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��ࡱ�>��	|����yz{��������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������[�	��JIbjbj����	4RΐΐO4�������aaaaa����uuu8�dlu�a�}�C(kkk�.� \L!0�]�]�]�]�]�]�]$�c�Uf>�]�a|!��|!|!�]aakkW�a"K"K"K|!�akak�]"K|!�]"K"K��Th�Uk����@�"��uh<�	�T�]�a0�aU��fbF��f �U�U0�fa�V�|!|!"K|!|!|!|!|!�]�]"K|!|!|!�a|!|!|!|!���������������������������������������������������������������������f|!|!|!|!|!|!|!|!|!	':	Effect of spacing and water control on the yield potential of rice in the system of rice intensification: A review
Joseph Xorse Kugbe1, 2, Daniel Asomning Odoom2, 3, Philip Ghanney2, Ebenezer Kofi Sam2, Sherif Mohammed Abubakari2, Albert Berjour 2, Karimu Musah2 
1Center for Development Research (ZEF), Division of Ecology and Natural Resources Management, University of Bonn, Walter Flex Str. 3, D53113, Bonn, Germany
2Faculty of Agriculture, University for Development Studies, Tamale-Ghana
3Ministry of Food and Agriculture, Tamale- Ghana
*Corresponding Author: Daniel Asomning Odoom:  HYPERLINK "mailto:odoom24@gmail.com" odoom24@gmail.com +233209353907
Abstract
This review paper examines the effects of spacing, water control and rice root dry matter on the yield potential of rice in the System of Rice Intensification (SRI). A range of practices and results are reviewed from field trials of SRI and based on the reports from groups, institutions and individuals who have tried SRI and on personal interviews with SRI practitioners and researchers. Sumant Kumar set a new world record in rice production of 22.4 tons per hectare using SRI, breaking the existing world record held by the Chinese scientist HYPERLINK "http://en.wikipedia.org/wiki/Yuan_Longping"Yuan HYPERLINK "http://en.wikipedia.org/wiki/Yuan_Longping"Longping by 3 tons. Aside from increased in yield gained, the SRI results in phenotypically distinct plants, which produce 30, 50 or even 70 tillers with correspondingly vigorous root growth. The experiences with SRI methods suggest that average rice yields can be about twice the present world average without requiring a change in cultivar or the use of purchased inputs. 

Keywords: Rice yield, Cultural practices, Water use efficiency, System of rice intensification 
Introduction 
Rice yield, obtained in the System of Rice Intensification (SRI) method is similar to the yields obtained under conventional systems although the SRI requires 50 percent less water. This is an important feature, since the water table depletion has become a global phenomenon, which is sending alarming signals to rice growing countries. Rice crop is being traditionally cultivated under continuous submerged conditions; requiring the need for alternative cultivation methods in order to minimize the water requirement of rice crops. The SRI cultivation method offers a wonderful approach to minimize water consumption for rice cultivation and to increase the productivity (Laulanie, 1993). 
A study involving simultaneous evaluation of root characteristics and grain yield, concluded that genotypes with a deep rooting habit had an advantage in stress conditions and also, genotypes that produce deep roots prior to the onset of stress shows improved productivity compared with genotypes that do not have the capacity to produce roots prior to the onset of stress (Kanbar et al., 2009). The study also suggested, based on Quantitative Trait Loci (QTL) mapping, that the loci for productivity are not congruent with those related to root morphology, except at one locus. Subsequently, studies conducted under contrasting moisture regimes, suggested that maximum root depth, root: shoot ratio, and root dry weight confers an advantage to grain yield under water stress (Kanbar et al., 2009).
Rice cultivation has two major land management systems, commonly referred to as upland and lowland. These two systems differ greatly in their yield potentials because of soil characteristics that affect root growth and plant response to drought.
To ensure food security in the rice-consuming countries of the world, these countries will have to produce 50% more paddy with improved quality to meet the demands of the consumers by 2025 (FAO, 2010). 
This additional rice will have to be produced on less land with less water, less labour, and little chemicals (Yonglu, 2012). Aside these perceived problems of future rice production, rice quality preferences are gradually receiving more attention. Crop improvement and management practices have played an important role in increasing the production of rice in the past. The System of Rice Intensification (SRI), developed in Madagascar over a 20-year period and synthesized in the early 1980s, offers opportunities to researchers and farmers to expand their understanding of potentials already existing in the rice genome (Uphoff et al., 2002). 
Experience with SRI methods suggests that average rice yields can be about twice the present world average without requiring a change in cultivars or the use of purchased inputs (Wang et al., 2002). Moreover, only about half as much water per season is required for these higher yields. Crop protection requirements are reduced because plants in the SRI are more resistant to damage by pests and diseases (Wang et al., 2002). To achieve a super high yield, the SRI methodology for rice production makes three main changes to irrigated rice cultivation: transplanting younger seedlings, preferably 8-15 days old before the plants enter their fourth phyllochron of growth, planting the seedlings singly rather than in clumps of 3-6 plants, and keeping the paddy soil moist but not continuously saturated during the plant�s vegetative growth phase (Uphoss et al., 2003).
It has become difficult to increase rice production by the traditional rice farming system. It requires extra labour and a lot of compost. Farming with modern methods is also expensive since very costly external inputs are required. With the conventional methods of rice production, it is only by the use of expensive inorganic fertilisers, pesticides and hybrid seeds that the farmer increases production. Nitrogen use efficiency is very low on average in conventional agricultural systems (Raun, 1999). This demonstrates a vast inefficiency in production. It is increasingly becoming more difficult for the ordinary resource poor farmer, particularly in Africa to afford all the inputs required for conventional rice production (FAO, 2010). Meanwhile, the chemicals used in such conventional systems are known to impact negatively on human health and the environment (Katherine and Hendrik, 2010).
Farmers in more than 20 countries are now using the SRI to raise their rice production, at a reduced negative effect on human health and the environment, while also reducing the use of external inputs and production costs. The SRI works by changing the management of the plants, soil, water and nutrients that are utilized in paddy rice production. Transplanting single young seedlings with wider spacing, carefully and quickly into fields that are not kept continuously flooded, and whose soil has more organic matter and is actively aerated (Randriamiharisoa et al., 2006). These practices improve the growth and functioning of rice plants, the root system and enhance the numbers and diversity of the soil biota that contribute to soil health and productivity (Mishra et al., 2006; Randriamiharisoa et al., 2006).
The SRI inculcates the principle of synergy that is derived through the �border-edge effect� in agronomic research, where plants that grow on the borders of experimental fields generally perform better and give more yield than plants located in the middle of the field. The high performance is attributed to prolonged exposure to sunlight and adequate air circulation (Gomez et al., 1971). By introducing optimally wider spacing, therefore, SRI methods achieve the �border-edge effect� for the whole field. Sato et al., (1983) reported, for example, that rice plants in border rows can give 45-49% more yield than those in the centre of a field. Besides rice, external rows of wheat and barley are also reported to produce 40% more grain than is harvested from the innermost rows (Romani et al., 1993).

Based on research and observed phenomena on paddy fields (Sato et al., 1983; Romani et al., 1993; Wang et al., 2002; Mishra et al., 2006; Randriamiharisoa et al., 2006), the basic concepts of SRI can be summarized as:
Use of young seedlings to preserve the growth potential of mature plants. 
Avoidance of trauma to the roots. Transplanting young seedlings as quickly as possible and in shallow waters (1-2 cm), with no inversion of seedling root tips that will delay the plants resumption of growth after transplanting. 
Giving plants optimally wider spacing � one plant per hill and in square pattern so as to achieve �the border-edge effect� for the whole field. 
Keeping the paddy soil sufficiently moist but not continuously flooded, mostly aerobic and not saturated. 
Actively aerating the soil as much as possible, using a rotary hoe or weeder to control weeds. 
Enhancement of soil organic matter as much as possible by the application of compost, mulch, manure, etc. Chemical fertilizers can also be used with SRI. The best results have, however, been observed with organic soil amendments.
 
The first three practices stimulate plant growth, while the latter purposefully enhance the growth and health of plant roots and soil biota. For optimum yield, some beneficial practices are recommended for use with SRI. These include the selection of most suitable varieties to grow under the prevailing moisture, soil, diseases and environmental conditions, selection of best seeds within the varieties, practice of seed priming and seedbed solarisation (Culman et al., 2005).

SRI impacts root morphological traits and affects rice yield
 Growth of the rice root; in terms of total dry matter, maximum root depth, and root length density, increases until the �owering stage and then decreases sharply to maturity (Yoshida et al., 1982). Kawata et al., (1974) indicated that roots produced after �owering may play an important role during the grain-�lling period. The shape of the root system differs greatly with soil texture, soil water status, and soil compaction (Hoshikawa, 1989). 
Rice is characterized by a shallow root system compared with other cereal crops (Angus et al., 1983), having limited water extraction below 60 cm (Fukai et al., 1988). The form of the rice root system also varies with cultivation methods (Yoshida and Hasegawa, 1982; Tuong et al., 2002). In upland conditions with direct sowing, the root system generally develops deeper than in transplanted plantings in lowland conditions.
The allocation of resources from the root is high at early vegetative stages but decreases markedly at �owering and is almost negligible after anthesis (Gregory et al., 1996). 
In their adaptation to growth in �ooded conditions, rice roots display a unique formation of apoplastic barriers compared with other crop plants. The role of suberin and the casparian band in limiting water uptake has received mixed reports. In drought-stressed lowland rice, the implications of apoplastic root barrier formation are complex since the soil at the start of the season is �ooded, then �uctuates or steadily decreases based on rainfall patterns. Hose et al. (2001) concluded that the extent and rate of casparian band and suberin lamella formation depend on environmental conditions (drought, hypoxia, salt, heavy metal, and nutrient availability). Typically, a greater degree of casparian band and suberin lamella development results in less water permeability of the root. Formation of an exodermis in rice is induced by growth in stagnant solution, and results in an effective barrier to radial oxygen loss in rice (Colmer et al., 1998).
Root water uptake ability and hydraulic conductance have been evaluated according to root pressure, determined by root system xylem sap �ux through cut stems. Xylem sap �ux of lowland varieties decreased with decreasing water availability in a water-saving trial, but did not change in two upland varieties (Beodin et al., 2009; Matsuo et al., 2009b). 
Chang et al., (1972) compared root traits of several upland and lowland varieties and found that drought resistance was associated with coarse, long roots, a dense root system, and a high root to shoot ratio. Yoshida et al., (1982) also reported genetic variation in root depth, with a tendency for upland rice cultivars to have deeper roots than lowland rice cultivars. Ingram et al., (1994) used cultivars belonging to different types of rice for root studies and found tropical japonica types to have larger root systems than indica types. In another study, La�tte et al., (2001) investigated the genotypic variation for root traits in different types of rice and reported that indica rice types had �ne, highly branched super�cial roots with narrow xylem vessels and low root to shoot ratio, whereas japonica types had coarse roots with wider vessels, less branched long roots, and a large root to shoot ratio. The large root to shoot ratio contributes to the comparatively higher yield observed in these rice varieties.
In rice, some mutants without seminal roots or without lateral roots have been identi�ed. Ten genes have been identi�ed in rice relating to root growth, out of which mutants of six genes, rm1 and rm2 (Ichii et al., 1997), rrl1 and rrl2 (Inukai et al., 2001a), srt5 (Yao et al., 2002), and srt6 (Yao et al., 2003), are involved in a reduction in seminal root elongation. Mutants of two genes, crl1 and crl2 (Inukai et al., 2001b), are involved in a reduction in number of nodal roots, one (rm109; Hao et al., 1999) is involved in blocking lateral root initiation, and one (rh2; Ichii et al., 2000) is involved in reducing root hairs. Other mutants include crl4 (Kitomi et al., 2008), lrt1 (lateral rootless phenotype; Chhun et al., 2003a), arm1 and arm2 (lateral root number decreases; Chhun et al., 2003b), and lrt2 (lateral rootless phenotype; Wang et al., 2006b). Most of these genes have not been identi�ed: except for crl1 (Inukai et al., 2008).
The relationships between root growth and grain yield under drought are complex. Positive associations between root length and grain yield have been documented in rice (Lilley et al., 1994). In contrast, Ingram et al., (1994) found no signi�cant association between the two traits. It may be that a simple correlation between root growth and yield could be expected only in well-de�ned target environments (Mambani et al., 1983a; Ekanayake et al., 1985a; Venuprasad et al., 2002).
A study involving simultaneous evaluation of root character and grain yield, concluded that genotypes with a deep rooting habit had an advantage in stress conditions and that those genotypes that had produced deep roots prior to the onset of stress showed improved productivity compared with genotypes that did not have the capacity to produce roots prior to the onset of stress. The study also suggested, based on QTL mapping, that the loci for productivity traits were not congruent with those related to root morphology, except at one locus. Subsequently, Toorchi et al., (2006) and Kanbar et al., (2009), based on canonical correlation studies conducted under contrasting moisture regimes, suggested that maximum root depth, root�shoot ratio, and root dry weight conferred an advantage to grain yield under stress.
Lowland and upland management systems affect rice yield
Rice cultivars that are adapted exclusively to upland conditions are typically characterized by a deep and coarse root system, tall stature, thicker stems, and fewer tillers (Ge, 1992; Ling et al., 2002), whereas lowland rice cultivars have shallow and �ner roots, a large number of roots, and many tillers (Lang et al., 2003). In upland �elds and during stress conditions, the major sources of water for growth and development are rain water that is retained by the soil and groundwater. Depending on soil and weather conditions, pre-plant N could leach below the crop rooting zone early in the season before peak N uptake (Cameron, 2013), therefore a coarse and deep root system, for soil penetration and access to water reserves deep in the soil, is considered valuable for improved drought resistance and efficiency in N uptake under upland conditions (O�Toole and Chang, 1979; Ling et al., 2002). Large variations are noticed among upland rice cultivars for root length density below 30 cm, which suggests that the effect of drought stress depends on the ability of plants to develop a deep root system. Chang et al., (1986) also found that rice with a deep root system avoids drought better than rice with a shallow root system. Advantages conferred by a deep root system depend on three major factors: duration of the drought period, availability of water at depth, and rate of water uptake. If water is not limited in upper layers of the soil, the plant may not bene�t from the formation of deep roots. In upland conditions, Puckridge and O Toole, (1981) found that deep-rooted cultivars extracted more water at a depth of 40 70 cm than shallow-rooted cultivars.
 A range of trends have been reported regarding root growth response to drought stress in upland rice, including both root growth inhibition and root growth promotion in drought stress treatments. Kato et al., (2006) reviewed the effects of various water regimes on deep root growth and biomass partitioning and concluded that while many studies report an increase in root: shoot ratio and deep root growth in response to drought, conditions such as the timing of the drought at seedling stage, very severe drought stress, and presence of hardpans have reduced resource partitioning to roots. Conclusions about drought effects on root growth may also differ because root mass and root length can show opposite trends, especially when root diameter decreases because of drought, resulting in greater length but less mass.


The mechanical breakdown of surface soil aggregates is the most common land preparation method for lowland rice in Africa. Hardpans that develop from puddling improve soil water retention capacity (Sharma et al., 1985) but hinder root penetration to reach moisture in deeper zones of soil after drying occurs (Ghildyal, 1978; Hasegawa et al., 1985; Yu et al., 1995; Clark et al., 2000, 2002; Babu et al., 2001; Samson et al., 2002). Across species, morphological and physiological changes in plant growth due to effects of hardpans on roots include a reduction in transpiration rate and leaf area expansion, and ultimately a decrease in dry matter accumulation (Masle and Passioura, 1987; Ludlow et al., 1989; Assaeed et al., 1990; Masle, 1992). These effects may be due to direct consequences of reduced root access to water and nutrients. The presence of a hardpan in shallow soil layers may further promote uneven moisture distribution in the soil pro�le, so that root system tends to be partially exposed to dry soil, causing stomates to close while the rest of the root system can access water (Siopongco et al., 2008, 2009). In addition to the presence of a hardpan and greater strati�cation of soil characteristics due to puddling, rainfed lowland soils differ from upland soils in that severe soil cracking occurs upon drying of the soil under the lowland situation. Soil cracking, which can penetrate hardpans, strongly in�uences rainfall in�ltration and water evaporation processes (Tuong et al., 1996). Sanchez (1973) reported that soil cracking impeded root development based on visual observations; otherwise very little is known of the effects of soil cracking on growth of rice root systems. 
Compared to other crops, rice shows less soil penetration. Iijima et al., (1991) indicated that maize roots were better able to penetrate hard soil than rice roots. In general, roots of dicotyledonous plants have a higher penetration rate than monocot roots (Materechera et al., 1992), and upland rice cultivars have greater penetration ability than lowland cultivars (Yu et al., 1995). Changes in soil conditions can greatly alter root distribution patterns. However, the mechanism by which roots penetrate compacted soil layers is not well understood. 
It remains to be demonstrated whether the penetrated root mass has any role in the uptake of moisture and increasing grain yield in lowland conditions. One root characteristic, which is of interest in rice production is root plasticity; de�ned as the ability of a genotype to adjust its root growth phenotype according to environmental constraints (O Toole and Bland, 1987). The timing of root growth in response to the resupply of water following a period of drought stress is a vital feature of root growth plasticity. Genetic variation has been observed in rice for plasticity in several root traits, including the ability of lateral roots to develop in response to rewatering after soil drying (Banoc et al., 2000b), and also in response to soil drying after �ooding (Suralta et al., 2010). Genetic differences have also been observed in the ability of seminal roots to continue elongation and form aerenchyma under �ooded conditions after drought (Suralta et al., 2008a,b, 2010). 
Under progressive drought (Kato et al., 2006), this type of plasticity may be valuable for improved rice growth under drought by allocating resources to increased root growth only when needed. Different genotypes have exhibited different responses in plasticity depending on type of drought stress (Kano et al., 2011). These abilities are quite important for rainfed lowland rice due to the unique soil environment as mentioned earlier, and thus the desirable root traits are not as simple as those for upland rice such as deep or coarse roots. 
Water control enhances water use efficiency (WUE) in rice
It is now well reported that high yield potential and high yield under water-limited conditions are generally associated with high WUE (Munoz et al., 1998) mainly because low levels of water are required to achieve desirable yield. Features linked to low yield potential, such as smaller plants (Martin et al., 1999) or short growth duration (Lopezcastaneda et al., 1994) are  ascribed high WUE because they reduce water use (Condon et al., 2002). In the SRI, water loss is greatly reduced as the continuous flooding of rice fields are avoided during cultivation. Dehydration avoidance as achieved by enhanced capture of soil moisture by roots has been found to be associated with low WUE in various rice varieties (Kobata et al., 1996). On the other hand, reduced evapotranspiration in rice have been associated with high WUE (Kobata et al., 1996; Tolk et al., 2003). Water-saving irrigation, with shallow or little standing water management during transplanting as seen in the SRI is desirable, since a large water supply during puddling and after transplanting could increase water loss through evapotranspiration, and the loss of fertilizers through percolation; thereby resulting into greater environmental pollution (Lu et al., 2000; Won et al., 2003). 
Water control impacts growth hormones in rice
Hormones play important roles at different stages during the growth and development of the rice plant (Chen et al., 2006; Colmer et al., 2006; Rzewuski and Sauter, 2008). 
Factors that affect the synthesis and transport of rice hormones consequently affect the development and growth of rice. In this sense, water management and timing of water needs in rice production are essential as they influence the production of auxin, abscisic acid, cytokinin and gibberellic acid. Auxin and abscisic acid promote lateral root formation in rice, root tip swelling, root hair formation, and water permeability in roots of rice (Chen et al., 2006). Cytokinin suppresses lateral root formation, while ethylene has interactions with auxin and may play a role in lateral root formation through cortical cell breakdown (Liu et al., 2003). Gibberellic acid acts with ethylene to promote adventitious root growth in �ooded rice. Studies of hormone effects on rice root growth report that ethylene mediates aerenchyma formation and adventitious root growth under �ooded conditions in rice (Rzewuski and Sauter, 2008), but is not involved in the formation of barriers to radial oxygen loss (Colmer et al., 2006). Use of an antisense transgenic indicated a positive role of cytokinins for rice root development (Liu et al., 2003). 
Adequate rice spacing and plant density impacts rice yield in SRI 
Rice plants largely depend on temperature, solar radiation, moisture and soil fertility for their growth and nutritional requirements (Baloch et al., 2002). A dense population of crops may have limitations in the maximum availability of these factors. It is, therefore, necessary to determine the optimum density of plants population per unit area for obtaining maximum yields (Baloch et al., 2002). Optimum plant spacing ensures that plants grow properly, both in their aerial and underground parts through different utilization of solar radiation and nutrients. The optimum plant density depends on different factors, including: plant characteristics, growth period, duration of maturity, planting time and methods, soil fertility, plant size, available moisture, sunshine, planting pattern and situation of weeds (Shirtliffe et al., 2002). Plant spacing is an important production factor in transplanted rice (Gorgy et al., 2010). Mohapatra et al., (1989) reported that rice plant spacing of 20 � 20 cm was better than those of 15 � 15cm or 15 � 20 cm under normal soils for rice productivity. Maske et al., (1997), reported that plant height, leaf area index, yield and yield components of rice with plant spacing of 15�10 cm were higher than of 15�15 cm or 15�20 cm. Patel (1999), observed that hill spacing of 20 � 20 cm when compared with 20 � 15 cm and 20 � 10 cm recorded increases in number of panicles per square meters, yield and straw yield. However, the number of grains per panicle and 100-grain weight was not affected by hill spacing (Patel, 1999). In a similar studies; IRRI (1967), reported that the yield of rice varieties did not change when the planting distance was maintained below 35x35 cm.
The number of seedling per hill is another important factor that plays important roles in boosting yield of rice in the SRI. This factor influences tiller formation, interception of solar radiation, total sunshine reception, nutrient uptake, rate of photosynthesis and other physiological phenomena and ultimately affects the growth and development of rice plant. In densely populated rice fields the interspecific competition between plants is high. 
This may result in gradual shading and lodging and thus increase production of straw instead of grain. It is, therefore, necessary to determine the optimum plant spacing and number of seedling per hill for high yield (Ghosh and Singh, 1998; Hossain et al., 2003). Faruk et al., (2009) reported a high grain yield from two seedlings per hill when compared with single seedling per hill. The System of Rice Intensification (SRI) recommends wider spacing (25 x 25 cm to 30 x 30 cm) for higher yield (Batuwitage, 2000). However, Ferraris et al., (1973) mentioned that the plant spacing do not have much effect on final grain yield of rice under high nitrogen application. The finding by Ferraris et al., (1993) is supported by Peng et al., (1998) who found that the final grain yield for hybrid rice depends more on panicle size and panicle number.  Damage to rice seedlings due to transplanting shock is inevitable in the process of pulling and transplanting of rice seedlings, and may lead to growth stagnation for several days or a week after transplanting (Matsuo et al., 1995). Avoidance of transplanting shock and enhancing rapid growth and development without stagnation requires that rice plants be directly and quickly transplanted without delay, and at the recommended planting density and spacing.

Flooding in rice fields impacts yield in the SRI
Although large amounts of N fertilizers are being used worldwide, the recovery efficiency by crops is relatively low, between 25% - 50% of the N applied (Dobberman et al., 2003; Chien et al., 2016) and is most susceptible to losses (HYPERLINK "http://ucanr.org/repository/cao/landingpage.cfm?article=ca.v065n02p80&fulltext=yes" \l "R13"Schnier 1995). The change in early-season water management in rice fields has direct implications for nitrogen fertility management. Current recommendations on SRI were developed for continuously flooded rice. The impact of early-season drain on nitrogen fertilizer dynamics; particularly the effect on potential nitrogen losses, are not well understood and remains a priority area for research (HYPERLINK "http://ucanr.org/repository/cao/landingpage.cfm?article=ca.v065n02p80&fulltext=yes" \l "R8"Linquist et al., 2006). As rice fields are drained and the soil becomes aerobic, ammonium, applied as fertilizer is nitrified into nitrate (NO3) and becomes susceptible to losses. 
The nitrate disappears from the rice rooting zone within a week or two of a soil being flooded (HYPERLINK "http://ucanr.org/repository/cao/landingpage.cfm?article=ca.v065n02p80&fulltext=yes" \l "R8"Linquist et al., 2006). The fate of nitrate in flooded soils is difficult to determine. Plants, including rice, can take up nitrate before it is lost by other means. The most likely cause of nitrate loss from African rice systems is denitrification. When the field is reflooded and the soil becomes anaerobic, microbes convert a portion of the nitrate into nitrogen gas (denitrification), which is lost to the atmosphere (HYPERLINK "http://ucanr.org/repository/cao/landingpage.cfm?article=ca.v065n02p80&fulltext=yes" \l "R4"Buresh et al., 1991). In some rice systems, nitrate leaching can be a significant loss as the negatively charged nitrate molecules are repelled by similarly charged soil particles (HYPERLINK "http://ucanr.org/repository/cao/landingpage.cfm?article=ca.v065n02p80&fulltext=yes" \l "R16"Zhu et al., 2000; HYPERLINK "http://ucanr.org/repository/cao/landingpage.cfm?article=ca.v065n02p80&fulltext=yes" \l "R14"Yoon et al., 2006). However, rice soils are very impermeable, and under such conditions it is likely that soil nitrate denitrifies before it leaches (HYPERLINK "http://ucanr.org/repository/cao/landingpage.cfm?article=ca.v065n02p80&fulltext=yes" \l "R1"Bowman et al., 2002). HYPERLINK "http://ucanr.org/repository/cao/landingpage.cfm?article=ca.v065n02p80&fulltext=yes" \l "R8"Linquist et al., (2006) reported that shortly after flooding for planting, most nitrates are lost from the soil plough layer, and most mineral nitrogen is in the form of ammonium. The nitrate present prior to flooding the fields for planting would most likely have been lost via denitrification (HYPERLINK "http://ucanr.org/repository/cao/landingpage.cfm?article=ca.v065n02p80&fulltext=yes" \l "R4"Buresh et al., 1991). Over time, however, this nitrogen moves laterally through the soil (HYPERLINK "http://ucanr.org/repository/cao/landingpage.cfm?article=ca.v065n02p80&fulltext=yes" \l "R11"Obcema et al., 1984), and subsurface nitrogen levels become less variable. Besides these water induced nitrogen losses, ammonium can be lost in water runoff from rice fields. These losses, though comparatively small, can be large when nitrogen fertilizers are applied just before or during a runoff period (Patrick et al., 1976; HYPERLINK "http://ucanr.org/repository/cao/landingpage.cfm?article=ca.v065n02p80&fulltext=yes" \l "R15"Zhao et al., 2009). Minimizing continuous flooding throughout the crops growing period and limiting puddling to stages when the rice plant requires the bulk of water consequently should reduce nitrogen losses and increase rice yield in the SRI. Besides, this practice conserves water that can be channelled into other agricultural ventures, while enabling rice production in water limited areas. 
Quantitative trait loci (QTL) and phenotyping 
A lot of studies on rice crops have indicated considerable genotype � environment (G � E) interactions for root traits, which should be expected, given the number of environmental factors (soil, physical, chemical, and biological factors) that affect root growth and development in the rice plant. In a study by Kondo et al., (2003), who investigated G � E interactions by using both upland and lowland varieties at three upland sites in the Philippines, where both site and nitrogen treatments contributed to environmental variation; Genotype accounted for the largest proportion of variation for speci�c root weight, nodal root number, and root to shoot ratio, whereas the environmental effect of nitrogen treatment was relatively high for total dry weight and deep root length ratio.
Traits of roots are generally controlled by many genes through quantitative trait loci (QTL). Since the earlier study by Champoux et al., (1995) to locate genes controlling rice root traits with molecular markers, many QTLs related to root traits have been identi�ed in rice using 12 different mapping populations, with QTLs, identi�ed and analyzed in rice for more than 30 root morphological parameters. The most studied root traits in all QTL mapping studies are maximum root depth, root diameter, and root to shoot ratio.
The G � E effect on rice root growth is particularly important for rice under drought conditions: with lowland soils being a complex layering of disaggregated soil over a hardpan and ranging from �ooded paddies to dry cracked soils over the same season. Understanding how growth and observation methods affect root QTL studies is thus key for using knowledge of QTLs to improve drought resistance in rice. Most QTL studies have measured root traits in containers under controlled conditions, although it has yet to be proven whether these results reflect true genetic differences (Steele et al., 2007).
Root traits that result in improved plant water status through a stress-prone growing season could confer non-stage-speci�c drought resistance. For example, conventional breeding for root-related drought resistance in rice was conducted using farmer-participatory plant breeding approaches (Shashidhar, 2008). Selecting the best variety, with adequate drought resistance and capable of tolerating the prevailing environmental conditions is thus a priority in the SRI.
Root dry matter impacts rice yield
Genetic variation exists in potential root length in rice plants, which is attainable under non-stress and non-restrictive soil conditions. However, when plants are exposed to drying soils, root morphology and growth can change to the extent that the potential root length, whether short or long, becomes irrelevant. In rice, drying hard topsoil resists the penetration and establishment of adventitious (crown) roots; while existing roots receive all transient assimilates and grow deeper (Blum et al., 1984; Asseng et al., 1998).
Shoot to root dry matter ratio increases under drought stress, not because of an increase in shoot mass but due to a relatively greater decrease in root mass. Root mass rarely increases under stress. However, root length and depth may increase in a drying soil even at a reduced total root mass. One major exception that constitutes a form of an effective dehydration tolerance mechanism in crop plants is stem reserve utilisation for grain filling under drought stress (Blum, 1998). This is a harmonised whole plant process that allows effective grain filling when whole-plant photosynthesis is inhibited by stress during grain filling. It is a tolerance mechanism that allows grain filling in dehydrated or over-heated cereal plants, which can account for up to 90% of total loss in grain weight under stressful conditions (Blum et al., 1994; Asseng et al., 2003). 
Stem reserve utilisation has been found to be an effective yield-supporting mechanism under drought stress (Hossain et al., 1990; Pheloung et al., 1991; Gavuzzi et al., 1997; Yang et al., 2002; Asseng et al., 2003; Plaut et al., 2004). The major condition for stem reserves for grain filling is sufficient carbohydrate storage before grain filling (Yang et al., 2002). Reserve mobilisation is noticeably induced by drought stress during grain filling (Blum et al., 1994; Palta et al., 1994; Yang et al., 2001 a; Plaut et al., 2004). Some stem reserve mobilisation may support grain filling under non stress conditions. The signal for the induction of reserve mobilisation under drought stress is not clear but likely to involve the hormones, gibberellin and abscisic acid as reported by Yang et al., (2001 b). Generally in rice, and besides the stem, these reserves are stored in the roots with a high correlation with the root dry biomass (Palta et al., 1994; Yang et al., 2001 a; Plaut et al., 2004). The higher the root mass, therefore, the higher the reserve and hence the corresponding high nutrient mobilization during stress conditions: resulting into comparatively high yields.

Conclusion 
Under the conventional system of rice production, flooding represents the ideal condition for growing rice whiles the system of rice intensification is based on improving aeration to the soil, which favours deep and proliferous root development that can support a high above-ground productivity. Water conservation in the SRI system is beneficial to the performance of the rice plant. The development of the rice root system is essential for a high yield in rice, and is favoured by a number of management interventions in the system of rice intensification: 
Intermittent irrigation allows adequate respiration of the rice roots.
Application of organic matter enriches the soils, stimulates soil microbial development and creates a better environment for the growth and development of the rice roots.
Early transplanting of seedlings minimizes transplanting shock as soil that is attached to roots during transplanting is kept in place, and helps to protect the roots.
Root systems can develop better when transplanted singly � thus eliminating competition that exists in the conventional planting method, where a number of seedlings are planted per hill at closer spacing. 
Use of a mechanical hand-pushed or motorized weeder contributes to soil aeration.
Roots, under flooded conditions, do not develop as those under the SRI and have a shorter life span. Under flooded conditions, root system activity declines after mid-season, while in the SRI, roots remain actively longer into the grain filling period. Under continuous flooding, about three-fourth of the roots degrade by the flowering stage (Satyanarayana et al., 2007). Under SRI, roots reach deeper and achieve twice the volume compared to plants in conventionally planted hills (Thakur et al., 2011).� With larger root systems, plants access water at deeper soil horizons, which makes the crop more resilient towards drought stress (Satyanarayana et al., 2007).
These more productive phenotypes under SRI are characterized by higher number of tillers per plant, increased plant height, longer and wider leaves, longer panicles, more grains per panicle and improved grain quality. Plant internal water-use efficiency is also improved, by assimilating more CO2 per unit area of water transpired.	




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