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Waterlogging and its impact on wheat
Ashwani Kumar1, Rajesh Kumar1, Jogendra Singh1, Pooja2 and Vijayata Singh1
1Central Soil Salinity Research Institute, Karnal-132001(Haryana)
2Sugarcane Breeding Institute, Regional Station, Karnal-132001(Haryana)
Waterlogging is a widespread problem for wheat production, especially in the
sodic/alkaline soils of India. Waterlogging is a well known problem in alkali and saline alkali soil
of wheat growing area of Indo-Gangetic plain of India. Around 3.77 mha of sodic soil along with
2.96 mha area adjoin to seepage fed area under extensively circulated irrigation canal network
are severely affected by undesirable waterlogging during wheat season and claim drastic yield
harvesting potential (Yaduvanshi et al., 2012).
Waterlogging is caused by the same processes as dry land salinity. The difference is that
salts do not accumulate at the soil surface, either because ground water is of very low salinity
(less than 6 dS/m) or it flows out of the soil (i.e. from a small spring), flushing the salts away.
Waterlogging problems can also be ephemeral, such as when a perched water table develops on
impermeable sub-soil in a wet season. Most of the information given in this chapter is also
directly relevant to management of waterlogging, except the references to salt accumulation and
its effects. Thus waterlogging will generally not be discussed separately.
Fig. 1: Farmers field affected by salinity, sodicity and waterlogging in India
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In agriculture, various crops need air(specifically, oxygen) to a greater or lesser depth in
the soil. Waterlogging of the soil stops air getting in. How near the water table must be to the
surface for the ground to be classed as waterlogged varies with the purpose in view. A crop's
demand for freedom from waterlogging may vary between seasons of the year, as with the
growing ofrice (Oryza sativa).
In irrigated agricultural land, waterlogging is often accompanied by soil salinity as
waterlogged soils prevent leaching of the salts imported by the irrigation water. From
a gardening point of view, waterlogging is the process whereby the soil blocks off all water and is
so hard it stops air getting in and it stops oxygen from getting in.
Hypoxia and anoxia
Hypoxia or oxygen depletion is a phenomenon that occurs in soil environments as
oxygen in soil air becomes reduced to a point below optimum level. In plant physiological
studies, the term hypoxia is reserved for situations in which the oxygen concentration is a
limiting factor (Morard and Silvestre, 1996). It is usual form of stress in soil that experiences
long-term flooding or waterlogging. It occurs in plants completely submerged by water, and
in deep roots below flood waters (Sairam et al., 2008).
How does waterlogging induce hypoxia and anoxia?
One of the most important properties of soil is soil aeration which relates to the ability of
soils to exchange gases with the atmosphere. This exchange is usually achieved primarily
through diffusion of gasses from and to the soil via pore spaces in the soil. In most well drained
soils, the air-filled pore spaces make up 10 to 40% of total soil volume. Waterlogging eliminates
these gas-filled pore spaces and cuts the supply of oxygen to the roots to a large extent
(Ponnamperuma, 1972). In the waterlogged soil, micro channels for gas diffusion among soil
particles or aggregations become sealed with water, which results the gas diffusivity in soil
104 times lower than in well-drained soil (Armstrong, 1979; Ponnamperuma, 1984).
The lower gas diffusivity between ambient air and waterlogged soil results in low O2
concentration (hypoxia) and high toxic gas concentration, such as CO2 and reduced gases
(Ponnamperuma, 1972, 1984). Moreover, gases formed by soil metabolism, including carbon
dioxide, start to accumulate near root surfaces (Setter and Belford, 1990). The gas exchange
http://en.wikipedia.org/wiki/Crophttp://en.wikipedia.org/wiki/Earth's_atmospherehttp://en.wikipedia.org/wiki/Oxygenhttp://en.wikipedia.org/wiki/Ricehttp://en.wikipedia.org/wiki/Irrigationhttp://en.wikipedia.org/wiki/Soil_salinityhttp://en.wikipedia.org/wiki/Soil_salinity_controlhttp://en.wikipedia.org/wiki/Sodium_chloridehttp://en.wikipedia.org/wiki/Gardeninghttp://en.wikipedia.org/wiki/Crophttp://en.wikipedia.org/wiki/Earth's_atmospherehttp://en.wikipedia.org/wiki/Oxygenhttp://en.wikipedia.org/wiki/Ricehttp://en.wikipedia.org/wiki/Irrigationhttp://en.wikipedia.org/wiki/Soil_salinityhttp://en.wikipedia.org/wiki/Soil_salinity_controlhttp://en.wikipedia.org/wiki/Sodium_chloridehttp://en.wikipedia.org/wiki/Gardening -
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between soil and atmosphere almost stops as soon as the waterlogging sets in. The soil microbes
and plant roots use up the oxygen trapped in the soil and therefore, the roots may become
exposed to complete lack of oxygen (anoxia) (Jackson and Drew, 1984). However, under natural
conditions, oxygen concentration decreases gradually, and hence, anoxia is always preceded by
hypoxia (Setter and Waters, 2003) in waterlogged environment.
Mechanisms of tolerance of wheat to waterlogging
Waterlogging tolerance is defined as the survival or the maintenance of plant growth at
high rates under waterlogged conditions relative to well drained conditions. It may be defined as
the maintenance of relatively high grain yields under waterlogged conditions relative to non- or
less-waterlogged conditions (Setter and Waters, 2003).
Morphological and Metabolic adaptation
A. Morphological adaptation
(i) Root growth
A common adaptation of plants to waterlogging is the survival and growth of seminal roots
and production of numerous adventitious roots with aerenchyma. The root growth in
waterlogging intolerant genotypes is drastically suppressed by waterlogging stress. However, thetolerant genotypes have the ability to continue their root growth under the stress in some extent.
The above hypoxic stress had no significant effect on the growth of seminal roots for tolerant
genotypes (Gore and Savannah). Total root dry mass was reduced for all genotypes except for
Savannah (Huang et al., 1994a). However, the waterlogging tolerance of a plant is determined
not only by its capability to undergo morphological adaptations, but also by the ability to recover
from transient waterlogging or hypoxia of the root system (Krizek, 1982; Huang et al., 1994a,
1997).
(ii) Aerenchyma formation and increased root porosity
Aerenchyma is a special tissue which consists of continuous gas filled channels or much
enlarged gas spaces, and root porosity is volume of gas-filled spaces in relation to the total tissue
volume. Aerenchyma provides a low resistance internal pathway for the movement of O2 from
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the shoots to the roots (Armstrong, 1979) Armstrong and Webb, 1985; Drew et al., 1985).
Aerenchyma tissue in roots allows the roots to respire aerobically and to maintain growth under
hypoxic conditions. Moreover, a part of oxygen transported to plant root tips through the
aerenchyma leaks out into the surrounding soil and results in a small zone of oxygenated soil
around the roots providing an aerobic environment for microorganisms that can prevent the
influx of potentially toxic soil components such as nitrites and sulphides of Fe, Cu and Mn.
Therefore, aerenchyma formation is thought to be one of the most important morphological
adaptations for the tolerance to hypoxic or anoxic stress. Under oxygen deficient condition,
ethylene production is accelerated which in turn stimulates aerenchyma formation in adventitious
roots and induces the growth of the roots (Drew et al., 1979; Jackson, 1989).
However, the conversion of ACC to ethylene requires oxygen and the conversion
reaction is blocked in an anaerobic root cell. The ACC is therefore, translocated from the
anaerobic root cells towards the more aerobic portions of the root or to the shoot. The lower
portions of the stems are usually the site of highest ACC accumulation and in the
presence of oxygen ethylene is released (Sairam et al., 2008). Formation of aerenchyma has
been observed in the roots of wheat when grown under low O2 concentrations.
Fig. 2: Main physico-chemical events taking place in the rhizosphere during soilwaterlogging and the resulting modifications in plant metabolism and physiology followed
by the initiation of adaptive responses.
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(iii) Barriers to radial oxygen loss
Oxygen in aerenchymatous roots may be consumed by respiration or be lost to the
rhizosphere via radial diffusion from the root. The flux of oxygen from roots to rhizosphere is
termed as radial oxygen loss (ROL) which usually oxygenates the rhizosphere of the plants
growing in waterlogged soils (Armstrong, 1979). However, ROL decreases the amount of O2
supply to the apex of roots that solely depends on aerenchymatous O 2 and, therefore, would
decrease the root growth in hypoxic or anoxic environment (Armstrong, 1979; Jackson and
Drew, 1984).
It is suggested that the loss in internal O2 contributes the poor growth of adventitious roots
and intolerance of wheat to waterlogged soil (Thomson et al., 1992). In contrary, waterlogging
tolerant rice not only has a larger volume of aerenchyma, but it also has a strong barrier to ROLin basal regions of its adventitious roots and therefore deeper root penetration into waterlogged
soil. However, some wheat genotypes can increase suberin or lignin on epidermis or exodermis
of root which may acts as barriers to ROL and results in increased tolerance to waterlogging.
B. Metabolic adaptation
The plant tissue under hypoxia or anoxia suffers from energy crisis (Gibbs and Greenway,
2003) due to reduced root respiration in both waterlogging-tolerant and intolerant plants
(Marshall et al., 1973; Lambers, 1976; Drew 1983, 1990). The tolerant plant species cope with
the energy crisis through metabolic adaptation to oxygen deficiency. The metabolic adaptations
to oxygen deficiency includes: anaerobic respiration, maintenance of carbohydrate supply for
anaerobic respiration, avoidance of cytoplasmic acidification and development of anti-oxidative
defense system
Anaerobic respiration
Plant cells produce energy in presence of oxygen through aerobic respiration which
includes glycolysis, TCA or Krebs cycle and oxidative phosphorylation (Fig. 1). In absence of
oxygen (under anoxic condition), Krebs cycle and oxidative phosphorylation are blocked, and
cells inevitably undergo anaerobic respiration to fulfill the demand for energy (Davies, 1980).
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Mechanisms of tolerance of wheat to waterlogging
Exposure of plants to most adverse conditions like hypoxia or anoxia causes oxidative
stress, which affects plant growth due to the production of reactive oxygen species (ROS) such as
superoxide radicals, hydroxyl radicals and hydrogen peroxide (Mittler et al., 2004). These ROSare very reactive and cause severe damage to membranes, DNA and proteins (Bowler et al.,
1992; Foyer et al., 1997). Hypoxia stress triggers the formation of ROS and induces oxidative
stress in plants.
The tolerance to the stress may be improved by increased antioxidant capacity. Many
recent attempts to improve stress tolerance in plants have been made by introducing and
expressing genes encoding enzymes involved in the antioxidative defense system. Moreover, the
end-products of glycolytic and fermentative pathway, such as ethanol, lactic acid and carbondioxide pose an additional hazard to the cell. It is well reported that the maintenance of an active
glycolysis and an induction of fermentative metabolism are adaptive mechanisms for plant
tolerance. This induction can improve or at least sustain the glycolytic rate in anoxic plants
contributing higher tolerance to anoxia.
Increased availability of soluble sugars
Due to shifting of energy metabolism from aerobic to anaerobic mode under hypoxia or
anoxia the energy requirements of the tissue is greatly restricted as very few ATPs are generated
per molecule of glucose. A high level of anaerobic metabolism in hypoxic or anoxic roots is
therefore very important to supply the energy charge high enough which can sustain metabolism
in roots for the survival of plants (Jackson and Drew, 1984). The roots of comparatively tolerant
genotypes contain greater sugar content (total, reducing and non-reducing sugar) than
in susceptible genotypes of wheat.
Adverse effect of water logging
The adverse effects of waterlogging on wheat are due to decrease availability of oxygen
and accumulation of phytotoxins, severe energy deficiency and ultimately death of plant. The aim
of this study was to evaluate the effect of flooding in field and micro plots of wheat growth stage
under alkali soil and normal soil.
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Depletion of oxygen in root zone and increase of CO 2 due to water logging. An aerobic
condition adversely affects micro-organisms while harmful organisms proliferate and restrict the
plant growth. Physical or chemical and biological activities in the soil are disturbed due to low
temp as a result of water logging. Thus pest and diseases infestation problem arises. Water
logging makes field operations difficult on impossible. The adverse effects of water logging get
accelerated when the capillary water brings salts from lower horizon of soil or they are present in
the ground water used for irrigation. Water logging adversely affect the soil water plant
relationship there by creating ecological imbalance. Crops yields reduced and some time crop
failure due to inadequate uptake of moisture and nutrients and due to the injurious effect of salts or
deteriorated soil condition. Fodders grown in slat-affected soils may contain high molybdenum in
or selenium and low amount of zinc. The nutritional imbalance may cause disease in live stock.
The adverse effects of waterlogging on plants are often ascribed to decreased availability
of O2 and accumulation of phytotoxins (Armstrong and Armstrong 2001). Oxygen deficiency
inhibits aerobic respiration, resulting in severe energy deficiency and eventually death (Greenway
and Gibbs 2003). In addition, waterlogging can also reduce the availability of some essential
nutrients, e.g. nitrogen, and increase the availability of nutrients, e.g. Fe and Mn (Ponnamperuma
1972). Such increases in micronutrients in soil and subsequently in shoots may affect plants both
during waterlogging and also after waterlogging during recovery, as higher micronutrient
concentrations in shoots have been reported during recovery period when soils have returned to
fully aerated conditions (Setter and Waters 2003).
Management options to control waterlogging in crops
1. Drainage is usually the best means of managing waterlogging. Other management options
include: choice of crop, seeding, fertilizer, weed and disease control. Typically, with
changes to crop rotations and management, major costs would include cost of buying seed
and extra fertilizer, and the costs of weed and insect control. Some species of grains are
more tolerant than others. Grain legumes and canola are generally more susceptible to
waterlogging than cereals andfaba beans.
2. Seeding crops early and using long-season varieties help to avoid crop damage from
waterlogging. Crop damage is particularly severe if plants are waterlogged between
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germination and emergence. Increase sowing rates in areas susceptible to waterlogging to
give some insurance against uneven germination, and to reduce the dependence of cereal
crops on tillering to produce grain. Waterlogging depresses tillering. High sowing rates
will also increase the competitiveness of the crop against weeds, which take advantage of
stressed crops.
3. Crops tolerate waterlogging better with a good nitrogen status before waterlogging occurs.
Applying nitrogen at the end of a waterlogging period can be an advantage if nitrogen was
applied at or shortly after seeding has been lost by leaching or denitrification. However,
nitrogen cannot usually be applied from vehicles when soils are wet, so consider aerial
applications.
4. If waterlogging is moderate (730 days waterlogging to the soil surface), then nitrogen
application after waterlogging events when the crop is actively growing is recommended
where basal nitrogen applications were 050 kg N/ha. However, if waterlogging is severe
(greater than 30 days to the soil surface), then the benefits of nitrogen application after
waterlogging are questionable. Weed density affect a crop's ability to recover from
waterlogging. Weeds compete for water and the small amount of remaining nitrogen,
hence the waterlogged parts of a paddock are often weedy.
5. Root diseases, particularly take-all of wheat and barley, are often more severe in
waterlogged crops because the pathogens tolerate waterlogging and low oxygen levels
better than the crops.
Concentrations of oxygen rapidly decreased after the commencement of waterlogging, but
increased again after drainage. If waterlogging is suspected in your crop:
Dig shallow holes 30 to 40 cm deep in winter to see whether any free water is within 30cm of the surface (if so, the soil is waterlogged).
Observe where soils are boggy and crops are yellow.
Mark out the areas that are affected, either with posts laid on the ground or on an accurate
map.
At harvest time observe where the crops are poor and check this against earlier
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Observations.
Survey seepage interceptor drains, placing them immediately above affected sites or
consider raised beds.
Install drains when the soils are moist (for example, after summer or autumn rains).
If installing raised beds, seek further advice from DAFWA or other professionals on
placement and design.
Reference
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