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Suggested citation:
Kirk, G J D, ed. (1994) Rice roots: nutrient and water use. Selected papers from the
International Rice Research Conference. International Rice Research Institute, P.O.
Box 933, Manila 1099, Philippines.
ISBN 971-22-0050-7
Contents
Foreword
The rice root-soil interface 1
G.J.D. KIRK, J.L. SOLIVAS, AND C.B.M. BEGG
Rice root traits for drought resistance and their genetic variation 67
K.T. INGRAM, F.D. BUENO, O.S. NAMUCO, E.B. YAMBAO, AND C.A. BEYROUTY
Use of molecular markers to exploit rice root traits for drought tolerance 79
H.T. NGUYEN, J.D. RAY, AND LONG-XI YU
Foreword
The rice plant invests up to 60% of its energy as carbon in its root system. However,
the proportion of research effort devoted to the root system has been much smaller than
this, and consequently our understanding of the rice roots and their function in the
capture of nutrients and water lags behind our understanding of the rest of the plant.
This is particularly so for rice, compared with other cereals, because the rice plant's
ability to grow under waterlogged soil conditions arises from morphological and
physiological adaptations in its roots. These adaptations have important consequences
for nutrient and water uptake.
A greater understanding of the root system is now needed to support efforts to
increase rice yield potential and improve the productivity of resources. As part of the
1992 International Rice Research Conference, a symposium on rice roots and the
uptake of nutrients and water was held to review present knowledge and to develop
recommendations for future research. The papers included here were reviewed and
revised on the basis of the discussions during the symposium. It is hoped that they will
serve as a benchmark on the topic for some time to come.
Klaus Lampe
Director General
The rice root-soil interface
G.J.D. Kirk, J.L. Solivas, and C.B.M. Begg
Models and experimental studies of the rhizosphere of rice plants growing in an-
aerobic soil show that two major processes lead to considerable acidification (1-2 pH
units) over a wide range of root and soil conditions. One process is the generation of
H+ in the oxidation of Fe 2+ by O2 released from the roots. The other is the release of
H+ from the roots to balance the excess intake of cations over anions, N being taken
up chiefly as NH 4+. CO2 exchange between the roots and soil has a much smaller
effect. The zone of root influence extends a few millimeters from the root surface.
Substantial differences are found along the root length and over time. The acidifica-
tion and oxidation cause increased sorption of NH4+ by soil solids, thereby impeding
the movement of N to absorbing root surfaces but may also cause solubilization and
enhanced uptake of soil P. Implications for nutrient management and the develop
ment of nutrient-efficient rice germplasm are discussed.
Rice roots growing in anaerobic soil can greatly modify the soil near them. As a result,
the soil as “seen” by the root is entirely different from the bulk soil, although it is the
latter whose properties we normally measure. Some important processes influenced by
conditions in the rice rhizosphere are listed in Table 1. This paper examines the extent
and dynamics of root-induced changes and their consequences for nutrient and toxin
dynamics.
Three main processes operate. First, release of O2 from roots and its reaction with
Fe2+ , generating acidity:
Secondly, because the roots take up a considerable excess of cations over anions (N
being taken up from anaerobic soil chiefly as NH 4+ ), they release H+ into the soil to
Table 1. Effects of rice rhizosphere conditions.
maintain electrical neutrality across the root-soil interface. Thirdly, because high
pressures of CO2 arise in anaerobic soil and within the roots, the roots may either
release CO2 or take it up from the soil, with corresponding changes in soil pH.
The root-influenced zone extends just a millimeter or so into the soil, but existing
experimental methods cannot analyze the soil with resolutions much finer than a
millimeter. Because of this and because the system involves the simultaneous opera-
tion of a series of complex, linked processes, we use simulation models in addition to
experiments.
2 Kirk et al
1. Simulated reactant profiles in the rhizosphere after different times. Parameter values given in
Kirk (1993).
2. Simulated pH profiles in the rhizosphere after 10 d. Root wall permeability and cortical O 2
concentration held constant giving mean O2 efflux from root = 0.2 nmol/dm 2 per s; H+ efflux
from root = 1 nmol/dm 2 per s. Other parameter values as in Figure 1.
4 Kirk et al
in Fe2+ oxidation and H+ released from the root to balance ion intake. The model
predicts that CO2 is lost from the root to the soil near the root tip, but taken up by the
root near the base; the resultant CO2 concentration profiles in the soil are small. There
is an important interaction between the two sources of acidity which results in their
combined effect being greater than the sum of their individual effects (Fig. 2). This is
explained as follows. pH changes are propagated through soil by the movement of
conjugate acid-base pairs: acids from regions of low pH to high and bases in the
opposite direction (Nye 1986). The most important pairs are generally H 3O+-H2O and
H2CO3 (derived from CO2)-HCO3- . The relative importance of the pair H3O+-H2O is
greater at low pH and that of H2CO3-HCO3 at high pH. Therefore the net rate of soil
acidity diffusion is minimal in the pH range in which H3O+ and HCO3- concentrations
are both low. In this pH range, a flux of acid or base through the soil results in steep
pH gradients.
The predictions in Figures 1 and 2 are for conditions around aerenchymatous
primary roots, but the model predicts similar changes in the rhizospheres of fine lateral
roots. Laterals are presumably responsible for the bulk of nutrient uptake in lowland
rice because they comprise the bulk of the total root length. (Indeed, the reason they
comprise the bulk of the root length is the slow rate of movement of N and other
nutrients to root surfaces in anaerobic soil and the consequent need for a large surface
area per unit root mass.) Thus, it is conditions in the rhizospheres of laterals that we are
most concerned with.
6 Kirk et al
of N to the root surfaces. For arable crops growing in fertile aerobic soil, the dominant
form of N taken up is NO3 - and it is sufficiently mobile in the soil that root morphology
and uptake properties do not limit uptake until soil N levels become very low (Drew
1990). But for lowland rice, the situation is entirely different: the dominant form of N
taken up is NH 4+, which is far less mobile.
The rate of NH4+ transport through the soil to root surfaces depends on its
distribution between the soil solid and solution, and this is largely determined by the
concentration of anions in solution (in accordance with electroneutrality), though
modified by the concentrations of other exchangeable cations. In most flooded soils,
HCO3- is the dominant anion (in aerobic soils, it is NO 3 - ) but its concentration will
decrease as the rhizosphere pH decreases, becoming negligible below about pH 5.5.
NH4 + mobility will be correspondingly decreased. This will be offset, to some extent,
by the effect of acidity on cation sorption, tending to decrease the NH4+ concentration
in the solid relative to that in solution, but exacerbated by the depletion of exchangeable
Fe 2+.
Phosphorus
In contrast to the effect on N uptake, rhizosphere oxidation and acidification may
enhance solubilization and uptake of P. This is important because, in many rice soils,
flooding results in a decrease in P mobility following an initial release, and rice plants
consequently depend on root-induced solubilization for the bulk of their P.
We measured concentration profiles of extractable P fractions in the soil near roots,
using the same root-plane technique. The fractions, measured sequentially, were anion
exchange resin, alkali (subdivided into inorganic and organic), acid, and residual. The
results showed a large depletion of acid-soluble and residual P in the zone of Fe
oxidation and acidification, 0-5 mm from the root plane (Fig. 5). Part of the solubilized
P appeared closer to the root plane in the alkali-soluble P pool, but the bulk was
absorbed by the roots. The quantity of P taken up by the plants during the experiment,
calculated from the increase in plant dry matter multiplied by the mean % P content,
was about 25 µmol. This was roughly matched by the net depletion of acid-soluble and
residual soil P. The quantity contained in the freely available resin pool was negligible
in comparison with plant demand.
In flooded soils, a large part of the P is often acid-soluble, being associated with
metal carbonates and hydroxides formed following soil reduction and with other
negatively charged surfaces through exchangeable cations. The alkali-soluble pool
would include that associated with Fe(OH)3 formed in the oxidation reaction and with
root-derived organic material. Evidently, the reimmobilization of acid-solubilized P
on Fe(OH)3 and root-derived organic matter did not prevent access of P to the root
surfaces.
By contrast, we found that the bulk of P taken up by rice growing in aerobic soil was
drawn from an alkali-soluble pool (Hedley et al 1994). To solubilize this P, the roots
would need an alternative mechanism. Current evidence suggests that this mechanism
involves the release of low molecular weight organic acid anions such as citrate. These
must leave the root cells in the dissociated salt form, not the acid form, because the
cytoplasmic pH (6-7) is well above the pKs of the acids in question. Therefore, from
charge-balance considerations, in anaerobic soil where N is taken up as NH4+ , export
of organic acid anions would be difficult.
8 Kirk et al
5. Profiles of extractable P fractions in flooded soil exposed to a planar layer of rice roots for
10 d. Numbers of curves are means of values in the soil bulk.
Discussion
For reasonable rates of nutrient uptake and O2 loss from rice roots, acidification and
Fe 2+ oxidation in the rhizosphere can result in impeded N uptake but enhanced P
uptake.
These findings have interesting consequences for the relation between root traits
and nutrient uptake efficiency under different conditions. In conditions in which N is
the limiting nutrient, root porosity and wall permeability should be just sufficient to
References cited
Ahmad A R, Nye P H (1990) Coupled diffusion and oxidation of ferrous iron in soils. I. Kinetics
of oxygenation of ferrous iron in soil suspension. J. Soil Sci. 44:395-409.
Armstrong W, Justin SHFW, Beckett PM, Lythe S (1991) Root adaptation to soil waterlogging.
Aquatic Bot. 39:57-73.
Bedford B L, Bouldin D R, Beliveau B D (1991) Net oxygen and carbon dioxide balances in
solutions bathing roots of wetland plants. J. Ecol. 79:943-959.
Drew M C (1990) Root function, development, growth and mineral nutrition. Pages 35-57 in The
rhizosphere. J.M. Lynch, ed. Wiley-Interscience, Chichester.
Hedley M J, Kirk G J D, Santos M B (1994) Phosphorus efficiency and the forms of soil
phosphorus utilized by upland rice cultivars. Plant Soil 158: 53-62.
IRRI—International Rice Research Institute (1991) Program report for 1990. P.O. Box 933,
Manila, Philippines. p. 203-205.
IRRI—International Rice Research Institute (1992) Program report for 1991. P.O. Box 933,
Manila, Philippines. p. 198-199.
Kirk G J D (1993) Root ventilation, rhizosphere modification, and nutrient uptake by rice. Pages
221-232 in Systems approaches for agricultural development. F.W.T. Penning de Vries, P.S.
Teng, and K. Metselaar, eds. Kluwer Academic Publishers, Dordrecht, The Netherlands.
Kirk G J D, Ahmad A R, Nye P H (1990) Coupled diffusion and oxidation of ferrous iron in soils.
II. A model of the diffusion and reaction of Fe 2+, H+ and HCO 3- in soils and a sensitivity
analysis of the model. J. Soil Sci. 44:411-431.
Kirk G J D, Hedley M J, Bouldin D R (1993) Phosphorus efficiency in upland rice cultivars.
Pages 279-295 in Papers and reports on the management of acid soils (IBSRAM/ASIALAND).
Network Document No. 6. International Board for Soil Research and Management, Bang-
kok.
Nye P H (1986) Acid-base changes in the rhizosphere. Pages 129-153 in Advances in plant
nutrition. Vol. 2. P.B. Tinker and A. Lauchli, eds. Praeger, New York.
Notes
Authors’ address: G.J.D. Kirk, J.L.Solivas, and C.B.M. Begg, International Rice Research
Institute, P.O. Box 933, Manila, Philippines
Citation information: Kirk G J D, ed. (1994) Rice roots: nutrient and water use. International
Rice Research Institute, P.O. Box 933, Manila, Philippines.
10 Kirk et al
Root growth and nitrogen
uptake in rice: concepts
for modeling
H.F.M. ten Berge, T.M. Thiyagarajan, B. Mishra,
K.S. Rao, and R.N. Dash
Data from field experiments are used to examine the rate-limiting step in nitrogen (N)
uptake by lowland rice. For the vegetative phase over a wide range of fertilizer levels,
N uptake per ha (N c) and uptake rates per ha (dNc/dt) increased with the amount of
N applied (A). dNc /dt did not reach an upper limit. Increases in dNc /dt with A were
more than in proportion to increases in root mass (Mr). After panicle initiation (PI),
there was no consistent relation between A, dNc /dt, and Mr. The root-shoot ratio
decreased with development stages and was independent of A at any stage. Up to
PI, Mr and the rate of N absorption per unit Mr both increased with A to the extent
that fertilizer N recovery increased as A increased. An approximate calculation
showed that changes in cumulative recovery with A could be explained by differences
in Mr. Two hypotheses to explain dNc /dt during the pre-PI stage were consistent with
the experimental data: 1) the uptake rate was diffusion-limited and proportional to
the concentration of NH4+ in the root zone bulk soil solution [NH4+ ] and to root length
density (RLD), and, 2) the uptake rate was independent of [NH4+ ] and governed by
the plant internal N content through a mechanism superimposed on the effect of Mr.
Total N uptake over the season was a linear function of A. This linearity was the
result of a positive feedback between Nc and dN c/dt during vegetative growth,
followed by a negative or neutral feedback after PI.
Much work on N in rice has focused on loss processes in relation to floodwater and soil
chemistry and biology (e.g., De Datta and Patrick 1986), and detailed physicochemical
models of these processes have recently become available (Rachhpal-Singh and Kirk
1993 a,b). Also, the utilization of N by the crop once absorbed and its effects on dry
matter production are now reasonably well-understood (Yoshida 1981, Kropff et al
1993). However, the process of N uptake by the root system has received relatively
little attention. An understanding of this is essential for the development of a complete
model of the fate of N in rice production systems.
The supply of N to a single root depends on the rate of transport from the bulk soil
solution to the root surface. For lowland rice, this transport is principally by diffusion
(Makarim et al 1991). The maximum rate of diffusion is reached for “zero-sink”
conditions in which the root maintains a negligibly low concentration at its surface,
thus maximizing the concentration gradient for a given bulk concentration (de Willigen
and van Noordwijk 1987).
For an entire root system, “supply” is a more complex group of processes. Roots are
unevenly distributed and the entire root system is not necessarily equally active. Good
mathematical models exist to integrate knowledge on the various physiological,
physical, and physicochemical processes affecting uptake. Taking single root behavior
as a starting point, the uptake model of de Willigen and van Noordwijk (1987, 1991)
assesses nutrient uptake rate as a function of soil conditions (water content, bulk
nutrient concentration, and nutrient diffusivity), root length density (RLD), and
parameters specifying “crop demand” and maximum flux density over the root surface.
But parameterization of such mechanistic models is difficult. The measurement of
RLD in rice under field conditions is virtually impossible due to very fine branching
(Yoshida 1981, Drenth et al 1991) and strong spatial RLD gradients near the soil
surface. Moreover, the roles of different root fractions in uptake are unknown
(Makarim et a1 1991), and we have little information about maximum nutrient flux
densities across rice root surfaces under either controlled or field conditions, or about
the dependence of root nutrient uptake characteristics on soil and crop conditions. Also,
the concept of “demand” is not well-defined, and if the crop’s nutritional status affects
root uptake characteristics, demand turns out to directly affect supply itself.
The purpose of this study was to determine what measurable variables must be
considered to describe N uptake at different stages of lowland rice crop development,
as a first step toward a mechanistic description of the system.
12 ten Berge et al
2. The PUAT set was collected at G.B. Pant University of Agriculture and Technol-
ogy, Pantnagar, Uttar Pradesh, during the 1987 WS. Twenty-nine-d-old seedlings
of Pant Dhan 4 were transplanted on 9 Jul at 20- x 100-cm spacing, at two
seedlings per hill, in a randomized block design with four replications. Nitrogen
was applied as urea at 0, 60, 120, 180, and 240 kg N/ha in three splits: 50% basal,
25% topdressed at 27 DAT (active tillering), and 25% topdressed at 51 DAT (PI).
The 60 kg N/ha treatment received no second topdressing, but received 50% of
the total dose at 27 DAT. Flowering occurred at 72 DAT and maturity at 108 DAT.
Plant samples were collected at 25, 49, 72, and 108 DAT. Dry mass and N contents
of leaves, stems plus sheaths, grains, and roots were measured for all harvests,
except root mass which was omitted at maturity.
3. The CRRI set was collected at the Central Rice Research Institute, Cuttack,
Orissa, during the 1990 dry season (DS). Thirty-eight-d-old seedlings of IR36
were transplanted on 25 Jan at 15- x 15-cm spacing, at two seedlings per hill, in
a randomized block design with four replications. Nitrogen was applied as urea
at 0, 50, 100, and 150 kg N/ha in three splits: 50% basal, 25% at 20 DAT, and 25%
at 53 DAT (1-2 wk after PI). The crop flowered at 70 DAT and was harvested at
92 DAT. Plant samples were collected at 20, 30, 40, 50, 70, and 92 DAT and
analyzed as above. Nitrogen in plant samples was determined only at 20, 50, and
70 DAT.
Results
N uptake as a function of N application
Tables 1-3 give the results for TNAU-TNFUU, PUAT, and CRRI, respectively. The
maximum amount of N accumulated in the crop Nc (kg/ha), as a function of the amount
1. Relation between maximum total crop (leaves, stems, roots, and panicles) N uptake (N,) and
N application in the three experiments. Maximum Nc occurred on 63 DAT at TNAU-TNRRI,
on 108 DAT at PUAT, and on 70 DAT at CRRI.
0 19 19 19 19 19
18 260 377 491 50 521
25 326 494 620 710 775
32 370 590 764 925 1,030
39 421 672 880 1,150 1,250
46 463 734 930 1,324 1,525
53 482 848 1,120 1,502 1,750
63 537 836 1,183 1,790 2,003
81 530 836 997 1,507 1,968
0 85 85 85 85 85
18 706 916 984 1,211 1,370
25 1,184 1,745 2,341 2,738 3,050
32 1,880 2,757 3,853 4,465 5,150
39 2,501 3,593 5,011 5,952 7,184
46 3,058 4,422 6,132 7,371 8,636
53 3,566 5,532 7,390 8,986 10,443
63 4,303 7,806 9,809 12,514 13,669
81 5,478 9,891 11,856 13,660 14,671
14 ten Berge et al
Table 2. Total crop (leaves, stems, roots, and panicles) N uptake and root and shoot mass on
different days after transplanting (DAT). PUAT, Pantnagar, 1987 WS.
0 13 13 13 13 13
25 317 349 414 479 520
49 638 976 1,034 1,276 1,500
72 1,016 1,434 1,566 1,806 2,041
108 ndb nd nd nd nd
0 32 32 32 32 32
25 853 1,187 1,481 1,797 2,043
49 2,599 4,295 5,393 6,813 7,774
72 6,502 9,570 10,887 11,722 12,485
108 4,910 6,737 7,741 9,035 9,978
2. Relation between maximum root mass M, and N application in the three experiments.
Maximum M, occurred on 63 DAT at TNAU-TNRRI, 72 DAT at PUAT, and 50 DAT at CRRI.
Discussion
We need to know what the rate-limiting step or steps in N uptake are in order to decide
what process or processes must be modeled in detail. At one extreme, the root system
might be so efficient at extracting N from the soil that the rate of uptake is essentially
independent of root and soil characteristics until the soil N concentration falls to a low
level. At the other extreme, the rate of transport of N to root surfaces might be so slow
that uptake is solely determined by the rate of transport-asgoverned by the soil
transport characteristics and the N concentration-and the root surface area or RLD.
16 ten Berge et al
3. Changes in root-shoot ratios with time at different N application levels in the three
experiments.
In between these extremes, root characteristics would have a greater or lesser effect on
uptake depending on how well the plant is able to regulate the root system’s
characteristics to match uptake with the rate of N transport through the soil. Clearly,
as the crop grows and soil N becomes depleted, the rate-controlling step may change,
and a complex model would be required to describe all eventualities. However, our
purpose here is to identify the most critical process or processes and thereby simplify
modeling as far as possible.
In the following sections, we use our experimental results to test various hypotheses
about rate-limiting steps. In the absence of detailed measurements of the changes in
Hypothesis 1: If [NH4+] is above a critical value, the uptake rate is independent of the
amount of N applied.
This hypothesis addresses the first extreme previously described. In this case, increases
in Nc with A would be due to N application extending the period over which [NH4+]
is above the critical value. To model this situation, we would only need information
on the period when [NH4+] exceeds the critical value in some part of the root zone.
We test the hypothesis by comparing rates of N uptake. Ideally, we need instanta-
neous uptake rates, but we necessarily only have average values over the time between
sampling events (sampling intervals). This is awkward because differences in time
averaged dNc/dt with A could be due to different durations of uptake at the level that
is independent of [NH4+], (dNc/dt)max , within the sampling interval, as opposed to
differences in (dNc/dt)max per se. The problem is avoided by comparing two consecu-
tive sampling intervals that are not interrupted by N application. If dNc/dt depends on
A in both such intervals, a variable duration of (dNc/dt)max is not responsible and
Hypothesis 1 can be rejected. Only the TNAU-TNRRI data set satisfies the require-
ments for such an analysis.
In the TNAU-TNRRI data, dNc/dt increases with A, both during the 0-18 and 18-
25 DAT periods (from data in Table 1). This finding is supported by other reported
data—e.g., Schnier et al (1990) for both direct seeded and transplanted rice and
Makarim et al(1991) for transplanted rice. We therefore reject Hypothesis 1.
Hypothesis 2: If [NH4+] is above a critical value, the uptake rate is proportional to RLD
but is otherwise independent of [NH4+].
18 ten Berge et al
4. N uptake rates per unit root mass at TNAU-TNRFU. Numbers on curves are N application
levels (kg N/ha).
(1)
and
(2)
Hypothesis 3: Uptake rates are linearly related to both [NH4+ ] and RLD.
5. Observed ratios of N uptake at two consecutive N application levels (R N ), compared with the
corresponding ratios of root mass (RM ).
20 ten Berge et al
6. Relation between total crop (leaves, stems, roots, and panicles) N uptake (Nc ) and N
application in the three experiments at different times.
the end of the time interval (t 2) are 25 and 46 DAT in Figure 6a, respectively, the
momentary recovery for A(t 2 ,i) is It is the
extra (over the i=0 treatment) uptake in treatment i as a fraction of the latest N split
applied preceding t2 . For the first sampling event, momentary and cumulative recov-
eries are identical.
The values for each of these recoveries are given in Table 4. In the early tillering
stage (18 and 20 DAT for TNAU-TNRRI and CRRI, respectively), recoveries were in
the 0-20% range for all treatments, but part of the applied N was still in the soil. No clear
trend with A t (i) was found at this stage for any of the recovery types. Later during the
tillering stage (25 DAT at TNAU-TNRRI and PUAT), recoveries of all three types
increased slightly with At (i). Because constant recovery across i is equivalent to
TNAU-TNRRI TNAU-TNRRI
N application (kg/ha) N application (kg/ha)
DAT
100 200 300 400 100 200 300 400
Incremental recovery
PUAT CRRl
N application (kg/ha) N application (kg/ha)
DAT DAT
60 120 180 240 50 100 150
Cumulative recovery Cumulative recovery
increasing absolute uptake rates, this response is consistent with the above observation
that dNc/dt increased with At (i). The generally low recoveries during the early
development stages are attributed to the limited size of the root system and possible
transplanting damage; N demand is unlikely to limit uptake because in that case
recovery would decrease with At (i).
Recoveries at PI (46, 49, and 50 DAT for TNAU-TNRRI, PUAT, and CRRI,
respectively) were generally high. Cumulative, momentary, and incremental recover-
ies all increased substantially with At (i) in each data set, except at the highest i. The
22 ten Berge et al
latter lack of response shows that Hypothesis 3 is not valid at these high application
levels. That does not contradict the earlier observation that Mr and (dNc /dt)/Mr
continued to respond to At(i) up to the highest i, but it does imply that these responses
were not strong enough to cause a sustained increase in recovery.
Momentary recoveries at PI exceeded 1.0 in all data sets, indicating that some N was
still available from the basal dressing, even after 3-4 wk. Even so, such high values
suggest that practically all N applied around early tillering had been absorbed by the
crop by PI. Incremental recoveries were near 1.0 for the two highest N application
levels in CRRI, and for i=3 in TNAU-TNRRI. The values of this variable were lower
at PUAT, but still reached 0.62 for i=2 and i=3.
Results were less consistent for the sampling intervals from PI to flowering.
Momentary recoveries were poor in all treatments at PUAT and CRRI. All recovery
types decreased from PI to flowering at PUAT, except for i=1. Moreover, recoveries
decreased here with increasing i. At CRRI, too, recoveries decreased with time. At
TNAU-TNRRI, recoveries remained fairly high and decreased only after flowering (63
DAT). Then, momentary recovery also decreased across i levels.
The decline in recovery after PI (or later in the TNAU-TNRRI case) leads us to
conclude that either Hypothesis 3 is invalid at this stage, or that the fraction of roots
active in N uptake decreases as time proceeds. For the latter to be a valid explanation,
however, we have to accept that this sencscence is more pronounced at higher i levels,
which is unlikely.
The larger root system resulting from N application makes the plant better able to
compete for N against loss processes, and hence a larger fraction of N in the root zone
is recovered. This was indeed observed during the vegetative phase, which supports
Hypothesis 3. The phenomenon of increasing recovery with A, however, may also be
caused by other mechanisms involving positive feedback between Nc and dNc /dt. In the
following section, we try to explain the observed trend in recovery vs A(i) on the basis
of observed root masses.
Assuming that the rates of N uptake and loss are both governed by first-order
processes, we obtain for the recovery fraction in treatment j as compared with that in
treatment i (Appendix 1)
(3)
Figure 7 shows cumulative recoveries during the vegetative period calculated with
Equation 3 from observed M rj/Mri and observed recovery at the lowest application
level. We conclude that cumulative recoveries at the higher A(i) levels can be
reasonably well-explained by differences in observed root mass, without invoking
other mechanisms. Therefore, Hypothesis 3 cannot be disproved on the basis of these
observations, except in the highest i of each experiment. The material presented earlier
in disproving Hypothesis 2 provides additional support for Hypothesis 3; this is
essentially independent of the recovery analysis.
This hypothesis addresses a situation in which the uptake rate per unit root length, or
the fraction of the total root length that is active in uptake, is regulated by the plant in
accordance with its internal N content. A high crop growth rate induced by Nc is likely
to be associated with a high carbohydrate supply to the roots, which would tend to
increase N uptake through a positive feedback loop. At high growth rates, possibly the
rate of detoxification of ammoniacal N via amino acid synthesis results in another
positive feedback loop between Nc and N uptake rate.
The present experimental results do not allow differential testing of Hypotheses 3
and 4.
Root-shoot ratio
The lack of sensitivity of the root-shoot ratio to N application observed in these
experiments is unusual in crops. Numerous studies, including the early work by
Brouwer (1962a,b, 1963), have shown that higher nutrient supply results in lower root-
shoot ratio. The concept of functional equilibrium evolved from such observations, and
although the mechanisms are often questioned (Lambers 1983, van Andel et al 1983),
the occurrence of shifts in dry matter partitioning induced by nutrient supply is widely
accepted (Wilson 1988). Partitioning models describe root and shoot growth in terms
of C and N balances, applying a resistance coefficient for transport of these compounds
within the plant (Thornley 1972) or assuming an optimal allocation strategy by the
plant (Johnson and Thornley 1987). On the basis of such concepts, there is no obvious
reason why rice should behave differently from other crops. Peculiar characteristics of
the lowland rice system include the low mechanical resistance to root growth; the root’s
O2 demand being met by diffusion through internal root gas channels; the uptake of N
is as NH4+ ions; and the particular rhizosphere chemistry of anaerobic soils. Kirk et al
24 ten Berge et al
(1993) state that NH4 + uptake may be hampered by rhizosphere acidification arising
from Fe2+ oxidation and NH 4+ uptake, resulting in increased NH4+ sorption by the soil
complex. A phenomenon possibly important in view of O2 supply is the formation of
surface root mats (Kirk and Bouldin 1991), the development of which was shown to
be enhanced by NH4 + as opposed to NO 3 – nutrition (Soezima and Kawata 1969).
Conclusions
Hypothesis 3 seems to be the easiest starting point for the development of an uptake
model. The most important variables would be [NH4+ ] as a function of (z,t), effective
NH4+ diffusivity, and RLD. Assessing the active fraction of RLD and its dependence
on (z,t) remains problematic. This research showed no consistent relationship after PI
among uptake, application level, and root mass.
The root-shoot ratio decreased with development stage and was independent of A t (i)
at every stage. The possibility that this may be related to root O 2 supply bears further
investigation.
Root mass and the rate of N absorption per unit root mass both increased with
application At (i), resulting in high recovery levels for N.
Instantaneous N uptake rate per unit root mass is likely to be roughly proportional
to [NH4 +] up to high [NH4 + ] values. Cumulative N uptake over a finite time interval is
higher than the first-order function of At (i) during vegetative growth. An approximate
calculation shows that the response of cumulative recovery to At (i) can be explained
by differences in root mass.
Nitrogen absorption by the crop over the entire season was shown to be a linear
function of A. This linearity is the result of a positive feedback between cumulative
uptake and uptake rate during vegetative growth, followed by a negative or neutral
feedback after PI.
Nitrogen uptake in the first 3 wk after transplanting was low, but proportional to
At (i), implying that demand did not limit N uptake. Ten to twenty percent of N applied
basally was absorbed, and momentary recoveries exceeding 1.0 found for the next
sampling interval indicate that this N was still available for absorption at the time of
the first topdressing.
References cited
Andel O M van, Soekarjo R, Verkaar H J PA, eds. (1983) Functional equilibrium between shoots
and roots. Neth. J. Agric. Sci. 31(4):
Brouwer R (1962a) Distribution of dry matter in the plant. Neth. J. Agric. Sci. 10:361-376.
Brouwer R (1962b) Nutritive influences on the distribution of dry matter in the plant. Neth. J.
Agric. Sci. 10:399-408.
Brouwer R (1963) Some aspects of the equilibrium between overground and underground plant
parts. Mededeling 213 van het IBS, 31-39.
De Datta S K, Patrick W H Jr, eds. (1986) Nitrogen economy of flooded rice soils. Developments
in plant and soil sciences. Kluwer Academic Publishers, Dordrecht, The Netherlands. 186 p.
Drenth H, ten Berge H F M, Meijboom F W (1991) Effects of growth medium on porosity and
branching of rice roots. Pages 162-175 in Simulation and systems analysis for rice production
(SARP). F.W.T. Penning de Vries, H.H. van Laar, and M.J. Kropff, eds. PUDOC, Wageningen,
The Netherlands.
Johnson I R, Thornley J H M (1987) A model of shoot:root partitioning with optimal growth.
Ann. Bot. 60: 133- 142.
Keulen H van (1977) Nitrogen requirements of rice with special reference to Java. Contrib. Cent.
Res. Inst. Agric. Bogor 30: 1-67.
Kirk G J D, Bouldin D R (1991) Speculations on the operation of the rice root system in relation
to nutrient uptake. Pages 195-203 in Simulation and systems analysis for rice production
(SARP). F.W.T. Penning de Vries, H.H. van Laar, and M.J. Kropff, eds. PUDOC, Wageningen,
The Netherlands.
Kirk G J D, Solivas J L, Begg C B M (1993) The rice root-soil interface. Pages in Rice roots and
nutrient and water use. G.J.D. Kirk, ed. International Rice Research Institute, P.O. Box 933,
Manila, Philippines.
Kropff M J, Cassman K G, van Laar H H (1993) Quantitative understanding of the irrigated rice
ecosystem for achieving high yield potential in (hybrid) rice. Pages in Hybrid rice. S.S.
Virmani, ed. International Rice Research Institute, P.O. Box 933, Manila, Philippines.
Lambers H (1 983) ‘The functional equilibrium,’ nibbling on the edges of a paradigm. Neth. J.
Agric. Sci. 31:305-311.
Makarim A K, Hidayat A, ten Berge H F M (1991) Dynamics of soil ammonium, crop nitrogen
uptake, and dry matter production in lowland rice. Pages 214-228 in Simulation and systems
analysis for rice production (SARP). F.W.T. Penning de Vries, H.H. van Laar and M.J.
Kropff, eds. PUDOC, Wageningen, The Netherlands.
Rachhpal-Singh, Kirk G J D (1993a) A model for predicting the fate of nitrogen fertilizer in
lowland ricefields. I. Theory. J. Soil Sci. (in press)
26 ten Berge et al
Rachhpal-Singh, Kirk G J D (1993b) A model for predicting the fate of nitrogen fertilizer in
lowland ricefields. II. Predicted dynamics of inorganic carbon, nitrogen and acidity in the soil
and floodwater. J. Soil Sci. (in press)
Schnier H F, Dingkuhn M, De Datta S K, Mengel K, Faronilo J E (1990) Nitrogen fertilization
of direct-seeded flooded vs transplanted rice. I. Nitrogen uptake, photosynthesis, growth and
yield. Crop Sci. 30:1276-1284.
Soezima M, Kawata S (1969) “Lion tail like root” formation in rice plant and soil conditions.
Proc. Crop Sci. Soc. Jpn. 38:442-446.
Thiyagarajan T M, Mohandass S, Palanisamy S, Kareem A A (1991) Effect of nitrogen on
growth and carbohydrate partitioning in rice. Pages 132-136 in Simulation and systems
analysis for rice production (SARP). F.W.T. Penning de Vries, H.H. van Laar, and M.J.
Kropff, eds. PUDOC, Wageningen, The Netherlands.
Thornley J H M (1972) A model to describe the partitioning of photosynthate during vegetative
plant growth. Ann. Bot. 36:419-430.
Willigen P de, van Noordwijk M (1987) Roots, plant production and nutrient use efficiency. Ph
D thesis, Wageningen Agricultural University, Wageningen, The Netherlands. 282 p.
Willigen P de, van Noordwijk M (1991) Modelling nutrient uptake: from single roots to complete
root systems. Pages 277-295 in Simulation and systems analysis for rice production (SARP).
F.W.T. Penning de Vries, H.H. van Laar, and M.J. Kropff, eds. PUDOC, Wageningen, The
Netherlands.
Wilson J B (1988) A review of evidence on the control of shoot:root ratio, in relation to models.
Ann. Bot. 61:433-449.
Yoshida S (1981) Fundamentals of rice crop science. International Rice Research Institute, P.O.
Box 933, Manila, Philippines. 269 p.
Notes
Authors’ addresses: H.F.M. ten Berge, Centre for Agrobiological Research, P.O. Box 14,6700
AA Wageningen, The Netherlands; T.M. Thiyagarajan, Tamil Nadu Agricultural University,
Aduthurai, Tamil Nadu 612101, India; B. Mishra, G.B. Pant University of Agriculture and
Technology, Pantnagar, District Nainital, Uttar Pradesh 263145, India; K.S. Rao and R.N. Dash,
Central Rice Research Institute, Cuttack, Orissa 753006, India.
Citation information: Kirk G J D, ed. (1994) Rice roots: nutrient and water use. International
Rice Research Institute, P.O. Box 933, Manila, Philippines.
(A1)
(A2)
The ratio in Equation A1 is equal to r/(l-r), where r is the recovery fraction. Writing
this ratio Xi for treatment i and X j for treatment j, we have
(A3)
Now, assuming that a) ci (z,t ) is proportional at any ( z,t ) to the applied amount A( t,i);
b) Lr is not affected by A; and c) mr (z,t ) varies across treatments by a factor Mri(t) /Mrj (t)
which is independent of t , then Equation A3 reduces to
(A4)
or
(A5)
28 ten Berge et al
Genetic variation in nitrogen
uptake by rice and the effects
of management and soil
fertility
P.C. Sta Cruz and G. Wada
Variation in nitrogen (N) uptake among rice genotype groupings (ecotype, hybrid,
inbred, modern, and traditional) and between plant types and growth types are
investigated for low to moderate fertilizer levels. Rice genotypes of varying growth
durations (GD) were evaluated in terms of N uptake and N use efficiency (NUE). Total
N in the plant at late spikelet initiation, flowering, and maturity increased with GD.
There were substantial differences in total N in the plant up to 30 d after transplant-
ing (DT), but not after maximum tillering (MT). Total N in the plant at 30 DT (NE) was
positively correlated with sink size, grain yield, and N harvest index (NHI), but not
with percent spikelet degeneration or percent filled spikelets in short- and medium-
duration genotypes. This suggests that NE is an important determinant of sink size
and grain yield in short- and medium-duration genotypes. Narrow spacing and high
basal N application shortened the duration of exponential N uptake and increased
the rate of uptake during this phase. A slow-release fertilizer and high soil fertility
increased the rate of N uptake during the middle and late growth stages. In general,
genotypes with low N uptake during the dry season at high N levels have relatively
low N uptake during the wet season at low N levels. The effects of cultural practices
and soil fertility status, shown in altered durations and rates of N absorption, were
reflected in changes in N uptake patterns.
One strategy for improving the utilization of soil and fertilizer N in rice production
systems is to exploit genotypic differences in N absorption ability. Such differences
can be sought among a) different genotype groupings: ecotypes, hybrids, inbreds, and
modem vs traditional cultivars; and b) different morphological and physiological
features: growth type, panicle type, and growth duration (GD). Genotypic differences
can also interact with cultural practices and environmental variables. This paper
examines the sources of variation in N uptake in various genotype groupings and their
interaction with cultural practices and environmental variables. The relationship
between N uptake efficiency and yield-determining parameters in various genotypes
is also considered.
Plant types
Generally, genotypes that perform well at low N levels are tall, have a low percentage
of productive tillers, have high dry matter production, are susceptible to lodging, and
have low panicle to straw weight ratios. They grow relatively fast in the early growth
stages and cover the fields even without N addition. At high N levels, higher growth
rates in the early growth stages result in mutual shading during the middle and late
growth stages. This reduces N uptake because net photosynthesis is decreased,
resulting in fewer tillers and weaker roots. By contrast, genotypes that respond well to
high N have short stature, high panicle to straw weight ratios, high percentage of
productive tillers, and slow growth in the early growth stages. This plant type shows
little mutual shading (Tanaka et al 1964).
Current improved lowland varieties are short- to medium-maturing and intermedi-
ate in panicle number type. Although they produce many tillers and have large leaf area
indices (LAI), they are short and have erect leaves. Mutual shading is avoided by
improved leaf canopy architecture. Yoshida (1981) identified short and stiff culms,
erect leaves, and high tillering as the most important characters for lodging resistance;
increased photosynthesis as a function of LAI; and yield increment as related to number
of panicles.
Hybrids
High total N uptake, dry weight, and leaf area have been observed in F1 hybrids (Blanco
and Akita 1988). This was more evident in japonica-indica crosses than in japonica-
japonica crosses. Similar results were obtained with F1 hybrids of IR varieties and lines
(Sta Cruz et al 1988). The F1 hybrids have higher N uptake compared with their inbred
parents, particularly at 3 wk after transplanting (WT). At 5 WT, differential N uptake
diminished except for the hybrid with medium GD (125 d). The medium-GD hybrids
had consistently higher N uptake until maturity. Likewise, we found that F1 hybrids
generally had higher N harvest index (NHI).
1. Total N in the plant at flowering in relation to growth duration of different rice genotypes under
two N levels. IRRI Farm, 1987 DS.
Total N uptake at critical plant growth stages was positively correlated with GD. At
zero and 90 kg N/ha fertilizer rates, the increase in GD was accompanied by a linear
increase in N uptake at flowering (Fig. 1), at the late stage of spikelet initiation (Y =
1.94 + 0.78x, r = 0.815**, 90 kg N/ha) and at maturity (Y = 2.07 + 0.069x, r = 0.789**,
90 kg N/ha). The short-duration genotypes generally had lower total N uptake than the
long-duration genotypes at flowering, late stage of spikelet initiation, and maturity.
The variation in N uptake among genotypes with different GD can be explained by
differences in the vegetative lag phase (VLP). The VLP is the difference in days
between the occurrences of panicle initiation (PI) and maximum tillering (MT). The
PI of short-duration genotypes occurred before or at the same time as MT when total
N in the plant was still low. By contrast, the PI of long-duration genotypes occurred
after MT when total N in the plant was already high (Fig. 2).
Table 2. Correlation coefficients for N uptake between 0 and 90 kg N/ha and between wet (WS)
and dry seasons (DS). a IRRl Farm, 1986 WS and 1987 DS.
Total N uptake
Parameter
30 DT Flowering Maturity
0 N vs 90 N
DS 0.835 0.905 0.887
WS 0.850 0.889 0.781
DS vs WS
0N 0.830 0.880 0.883
90 N 0.815 0.771 0.841
a All values are significant at the 1% level. n = 60, mean of two replications.
Variation in N uptake among genotypes was apparent during the early growth stages
(30 DT) and MT (30-35 DT). Little variation in N uptake was noted from MT to
flowering and from flowering to maturity (Table 1). This observation can be explained
by the availability of NH4 -N in the plow layer as reported by Wada et al(l989). The
NH4 -N in the plow layer decreased exponentially from 7 DT until 35 DT and remained
constant at a very low level after 35 DT (Fig. 3). During the early growth stages,
NH4-N in the plow layer was not limiting, and variation in N uptake could only have
been due to differences in N absorption abilities between the genotypes. The availabil-
ity of NH4 -N in the plow layer did limit N uptake after MT.
Sta Cruz (1990) found that there were positive correlations between 0 and 90 N
treatments and between the wet season (WS) and dry season (DS) for total N uptake
at 30 DT, flowering, and maturity (Table 2). The genotypes with low total N uptake
under low N level also had low total N uptake under high N level (Fig. 4). The same
relationship was obtained for N uptake between WS and DS (Fig. 5).
5. Relationship between total N uptake at maturity in dry-season (DS) and wet-season (WS)
croppings in rice genotypes with different growth durations. IRRI Farm, 1986 WS and 1987 DS.
Growth
duration Y S DEG S R NHI
(d)
1987 DS
1986 WS
genotypes (Table 3). This result suggests the importance of NE as a parameter for
increasing both sink size and yield of short- and medium-duration genotypes, without
increasing the percent unfilled spikelets and degenerated spikelets.
The NHI increased with NE in short- and medium-duration genotypes. This further
suggests the importance of NE as a parameter in increasing the yield and NUE of short-
and medium-duration genotypes.
In long-duration genotypes, NE was not correlated with sink size, grain yield, and
NHI. Although the proportion of filled spikelets was not affected by NE, degenerated
spikelets increased with NE (Table 3). This result suggests that N uptake during the
middle and late growth stages is an important yield determinant in long-duration
genotypes.
6. N absorption patterns of rice plants grown at different plant spacings (Wada et al 1989).
8. N absorption of rice plants grown with two N fertilizer sources (Wada et al 1989).
Conclusions
There is substantial variation in N uptake between rice ecotypes, plant types, hybrids,
and plants with different GD. Plant type (based on leaf erectness and panicle number)
and GD are the most important factors.
Nitrogen uptake during late spikelet initiation, flowering, and maturity increases
with GD. The variation in N uptake among genotypes with different GD is due to
differences in the duration of VLP. Short-duration genotypes generally have shorter
VLP than long-duration ones and consequently have lower total N uptake.
Variation in N uptake among genotypes of the same GD is apparent during the early
growth stages (30 DT). During these stages, NH 4 -N in the plow layer is likely to be
high, so N uptake is controlled by the N absorption ability of the genotype. During the
middle and late growth stages, the availability of NH4-N in the plow layer is low and
therefore may limit N uptake. Hence, differences in N uptake among rice genotypes
during the middle and late growth stages were not observed.
References cited
Blanco L C, Akita S (1988) Physiological mechanism of heterosis in seedling growth of
japonica-indica F1 rice hybrids. Jpn. J. Crop Sci. 57 (extra 1):99-100.
Cook M G, Evans L T (1983) Nutrient responses of seedlings of wild and cultivated Oryza
species. Field Crops Res. 6:205-218.
Ehara H, Tsuchiya M, Ogo T (1990) Fundamental growth response to fertilizer in rice plants.
Varietal difference in the growth rate at seedling stage. Jpn. J. Crop Sci. 59(3):426-434.
Maruyama S, Tajima K (1988) Growth response to nitrogen in Japonica and Indica rice
varieties. II. Differences in the rate of increase in culm length and leaf area due to nitrogen
fertilization. Jpn. J. Crop Sci. 57(4):692-698.
Miyagawa S (198 1) Nitrogen balance in paddy fields planted with different Indica varieties. Jpn.
J. Trop. Agric. 25:107-114.
Sta Cruz, P C (1990) Sink development in rice genotypes of different growth durations as
affected by nitrogen nutrition, plant spacing, and shading. Ph D dissertation, University of
the Philippines at Los Baños, Laguna, Philippines. 135 p.
Sta Cruz P C, Wada G, Virmani S S (1988) Nitrogen response of F1 hybrid of rice plants. Jpn.
J. Crop Sci. 57 (extra 1):247-248.
Tanaka A, Navasero S A, Garcia C V, Parao F T, Ramirez E (1964) Growth habit of the rice plant
in the tropics and its effect on nitrogen response. IRRI Technical Bulletin 3. International
Rice Research Institute, P.O. Box 933, Manila, Philippines.
Takahashi J, Wada G, Shoji S (1976) The fate of fertilizer nitrogen applied to the paddy field and
its absorption by rice plants. VI. Influence of a thermal factor on the soil ammonium nitrogen
and the absorption of nitrogen by rice plants. Proc. Crop Sci. Jpn. 45:213-219.
Wada G, Aragones D V, Aragones R C (1989) Nitrogen absorption pattern of rice plants in the
tropics. Jpn. J. Crop Sci. 58(2):225-231.
Wada G, Sta Cruz PC (1989) Varietal differences in nitrogen response of rice plants with special
reference to growth duration. Jpn. J. Crop Sci. 58(4):732-739.
Wada G, Sta Cruz P C (1990) Nitrogen response of rice varieties with reference to nitrogen
absorption at early growth stage. Jpn. J. Crop Sci. 59(3):540-547.
Yoshida S (1981) Fundamentals of rice crop science. International Rice Research Institute, P.O.
Box 933, Manila, Philippines. 269 p.
N balance method
N balance is the terminal N content of the soil minus the initial soil N content following
a given treatment. An N balance sheet gives the sum of N gains and losses. An
experiment by Ventura and Watanabe (1983) with 15N-labeled soil showed that gains
in N balance were largely due to differences in N 2 fixation.
In an N balance study with 76 rice genotypes and 8 wild rices grown in pots, App
et al (1986) found wide variation in ability to stimulate N gain (Fig. 1). Palawan, a
traditional variety, ranked low; IR42, an improved variety, ranked high among the
long-duration varieties tested. Extrapolated to a per hectare basis, N gain differences
ranged from 16 to 70 kg N/ha per crop. High positive correlation coefficients were
obtained between N gain and total N uptake, N gain and total dry matter production,
44 Wu et al
1. Differences in N gains of several rice genotypes (App et a1 1986).
and daily dry matter production. However, because N balance experiments are time-
consuming, they are not suitable for large-scale screening.
2. Differences in ARA/plant and ARA/g total plant dry weight of 37 short-, medium-, and long-
duration genotypes (Ladha et al 1988).
46 Wu et al
Table 1. Correlations of plant traits with plant-associated ARA and N uptake at maturity in
37 rice genotypes of different growth durations (Ladha et ai 1988). a
r
Plant traits vs
ARA N uptake at maturity
ARA 0.425**
Root + submerged portion 0.842** 0.294
dry weight (heading)
Shoot dry weight (heading) 0.735** 0.413*
N uptake (heading) 0.735** 0.496**
Total dry weight (maturity) 0.617** 0.871**
N uptake (maturity) 0.425**
Grain yield 0.266 0.707**
Grain N 0.385* 0.822**
Grain yield/N 0.108 0.350*
a
*, ** = significant at the 5 and 1% level, respectively. ARA = acetylene reduction assay.
Although ARA has many advantages, a large plant-to-plant variation and the need
to assay the plant several times during the growth cycle limited its wide use.
15 N dilution method
The 15N dilution method has recently gained acceptance among researchers because it
is relatively simple and provides integrated values over the whole growth duration
(Chalk 1985). The importance of using the right nonfix reference plants to obtain
reliable N 2 fixation estimates and the difficulties in finding suitable ones have been
extensively reviewed (Chalk et a1 1983, Hauck and Weaver 1986, Witty 1983, Danso
1986). However, the need for a perfect match of nonfixing and fixing plants becomes
less important if the soil organic N is labeled with 15N and, on mineralization, a constant
l5N- 14 N ratio in plant available N is produced (Pareek et a1 1990, Rarivoson and Ladha
1992). Labeling and maintaining the stability of 15 N are easier in flooded soils than in
upland soils. This is because the plow sole delineates an Ap horizon where most roots
are located and which is relatively easy to homogenize.
The heterogeneity of added or natural 15N abundance in field soil and the differences
in rooting pattern between genotypes can be major sources of error in this kind of
research. Therefore, six rice genotypes of different growth durations were grown in
well-mixed soil in pots and their 15N contents in grain and straw were measured
(Watanabe et a1 1987). Differences in 15N values were small but significant. The
difference between the maximum and minimum 15N values in the grains was 22% of
the mean value (Table 2).
d 15N
Variety Growth duration
(d) Grain Straw
Table 3. Group of parameters for screening rice ARA and NUE (Ladha et al 1988).
Nf, WP/Ns, Y/Nt, DM/Ns, WP/Nt, and DM/Nt. Significant variations in several
parameters among 24 genotypes were found. Parameters WP/Nt and Y/Nt have been
suggested as the most useful for assessing NUE. Three groups of variables were used
to rank 21 genotypes with different growth durations during the 1986 and 1987 DS
(Table 3). Rankings based on group 1 variables did not correlate significantly with
rankings based on variables of groups 2 and 3. Rankings based on groups 2 and 3
significantly and positively correlated, suggesting that a genotype with high N2
fixation may not always be high in N uptake and efficient utilization, but that a
genotype with higher N uptake may have higher utilization efficiency.
Eleven genotypes were selected from previous field screening trials based on their
divergent BNF, N uptake, and NUE (the yield/Nt absorbed) characteristics (Watanabe
et al 1978; App et al 1986; Ladha et al 1987, 1988). These genotypes were subjected
to preliminary RFLP analysis for evaluating the frequency of polymorphism detected
among them. Four enzymes were used: EcoRV, HindIII, ScaI, and Xba I. Forty-six
mapped DNA from both a rice genomic library (RG) and an oat cDNA library (CDO)
were selected based on their distribution throughout the 12 chromosomes of rice. The
results showed that the combination of Cigalon and IR42 had the highest percentage
of polymorphism (78%), followed by IR42 and OS4 (73%), Cigalon and IR13429-150-
48 Wu et al
Table 4. Percent polymorphism of possible combinations differing in associative N 2 -flxing ability
and NUE (46 probes surveyed using 4 enzymes).a
Cigalon 67 69 67 78
OS4 56 58 58 73
ClCA 58 60 52 45
C66-2803 52 41 43 45
Palawan 67 43 58 67
DR33 54 65 54 45
BR40-300-2-1-E 50 54 41 37
3-2-1-2-(69%), IR42 and Palawan (67%), BG367-4 and Cigalon (67%), and BR51-
46-CI and Cigalon (67%) (Table 4). Considering the good level of divergence
measuring in phenotypic values and the percentage of polymorphism detected, IR42
and Palawan were selected for further study. They are both long-duration varieties
(125-130 d). IR42 consistently demonstrated superior performance. BG367-4 and
IR13427-40-2-3-3-3-3 (100 d) were also selected as test materials. BG367-4 exhibited
higher N uptake and higher NUE than IR13427-40-2-3-3-3-3 (Broadbent et al 1987,
Ladha et al 1988).
Correlation coefficient
Experiment a
Yield/Nt Yield/NUE Nt/NUE
Table 6. Correlation between chlorophyll content of flag leaf at heading stage and atm % 15N
excess of straw samples.
an = 8 for field experiment, n = 12 for open and Leonard's pot experiments. b Y = atm % 15 N excess of straw samples.
50 Wu et al
Chlorophyll content of flag leaf. A close relationship between associative N2
fixation activity and photosynthesis has previously been established for wheat (M.I.
Chumakov, Russian Academy of Sciences, pers. commun.). When the chlorophyll
content of the flag leaf was measured in our experiments, significant variation existed
among varieties, with similar results found using an open pot and a modified Leonard’s
system. Correlations between atom % 15 N excess and chlorophyll content were found
to be negative (Table 6), suggesting that chlorophyll content and associative N2
fixation should be positively correlated. Thus, high chlorophyll content appears to be
an indication of the ability of the rice genotype to stimulate associative N2 fixation.
Eco Rl 11
Eco RV 25
Sca l 26
Hind III 31
Xba l 33
Dra l 33
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Khush and G.H. Toenniessen, eds. CAB International, Wallingford, UK.
Murty M G, Ladha J K (1987) Differential colonization of Azospirillum lipoferum on roots of
two varieties of rice ( Oryza sativa L.). Biol. Fert. Soils 4:3-7.
Nishizawa N, Yoshida T, Arima Y (1983) Electron microscopic study of associative N2 -fixing
bacteria in roots of rice seedling. Soil Sci. Plant Nutr. 29:261-270.
Pareek R P, Ladha J K, Watanabe I (1990) Estimation of N2 fixation by Sesbania rostrata and
S. cannabina in lowland rice soil by 15N dilution method. Biol. Fert. Soils 10:77-88.
Paterson A H, Lander E S, Hewitt J D, Peterson S, Lincoln S E, Tanksley S D (1988) Resolution
of quantitative traits into Mendelian factors by using a complete RFLP linkage map. Nature
335:721-726.
Rarivoson C, Ladha J K (1992) Estimation of N2 fixed by stem and root inoculated Sesbania
rostrata using 15N enrichment of soil ammonium-N as a reference. Biol. Fert. Soils. (in press)
Reiter R S, Coors J G, Sussman M R, Gabelman W H ( 1991) Genetic analysis of tolerance to low-
phosphorus stress in maize using restriction fragment length polymorphisms. Theor. Appl.
Genet. 82:561-568.
Sano Y, Fujii T, Iyama S, Hirota Y, Komagata K (1981) Nitrogen fixation in the rhizosphere of
cultivated and wild rice strains. Crop. Sci. 21:785-761.
Tirol-Padre A, Ladha J K, Gloria C P, Watanabe I (1988) A plant sampling procedure for
acetylene reduction assay to detect rice varietal differences in ability to stimulate N2 fixation.
Soil Biol. Biochem. 20 (2):175-183.
Ventura W, Watanabe I (1983) 15 N dilution technique of assessing the contribution of nitrogen
fixation to rice plant. Soil Sci. Plant Nutr. 29(2):123-131.
Watanabe I, Kee, Alimano B V (1978) Seasonal change of N2-fixing rate in rice field assayed
by in situ acetylene reduction technique. Soil Sci. Plant Nutr. 24:1-13.
Watanabe I, Yoneyama T, Padre B, Ladha J K (1987) Difference in natural abundance of 15N
in several rice ( Oryza Sativa L.) varieties:application for evaluating N2 fixation. Soil Sci.
Plant Nutr. 33(3):407-415.
Witty J F (1983) Estimating N2 fixation in the field using 15 N-labeled fertilizer: some problems
and solutions. Soil Biol. Biochem. 15:631-639.
54 Wu et al
Rainfed lowland rice roots:
soil and hydrological effects
Pradeep K. Sharma, G. Pantuwan, K.T. Ingram, and S.K. De Datta
Rice root systems differ greatly among cultivars, soil conditions, and hydraulic
regimes under rainfed lowland conditions. Unsaturated, light-textured soils should be
used for evaluating varietal differences in root systems for drought tolerance. Future
research to increase drought resistance through improved root systems must
combine germplasm improvement and soil physical management.
Rainfed rice environments are diverse and unpredictable and insufficient or excessive
water is often the major factor limiting productivity. Because root characteristics are
closely associated with drought resistance, they are particularly important under
rainfed conditions.
Root characteristics are basically genetically controlled but they are also strongly
affected by soil conditions and crop management practices (Ghildyal and Tomar 1982,
Cruz et a1 1986, Sharma et al 1987, Mambani et al 1990, Thangaraj et a1 1990). The
occurrence of drought or flood depends on the frequency and amount of rainfall, daily
evapotranspiration, and soil physical properties governing infiltration, water retention
capacity, and drainage. These factors in turn affect rice root growth. Soil nutrient
deficiencies and toxicities also affect root growth. In this paper, we focus on the effects
of soil texture, hydrology, and mechanical impedance on rice root growth.
textured upland soils may grow as deep as 1 m or more. But roots of rice cultivars in
lowland soils rarely grow deeper than 40 cm; about 90% of the total root system is
restricted to the top 20 cm of the soil layer (Table 1).
Hydrological effects
We classify ricefield hydrology into three groups depending on topographic location
and water supply: i) pluvial, also called upper paddy, where rainfall is the only source
of water and surface water accumulation is minimal; ii) phreatic, also called middle
paddy, where rainfall, seepage, water table, and runoff provide water to the root zone
with moderate surface water accumulation; and iii) fluxial, also called the lower paddy,
where rainfall, seepage, water table, and runoff provide water to the root zone with
reliable surface water accumulation. Most rainfed lowland rice areas fall in the fluxial
and phreatic toposequence positions, and only some are in the pluvial category
(O’Toole and Chang 1978, Greenland and Bhuiyan 1982). Drought is the major
56 Sharma et al
Table 2. Influence of soil moisture condition on oxygen diffusion rate (ODR) of soil, and root
porosity, root length, and root mass of IR8 (Pradhan et al 1973).
toposequence position, KDML 105 produced the greatest RMD in the 0-30 cm soil
layer, but at the low toposequence position, RD9 had the greatest RMD in the 10-30
cm soil layer (Fig. 4). The differences in RMD among cultivars in other soil layers were
not significant.
RD9 and IR46 produced relatively more root mass in the 10-40 cm soil layer under
the high-moisture regime; no such differences were detected for KDML 105 in either
toposequence position (Table 3). In general, root mass decreased exponentially with
soil depth (Fig. 5). Maximum depth of root penetration was similar for all cultivars at
both toposequence positions.
58 Sharma et al
3. Effect of moisture regime on root mass density of three rice cultivars at two toposequence
positions in loamy sand soil.
Table 3. Hydrological effects on percent distribution of root mass of three rice cultivars in loamy
sand soil.
5. Depthwise distribution of root mass of three rice cultivars in loamy sand soil at two
toposequence positions.
60 Sharma et al
in the 10-30 cm layer (Fig. 7). Therefore, soil texture must be considered when
evaluating differences in root systems among rice cultivars.
6. Effect of soil texture on root mass density of three rice cultivars grown at low toposequence
positions.
uniformly under flooded conditions until 42 days after transplanting (DT). From 42 to
61 DT, different moisture regimes were created by using a line-source sprinkler
irrigation system. The crop was reflooded uniformly after that. Soil mechanical
impedance increased from 0 to 1.0 MPa as gravimetric moisture content declined from
160 to 65%. Root length density decreased as a power function of soil mechanical
impedance (Fig. 8). Mechanical impedance as little as 0.05 MPa reduced root length
density by 60% as compared with the continuously flooded control. Mechanical
impedance greater than 0.3-0.5 MPa decreased root growth and root extension by 75%.
Although root mass and root length densities responded similarly to mechanical
impedance, effects on root length were of greater magnitude than those on root mass
density. Thus, roots became shorter and thicker in response to increased mechanical
impedance.
The effects of bulk density and mechanical impedance on root length density of
IR36 in a clay soil under flooded conditions were studied (Sharma and De Datta 1986).
Soil bulk density and mechanical impedance were modified by applying different
tillage treatments. Root length density decreased as a power function of bulk density
or mechanical impedance (Fig. 9). Similar observations were made by Mambani et al
(1989) for IR20 in soils of different texture using different tillage practices. They
concluded that mechanical impedance of 0.57 MPa was the critical limit for root
growth of rainfed lowland rice.
62 Sharma et al
8. Relation between root length density of IR54 relative to control and soil mechanical
impedance in the drying puddled lowland soil at 0-10 cm soil depth (Thangaraj et a1 1990).
Table 4. Effect of interaction between submerged soil temperature regime and bulk density
of a lateritic sandy loam soil on root growth of rice (Kar et al 1976).
Conclusions
Growing rice cultivars having extensive deep root systems that are capable of
extracting water from deep soil layers is a means of achieving more efficient use of
rainwater under drought-prone rainfed situations. The rice root system differs strongly
between cultivars, soil conditions, and hydraulic regimes. Hence, it is difficult to make
generalizations about the effects of individual variables. We found that differences
between cultivars in root growth generally decreases with a change from unsaturated
to saturated conditions and with texture from loamy sand to clay. Unsaturated, light-
textured soils rather than saturated clay soils should be used for evaluating differences
in root systems among rice cultivars. Effects of mechanical impedance on root growth,
however, appeared to be similar under saturated and unsaturated conditions; root
length and root mass densities decreased as a power function of soil mechanical
64 Sharma et al
impedance for different rice cultivars. KDML 105 appeared relatively more responsive
to hydrology and soil texture changes than RD9 and IR46. Thus, KDML 105 may be
more widely adapted to rainfed lowland situations than the other cultivars. Future
research should combine germplasm improvement and soil physical management to
increase drought resistance through an improved rice root system.
References cited
Cruz R T, O’Toole J C, Dingkuhn M, Yambao E B, Thangaraj M, De Datta S K (1986)
Shoot and root responses to water deficits in rainfed lowland rice. Aust. J. Plant Physiol.
13:567-575.
Das D K, Jat R L (1977) Influence of three soil water regimes on root porosity and growth of four
rice varieties. Agron. J. 69:197-200.
Ghildyal B P, Tomar V S (1982) Soil physical properties that affect rice root systems under
drought. Pages 83-96 in Drought resistance in crops wih emphasis on rice. International Rice
Research Institute, P.O. Box 933, Manila, Philippines.
Greenland D J, Bhuiyan S I (1982) Rice research strategies in selected areas: environment
management and utilization. Pages 239-262 in Rice research strategies for the future.
International Rice Research Institute, P.O. Box 933, Manila, Philippines.
Hasegawa S, Thangaraj M, O’Toole J C (1985) Root behavior: field and laboratory studies.
Pages 383-395 in Soil physics and rice. International Rice Research Institute, P.O. Box 933,
Manila, Philippines.
Kar S, Varade S B, Ghildyal B P (1979) Pore size distribution and root growth relations of rice
in artificially synthesized soils. Soil Sci. 128:364-368.
Kar S, Varade S B, Subramanyam T K, Ghildyal B P (1974) Nature and growth pattern of rice
root system under submerged and unsaturated conditions. I1 Riso 23:173-179.
Kar S, Varade S B, Subramanyam T K, Ghildyal BP (1976) Soil physical conditions affecting
rice root growth: bulk density and submerged soil temperature regime effects. Agron. J.
68:23-26.
Mambani B, De Datta S K, Redulla C A (1989) Land preparation requirements for rainfed rice
as affected by climatic water balance and tillage properties of lowland soils. Soil Till. Res.
14:219-230.
Mambani B, De Datta S K, Redulla C A (1990) Soil physical behaviour and crop responses to
tillage in lowland rice soils of varying clay content. Plant Soil 126:227-235.
Maurya P R, Ghildyal B P (1975) Root distribution pattern of rice varieties evaluated under
upland and flooded soil conditions. I1 Riso 24:239-244.
Ogunrerni L T (1991) Influence of bulk density and moisture regime of a permeable soil on the
performance of a lowland rice, Oryza sativa L. Trop. Agric. 68:129-134.
O’Toole J C, Chang T T (1978) Drought and rice improvement in perspective. IRRI Res. Pap.
Ser. 14.
Pradhan S K, Varade S B, Kar S (1973) Influence of soil water conditions on growth and root
porosity of rice. Plant Soil 38:501-507.
Sharma P K, De Datta S K (1986) Physical properties and processes of puddled rice soils. Adv.
Soil Sci. 5:139-178.
Sharma P K, De Datta S K, Redulla C A (1987) Root growth and yield response of rainfed
lowland rice to planting method. Exp. Agric. 23:305-313.
Thangaraj M, O’Toole J C, De Datta S K (1990) Root response to water stress in rainfed lowland
rice. Exp. Agric. 26:287-296.
Notes
Authors’ addresses: Pradeep K. Sharma, Department of Soil Science, HPKV, Palampur 176062
H.P., India; G. Pantuwan, Ubon Rice Research Center, P.O. Box 65, Ubon Ratchathani 34000,
Thailand; K.T. Ingram, Agronomy, Plant Physiology, and Agroecology Division, International
Rice Research Institute, P.O. Box 933, Manila, Philippines; S.K. De Datta, College of
Agriculture and Life Sciences, Virginia Polytechnic Institute and State University, Blacksburg,
VA 2401-0334, USA.
Citation information: Kirk G J D, ed. (1994) Rice roots: nutrient and water use. International
Rice Research Institute, P.O. Box 933, Manila, Philippines.
66 Sharma et al
Rice root traits for drought
resistance and their genetic
variation
K.T. lngram, F.D. Bueno, O.S. Namuco, E.B. Yambao, and C.A. Beyrouty
The root traits that confer drought resistance are not well-defined. Glasshouse
research has shown that the root xylem vessel cross-sectional area does not explain
water relation differences across genotypes. In field experiments under upland
conditions, root length density (RLD) was negatively correlated with grain yield of
three rice genotypes grown under different levels of water deficit stress. On the other
hand, in minirhizotron experiments under rainfed lowland conditions, only RLD in the
10-20 and 20-30 cm soil layers were related to relative yields (30-d vegetative phase
stress yield/continuously flooded control yield) in 12 rainfed lowland genotypes and
only on 2 sampling days (86 and 89 d after sowing). Two root characteristics were
common to drought-resistant rainfed lowland genotypes: 1) rapid root responses to
changing soil moisture level in the 10-20 and 20-30 cm soil layers, and 2) greater
absolute RLD below 20 cm.
Capacity for water uptake may limit rice productivity even in flooded soils. Thus,
midday stomatal closure (a symptom of leaf water deficit) has been observed in rice
growing in flooded soils (Dingkuhn et al 1990). However, water uptake and transport
by rice roots are most important and have been primarily studied in nonflooded soils,
especially as they affect yields under water-limited conditions, i.e., drought resistance.
Drought is not a single stress. Rather, as suggested by O’Toole and De Datta (1986),
it is a syndrome. The three major dimensions of the drought syndrome are the timing,
duration, and intensity of water deficit within the crop life cycle. Stress effects in each
of these dimensions result from a complex interaction between crop, soil, and
atmospheric factors.
Just as drought is a syndrome rather than a single stress, plant adaptations that
maximize yields under water-limited conditions are numerous and diverse. O’Toole
and Chang (1979) identified four components of drought resistance: tolerance,
avoidance, escape, and recovery. The second component, avoidance, can be subdi-
vided by its mechanisms—those that allow the plant to maintain a high internal water
status when available water is less than evaporated demand—into a) water conserva-
tion and b) water collection. Plant mechanisms for water conservation include leaf
rolling and stomatal closure; which tend to reduce growth and yield. Mechanisms for
water collection result in increased capacity for water uptake (e.g., through deeper
rooting or more efficient water extraction from soils); which should reduce yield
losses.
This paper focuses on water collection mechanisms in roots. We will describe recent
research on genetic variation and heritability of rice root traits as well as the
interrelationships among root characters, drought resistance, and water use.
Diameter/thickness
Water extraction
Maximum depth Japonica > indica
Deep root-shoot ratio
Xylem vessel diameter
NSH (%)
68 lngram et al
Geneticists have estimated narrow sense heritability from observations in F1, F2, or
F3 populations growing in either soil or aeroponic media (Table 2). For most traits,
inheritance was shown to be additive and polygenic. Overall, the root traits studied
show strong potential for improvement through breeding, with the greatest breeding
limitation being the large quantity of labor required for screening most root traits.
For much of the above research, genetic variation and heritability were estimated
from indica-japonica comparisons. Such a comparison ignores the fact that drought
occurs over widely varying rice environments, since indica rices are grown primarily
in the rainfed lowland ecosystem and japonica rices in the uplands. Thus, differences
in root traits between rice ecotypes may not be of practical use in breeding for drought
resistance. Focusing on the genetic variation and heritability within a single rice
ecotype instead might more quickly yield progress toward drought resistance. Simi-
larly, as we consider whether root traits can be transferred from wild Oryza species or
from one rice ecotype to another, we need to recognize that a trait that confers drought
resistance in one soil and rainfall environment may not be useful in another.
1. Total root length of wet seeded and transplanted rice through a plant and ratoon crop cycle.
Each point is the average of four cultivars. Vertical bars show significant differences by LSD at
P < 0.05. PI = panicle initiation, F = flowering, H = harvest.
Rice root traits for drought resistance and their genetic variation 69
2. Total root length of four rice cultivars through a plant and ratoon crop cycle. Each point is the
average of two establishment methods. Vertical bars show significant differences by LSD at
P < 0.05.
Table 3. Grain yields (av of 24 rainfed lowland rice genotypes) in response to vegetative water
deficit under transplanted and wet seeded crop establishment.
a Means followed by the same letter are not significantly different at P < 0.05 by LSD.
panicle. Similarly, the increase in TRL beginning about 15 d after panicle emergence
was probably associated with decreased competition for assimilates as grains reached
physiological maturity.
Although TRL changed greatly through the season, once roots had reached their
maximum depth, there were no significant changes in root distribution through the
profile (IRRI 199 1). Changes in TRL probably result mostly from death and initiation
of branch roots rather than from changes in primary root axes.
These results have important implications for both nutrient uptake and drought
resistance. Greater TRL and deeper roots may allow greater or more efficient nutrient
uptake in WSR than in TPR. These root system differences also explain differences in
drought resistance observed when TPR and WSR are subjected to water deficit. For 24
rainfed lowland rices, water deficit significantly reduced yields of TPR but not of WSR
(Table 3, IRRI 1988).
70 lngram et al
Root relationships with drought resistance and water use
When upland rice is grown in a range of soil moisture conditions, grain yields are
negatively related to RLD (Fig. 3). In contrast, Mambani and Lal (1983) reported that
for upland rice growing across a toposequence, grain yields are related to RLD. Several
researchers have reported that rice root growth under aeroponic or hydroponic culture
is well-correlated with root growth under upland field conditions (Ahmadi 1983,
Chang and Loresto 1986).
Unfortunately, root traits observed in aeroponic culture were not at all related to
drought resistance under rainfed lowland conditions (Ingram et al 1990). Also, neither
total RLD nor RLD by soil layer were significantly correlated with yields of 30 rainfed
lowland rices grown in a farmer’s field in Tarlac, Philippines (data not shown). The
lack of correlation between aeroponic and rainfed lowland field root growth under-
scores the need to consider the soil environment for which germplasm improvement
is targeted.
One hypothesis is that thick roots confer drought resistance because they have
greater xylem vessel radii, lower axial resistance to water flux, and hence, greater
capacity for water uptake from deeper soil layers (Yambao et al 1992). This hypothesis
is the opposite of that proposed for wheat grown on residual soil moisture—i.e., that
smaller root vessels confer drought resistance (Richards and Passioura 1989). In rice,
the axial resistance to water flux can be estimated by summing the individual
resistances for different root vessels, assuming that the vessels act as capillaries and
obey the Poiseuille-Hagan equation (Yambao et al 1992). Root box and hydroponic
research showed that differences in axial resistance to water flux explained shoot-water
3. Relationship between root length density and grain yield of upland rice.
Rice root traits for drought resistance and their genetic variation 71
4. For IR36, the response surfaces for relationships between equivalent root vessel radius,
transpiration, and leaf water potential as estimated by a) Poiseuille-Hagan equation or b) the
best-fitting curve. Variation in root vessel radius was achieved through root pruning in
hydroponic culture.
relations for a single cultivar (Fig. 4) but did not explain differences across cultivars
(Fig. 5). Clearly, factors other than axial resistance explain differences among cultivar
responses to water deficit.
Thick roots may also be hypothesized to confer drought resistance because
branching is directly related to root thickness (Fitter 1991). Thick roots persist longer,
produce more and larger branch roots, thereby increasing RLD and water uptake
capacity.
72 lngram et al
5. For 5 cultivars, the response surfaces for relationships between equivalent root vessel radius,
transpiration, and leaf water potential as estimated by a) Poiseuille-Hagan equation or b) the best-
fitting curve. Variation in root vessel radius was achieved through different cultivars and through
root pruning in hydroponic culture.
Rice root traits for drought resistance and their genetic variation 73
6. Total root length (0-70 cm soil layer) of IR46 and IR20 when continuously flooded or
nonflooded at 25-55 DAS throughout the full crop season.
conditions in the 10-30 cm soil layer, grading them high, medium, and low. We further
considered yield potential and effect of stress on maturity (Table 4).
The most important finding was that there was great genetic variation in root system
plasticity. In all genotypes, root length of stressed plants rapidly dropped below that
of continuously flooded plants when the plots were initially drained. There was also a
sharp dip in root length when stressed plants were reflooded (Fig. 6). After each dip,
genotypes with good root systems (e.g., IR46) produced greater root length in the
stressed than in the well-watered plots and had maximum RLDs greater than 0.5 cm
cm3 in the 20-30 cm soil layer. Genotypes with poor root systems (e.g., IR20) had
greater root length in the control than in the stressed plots and had RLDs less than 0.5
cm/cm 3.
The dynamic changes in root length are shown as the ratios of root length in stressed
and control treatments (Fig. 7). We hypothesize that these dynamic root responses to
74 lngram et al
Table 4. Root length and other characteristics related to drought resistance of 12 rice genotypes
under rainfed lowland conditions. a
a C = continuously flooded control: S = nonflooded 2555 DAS: +, ++ = less than 5 d and greater than 8 d delay
in maturity by stress, respectively.
7. Idealized relative root length density (root length stress divided by root length control) for
drought-resistant and drought-susceptible rice cultivars through a full crop season.
Rice root traits for drought resistance and their genetic variation 75
soil moisture are mostly a result of the shedding of old branch roots and the production
of new ones. According to our hypothesis, branch roots produced during stress are
adapted to aerobic conditions and have little or no aerenchyma; those produced after
reflooding are adapted to anaerobic conditions, with aerenchyma.
References cited
Ahmadi N (1983) Variabilite genetique at heredite do mechanismes de tolerance a la secheresse
chez le riz Oryza sativa L.: I. Development du system racinaire. Agron. Trop. 38:110-117.
Armenta-Soto J, Chang T T, Loresto G C, O’Toole J C (1983) Genetic analysis of root characters
in rice. SABRAO J. 15:103-116.
Chang T T, Loresto G C (1986) Genetic analysis of rice root systems by the aeroponic technique
and applications in plant breeding. Pages 123-220 in New frontiers in breeding research. B.
Napompeth and S.S. Abandhu, eds. Kasetsart University, Bangkok.
Chang T T, Loresto G C, O’Toole J C, Armenta-Soto J L (1982) Strategy and methodology of
breeding rice for drought-prone areas. Pages 217-244 in Drought resistance in crops with
emphasis on rice. International Rice Research Institute, P.O. Box 933, Manila, Philippines.
Dingkuhn M, Schnier H F, De Datta S K, Wijangco E J, Dorffling K (1990) Diurnal and
developmental changes in canopy gas exchanges as related to growth in transplanted and
direct seeded lowland rice. Aust. J. Plant Physiol. 17:119-134.
Ekanayake I J, Ganity D P, Masajo T M, O’Toole J C (1985a) Root pulling resistance in rice:
inheritance and association with drought resistance. Euphytica 34:905-913.
Ekanayake I J, O’Toole J C, Ganity D P, Masajo T M (1985b) Inheritance of root characters and
their relations to drought resistance in rice. Crop Sci. 25:927-933.
Fitter A H (1991) Characteristics and functions of root systems. Pages 3-25 in Plant roots, the
hidden half. Y. Waisel, A. Eshel, and U. Kafkafi, eds. M. Dekker, Inc., New York.
Ingram KT, Real J G, Maguling M A, Obien M A, Loresto G C (1990) Comparison of selection
indices to screen lowland rice for drought resistance. Euphytica 48:253-260.
IRRI—International Rice Research Institute (1988) Annual report for 1987. P.O. Box 933,
Manila, Philippines. 640 p.
76 lngram et al
IRRI—International Rice Research Institute (1991) Program report for 1990. P.O. Box 933,
Manila, Philippines. 317 p.
Mambani B, Lal R (1983) Response of upland rice varieties to drought stress. II. Screening rice
varieties by means of variable moisture along a toposequence. Plant Soil 73:73-94.
O’Toole J C, Bland W L (1987) Genotypic variation in crop plant root systems. Adv. Agron.
41:91-145.
O’Toole J C, Chang T T (1979) Drought resistance in cereals rice: a case study. Pages 373-405
in Stress physiology in crop plants. H. Mussel andR.C. Stables, eds. John Wiley & Sons, New
York.
O’Toole J C, De Datta S K (1986) Drought resistance in rainfed lowland rice. Pages 145-158 in
Progress in rainfed lowland rice research. International Rice Research Institute, P.O. Box
933, Manila, Philippines.
Richards R A, Passioura J B (1989) A breeding program to decrease the diameter of the major
xylem vessel in the seminal roots of wheat and its effect on grain yield in rainfed
environments. Aust. J. Agric. Res. 40:943-950.
Yambao E B, Ingram K T, Real J G (1992) Root xylem influence on the water relations and
drought resistance of rice. J. Exp. Bot. 43(252):925-932.
Notes
Authors’ addresses: K.T. Ingram, F.D. Bueno, O.S. Namuco, and E.B. Yambao, Agronomy,
Plant Physiology, and Agroecology Division, International Rice Research Institute, P.O. Box
933, Manila, Philippines; C.A. Beyrouty, Depatment of Agronomy, University of Arkansas,
Fayetteville, Arkansas 72701, USA.
Citation information: Kirk G J D, ed. (1994) Rice roots: nutrient and water use. International
Rice Research Institute, P.O. Box 933, Manila, Philippines.
Rice root traits for drought resistance and their genetic variation 77
Use of molecular markers
to exploit rice root traits
for drought tolerance
H.T. Nguyen, J.D. Ray, and Long-Xi Yu
Plants have various means by which they escape or resist periods of limited water
availability (drought) (Ludlow and Muchow 1990). In this paper, we consider the role
of the root system in the response of rice to drought and the use of molecular markers
as selection tools for incorporating root traits in improved rice cultivars.
80 Nguyen et al
rice genotypes had a greater penetration ability than lowland rice genotypes. Moreover,
root penetration ability was positively correlated with root thickness (unpubl. data).
Use of molecular markers to exploit rice root traits for drought tolerance 81
changes in the highly specific recognition site of the endonuclease; DNA remangements
resulting in the relocation of the site; or to insertion/deletion events within one of the
fragments (Botstein et al 1980; Beckmann and Soller 1983, 1986).
Botstein et al (1980) first proposed the use of RFLPs as genetic markers in humans.
By 1983, the potential applications of RFLPs in plants were being suggested (Burr et
al 1983, Soller and Beckmann 1983, Tanksley 1983), and by the mid-l980s, RFLP
linkage maps were published for maize and tomato (Bernatzky and Tanksley 1986,
Helentjaris et al 1986). In varying degrees of resolution, RFLP linkage maps are now
available for a large number of species including rice (McCouch et al 1988, McCouch
and Tanksley 1991).
High-resolution linkage maps increase the possibility of identifying markers linked
to traits of interest, including quantitative trait loci (QTL) (McCouch et al 1988, Knapp
et al 1990, McCouch and Tanksley 1991). The rice RFLP linkage map is presently of
sufficient resolution (about 1 cM, McCouch and Tanksley 1991) to permit mapping of
QTLs. Identification of RFLP markers linked to desirable quantitative root character-
istics would greatly facilitate their selection in breeding programs (Hanson et al 1990).
Identification of RFLP markers would also minimize the requirement for exhaustive
experiments to characterize root systems since REPS analyze the DNA directly and
are tissue-independent.
82 Nguyen et al
primers in constructing linkage maps, such as the dominant nature of RAPD markers
which prevents direct determination of heterozygosity (Devos and Gale 1992, Halward
et al 1992).
Conclusions
Incorporation of selection for root characteristics such as root penetration ability into
breeding programs offers the possibility of developing genotypes better able to exploit
soil water. Better exploitation of soil water reserves would allow plants to avoid the
adverse effects of periods of low rainfall. Phenotypic selection of root characteristics
in large breeding programs is difficult and requires too much time and labor, thus
limiting progress in developing improved genotypes. One solution is to use indirect
selection criteria based on the correlation between root characteristics and more readily
identifiable shoot phenotypes. However, this may not always be feasible or reliable. A
more direct approach is through the use of molecular genetic markers tightly linked to
the root characteristic of interest. The first step in using molecular markers in breeding
programs is to identify markers associated with desirable root characteristics. We have
begun this process in our laboratory with the tentative identification of four RFLP
molecular genetic markers associated with root penetration ability in rice (Texas
Technical University, unpubl. data). The identification of such markers will provide
breeders with better selection criteria for incorporation of desirable root characteristics
into improved cultivars.
Use of molecular markers to exploit rice root traits for drought tolerance 83
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Notes
Authors’ addresses: H.T. Nguyen, J.D. Ray, and Long-Xi Yu, Plant Molecular Genetics
Laboratory, Institute for Biotechnology and Department of Agronomy, Horticulture, and
Entomology, Texas Technological University, Box 42122, Lubbock, Texas 79409, USA.
Citation information: Kirk G J D, ed. (1994) Rice roots: nutrient and water use. International
Rice Research Institute, P.O. Box 933, Manila, Philippines.
86 Nguyen et al