Waterlogging and its impact on wheat

Ashwani Kumar1, Rajesh Kumar1, Jogendra Singh1, Pooja2 and Vijayata Singh1
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Central Soil Salinity Research Institute, Karnal-132001(Haryana)

Sugarcane 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

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 of rice (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
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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, the tolerant 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

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 soil waterlogging 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 O 2 supply to the apex of roots that solely depends on aerenchymatous O 2 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 ROL in 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). and, therefore, would decrease the root growth in hypoxic or anoxic environment (Armstrong, 1979; Jackson and

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 ROS are 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 carbon dioxide 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.

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 and faba 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

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 30 cm 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
Armstrong J, Armstrong W. (2001). An overview of the effects of phytotoxins on Phragmites australis in relation to die-back. Aquatic Botany. 69, 251-268. Fried A, Smith N. (1992) Soil structure deficiency in extensive croplands of northern Victoria. Land Degradation Study Group. Soil and Water Group Association of Victoria. Gill KS, Qadar A, Singh KN. (1992). Effect of wheat ( Triticum aestivum) genotypes to sodicity in association with waterlogging at different stages of growth. Indian Journal of Agricultural Sciences. 62, 124-128. Greenway H, Gibbs J. (2003). Mechanisms of anoxia tolerance in plants. II. Energy requirements for maintenance and energy distribution to essential processes. Functional Plant Biology 30, 999- 1036. Gupta VK. (2004). Soil analysis for available micronutrients. In Methods of Analysis of Soils, Plants, Watersand Fertilizers. (Ed HLS Tondon) pp. 36-48. (Fertilizer Development and Consultation Organization:New Delhi). Hamilton G, Bakker D, Houlebrook D, Spann C. (2000). Raised beds prevent waterlogging and increaseproductivity. Western Australia Journal of Agriculture. 41, 3-9. Lindsay WL, Norvell WA. (1978). Development of a DTPA soil test for zinc, iron, manganese and copper. Soil Science Society of America Journal. 42, 421-428. McFarlane D. (1990). Agricultural waterlogging A major cause of poor plant growth and land degradation in Western Australia. Land and Water Research News 7, 5-11. National Remote Sensing Agency (NRSA) and Associate (1996) Mapping salt affected soils of India, 1:250,000 mapsheets, Legend. (NRSA: Hyderabad, India). Patrick WH. (1964). Extractable iron and phosphorus in a submerged soil at controlled redox potentials. In: Trans. 8th Int. Congr. Soil Sci., Bucharest, Romania. pp. 605-609. (Acad. Socialist Rep. Romania).
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Silberstein RP, Vertessy RA, Morris JD, Feikema PM. (1999b). Modelling the effects of soil moisture and solute conditions on long-term tree growth and water use: a case study from the Shepparton irrigation area, Australia. Agric. Water Manage. 39, 283315. Vertessy RA, Hatton TJ, OShaughnessy PJ, Jayasuriya MDA. (1993). Predicting water yield from a mountain ash forest catchment using a terrain analysis-based catchment model. J. Hydrol. 150, 665 700. Vertessy RA, Hatton TJ, Benyon RG, Dawes WR. (1996). Long-term growth and water balance predictions for a mountain ash (Eucalyptus regnans) forest catchment subject to clear-felling and regeneration. Tree Phys. 16, 221232. Ward PR, Dunin FX, Micin SF. (2001). Water balance of annual and perennial pastures on a duplex soil in a Mediterranean environment. Aust. J. Agric. Res. 52, 203209. Zaslavsky D, Sinai G. (1981a). Surface hydrology. III. Causes of lateral flow. J. Hydraulics Division, ASCE. 107, 3752. Zaslavsky D, Sinai G. (1981b). Surface hydrology. IV. Flow in sloping layered soil. J. Hydraulics Division, ASCE. 107, 5364.

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