 corn
genotypes when K levels were 0 versus 200 mg kg 1 of soil (Baligar et al.,
1998). Data in Table 8 show that at 0 mg K kg 1 of soil root dry weight of
common bean (P. vulgaris L.) genotypes varied from 1.54 to 3.14 g per
3 plants, a variation of twofold. At the 200 mg K kg 1 level, root dry weight
varied from 1.50 to 2.30 g per 3 plants, a variation of 1.5-fold. Similarly,
maximum root length varied from 42 to 46 cm at low K level and 3244 cm
at higher K level. At the higher K level, there was a slight decrease in the
root length of all the genotypes, and the root weight of three genotypes also
decreased at the higher K level. However, at the higher K level, there were
more root hairs than at the low K level (visual observations).
There is widespread evidence for genotype diversity in the root characteristics of many crops in response to the environment and increasing
interest in using this diversity to improve agricultural production and
consequently, nutrient-use efficiency (Barber, 1994; Gregory, 1994).
Mullins et al. (1994) studied K placement effects on the root growth of
cotton grown on a fine sandy loam soil. Root density measurements taken
in-row showed that root growth at depths >20 cm was improved with inrow subsoil additions of K. Tupper (1992) also observed increased cotton
taproot length when K fertilizer was band-applied in the subsoils of
Table 8 Root dry weight and maximum root length of six common bean genotypes as
influenced by potassium levels applied to a Brazilian Oxisol
0 mg K kg 1
200 mg K kg 1
Maximum
Root dry weight root length
Genotypes (g per 3 plants) (cm)
Maximum
Root dry weight root length
(g per 3 plants) (cm)
Apore
Perola
Ruda
IAC
Carioca
Jalo
Precoce
Safira
Average
1.54
1.97
1.94
3.14
45
42
44
45
1.67
2.04
2.30
1.70
32
39
35
38
2.24
42
1.67
36
1.77
2.10
46
44
1.50
1.81
44
37
Mississippi soils with low soil-test K. On the other hand, Hallmark and
Barber (1984) and Yibrin et al. (1993) reported that localized applications of
K did not promote root growth. However, K has been shown to promote
the root growth of some vegetable crops (Zhao et al., 1991). Fageria (1992)
determined the root growth of rice grown in nutrient solution as well as in
an Oxisols at the stress and nonstress levels of K. At the stress levels of K, rice
root growth was reduced compared with nonstress levels. Alfalfa herbage
yield, root weight, and root total nonstructural carbohydrates increased
with increasing K fertilizer (Kitchen et al., 1990).
12.4. Calcium
Absolute Ca deficiency is difficult to identify on plants grown in acidic soils
(Kamprath and Foy, 1985). Most acidic soils contain adequate total Ca for
most plants, and Ca-deficiency symptoms are rarely observed in the field.
Only in highly leached, acidic, low-cation exchange soils (Oxisols and
Ultisols) would absolute deficiencies be likely to occur (Garrity et al.,
1983). Levels of Ca required for essential growth functions are so low as
to approach those of micronutrients. Hence, the major role of Ca in soils
and in plants is to exclude or detoxify other elements such as Al, Mn, and
heavy metals that might otherwise become toxic (Garrity et al., 1983).
Gonzalez-Erico et al. (1979) evaluated the maize response to deep
incorporation of limestone on an Oxisol. They reported that incorporation
12.5. Magnesium
Magnesium is an essential macronutrient for plant growth. The most wellknown and important role of Mg is its occurrence in chlorophyll molecules.
In addition to this, Mg is required for many essential physiological reactions,
especially phosphorylation reactions (Mengel and Kirkby, 1978).
Fageria and Souza (1991) determined the effects of Mg levels on root
weights of rice, common bean, and cowpea grown in an Oxisols of Central
Brazil (Fig. 38). Dry weights of rice roots were higher at the lowest Mg
concentration compared with the highest soil Mg concentration. Initial
exchangeable Mg levels of surface soils were 0.1 cmol kg 1. They increased
to 0.3 cmol kg 1 within 3 days after liming and to 0.75 cmol kg 1 at harvest
time (33 days after sowing). The lack of growth responses to applications of
Mg indicated that this level of exchangeable Mg was adequate to meet Mg
0 Mg lime ha1
Figure 37
12 Mg lime ha1
3
Upland rice
2
Y = 1.0752 0.08852X
R 2 = 0.85*
0
Common bean
4
3
2
Y = 2.4497exp(0.3009X 0.05047X 2)
R 2 = 0.88*
1
0
Cowpea
4
3
2
Y = 2.6658exp(0.3007X 0.05983X 2)
R 2 = 0.80**
1
0
0
Figure 38 Root dry weights of upland rice, common bean, and cowpea grown with
different Mg levels in an Oxisol (adapted from Fageria and Souza, 1991).
requirements of upland rice grown in this limed soil. Dry weight of roots of
common bean increased with Mg application up to 3 cmol kg 1 of soil.
Similarly, significant responses of cowpea root growth to soil Mg levels
were observed, and maximum root weight was achieved at
2.5 cmol Mg kg 1 of soil.
12.6. Sulfur
Sulfur has long been recognized as an essential element for plant growth and
development and classified as a macronutrient. Crop responses to applied
sulfur have been reported in a wide range of soils in many parts of the world
(Fageria, 2009). Sulfur plays many important roles in the growth and
development of plants. Fageria and Gheyi (1999) summarized important
functions of the sulfur in the plant. It is an important component of two
amino acids, cysteine and methionine, which are essential for protein
formation. Since animals cannot reduce sulfate, plants play a vital role in
12.7. Micronutrients
Micronutrients have also been called minor or trace elements, indicating
that their required concentrations in plant tissues are small compared to the
macronutrients (Fageria et al., 2002; Mortvedt, 2000). Based on physicochemical properties, except B and Cl, the essential micronutrients are
metals. Even though micronutrients are required in small quantities by
field crops, their influence can be as great as that of macronutrients in
crop production. Micronutrients are normally constituents of prosthetic
groups that catalyze redox processes by electron transfer (such as with
the transition elements Cu, Fe, Mn, and Mo) and form enzymesubstrate
complexes by coupling enzymes with substrates (Fe and Zn) or enhance
enzyme reactions by influencing molecular configurations between
enzyme and substrate (Zn) (Fageria et al., 2002).
Micronutrient deficiencies in crop plants are widespread because of (i)
increased micronutrient demands from intensive cropping practices and
adaptation of high-yielding cultivars which may have higher micronutrient
demand, (ii) enhanced production of crops on marginal soils that contain
low levels of essential micronutrients, (iii) increased use of high-analysis
fertilizers with low amounts of micronutrients, (iv) decreased use of animal
manures, composts, and crop residues, (v) use of many soils that are inherently low in micronutrient reserves, (vi) use of liming in acid soils, and (vi)
305
Figure 39
Figure 40
Table 9 Root dry weight of upland rice and root length of corn as influenced by zinc
fertilization
Zn rate (mg kg 1)
0
5
10
20
40
80
120
R2
1.08
1.33
1.48
1.55
1.73
1.45
1.18
0.75**
55
54
52
55
57
58
52
0.24*
10
40
307
80 mg Zn kg1
Soja
120
5
mg Zn kg1
Dry
bean
Figure 42
of dicots are generally higher than monocots (Fageria, 2000). Fageria (2000)
reported that maximum root growth of upland rice can be achieved with
the application of 0.4 mg B kg 1 soil, whereas maximum root growth of
dry bean required 1.9 mg B kg 1 of soil. Figure 43 shows that B requirements for root growth varied among the crop species. Application of
24 mg B kg 1 soil decreased the root dry weight of upland rice and corn
in greenhouse studies. However, application of B at lower rates to the same
soil increased the root dry weight in dry bean, soybean, and wheat (Fig. 43).
0.3
Upland rice
0.2
0.1
Y = 0.26810.00806X
R 2 = 0.83*
0.0
Corn
2
1
Y = 2.0432exp (0.05957X+0.00105X 2)
R 2 = 0.93**
Common bean
Y = 1.3058exp (0.03466X0.00961X 2)
R 2 = 0.98**
2
1
0
Soybean
0.6
0.3
Y = 0.7473exp (0.0991X0.00652X 2)
R 2 = 0.91*
0.0
Wheat
0.3
0.2
Y = 0.2793exp (0.01993X0.00328X 2)
R 2 = 0.90*
0.1
0.0
10
15
20
Boron applied (mg kg1)
25
Figure 43 Root dry weights of upland rice, maize, common bean, soybean, and wheat
grown with different B levels on an Oxisols of central Brazil (adapted from Fageria,
2000).
12.7.3. Copper
Copper deficiency is often observed on plants grown in soils inherently low
in Cu (coarse-textured and calcareous soils) and in soils high in OM, where
Cu complexes with organic substances (Fageria et al., 2002). Higher-thannormal Cu supplies usually inhibit root growth more than shoot growth
(Lexmond and Vorm, 1981). Use of Cu-containing fungicides and anthelmintic compounds in agriculture has resulted in Cu toxicity in some plants,
but naturally occurring Cu toxicity is relatively uncommon (Welch et al.,
1991). Root dry weight of wheat and root length of dry bean were significantly increased by the application of copper fertilization (Table 10).
12.7.4. Iron
Iron deficiency is a worldwide problem and occurs in numerous crops
(Fageria et al., 2002; Marschner, 1995). Iron deficiency occurs not because
of Fe scarcity in soil or plants, but because various soil and plant factors affect
Fe availability, inhibit its absorption, or impair its metabolic use (Marschner,
1995; Welch et al., 1991). Plant species that commonly become Fe-deficient
are peanut (A. hypogaea L.), soybean (G. max L.), sorghum (Sorghum bicolor
L. Moench), and upland rice (O. sativa L.). Iron deficiency reduces root
growth (Table 11). Iron toxicity (indicated by leaf bronzing) can be serious
for production of crops in waterlogged soils. For wetland rice, Fe toxicity is
the second most severe yield-limiting nutrient disorder, and it has been
reported in South America, Asia, and Africa (Fageria et al., 2008b). Iron
toxicity decreases the root growth of lowland rice (Fageria et al., 2008b);
however, genotypic differences exist (Fig. 44).
Table 10 Root dry weight of wheat and root length of dry bean as
influenced by copper fertilization
Cu rate (mg kg 1) Wheat (g per 4 plants)a Dry bean (cm)a
0
2
4
8
16
32
64
96
R2
0.53
0.60
0.50
0.48
0.47
0.47
0.43
0.17
0.88**
25
30
27
28
28
24
30
14
0.42**
20 days age
60 days age
0.0
2.5
5.0
10.0
20.0
40.0
80.0
160.0
Average
0.03
0.09
0.13
0.10
0.08
0.11
0.07
0.02
0.08
0.64
1.19
1.05
1.07
1.18
1.24
0.84
0.12
0.92
CNA810208
CNA808951
Fe100 mg L1
311
313
4.1
4.7
5.3
5.9
6.6
7.0
R2
1.90
3.58
4.67
5.40
4.73
3.80
0.99**
pH 4.9 pH 5.9
Dry
bean
Lowland soil
Figure 45 Root growth of dry bean grown on a Brazilian lowland soil (Inceptisol;
Fageria and Stone, 1999).
Much of the Al displaced by Ca can be leached from the root zones. This
practice works well in sandy soils or Oxisols with clay loam aggregates
which behave hydrologically like sands (Foy, 1992). Poor root growth of
crop plants has been frequently observed in highly weathered acid soils in
various countries (Alcordo and Rechcigl, 1993). The chemical factor identified as most responsible for poor root growth is excess soluble Al (Alcordo
and Rechcigl, 1993; Foy, 1992). Excess Al3 has been reported to inhibit
root growth by binding to the PO4 portion of DNA in the root cell nuclei,
reducing template activity and thus cell division (Matsumoto and
Morimura, 1980). In legumes, it has been shown to impair the growth of
root hairs and rhizobia, reducing root nodule initiation and function
(Munns and Franco, 1982). Excess Al may also adversely affect the root
and overall plant growth in nonphytotoxic ways by competing with Ca and
Lowland soil
pH 4.9
pH 5.9 pH 6.4
Soybean
pH 6.7 pH 7.0
Mg for uptake by plants (Rengel and Robinson, 1989). The use of gypsum
can neutralize subsoil acidity by leaching CaSO4 and forming AlSO4
which is not toxic for root growth (Alva and Sumner, 1989). Ritchey
et al. (1980) reported that application of gypsum increased subsoil Ca and
Mg while decreasing Al and improving the root growth of corn in Brazilian
Oxisol.
13.1.3. Maintenance of adequate amounts of organic matter
The benefits of organic matter addition to soils include improving nutrient
cycling and availability to plants through direct additions as well as through
modification in soil physical and biological properties. The complementary
use of organic manures and chemical fertilizers has proved to be the best soil
fertility management strategy in the tropics (Fageria and Baligar, 2005).
Enhanced soil organic matter increases soil aggregation and water-holding
capacity, provides an additional source of nutrients, and reduces P fixation,
toxicities of Al and Mn, and leaching of nutrients (Baligar and Fageria,
1999). Build-up of organic matter through additions of crop and animal
residues increases the population and species diversity of microorganisms
and their associated enzyme activities and respiration rates (Fageria, 2002c).
The use of organic compost may result in a soil that has greater capacity to
resist the spread of plant pathogenic organisms. The improvement in the
overall soil quality may produce more vigorous root systems and higher
crop yields (Fageria, 2002c).
315
Dd ECa
;
Da ECd
where Dd and Da are depths of drainage water and applied water, respectively, and ECa and ECd are electrical conductivity of applied water and
drainage water, respectively.
13.1.8. Integrated cropping systems and pest management
Reduced tillage, N side-dressing, and early planting can be included in
integrated pest management programs with no risk of increasing potential
for root damage from western maize rootworm (Roth et al., 1995). Miltner
et al. (1991) reported that cyst nematodes suppressed soybean root growth
on susceptible cultivars, whereas root growth of tolerant cultivars was
317
14. Conclusions
The study of plant roots is one of the most promising, but least
explored, areas of research related to mineral nutrition. Roots are poorly
studied primarily because of their physical location and growth habit.
Understanding plant root growth is important for improving productivity
of annual crops in agroecosystems. Root systems are important plant organs
because they absorb water and nutrients and also provide mechanical
support to the plant. In addition, roots synthesize growth substances and
hormones such as cytokinins that are important in leaf function and possibly, grain development. In plant growth analysis, the role of roots is
generally ignored due to the difficulty in getting accurate root growth
data under field condition. The principal structure of the root system
includes four developmentally distinct classes of roots. These are taproots,
basal roots, lateral roots, and shootborn roots. Current evidence suggests
that the four root classes that make up the primary and secondary root
systems are physiologically distinct from each other. Rooting depth and
spreading capacity are important traits for uptake of water and nutrients.
Root growth varies among plant species and cultivars within species and can
be modified by environmental factors. Genotypic differences in root growth
among crop species and genotypes of the same species under similar and
variable environmental conditions are now well demonstrated, and the
possibility of developing genotypes of desirable root systems to soil properties offers exciting prospects for the future. Root number, maximum length,
and root dry weight increase with increasing levels of macro- and micronutrients in the soil to a point beyond which root growth is suppressed.
Plant roots together with their associated bacteria and fungi play an important role in the formation, maintenance, and turnover of soil aggregates.
Currently, the techniques available to measure root systems are laborious
and time consuming, and this limits their use in plant physiological research.
Hence, it is necessary to develop root measurement techniques that are
simple, cheap, and less time consuming.
ACKNOWLEDGMENT
The authors thank Dr. Charles Allan Jones, Professor, Texas-A&M University, Texas
AgriLife Research and Extension Service, Dallas, Texas for their peer review of the chapter
and for giving valuable suggestions for its improvement.
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