Root plasticity and its implications for enhancing
crop productivity
It is seen from these research findings that rice roots
demonstrate significant morphological and physiological
variability under altered soil environments, whether due to
varying water regimes, soil microbial density, or different
planting patterns. In the nursery seedbed, the most important
factor that influenced root growth was the presence of
adequate water, but not so much water as to make the soil
hypoxic. In our study, oxygenated soil promoted development
of nodal roots (Fig. 1). Further, it was seen (as
detailed in Mishra and Salokhe 2008a) that root elongation
was not restricted under flooded soil conditions, but with
flooding, i.e., under hypoxic soil conditions, there was
restriction of root initiation which is important for developing
a greater number of nodal roots.
This might be due to the nitrogen status of the soil,
which has an influence on the rooting pattern. It is known
that a higher ratio of NH4
?:NO3
- favors greater cytokinine
production, which in turn helps support elongation of
roots, whereas a higher NO3
-:NH4
? ratio conversely
favors the production of auxins, which support root
initiation (Debi et al. 2005). That is why NO3
- is seen to
induce root development (Friend et al. 1990). It also
suggests that alternating flooding and wetting of paddy
soils could have a beneficial effect on whole-plant
growth.
The proliferation of lateral roots within nitrate-rich soil
patches has been shown by Zhang and Forde (1998) and
has been recently reconfirmed by Krouk et al. (2010). The
latter found that under high soil-nitrate conditions, a
transporter protein (NRT1.1) is compelled to transport
nitrate, thereby preventing auxin uptake. Auxin therefore
accumulates at the tip of the root, stimulating primordium
outgrowth, and this promotes lateral root growth. This
could be a reason why, under dry seedbed conditions
compared to wet seedbed, one finds a greater number of
nodal roots, with a higher ratio of highly branched nodal
roots (Fig. 1).
SRI methods achieved better nodal root development
than conventional methods at the initial growth stage when
soil nutrients were not a limiting factor. The same effect
was seen for shoot length and dry matter production,
indicating that a drained soil favors shoot and root development
when soil nutrients are not limiting. At the same
time, the seedbed study indicated that SRI advantages are
not seen under non-flooded water regimes when older
seedlings are used, especially when older seedlings are
transplanted into a flooded soil environment.
Farmers sometimes report difficulties in transplanting
very young seedlings as these are more difficult to handle.
However, once they realize that raising and transplanting
younger seedlings will have a positive impact on their
production and yield—with reduced water application—the
acceptability of SRI management should spread. Skill in
handling smaller young seedlings is something that can be
acquired with practice, and this skill could become
appreciated once farmers see the impact that it can have on
crop profitability.
It was seen in the second set of trials that under alternative
water regimes, the soil’s microbial density similarly
plays an important role in influencing root length density,
and also the sink capacity of the plant. This response is
affected by the nutrient status of the plant, the rate of
nutrient demand, and the soil’s fertility status.
With both treatments studied together—water regime
and soil microbial condition—the most important soil
factor influenced by both treatments was the soil nitrogen
status that in turn influenced root growth and physiology.
On a short time scale, soil microorganisms do compete
with plants for added nitrogen (N). This could account for
the faster senescence of plants under EMS treatment
compared to autoclaved and normal soil at the later growth
stage, i.e., during reproduction when demand for N is
higher.
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However, it is possible that after the initially rapid
capture of N and the assimilation of inorganic N by soil
microorganisms, the microbial population can reach a
steady state. One possible explanation for this could be that
the microbial population again becomes limited, by carbon
rather than by nitrogen. As the microbial population turns
over, there is insufficient carbon to maintain its fast growth,
and therefore N is released into the soil and becomes
available to plants. This will be possible on long time scale
only. In any case, it is important to have sufficient organic
matter in the soil to capitalize on the benefits associated
with soil microbial enhancement, either through mineralization
or immobilization.
The greater root length density under treatments with
enhanced microbial density at flowering (Table 3) indicates
that initially there was, relatively, a higher mineralization
rate with EMS treatment compared to other soil
conditions, and more mineralized N was present in EMtreated
soil. This root proliferation, supported through
microbial processes if maintained for a sufficient duration,
can have two ecologically relevant effects: it can enhance
N capture for the plant; and it can reduce N capture by
competitors. However, proliferation and prolongation of
roots depends on the relative costs and benefits to the plant.
This balance will depend on soil nutrient availability,
duration, and photosynthetic supply.
If supply of photosynthate to the roots, which comes
mostly from the lower leaves of the plant, is restricted,
or if the soil is limited in its nutrient availability and
roots are unable to supply nutrient to the above-ground
parts, the plasticity of response of plants’ roots—either
morphological proliferation or higher physiological
activity—will be a burden for the plant. Ultimately, the
cost to the plant will depend on what is actually limiting
its growth, whether nutrients or photosynthate supply.
This is evident from the relationships presented in
Figs. 3 and 4, where:
• Duration of grain filling is significantly correlated to the
chlorophyll content of the flag leaf;
• The chlorophyll content of the lower leaves is significantly
correlated to root oxidizing activity rate; and
• The root oxidizing activity rate is correlated to soilavailable
nitrogen.
If the soil has limited N availability, decomposition of
chloroplasts will take place in the lower leaves to supply N
to the flag leaf. Consequently, the photosynthetic ability of
the lower leaves will decrease, and they are less able to
supply sugar and oxygen to the roots. Therefore, to capitalize
on the benefits associated with enhanced soil
microbial density, and to insure both nutrient availability
and in turn photosynthate supply to the roots for longer
duration, application of organic matter (as much as
possible) is crucial. This is why SRI is consistent with the
recommendation to farmers: feed the soil, so that the soil in
turn can feed the plant, rather than focus solely on feeding
the plant directly.
Another factor that would limit the benefits of root
proliferation is shading of leaves. If shading simply
reduced the nutrient demand by the plant due to reduced
photosynthetic capacity of its leaves, nutrient capture
would be lower, irrespective of higher root length density.
In that case, root proliferation would be a costly affair for
the plant, and the plant would try to reduce this cost, so as
to come closer to an optimizing balance between supply
and demand.
One should not be surprised to see, therefore, that with
the crowding of plants in hills or of hills together, root
degeneration is faster compared to sparser planting, even
under alternative water regimes where oxygen is not a
limiting factor (Fig. 5). Hence, it is important to maintain
the source–sink demand simultaneously to realize the
benefits of adaptive root response.
As evident from the results shown in Fig. 5a and b,
reduced intra-hill competition and wider spacing favored
the development of more lateral roots. More laterals
helped the root to achieve a greater surface-absorption
area, with higher cation exchange capacity favoring
higher nutrient absorption from the soil, and hence higher
yield. This was the reason for the P2 planting pattern
realizing a higher yield compared to P3, which had many
more plants m-2.
To realize the potential benefits in terms of crop yield
per hectare, spacing is a factor that always needs to be
optimized, not necessarily maximized or minimized.
Depending on soil and varietal characteristics such as
maturity period, tillering ability, and photoperiod sensitivity,
location- and variety-specific practices need to be
identified empirically, integrating the effects of many factors,
such as those examined above which affect above- and
below-ground dynamics.
Further agronomic research evaluating SRI practices,
both respectively, and in combination, should be undertaken
for achieving better yield and profitability. This
applies to fine-tuning of water management regimes as well
as optimizing the age of seedlings, inter-row and inter-plant
spacing, weeding frequency, etc., weighing the costs and
benefits of the respective practices. Such adaptations and
adjustments are considered as part of the principles and
philosophy of SRI management (Stoop et al. 2002).
However, the practical constraints for making adaptations
and adjustments for achieving better soil–plant–water and
environmental relationships should not be underestimated.
These constraints can be minimized if SRI is regarded and
used as a heuristic vehicle for linking rice research with
farmers’ participatory research programmes.
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