1 s2.0 S0065211303800032 Main

NUTRIENT MANAGEMENT FOR IMPROVING LOWLAND RICE PRODUCTIVITY AND SUSTAINABILITY
N.K. Fageria,a N.A. Slatonb and V.C. Baligarc

National Rice and Bean Research Center of EMBRAPA, Caixa Postal 179, Santo nio de Goia s, Goia s, CEP 75375-000, Brazil Anto b University of Arkansas, 1366 W. Altheimer Dr. Fayetteville, Arkansas 72704, USA c USDA-ARS-Alternate Crops and Systems Research Laboratory, Beltsville Agricultural Research Center, Beltsville, Maryland 20705-2350, USA
a
I. Introduction II. Soils Used for Rice Production III. Management Practices to Improve Nutrient Use Efciency A. Liming B. Nitrogen C. Phosphorus D. Potassium E. Calcium, Magnesium, and Sulfur F. Micronutrients IV. Conclusions Acknowledgments References Rice (Oryza sativa L.) is an important food crop for a large proportion of the worlds population. Total rice production will need to increase to feed an increasing world population. Rice is produced under both upland and lowland ecosystems with about 76% of the global rice produced from irrigated-lowland rice systems. The anaerobic soil environment created by ood-irrigation of lowland rice creates a unique and challenging environment for the efcient management of soil and fertilizer nutrients. Supplying essential nutrients in adequate rates, sources, application methods, and application times are important factors that inuence the productivity and sustainability of rice. This review emphasizes our current, research-based knowledge of N, P, K, Ca, Mg, S, B, Fe, Mn, and Zn management in regards to the efciency and sustainability of lowland rice production and identies where additional research is needed to bridge information gaps. Our goal is to provide a comprehensive review describing the nutritional problems, nutrient use efciencies, and the production strategies used for efcient nutrient use and production of lowland rice. While the soils, climatic environments, cultivars, and degree of mechanization may vary considerably among the rice
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Advances in Agronomy, Volume 80 Copyright q 2003 by Academic Press. All rights of reproduction in any form reserved 0065-2113/03$35.00
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N. K. FAGERIA, N. A. SLATON AND V. C. BALIGAR producing regions of the world, the basic principles governing efcient nutrient use by ood-irrigated rice are relatively constant. A summation of best management practices should help scientists develop practical, integrated recommendations that improve nutrient use efciency in lowland rice production systems. q 2003 Academic Press.
I. INTRODUCTION
Rice (Oryza sativa L.) is the staple food in the diet of about one-half of the worlds population, who live mostly in developing countries, and is arguably the most important crop worldwide. In 2001, rice was grown on 151.5 million hectares, which represents approximately 23% of the worlds total area seeded to cereals (FAO, 2001). Rice provides 35 60% of the dietary calories consumed by nearly 3 billion people (Guerra et al., 1998). By the year 2025, it is estimated that it will be necessary to produce about 60% more rice than what is currently produced to meet the food needs of a growing world population (Balasubramanian et al., 1998). Enhancement of rice production and sustainability are important features of grain production to benet the worlds 3 billion people who depend on rice for their livelihood and as their basic food. Adequate amounts of essential nutrients are needed by modern rice cultivars with improved cultural practices to obtain higher yields. In this context, efcient use of inputs is vital to safely produce the additional food from limited resources with minimal impact on the environment. Rice, like barley (Hordeum vulgare L.), oats (Avena sativa L.), rye (Secale cereale L.), and wheat (Triticum aestivum L.), belongs to the Gramineae family. Rice was rst domesticated about 10,000 years ago; however, its exact origin of domestication is not known. The domestication of rice could have occurred independently at several places in a broad belt from the foothills of the Himalayas to Vietnam and southern China (Chang, 1975). The geographical dispersal and selection pressures of farming led to a large number of varieties of Oryza sativa, the Asian species. Another species, Oryza glaberrima, was later domesticated in western Africa (Hargrove, 1988). Before discussing nutrient management practices, the different types of cultural systems used to produce rice should be dened since the majority of the discussion here will focus on a specic production system. There is no true consensus on the terminology used to describe the different rice growing systems and environments. Upland, lowland, dryland, and wetland are often used to describe the different production systems, but these general terms frequently have regional connotations and may share some common characteristics. The International Rice Research Institute (IRRI, 1984) classied rice into ve
NUTRIENT MANAGEMENT FOR IMPROVING LOWLAND RICE 65
groups based on the seasonal water regime (decit, excess, or optimal), drainage (poor or good), air temperature (optimum or low), soils (normal or problem), and topography (at or undulating). These groups are: (i) irrigated lowland, (ii) rainfed lowland, (iii) deep water, (iv) upland, and (v) tidal wetlands. In this discussion, we primarily consider irrigated-lowland rice, which is cultivated on relatively at lands with water control so that rice is ooded for all or part of the growing season. The presence of oodwater for part or all of the growing season requires that the rice root system is adapted to largely anaerobic soil conditions. The rice plant has adapted to this environment by transporting oxygen from the aerial portions of the plant to the root system via aerenchyma tissues (Yoshida, 1981). A secondary adaptive mechanism is the development of an extensive lateral, brous root system located in the surface 1 2 mm of oxidized soil at the soil water interface. Oxygen diffusing through the water layer allows this zone of soil to remain oxidized. For these reasons, ooded rice normally has a shallow, brous root system (Wells et al., 1993). The aquatic environment not only inuences the development of the root system, but also alters the availability of several essential nutrients, affects nutrient uptake and use efciency, fertilization practices, and makes rice especially unique among crop production systems. The nutrient management practices discussed here will address the essential nutrients in terms of their specic functions pertaining to rice growth in an aquatic environment, nutrient deciency, nutrient toxicity, and potential environmental implications. Finally, we will review the fertilization practices (i.e., fertilizer sources, rates, timing, and methods of application) that inuence nutrient use efciency and provide insight on how these practices may be improved upon in the future.
II. SOILS USED FOR RICE PRODUCTION

Rice may be the worlds most diverse crop. Rice is grown in both dry and wet conditions, over a wide range of latitudes, and across a wide range of soil, climatic, and hydrological conditions. For example, rice tolerates water as deep as 5 m in the ood-prone areas of south and southeast Asia and grows well in the drought-prone upland areas of Asia, Latin America, and Africa. A major portion of the rice produced in Brazil is grown under upland, or dryland, conditions much like wheat or corn (Zea mays L.). Rice grows as far North as Czechoslovakia and Manchuria, and as far South as Uruguay and New South Wales, Australia (Hargrove, 1988). The geographic range of worldwide rice production is from the equator to the temperate areas of northern Japan and southern Australia, and from sea level to altitudes of more than 2500 m. The pedogenetic and morphological characteristics of soils used to grow rice also vary considerably. The soils used for rice production worldwide are
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N. K. FAGERIA, N. A. SLATON AND V. C. BALIGAR
distributed over the 10 soil orders (Moormann, 1978; Hudnall, 1991). Moormann (1978) summarized that, worldwide, rice is grown on all soil orders identied in the soil classication system (USDA, 1975). Worldwide, the wide array of soils used to produce rice results in an equally diverse assortment of management practices implemented for successful rice production on these soils. Murthy (1978) reported that the soils on which rice grows in India are so extraordinarily varied that there is hardly a type of soil, including salt-affected soils, on which it cannot be grown with some degree of success. In Brazil, ooded rice is mainly grown on Alsols, Vertisols, Inceptisols, Histosols, and Entisols (Moraes, 1999). In Sri Lanka, rice is grown on Alsols, Ultisols, Entisols, Inceptisols, and Histosols (Panabokke, 1978). In Indonesia, the main rice soils are Entisols, Inceptisols, Vertisols, Ultisols, and Alsols (Soepraptohardjo and Suhardjo, 1978). Raymundo (1978) reported that in the Philippines the soils used for wetland rice production are mainly Entisols, Inceptisols, Alsols, and Vertisols. In Europe, rice is planted on limited areas in Albania, Bulgaria, France, Greece, Hungary, Italy, Portugal, Romania, Spain, and Yugoslavia where the predominate soil orders are Inceptisols, Entisols, and Vertisols (Matsuo et al., 1978). In the United States, rice is grown primarily on Alsols, Inceptisols, Mollisols, and Vertisols (Flach and Slusher, 1978). However, in Florida, a small hectarage of rice is produced on Histosols. Most of the soils used for rice production in the United States and some other geographic areas have properties that make them ideally suited for ood-irrigated rice. The soils are relatively young, contain signicant amounts of weatherable minerals, and have relatively high base saturations despite the fact that some are in areas of high precipitation (Flach and Slusher, 1978). Soil parameters for optimum rice yields are optimum soil depth, compact subsoil horizon, good soil moisture retention, good internal drainage, good fertility, and a favorable soil structure. Clayey to loamy clay texture soils are appropriate for lowland rice production. Permeable, coarse-textured soils are less suitable for ood-irrigated rice production because they have low water or nutrient holding capacities. In Brazil, there are about 35 million hectares of poorly drained soils, known locally as Varzea, distributed throughout the country. At present about 1.8 million hectares of these soils are cultivated, primarily to lowland rice, during the rainy season (IICA, 2000). Generally, Varzea soils have good initial soil fertility, but after 2 3 years of cultivation, the fertility level is known to decline (Fageria and Baligar, 1996). Farming systems need to be developed with improved soil management technology to bring these areas under successful crop production. A sufcient supply of nutrients is one of the key factors required to improve crop yields and maintain sustainable agricultural production on these soils. Flood-irrigated rice is an important crop that needs to be included in the cropping system of these poorly drained areas during the rainy seasons. During dry periods, other crops can be planted in rotation, provided there is proper drainage. These soils generally have an
adequate natural water supply throughout the year, but are acidic and require routine applications of lime if legumes are grown in rotation with rice.
III. MANAGEMENT PRACTICES TO IMPROVE NUTRIENT USE EFFICIENCY

Modern production agriculture requires the implementation of efcient, sustainable, and environmentally sound management practices. In this context, fertilization is an important factor required to achieve optimum yields of all crops, including rice (Fageria et al., 1997a). Fertilizer is, however, one of the most expensive crop production inputs and, if used improperly, can pollute the environment or decrease production efciency. Increasing the rice yield per unit area by the use of appropriate nutrient management practices has become an essential component of modern rice production technology. Management practices to improve the nutrient use efciency of crops include management of soil pH, use of the proper inorganic and organic nutrient sources, correct method and rate of fertilizer application, appropriate application timing of fertilizers, water management, pest management, and the use of high-yielding cultivars adapted to a given environment.
A. LIMING
Soil acidity is a major factor limiting crop yield in vast areas of the world (Shainberg et al., 1989; Farina et al., 2000). Liming is an effective and widely used means of manipulating soil pH. Several acidity indices are used to diagnose and correct the problem. These include pH, cation or base saturation, and aluminum saturation. Crop response to soil pH is most commonly used to determine the quantity of lime required to reduce the adverse effects of soil acidity on productivity. The quantity of lime required depends on crop species, and sometimes, the cultivar within species, soil clay content, soil organic matter content, soil pH, concentration of soil Ca2, Mg2 and Al3, and the quality of liming material (Adams, 1984; Eckert, 1987). The potential advantages or disadvantages of liming soils used for lowland rice production are a controversial issue. The reduction processes that occur in ooded soils cause the soil pH to increase in acid soils and decrease in alkaline soils to near neutrality (Ponnamperuma, 1972; Reynolds et al., 1999). During the rst 2 3 days after ooding an acid soil, the pH decreases due to biotic transformation of easily degradable organic compounds to organic acids and CO2 (Motomura, 1962). After an extended period of anaerobic soil conditions, the pH increase is related to the reduction of Fe3 and Mn4 oxides (Ponnamperuma, 1972). However, factors such as the initial soil pH, organic matter content, soil
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texture, and soil iron content govern whether the soil pH increases or decreases after ooding. Patella (1976) reported a positive effect of liming on lowland rice yield in the State of Rio Grande do Sul of Brazil when the initial soil Ca Mg content was 2.5 cmolc kg21. He also reported that liming improved N uptake by rice. Fageria (2002) conducted a greenhouse experiment to evaluate crop tolerances to soil acidity of an Inceptisol and found a signicant P , 0:01 quadratic relationship between the shoot weight of lowland rice and a range of soil basic cation saturations (56 98%). Maximum shoot weights were produced at the lowest basic cation saturation. Crop tolerance to soil acidity of the ve crops used in this study, in order from least sensitive to most sensitive, was rice . corn . soybean [Glycine max (L.) Merr.] . common bean (Phaseolus vulgaris L.) . wheat. The tolerance of rice to soil acidity, and the intolerance of soybean, common bean, and wheat to soil acidity have been previously reported (Fageria et al., 1997a; Fageria and Gheyi, 1999). In India, rice seldom shows signicant, positive responses from lime applications to acid soils (Goswami and Banerjee, 1978). Liming is generally for the benet of other crops grown in rotation with rice because rice grows well in acid soils. If liming is necessary to optimize the growth and yield of other crops in the rotation, it should be applied immediately after the rice crop and prior to the production of other crops in the rotation, not immediately ahead of the rice crop (Wilson et al., 2001). Research in Arkansas has shown that rice yields may benet from low to moderate rates of lime application to soils with pH near or below 5.0 (Ntamatungiro et al., 1999). Wang (1971) reported that 5 t lime ha21 applied to a submerged latosolic soil (pH 4.8) low in base saturation increased rice grain yield by 23% and straw yield by 27%. Ponnamperuma (1960) observed an increase in rice yields from lime applications to the very acidic lateritic soils of Sri Lanka. Likewise, Abichandani and Patnaik (1961) and Mandal et al. (1966) also increased rice yields by liming. The decision to lime acid soils used for rice production should consider the crops grown in rotation with rice. Management of soil acidity is often essential to improve or maintain the yields of important legume crops such as soybean and common bean (Fageria and Gheyi, 1999) and grain crops like wheat and corn (Z. mays L.) produced on acidic lowland soils (Fageria and Baligar, 1999b).
B. NITROGEN
Nitrogen is usually the most yield-limiting nutrient in lowland rice production. The exception is for rice grown on organic soils that have ample N release to supply seasonal crop N requirements. Research has shown that ood-irrigated rice can be one of the most efcient or inefcient crops in regards to N fertilizer use efciency. The relative efciency of rice utilization of N fertilizer is directly
related to water management, rice growth stage at N application, N source, and the chemical transformations that occur to N after it is applied to the soil. Intensive agricultural production systems have increased the use of N fertilizer in effort to produce and sustain high crop yields. Consequently, N losses into the environment have also increased (Schmied et al., 2000). Environmental N losses can occur from NO3 leaching, NH3 volatilization, surface runoff, nitrication, and denitrication. Release of N2O from soils occurs during biological and chemical denitrication (chemodenitrication) and contributes to global warming from the emission of greenhouse gasses. (Nelson, 1982; Hutchinson and Davidson, 1993; Thorn and Mikita, 2000). Over the last 20 years the concentration of N2O in the atmosphere has increased by , 0.25% yr21 (Elkins and Rossen, 1989). Anthropogenic inputs of all forms of N into the global N cycle are an estimated 145 Tg yr21 (Galloway et al., 1995). Of this total, N losses from N fertilizers are estimated at , 80 Tg yr21. Thus, fertilizer usage is considered as a major contributing factor of this increase, compared with other sources of N2O such as biomass burning, fossil fuel combustion, deforestation, increasing amounts of wet and dry NOx deposition, and NH4 in rainwater. Raun and Johnson (1999) estimated the worldwide fertilizer N recovery efciency by cereal grains (rice, wheat, sorghum (Sorghum bicolor L.), millet (Pennisetum glaucum (L.) R. Br.), barley, corn, oat, and rye) is only , 33%. The 67% of unaccounted for N represented a US$15.9 billion annual loss of N fertilizer. Thus, when N fertilizers applied to all crops, including rice, are not used efciently, crop production costs increase as more N fertilizer must be used to compensate for losses; however, more importantly this lost N contributes to air and water pollution. The development, demonstration, and eventual grower acceptance of efcient N fertilization practices can reduce N use, maintain or increase current rice yields, and minimize the potential for N losses into the environment (Kundu et al., 1996). Improving the N use efciency of rice is an important goal in sustainable production systems. Various management practices can be employed to balance the supply of N required for optimum crop production while minimizing potential losses into the environment. The successful adoption of efcient N management practices for ood-irrigated rice requires an understanding of the relationships among rice growth and development, the biological and chemical transformations of fertilizer and native soil N, soil chemical and physical properties, and management of irrigation water. 1. Chemistry in the Soil
Lowland ood-irrigated rice grows in a complex, dynamic soil environment during the course of a complete growing season. Understanding nutrient use efciency requires that the seasonal soil environment be described. Most of
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the soils used for ood-irrigated rice production are subjected to alternate aerobic anaerobic cycles because rice is drained for harvest and may be grown in rotation with upland crops. The soil is generally ooded immediately before rice is seeded (i.e., direct waterseeded culture); transplanted; or when seedlings, established by direct, dry-seeded methods (i.e., broadcast or drill-seeded delayed ood culture) reach the 5-leaf stage and begin to tiller. Once established, the ood is commonly maintained until physiological maturity when it is drained for harvest. In some instances, the ood is intentionally drained to allow the soil to completely dry for specic management reasons (i.e., straighthead control and pest control) or is lost due to lack of short-term water availability. Flooding an oxidized soil affects the rice utilization efciency of both native soil and fertilizer N. Under aerobic conditions, if environmental conditions are not limiting, NH4 is oxidized to NO3 almost as rapidly as it is formed (Schmidt, 1982). At the time of 2 ooding, nearly all the NO2 2 and NO3 present in the soil is lost to denitrication, leaching, or both, depending on the soil properties (Patrick and Wyatt, 1964). Thus, the mineralized organic N that undergoes nitrication under aerobic soil conditions, during the off-season, or while upland rotation crops are grown is ultimately lost to denitrication before signicant uptake by ood-irrigated rice occurs. One benet of ooding is that organic N continues to be mineralized under anaerobic conditions and remains as NH4, which is available for plant use rather than being nitried to NO3 and lost as N2. The organic N mineralized after ooding plays an important role in determining the rate and times that fertilizer N is required to produce optimum rice yields. Rice production systems that lack controlled water management and undergo several alternating aerobic and anaerobic cycles during a single season are highly inefcient in regards to native soil N use (Patrick and Wyatt, 1964). The majority of fertilizer-N losses from all ood-irrigated rice production systems are usually attributed to NH3 volatilization and denitrication (Freney et al., 1990). Once a soil is ooded, several distinct zones of varying oxygen status exist and play important roles in determining soil and fertilizer N use efciencies (Fig. 1). The ood slows oxygen diffusion to the soil surface and prevents aerobic N transformations from occurring in the majority of the soil beneath the oxygenated soil surface oodwater interface. The ood depth is generally 5 20 cm. The oxygenated oodwater soil surface layer is often depicted as being about 1 cm thick, but its depth varies with soil physical properties (Mikkelsen, 1987). Below
Figure 1 An illustration of the various N chemical forms, transformations and behavior in the ooded soil environment in which rice is grown. Nitrogen sources are in blocks, N chemical forms are in circles and the mechanisms responsible for the various N transformations or behavior are located on the arrowed lines. (Reprinted with permission of R. J. Norman, C. E. Wilson, Jr. and N. A. Slaton).
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the oxygenated soil surface layer, the soil is highly anaerobic except for the oxidized root rhizosphere. The oxygenated soil surface is a dynamic layer where N transformations occur that inuence N uptake and the numerous N loss pathways. The interaction of the anaerobic soil environment with N fertilizer sources, application times, and placement will be further scrutinized when fertilization practices are later reviewed, but will be mentioned briey in describing the various N loss mechanisms in the following sections. Understanding the relationships among various N loss mechanisms, crop nutrient uptake, and crop management practices is vitally important for adopting management practices that improve N use efciency. Ammonia-based fertilizers, such as urea, are the most abundant forms of nitrogenous fertilizer applied to agricultural soils (Eichner, 1990). The most fundamental N recommendation for ood-irrigated rice is to use an ammonical form of N fertilizer. Ammonia N may be lost through volatilization, xed by clay minerals, or immobilized by soil organic matter. Use of NO2 3 fertilizer sources for preood N application is not recommended because NO3-N is subject to denitrication after ooding and seedling rice is sensitive to NO3 salts (Baser and Gilmour, 1982). Denitrication is the anaerobic, microbial process that reduces NO3 to N2 gas which subsequently escapes into the atmosphere. Denitrication occurs very rapidly after a soil is ooded. Nitrate present at the time of ood establishment is lost within 8 10 days after ooding (Patrick and Wyatt, 1964). Nitrication may occur in the oxidized soil water interface from the upward movement of mineralized soil NH4 and NH4 fertilizer. For these reasons, NO3-N contributes very little towards the total N requirement of ood-irrigated rice. The goal of preood or early N applications is to keep the soil and fertilizer N in the NH4 form to prevent denitrication after ooding. The NH 4 is a cation which attaches to the soils cation exchange complex and is held by the soil very close to its point 2 of placement. In comparison to NH 4 , NO3 is an anion and is mobile in the soil. Thus, NO3 is also susceptible to leaching (on permeable, sandy soils) or may possibly be transported in the general direction of water movement across a eld. Nitrate that is not leached out of the soil prole or absorbed by rice is quickly denitried upon its movement into the anaerobic soil zone. In soil, N availability for plant uptake and leaching processes is related to the NH4 adsorption and desorption capacity of the soil. Wang and Alva (2000) reported that the potential NH4 buffering capacity and labile NH4 for sandy soils were much lower than those for clay and silt loam soils. This means that NH4 xation capacity of a soil should be taken into account when strategies for N fertilization are developed to maximize N use efciency. Further, the adsorption and desorption of NH4 are also related to soil organic carbon. Organic C is positively correlated with the potential NH4 buffering capacity and labile NH4 content for soils (Wang and Alva, 2000). Figure 1 summarizes the physical, chemical and biological transformation, plant uptake, and loss pathways of N in a ooded rice ecosystem.
2.
Functions and Deciency Symptoms
Nitrogen is one of the major elements required for plant growth. It is a constituent of numerous important compounds found in living cells, including amino acids, proteins (enzymes), nucleic acids, and chlorophyll (Traore and Maranville, 1999). Nitrogen fertilizers are routinely applied to rice, but N deciency symptoms are frequently observed because of the many loss mechanisms that inuence fertilizer N use efciency and the amount of N required to produce high yields. Nitrogen is a mobile element inside the plant; hence, deciency symptoms appear on the older leaves rst. The lower leaves of N decient rice turn yellow, but the whole plant may become chlorotic if the N deciency is not corrected. The chlorosis begins near the tip of the leaf blade and advances towards the leaf base until the whole blade is yellow. The dominant symptom of N deciency in rice is the development of the pale yellow color on the lower leaves. Nitrogen deciency reduces plant height, tillering, leaf area index, leaf area duration, and crop photosynthetic rate which leads to lower radiation interception and lower radiation use efciency. Color photographs of the N deciency symptoms of rice have been presented by Mueller (1974), Cheaney and Jennings (1975), Ishizuka (1978), Yoshida (1981), Fageria (1984), and Fageria and Barbosa Filho (1994).
3.
Forms of N Uptake
Plants absorb N from the soil solution as NO2 3 or NH4 . Most plants can absorb both forms of N equally; however, the form of N uptake is mainly determined by its abundance and accessibility. In well-drained soils, NO3 dominates, while under anaerobic conditions or in cold climates NH4 is the dominant form. In general, under agricultural conditions, soil NO3 concentrations range between 0.5 and 10 mM (7 140 mg NO3 kg21), while NH4 concentrations are 10 1000 times lower and reach the millimolar range only in exceptional cases, such as after fertilization (Wiren et al., 1997). Although both forms of N can be absorbed by rice, Fried et al. (1965) showed that excised roots from rice seedlings absorbed NH4 5 20 times faster than NO3, with the rate being somewhat dependent on the solution pH. The majority of the published literature suggests that rice prefers NH4 compared to NO3 (Mengel and Viro, 1978). When NO3 is absorbed, it must be reduced for assimilation and transport within the plant, with the reduction process requiring energy. Takenaga (1995) suggested that rice may prefer one N form over another based on plant growth stage. During vegetative growth, NH4 was absorbed more effectively while NO3 is preferentially absorbed during reproductive growth. Moristsugu (1980) concluded that rice shoot and root dry weights were higher when rice was grown in an NH4-N culture solution compared to a NO3-N
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culture solution. Rice that is ooded continuously, from early vegetative growth until physiological maturity, does not absorb large amounts of NO3 because the NO3 is rapidly lost via denitrication after ooding. For this reason, soil or fertilizer NO3 is of little or no benet to lowland rice. In most cases, an NH4 forming N fertilizer must be used along with management practices that largely prevent nitrication to optimize N uptake efciency. Ammonium sulfate and urea [CO(NH2)2] are generally considered equally effective fertilizer sources for rice (Bufogle et al., 1998; Gaudin and Dupuy, 1999). The initial soil pH, due its inuence on potential NH3 volatilization losses, and the length of time required to ood the soil may inuence the efciency of uptake of these two N sources (Wells and Turner, 1984; Clark et al., 2001). Ammonium sulfate may be more efcient when applied on alkaline soils or when the N is not incorporated immediately by ooding. Urea, following application to the soil, undergoes hydrolysis to eventually form NH4. However, under certain soil and environmental conditions, N in the form of NH3, a gas, may be lost by volatilization. Ammonia volatilization may be a pathway of signicant N loss on alkaline soils. Marschner (1995) reported that calcifuges, or plants adapted to acid soils, and plants adapted to low soil redox potential (e.g., lowland rice), prefer NH4. In contrast, calcicoles, or plants with a preference for calcareous soils, utilize NO3 preferentially. De Datta (1981) reported that the N applied at planting should be in the NH4 form. The N used as topdressing is less critical as NH4 and NO3 forms appear to be equally effective (Wilson et al., 1994). When the crop is fully established (i.e., panicle initiation), the NO2 3 form of N is rapidly taken up before it can be leached down to the reduced soil layer where it could be lost through denitrication. This may account for the equal performance of NO3 containing fertilizers such as NH4NO3 as compared to other NH4 containing or forming N sources such as ammonium sulfate or urea (De Datta, 1981).
4.
Seasonal N Uptake and Partitioning
In order to understand the nutritional requirements of rice a general knowledge of rice growth and development and its inuence on the demand for nutrients is required. Dry matter and grain yield production are perhaps the best growth parameters to show how plant growth relates to nutritional requirements. Between emergence and beginning tillering, a gradual linear increase in plant dry matter occurs (De Datta, 1981; Fageria et al., 1997a). Dry matter increases very rapidly between the onset of tillering and beginning internode movement due to tiller formation and an increase in leaf size. This rapid growth produces a sigmoidal shaped growth curve. In long-season cultivars, a plateau or midseason lag phase may appear at the end of vegetative growth just prior
to panicle initiation, which is the onset of reproductive growth. The lag phase is followed by another rapid increase in dry matter accumulation as the panicle develops, emerges, and the grain lling process is completed. Very short-season cultivars may not have a distinct lag phase resulting in a continuous rapid growth curve from beginning tillering until the grain lling process is completed. For optimum growth and yield, rice requires that adequate N be available at the beginning of the rapid growth (tillering) period. The number of panicles per unit area is determined by either stand density or tiller development during vegetative growth and is the rst yield component determined (Stansel, 1975). By beginning internode elongation, or panicle initiation, the maximum tiller number has been reached. The second yield component, the potential number of grains per panicle, is determined at panicle initiation and is inuenced by the plants nutritional status during vegetative growth. Wells and Faw (1978) showed that under optimum stand densities increasing the N rate did not signicantly increase the number of tillers per unit area, but when stand density was constant, the number of orets (potential grains) per panicle signicantly increased with increasing N rate. The number of blanks per panicle also increased with increasing N rate, but grain yield increased too. Counce and Wells (1990) found that at extremely low stand densities (, 30 seedlings per square meter) increasing the preood N rate stimulated tillering, increased the number of panicles, decreased the number of grains per panicle, and increased grain yield. At low stand densities, the most yield-limiting factor is panicles per unit area. Thus, increasing the preood N rate on elds with below optimum stands increases tillering (potential panicles) which has the greatest impact on the number of grains per unit area. Grain weight, the third and nal yield component, is primarily determined by genetics and is not inuenced by N rate (Counce and Wells, 1990). Rice grain yield is a function of the panicles per unit area, number of spikelets per panicle, 1000-grain weight, and spikelet sterility or lled spikelets (Fageria et al., 1997b). Therefore, it is very important to understand the management practices that inuence yield components and consequently grain yield. The 3-year average of these yield components, as affected by N rate, on a Brazialian Inceptisol are presented in Fig. 2 (Fageria and Baligar, 2001). Application of N rates up to 210 kg N ha21 signicantly increased panicle length P , 0:01 and the relationship between N rate and panicle length was linear (Fig. 2). The panicle number per square meter and 1000-grain weight also increased signicantly and quadratically as N fertilizer application rate increased. Spikelet sterility, however, decreased signicantly and linearly with increasing N rates. Nitrogen rate accounted for about 96% of the variation in panicle length, 91% of the variation in panicles per square meter, 75% of the variation in spikelet sterility, and 73% of the variation in 1000-grain weight. Yoshida (1981) also reported that panicles per unit area, lled spikelet percentage, and 1000-grain weight were major contributors to increased grain yield in modern high-yielding rice cultivars.
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Figure 2 Inuence of N fertilizer rate on panicle length, panicle number per square meter, spikelet sterility, and 1000-grain weight of lowand rice. Values are the average of 3 years of eld experimentation. (Reproduced with permission from Fageria, N. K. and Baligar, V. C. 2001. Lowland rice response to nitrogen fertilization. Commun. Soil Sci. Plant Anal. 32: 14051429. (Copyright Marcel Dekker, New York).).
Compared to the untreated control, application of 210 kg N ha21 increased panicle length by 9%, increased panicle number per square meter by 24%, increased 1000-grain weight by 6%, and decreased spikelet sterility by 14%. Low spikelet sterility at high N application rates is considered one of the important selection criteria for N responsive rice cultivars (Yoshida, 1972). During the course of the growing season, whole plant tissue N concentration gradually declines as plant dry matter increases, regardless of N rate (Table I). Although tissue N concentration decreases, total N uptake increases as N fertilizer rate increases (Table II). Approximately one-half of the total dry matter of a mature rice crop is produced by panicle differentiation or the early boot stage (Sims and Place, 1968; Guindo et al., 1994b). Total N uptake tends to follow the same general pattern as dry matter accumulation. Under optimum fertilization, one-half of the total N is absorbed before one-half of the total dry matter is produced. The remaining one-half to one-third of the N, not in the plant by panicle differentiation, is obtained either from topdressed N fertilizer, soil N released via N mineralization, or both. The total N content of rice straw and grain at maturity usually contains 20 40% more N than that supplied by fertilizer and represents plant uptake of native soil N (Moore et al., 1981; Norman et al., 1992b).

Table I Whole Plant Concentration of N in the Shoots of Flooded-Irrigated Rice Cultivar Metica 1 During the Growing Season. Concentrations are the Average of Three Crop Years (Fageria, unpublished data, 2001) Days after seeding (growth stage) (g N kg21) N Fertilizer rate (kg N ha21) 0 30 60 90 120 150 180 210 R2
22 (IT) 40 42 43 44 44 45 46 45 0.95**
35 (AT) 28 28 30 31 34 32 34 33 0.85**
71 (IP) 12 11 12 12 13 13 13 15 0.84**
97 (B) 9 9 10 11 10 12 13 13 0.89**
112 (F) 8 7 9 8 9 9 10 10 0.72*
140 (PM) 5 6 6 6 6 6 7 7 0.75**
*,**Signicant at the 5 and 1% probability levels, respectively. IT, initiation of tillering; AT, active tillering; IP, initiation of panicle; B, booting; F, owering, PM, physiological maturity.
Table II Whole Plant Content (Accumulation) of N in the Shoots of Flooded-Irrigated Rice Cultivar Metica 1 During the Growing Season. Contents are the Average of Three Crop Years (Reproduced with Permission from Fageria, N. K. and Baligar, V. C. 2001. Lowland Rice Response to Nitrogen Fertilization. Commun. Soil Sci. Plant Anal. 32: 1405 1429. Copyright Marcel Dekker, New York) Days after seeding (growth stage) (kg N ha21) N Fertilizer rate (kg N ha21) 0 30 60 90 120 150 180 210 R2
22 (IT) 13 13 15 17 17 19 16 16 0.78**
35 (AT) 24 26 37 33 42 40 44 39 0.85**
71 (IP) 35 41 45 52 57 63 67 87 0.96**
97 (B) 50 62 82 88 91 113 137 130 0.95**
112 (F) 57 66 95 89 122 113 135 130 0.92**
140 (PM) 28 37 40 41 48 52 59 66 0.98**
Grain 36 41 55 55 66 67 71 74 0.97**
**Signicant at the 1% probability level. IT, initiation of tillering; AT, active tillering; IP, initiation of panicle; B, booting; F, owering, PM, physiological maturity.
78
Nitrogen is a mobile element within the plant. Nitrogen stored in the stems and bottom, older leaves can be translocated up to younger leaves when N is decient or to the developing panicle at heading. When individual leaves are analyzed for N, a concentration gradient can be observed among the leaf positions from top to bottom. The youngest (top) leaves have a higher N concentration than older leaves (Westfall et al., 1973). At panicle differentiation, the top two leaf blades, top two leaf sheaths, remaining bottom leaf blades, and remaining leaf sheaths and stems contain near equal amounts of N (Norman et al., 1992b). The top two leaf blades contain the largest amount of N. As rice development progresses towards heading, N is rst translocated from the bottom leaf sheaths and stems to the developing panicle which may cause some senescence of the older leaves. Once the panicle emerges, the total N content of all plant parts tends to gradually decline at the same rate until physiological maturity. Based on total N content, about 60 70% of the aboveground N is located in the rice panicles at maturity (Table II; Moore et al., 1981; Norman et al., 1992b). Norman et al. (1992b) found that 86% of the total N in the panicles was translocated from rice leaf blades (58%) and sheaths and stems (28%) to the panicles. The remaining 14% of the panicle N was obtained from N uptake by the rice root system, presumably from N released by mineralization from the ooded soil. Upon recognizing that N was lost from the rice leaves during grain lling it was estimated that about 30% of the panicle N was actually obtained from root uptake after heading. Regardless, the majority of N residing in the rice grain at harvest is initially absorbed between rice emergence and heading, stored in the vegetative plant tissues, and eventually translocated to the panicle. For this reason, proper preood N management and efcient plant use of early N fertilizer is very important to high yield production. The total N content of the straw and grain at maturity is equaled or surpassed only by K and Si. At physiological maturity, the aboveground biomass, including both straw and grain, of a healthy high-yielding rice crop may weigh between 16,000 and 28,000 kg ha21 and contain approximately 160 280 kg N ha21 (Moore et al., 1981; Guindo et al., 1994a; Wilson et al., 1994). The proportion of total plant N partitioned to the grain is called the N harvest index (NHI). Nitrogen in the roots has little inuence on the efciency of N partitioning (McNeal et al., 1966; Fawcett, 1980), the NHI ratio refers only to N in the aboveground parts of the plant. A high NHI is associated with efcient utilization of N (Fawcett and Frey, 1983; Rattunde and Frey, 1986) and grain protein yields (Cox and Frey, 1978; Welch and Yong, 1980), but only rarely with grain protein percentage (Peterson et al., 1975) and is inversely related to mean straw yield (Rattunde and Frey, 1986). Selecting for high NHI may give simultaneous improvement for grain yields and grain protein percentage (Fawcett and Frey, 1983), or an increase in grain yield with constant grain protein percentage (Lofer and Bush, 1982). The magnitude of NHI is inuenced by environmental factors. The efciency of N translocation can be depressed by high application rates of N (Fawcett and Frey, 1983). Dingkuhn et al. (1991) reported
NHI values ranging from 0.60 to 0.72 for three semidwarf rice cultivars differing in growth duration. Guindo et al. (1994a,b) also reported NHI values of 0.58 and 0.62 in two lowland rice cultivars. Mature rice straw has a N concentration of 5.0 8.0 g N kg21 and the reported C:N ratio ranges from 50:1 (Norman et al., 1990) to 71:1 (Eagle et al., 2001). The C:N ratio of rice roots is slightly higher than straw at 98:1 (Norman et al., 1990). A rice straw yield of 11,200 16,800 kg ha21 contains 60 84 kg N ha21. In comparison, soybean residue (stubble) remaining after harvest may weigh 5600 6720 kg ha21 and contain 112 224 kg N ha21 (Hanaway and Johonson, 1985; Norman et al., 1990). The N concentration of rough rice grain averages about 10.0 g N kg21 (Moore et al., 1981). Harvest index, the weight of harvested grain divided by the total aboveground biomass (straw and grain), is an indication of how efcient a crop is at producing grain. A crop harvest index could range from 0 to 1.0 depending on the crop and the plant part that is harvested. A high harvest index is desired for rice. Published harvest index values for rice usually range from 0.35 to 0.55 (Counce et al., 1992; Norman et al., 1992a; Roberts et al., 1993; Wilson et al., 1994). The maximum harvest indexes reported for rice are 0.60 0.65 (Roberts et al., 1993). Advances in pest management and breeding have increased the harvest index for rice. Roberts et al. (1993) showed that semidwarf cultivars had a higher harvest index than the taller cultivars that were grown before the development of semidwarf cultivars. Although the biological yield between semidwarf and tall cultivars did not vary, the harvest index decreased more rapidly for tall cultivars as N rate increased. Norman et al. (1992a) found that harvest index increased when preood N application was delayed for more than 7 days past the normal recommended time (5-leaf stage). Although, delayed fertilization and ooding reduced straw yield it did not decrease grain yields. 5. Critical Level in Plant The rate of nutrient uptake by rice changes during growth and development. The rate of nutrient uptake is inuenced by the nutrient; fertilizer application source, time, and rate; nutrient availability in the soil; and plant growth characteristics. The absorption of most nutrients is generally vigorous during vegetative growth, but is limited by the root system. Thus, in general, nutrient uptake rate reaches a peak before heading (Takenaga, 1995). Nitrogen, P, and K are the rst to reach the highest uptake rate in rice, while Mn and Si are the last in this respect. By heading, rice has absorbed about 60 80% of the total amount of each nutrient to be absorbed (Ishizuka and Tanaka, 1952a). The use of tissue N concentration of rice plants as a means of evaluating the N nutritional status of the rice crop must be carefully interpreted. The N concentration of rice tissues does not always increase as the rate of N fertilizer
80
increases. Nitrogen stimulates vegetative growth which dilutes N in the plant tissue rather than increasing tissue N concentration. Plant N content is often a more meaningful measurement than tissue N concentration. The N concentration of rice straw decreases throughout the growing season as plant biomass increases and the developing panicle acts as a strong sink for N and other nutrients. Fageria and Baligar (2001) reported that the critical N concentration in the rice plant increased quadratically with increasing N application rate and decreased with an increase in plant age due to the dilution effect (Table I). Yoshida (1981) reported that the tillering rate in rice increased linearly as leaf blade N concentration increased up to 50 g N kg21. Tiller development stopped when leaf blade N concentration was , 20 g N kg21. De Datta (1981) reviewed the critical N concentrations proposed by various sources and suggested 25 g N kg21 in the leaf blade of rice plants at the active tillering stage was the critical value. Wells et al. (1993) suggested a critical range of 25 32 g N kg21 in the Y-leaf at panicle initiation. Fageria et al. (1997a) reported adequate N concentrations ranged from 26 to 42 g N kg21 in the uppermost mature leaves at heading. 6. Nitrogen Use Efciency
Nutrient use efciency by crops can be expressed and interpreted in a number of ways. The majority of research on N fertilization of rice in the USA and many other rice growing regions has focused on calibration of N fertilizer rates, plant uptake and utilization, and the magnitude of N loss pathways among various N fertilizer sources, application times, and application methods. Information generated from these types of studies help to develop best management practices for the production of high yields while minimizing N losses and costs associated with N fertilization. Nutrient use efciency can also be expressed in terms of biomass or grain production efciency per unit of nutrient uptake or application. Regardless of how nutrient use efciency is expressed, the ultimate goal of this information is to enhance our knowledge of crop growth and production efciency. Numerous research studies have been conducted to measure the fate (i.e., plant uptake and N loss pathways) of N fertilizer under various cultural management systems. These studies have focused on accounting for N fertilizer use efciency or uptake by rice; N lost via leaching, NH3 volatilization, and denitrication; and microbial biomass use of N fertilizer in various cultural management systems, straw management, environments, and crop rotations. Rice may be one of the most efcient or inefcient crops in regards to N fertilizer use efciency. The N fertilizer source, the N application time, or both usually dictate at which end of the efciency spectrum N fertilizer use by rice resides. The recovery efciency of N fertilizer by rice generally ranges from 20 to 80% (Table III) with an average of about 30 40% (Cassman et al., 1993). Recovery efciency is generally dened
Table III Nitrogen Recovery Efciencies of Basal, Topdressed, or Basal and Topdressed N Applications from Various Geographic Regions Recovery efciency % of applied 15N fertilizer Reference Bollich et al. (1994) De Datta et al. (1988) De Datta et al. (1988) Norman et al. (1992b) Norman et al. (1989) Norman et al. (1989) Norman et al. (1989) Patrick and Reddy (1978) Patrick and Reddy (1978) Westcott et al. (1986) Westcott et al. (1986) Westcott et al. (1986) Westcott et al. (1986) Wilson et al. (1989) Wilson et al. (1989)
a
Cultural system Dry Water Transplant Dry Dry Dry Dry Dry Dry Water Dry Water Dry Dry Dry
15
N Application time
Plant 56 41 33 54 57 46 58 28 47 48 44 42 30 42 61
Root 35a 35a 26a 32a 11 6
Soil 33
Unaccounted for N % of applied 15 N fertilizer 11 25 33 26 29 35 33 46 21 44 48 46 60 25 16
Basal Topdress Basal Topdress Basal Topdress Basal Basal Topdress Basal Topdress Basal Topdress Topdress Topdress Basal Basal Basal Topdress
20 14 19 9
8 8 12 10 22 17
A single value denotes that soil and root N were grouped together.
82
as the total N accumulation in the aboveground biological yield (grain straw) per unit of applied N fertilizer. Recovery efciencies have been determined by use of 15N labeled fertilizers and by the difference method. Fertilizer N recovery measurements are affected by the method of determination, scope of sampling (i.e., plant and soil), and the time of sampling. Fertilizer N recovery by difference usually yields higher uptake efciencies than studies using 15N labeled fertilizers (Reddy and Patrick, 1976; Eagle et al., 2001). The differences between these two methods may either be minimal (Reddy and Patrick, 1976) or signicant (Reddy and Patrick, 1976; Eagle et al., 2001). The difference method usually yields higher recovery efciencies because of the priming effect of N fertilization, stimulation of N mineralization in the presence of fertilizer N, and root exploration is greater in fertilized plots (Westerman and Kurtz, 1974; Olson, 1980; Olson and Swallow, 1984; Rao et al., 1991; Yamaguchi, 1991; Cassman et al., 1993; Schnier, 1994). Regardless of the method of determination, the balance of N fertilizer not accounted for by N recovery efciency should not be interpreted as N that is initially lost without utilization from the soil plant system. A portion of the unaccounted for N may certainly be lost to denitrication, leaching, and ammonia volatilization, but a signicant portion of this N may be incorporated into the microbial biomass, reside in the rice root system, or be lost from senescing rice leaves and should not be ignored when performing N use efciency studies. When measured at plant maturity, N fertilizer uptake and recovery in the aboveground biomass alone may overestimate fertilizer N losses and underestimate fertilizer recovery by as much as 60% when soil, root, and gaseous N losses from senescing rice leaves are not accounted for. Wilson et al. (1989) consistently recovered 11% of the total applied N fertilizer in the rice root system at maturity. Furthermore, between 16 and 25% of the total applied N fertilizer has been recovered in the soil organic fraction between panicle differentiation and maturity (Wilson et al., 1989; Norman et al., 1992b; Bollich et al., 1994). Measurement of N fertilizer recovery in the aboveground biomass at maturity underestimates fertilizer N recovery by 14 25%. Norman et al. (1992a) initially observed that fertilizer N recovery was highest when measured at owering and declined during the grain ll process, presumably from gaseous N losses from the senescing leaves. The decline in the total N content of aboveground biomass during the ripening phase was later conrmed by Guindo et al. (1994a,b). The magnitude of N loss from rice leaves is inuenced by the cultivar and environment (Stutte and da Silva, 1981). Even when all possible sources of plant and soil N are considered, native soil and fertilizer N loss occurs in all oodirrigated rice cultural systems. Use of appropriate crop and fertilizer management strategies is important to maximize N recovery efciency by rice. The fertilizer N recovery efciency in the aboveground biomass of oodirrigated rice depends on the source, time of application, and method or placement of N fertilizer. The time that N fertilizer is applied to rice varies
slightly among the various cultural management systems (e.g., transplanted, direct-dry seed, or direct-water seed) used to produce ood-irrigated rice. However, many of the principles that govern N use efciency are the same. Nitrogen fertilizer is typically applied in at least two or more split applications in most production systems. The rst application is often termed basal or preood. The basal application generally represents the largest proportion (50 75% of the total N requirement) of N fertilizer applied in any single N fertilizer application during the growing season because of its inuence on grain yield potential. In waterseeded and transplanted cultural systems, an NH4 forming N fertilizer source is generally applied to the soil surface and mechanically incorporated before the eld is ooded, applied to the soil surface and incorporated by the ood, or injected into the soil. In the dry-seeded, delayed ood system, the basal N application is made during early vegetative growth (i.e., 5-leaf stage) and followed by establishing the ood. Regardless of the cultural system, the most fundamental components of basal N management are to use an ammonium forming N fertilizer source, apply the N fertilizer on a dry soil surface, and immediately ood the soil. Proper management of the oodwater is critical for efcient uptake of basal fertilizer N. Draining the ood before rice uptake of N may result in nitrication resulting in leaching of NO3 and denitrication if the soil is reooded before N uptake is complete. In the dry-seeded, delayed ood system where basal N is applied at the 5-leaf stage, maximum uptake is achieved in about 3 weeks (Wilson et al., 1989). The time requirement for maximum uptake of basal N is likely to increase as basal N rate increases, but has not been fully evaluated. The recovery efciency of basal N (urea) in aboveground rice biomass in the dry-seeded, delayed ood system may approach 70% under optimum conditions (Norman et al., 1992a; Wilson et al., 1994). Research consistently shows that recovery efciency of basal N in this system is 40 50% (Reddy and Patrick, 1976; Norman et al., 1989, 1992b; Bollich et al., 1994; Guindo et al., 1994a,b). However, when basal N is applied to a wet soil or the ood is mismanaged the recovery efciency declines. In the delayed ood system, the recovery efciency of preplant incorporated urea is only about 25 30%, which is considerably less efcient than urea application to the soil surface immediately before ooding (Norman et al., 1989). Fageria and Baligar (2001) found N recovery efciencies ranging from 30 to 50% when basal N was applied before seeding and ooded 30 days after seeding. The recovery efciency decreased as the basal N rate increased suggesting the N loss exceeded the rate of plant uptake after ooding. When basal N is applied far in advance of ooding the concentration of soil NO3 increases until ooding and is followed by rapid denitrication (Norman et al., 1988). Considerably less research has been published on the recovery efciency of preplant incorporated or injected basal N applications in the waterseeded cultural system. Most of the published research suggests that uptake of preplant, basal N, applied as urea, by waterseeded rice is about 35 40% (Eagle et al., 2001).
84
Westcott et al. (1986) showed that the recovery efciency of basal applied N fertilizer of waterseeded rice (42%) rice was greater than that of dry-seeded rice (30%). However, the majority of published research suggests that N recovery efciency between these two seeding methods is equal or greater for dry-seeded rice. In general, recovery efciency of basal N applied to 5-leaf rice in the delayed ood system is more efcient than preplant incorporated basal N in waterseeded systems because of the shorter interval between N application and eventual plant uptake. De Datta et al. (1988) showed that basal N applications made to transplanted rice have similar recovery efciencies as those cited for waterseeded rice. Regardless of the cultural system, broadcast basal N applications are relatively efcient as long as the N is incorporated immediately after application with tillage or irrigation water and maintained in the NH4 form. The amount of N lost to ammonia volatilization or that is nitried and eventually lost to denitrication increases as the time between application and ooding increases. Numerous researchers have found that split applications of N fertilizer applied into the ood are utilized more efciently than basal applications (Reddy and Patrick, 1976, 1978; Westcott et al., 1986; Wilson et al., 1998). However, the recovery efciency of N topdressed into the ood is dependent on the rice growth stage at the time of application. Nitrogen fertilizer topdressed into the ood from panicle initiation to the early boot stage is absorbed rapidly and efciently by rice, regardless of the cultural seeding method. The recovery efciency of N topdressed at panicle initiation ranges from 45 (Westcott et al., 1986) to 82% (Wilson et al., 1994). The high recovery efciency of topdressed N is due to the application of relatively low rates of N fertilizer (, 1/3 of total seasonal N fertilizer applied), a highly developed rice root system, and the high plant N requirement at this stage of plant development. Wilson et al. (1989) showed that maximum absorption of N fertilizer topdressed at panicle differentiation occurred during the rst 3 days after application. Both NH4-N and NO3-N are absorbed efciently when topdressed at the appropriate time, but NH4-N is still more efcient (Wilson et al., 1994). Nitrogen fertilizer topdressed into the ood before panicle initiation is less efcient because the plant root system is not fully developed (Norman et al., 1993, 1994). The rate of N loss, via N loss pathways, exceeds the rate of N uptake when N is topdressed into the ood when rice is at the seedling or tillering stages. Recovery efciency primarily concerns the efcient use of fertilizer N as a means of determining best management practices to produce high yields with a minimum of fertilizer inputs and N losses into the environment. Various other means are employed to examine the internal nutrient use efciency by a crop and include agronomic efciency (AE, D kg grain kg21 fertilizer-nutrient added), physiological efciency (PE, D kg grain D kg21 nutrient uptake), agrophysiological efciency (APE, D kg grain D kg21 total nutrient uptake) and apparent recovery efciency (ARE, %) (Fageria, 1992). All four N use efciencies shown in Table IV were calculated using the formulas from Fageria et al. (1997a). The
Table IV Nitrogen Use Efciencies as Affected by N Fertilizer Rate (Reproduced with Permission from Fageria, N. K. and Baligar, V. C. 2001. Lowland rice response to nitrogen fertilization. Commun. Soil Sci. Plant Anal. 32: 14051429. Copyright Marcel Dekker, New York) Agronomic efciency (Dkg grain kg21 N fertilizer added) 35 32 22 22 18 16 13 23 0.93**
N Fertilizer rate (kg N ha21) 30 60 90 120 150 180 210 Average R

2
Physiological efciency (Dkg grain Dkg21 N uptake) 156 166 182 132 146 126 113 146 0.62*
Agrophysiological efciency (Dkg grain Dkg21 N uptake) 72 73 75 66 57 51 46 63 0.87**
Apparent recovery efciency (%) 49 50 37 38 34 33 32 39 0.82**
*,**Signicant at the 5 and 1% probability levels, respectively.
86
use of appropriate management practices to attain high recovery efciency of applied nutrient sources is desirable when examining the internal nutrient use efciency by a crop. In any experiment with a nutritional variable, plants grown at the lowest nutrient concentrations will inevitably have the highest utilization quotient because of a dilution effect (Jarrell and Beverly, 1981). All N use efciencies tend to decrease as N fertilizer rate increases (Fageria et al., 1997a; Carreres et al., 2000), especially when the fertilizer rate required to produce maximum yields is exceeded (Table IV). Fageria et al. (1997a) found that all N use efciencies, except PE, signicantly decreased as N rate increased. In most experiments with a nutritional variable, plants grown at the lowest nutrient concentrations will inevitably have the highest utilization quotient because of this dilution effect (Jarrell and Beverly, 1981). The AE for N of lowland rice in the tropics ranges from 15 to 25 kg grain kg21 N fertilizer (Yoshida, 1981; Peng et al., 1996). The PE (113 182 kg dry matter kg21 N) is higher than the APE (46 75 kg grain kg21 N uptake) because total dry matter, rather than grain, is used in the calculation. Singh et al. (1998) reported an APE of about 64 kg grain kg21 N uptake and an AE of 37 kg grain kg21 N fertilizer for 20 lowland rice genotypes which are comparable to values listed in Table IV. The ARE of N by lowland rice is related to N losses from soil via nitrication denitrication, NH3 volatilization, leaching, or a combination of these loss mechanisms (Craswell and Vlek, 1979). The efciency of utilization for grain production in the tropics is about 50 kg grain kg21 total N uptake and is relatively constant regardless of rice yield (Yoshida, 1981). Wada et al. (1986) suggested 15 17 kg N t21 grain was required to produce an average yield of 5 6 t ha21 and 19 kg N t21 was needed for higher yields. However, the absolute amount of N absorbed is not the only factor to be considered in high-yielding rice. Rather, the N supply pattern and uptake process of N throughout the whole life cycle of the plant are equally important. This is because the formation (number, size, or extent) of each rice yield component is dependant on the amount of N supply at each crucial stage for the respective yield components (Mae, 1997). 7. Nitrogen Fertilization Practices
Adequate, but not excessive, rates of N fertilizer must be applied to produce high yields, minimize production costs, and reduce the potential for polluting the environment. The required N rate is governed by yield level, soil properties, organic matter content, cropping system, disease pressure, water management, weed control, socioeconomic condition of farmers, and the price of rice. Figure 3 shows the response of lowland rice to N fertilization on a Brazilian Inceptisol (Fageria and Baligar, 2001). Rice grain yield increased as N fertilizer rate
Figure 3 Response of ooded lowland rice to N fertilizer rate on a Brazilian Inceptisol. (Reproduced with permission from Fageria, N. K. and Baligar, V. C. 2001. Lowland rice response to nitrogen fertilization. Commun. Soil Sci. Plant Anal. 32: 1405 1429. (Copyright Marcel Dekker, New York).).
increased to a point showing a signicant P , 0:01 quadratic response in each of the 3 years. The 3-year average showed that the agronomic maximum grain yield of 6465 kg ha21 was produced by application of 171 kg N ha21. Singh et al. (1998) also reported that maximum average grain yield of 7700 kg ha21 of 20 lowland rice genotypes was obtained at 150 200 kg N ha21 at the International Rice Research Institute in the Philippines. However, in fertilization experiments 90% of the maximum yield is often considered as an economical rate (Fageria et al., 1997a). Fageria and Baligar (2001) found that application of 84 kg N ha21 produced 90% of the maximum agronomic yield, but stated that most Brazilian farmers apply only about 60 kg N ha21 due to economic reasons. Maskina et al. (1988) and Aulakh et al. (2000) reported that ooded rice responded to N rates up to 120 kg ha21 on sandy loam soils in India. In the Philippines, Dobermann et al. (2000) reported 80 100 kg N ha21 was used for
88
maximal yields in eld experiments during the wet-season (rainy period), and 120 150 kg N ha21 was used during the dry-season. Two-thirds of the N was incorporated into the soil at transplanting and one-third was topdressed before panicle initiation. These authors also reported that N fertilization rates, from 1992 onward, were increased from 108 to 120 kg N ha21 during the wet-season and from 190 to 216 kg N ha21 during the dry-season. The number of split applications was also changed from two to three or four to improve N use efciency. During the dry-season, an average 60 kg N ha21 was applied as basal, 60 kg N ha21 at midtillering, 60 kg N ha21 at panicle initiation and 30 35 kg N ha21 at owering. In the wet-season, average N application rates were 40 kg N ha21 as basal, 40 kg N ha21 at midtillering, and 30 kg N ha21 at panicle initiation. Berge and Riethoven (1997) reported, based on predictions from a N fertilization model, that the optimum economic N rate for medium-duration cultivars was 100 150 kg N ha21 and 150 200 kg N ha21 for short-duration cultivars. They also reported that estimated yield levels were higher in medium-duration cultivars, but the yield difference between the two types of cultivars decreased with increasing N inputs. The simulations indicated that longer duration genotypes absorbed more soil-derived N and were less efcient with fertilizer N. In most Asian countries N fertilizer recommendations are provided for specic soils, growing seasons, and geographic regions and include information on the rates and times to apply N fertilizer (Cassman et al., 1998). In the USA, N application rates range from about 120 to 180 kg N ha21 (Hill et al., 1992; Norman et al., 1997; Wilson et al., 2001). The recommended N rate is usually specic for cultivars, soil properties, previous crop, combinations of these factors, and are determined by each rice-producing state. Cassman et al. (1998) indicated that Philippine farmers did not adhere to some N fertilizer use recommendations meant to increase N use efciency. Farmers in all rice producing areas occasionally use N fertilization methods that are known to result in poor N use efciency. This suggests that the extension, demonstration, and successful adoption, or implementation, by growers of efcient, researchbased recommendations may be more limiting than our current knowledge of N use management. In some highly mechanized areas, growers may be required to manage nutrients, like N, efciently to comply with nutrient management regulations. Nitrogen fertilizers are mostly applied in bands or broadcast. Urea and ammonium sulfate are the two main sources of inorganic N fertilizer for lowland rice. There has been an increasing interest in the use of livestock manures and green manures in crop production, because soil organic matter is one of the most important soil components for productive and sustainable agriculture as well as a main nutrient source of C, N, P, and S (Allison, 1973). Use of organic manures not only contributes to the nutrient pool, but also indirectly inuences soil chemical and physical properties. Increases in soil organic matter content occur slowly, generally taking several years to detect (Wander et al., 1994), but can
have a dramatic effect on long-term productivity (Tiessen et al., 1994). The use of livestock manure can increase rice yields and enhance the mineralization of native N (Eneji et al., 2001). Fageria and Baligar (1996) showed that use of pigeon pea (Cajanus cajan L. Millspaugh) as green manure, in combination with inorganic fertilizers, increased lowland rice yields by 15% as compared to inorganic fertilizers alone. 8. Use of N Efcient Genotypes
Nitrogen use efciency has been dened in various ways and the denitions generally account for the quantity of N accumulated in the plant, known as uptake efciency, and the quantity of N utilized for grain production, known as utilization efciency. The N utilization efciency is more important than N uptake efciency when evaluating the genetic potential among cultivars for efcient grain production, especially on soils that require high rates of N to maximize yield (Moll et al., 1982). Utilization efciency coupled with economic yield is a desired characteristic in crop plants if minimum depletion of soil N is a goal. Moll et al. (1982) recommended the development of genotypes with both high uptake and utilization efciencies. Use of N efcient genotypes is an important N management strategy for oodirrigated rice. An ideal genotype could be the one that absorbs relatively large amounts of N from soil and fertilizer, produces a high grain yield per unit of absorbed N, and stores relatively little N in the straw (Isfan, 1993). Rice cultivars can differ in root growth and morphology (Slaton et al., 1990) and nutrient uptake rates (Teo et al., 1995). Fageria and Barbosa Filho (2001) evaluated N use efciency of eight lowland rice genotypes grown on an Inceptisol of Brazil. Nitrogen uptake in grain, NHI, and N use efciency were signicantly different among genotypes. Nitrogen harvest index is a measure of N partitioning in rice, which provides an indication of how efciently the plant utilized the acquired N for grain production. Genotypes that produced the lowest grain yield had the lowest NHI, whereas genotypes that produced the highest grain yields also had the highest NHI. Genetic variability for NHI exists within the small grain genotypes and high NHI is associated with efcient utilization of N (Rattunde and Frey, 1986). Thus, the variation in NHI is a characteristic of genotype and this trait may be useful in selecting rice genotypes for higher grain yield. Nitrogen use efciency varies signicantly among lowland rice genotypes (De Datta and Buresh, 1989; Fageria and Barbosa Filho, 2001). Many researchers have found signicant variations of N use efciency among lowland rice genotypes (Broadbent et al., 1987; Buresh et al., 1988; Fageria et al., 1997a). Such differences may be related to many physiological processes such as absorption, NO3 reduction efciency, N remobilization, translocation, assimilation, and storage (Isfan, 1993).
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Nutrient uptake efciency and nutrient utilization efciency are governed by different physiological mechanisms (Baligar and Fageria, 1997, 1999; Baligar et al., 2001). Plant physiological factors associated with N use efciency include (i) root proliferation; (ii) NO3 uptake efciency; (iii) N translocation efciency; and (iv) nitrate reductase enzyme efciency (Duncan and Carrow, 1999). Each stage of N assimilation is under genetic control. Nitrate is reduced to NH3 by nitrate (NR) and nitrite reductase (NiR). These metalloenzymes require cofactors: molybdenum-pterin for NR and Fe-containing hydrochlorine (Siroheme) for NiR. The NR enzyme is localized in the cytosol, whereas NiR is found in leaf and root tissue plastids (Duncan and Carrow, 1999). Rice hybrids have a 10 15% yield advantage over conventional rice cultivars (Yang and Sun, 1988). The yield advantage of hybrid rice is presumably related to a greater total N uptake and internal use efciency of N. The total N uptake by hybrid rice is greater than that of conventional cultivars, especially from transplanting to tillering and from panicle emergence to grain lling stages (Yang, 1987). Hybrid rice takes up about 15 20% of its total N after heading and consistently responds to N fertilizer applications made at owering compared to only 6 7% for conventional cultivars (Yang, 1987). Hybrid rice had a greater AE than conventional rice (Lin and Yuan, 1980; Yang, 1987). This increased N use efciency is not due to greater internal N uptake in dry matter production (dened as dry matter produced per unit N accumulated in the plant) (Yang et al., 1999). The primary factors contributing to the higher N use efciency of hybrid rice are higher N recovery efciency, greater root N absorption potential, greater shoot-N use capacity, and greater N remobilization efciency.
C. PHOSPHORUS
Phosphorus deciency is the second most important nutritional disorder of lowland rice, especially in the highly weathered acidic soils of the tropics that contain large quantities of Al and Fe oxides (Wells et al., 1993; Fageria and Baligar, 1996; Baligar and Fageria, 1997; Fageria et al., 1997a,b; Seneweera and Conroy, 1997). The highly weathered tropical soils are primarily Oxisols and Ultisols that have low total and available P content, and also have a high P xation capacity (Fageria et al., 1991). Plants rarely absorb more than 20% of the total fertilizer P applied due to xation (Friesen et al., 1997). The uptake of P by plants is governed by the ability of a soil to supply P to plant roots and by the desorption characteristics of the soil (Roy and De Datta, 1985). The supply of P to plant roots depends on the concentration of inorganic P in the soil solution and on the capacity of the soil to maintain this concentration. Application of P is often essential for protable agricultural production. However, accumulation of soil P in excess of crop needs has the potential to enrich surface runoff with P that can cause eutrophication. Eutrophication has
been identied as the main cause of impaired surface water quality (US Environmental Protection Agency, 1996). Eutrophication restricts water use for sheries, recreation, industry, and drinking due to the increased growth of undesirable algae and aquatic weeds and also due to oxygen shortages caused by their death and decomposition (Abrams and Jarrell, 1995; Sharpley et al., 1999). Hence, appropriate management of P is an important aspect not only for higher rice yields, but also for environmental protection. 1. Chemistry in the Soil
The transformations and chemistry of P in ooded soils have been thoroughly reviewed by Ponnamperuma (1972) and Sanyal and De Datta (1991); hence, only the primary factors affecting P availability will be discussed briey here. Soil P can be divided into the organic and inorganic fractions with the inorganic P fraction being recognized as the pool that controls P availability to plants. The inorganic P fraction is made up of at least ve basic categories including aluminum phosphates (Al-P), iron phosphates (Fe-P), calcium phosphates (Ca-P), reductant soluble or the occluded phosphates (RS-P) and the readily available 2 22 2 orthophosphate forms (PO3 4 , HPO4 , or H2PO4 ) of P (Chang and Jackson, 1957). The predominate form of orthophosphate present in the soil solution changes with soil pH. The reduction process that occurs in the soil following ooding normally increases the P availability to rice. Therefore, on many soils P availability is not a yield-limiting factor for rice. Rice yields may not respond to P fertilization, but upland crops, like corn and soybean, growing on the same soil might show dramatic responses to P fertilization. Phosphorus deciency of upland crops following rice in the rotation has been reported for several crops (Willett, 1982; Wells et al., 1995). The alternate anaerobic aerobic soil conditions reduce the availability of P to upland crops following rice in the rotation. The soils P sorption capacity and the bonding energy of P increases under alternate anaerobic aerobic conditions (Sanyal and De Datta, 1991). Flooding decreases the crystallinity of ferrous hydroxides, which increases their sorption capacity, increases the insoluble Fe-P fraction, and reduces P desorption. Iron phosphates are the primary source of P to lowland rice because their availability is quickly affected by the anaerobic conditions created by ooding (Patrick and Mahapatra, 1968; Ponnamperuma, 1972; Goswami and Banerjee, 1978). Willett (1986, 1989) reported that P availability increases after ooding from the (1) reductive dissolution of ferric oxides; (2) the liberation of sorbedand RS-P; (3) changes in soil pH that increase the solubility of Fe-, Al-, and Ca-P; and (4) the desorption of surface P. The relatively insoluble ferric phosphates are reduced to the more soluble ferrous phosphates resulting in hydrolysis of P compounds. On acid soils, soil pH generally increases following soil
92
submergence and increases the solution concentrations of Fe and Al phosphates. Flooding the soil usually reduces hydrated ferric oxides to ferrous hydroxides, which releases part of the RS-P (Patrick and Mikkelsen, 1971). Turner and Gilliam (1976) illustrated that ooding the soil signicantly inuenced P availability on calcareous soils where Ca-P predominates over Fe-P. They found that increased P availability in ooded or saturated soils occurred, at least partially, from the decreased tortuosity which increased P diffusion rate by 10- to 100-fold. However, Sah and Mikkelsen (1986a) observed the Ca-P fraction of some, but not all, clay soils in California increased for several weeks after ooding from the formation and precipitation of insoluble Ca-P, which would decrease the availability of P after ooding. An interesting aspect of the P chemistry of ooded soils is that more P is released from the soil into the solution under reduced conditions than under oxidized conditions if the soil solution is initially low in phosphorus, but reduced soils also have a greater sorption capacity for P (Patrick and Reddy, 1978). Khalid et al. (1979) and Roy and De Datta (1985) suggested that rice required 0.12 0.20 mg P L21 in the soil solution for optimum growth. Hossner et al. (1973) concluded that the minimum soil solution P concentration required to produce 90% rice yield was . 0.10 mg P L21. Several forms of soil Fe are highly correlated with P sorption (Willett and Higgins, 1978; Khalid et al., 1979). Soil P released under reduced conditions has been related to oxalate-extractable Fe (Khalid et al., 1979; Shahandeh et al., 1994). Fox and Kamprath (1970) and Evans and Smillie (1976) reported that soil clay and available Fe content strongly inuenced P sorption under aerobic soil conditions. Ammonium oxalate extractable Fe was highly correlated with P sorption under anaerobic conditions (Willett and Higgins 1978; Khalid et al., 1979; Shahandeh et al., 1994). Shahandeh et al. (1994) reported that 84% of the added P was sorbed under anaerobic conditions when ammonium oxalate extractable Fe was , 3000 mg Fe kg21. Oxalate extracts poorly crystalline Fe oxides (Campbell and Schwertmann, 1984), which are the most reactive Fe oxides in the soil because of their small size and high surface area (Shahandeh et al., 1994). Although our understanding of P availability to rice grown under ooded soil conditions has increased during the past 40 years, this remains an area that requires additional research. The general relationships of P availability are well characterized, but need to be better dened in regards to specic soil physical and chemical properties and correlated to plant uptake of P. 2. Functions and Deciency Symptoms
Phosphorus is one of the major essential nutrients needed for the growth and reproduction of higher plants. Phosphorus is required for the synthesis of phospholipids, nucleotides, adenosine triphosphate (ATP), glycophosphates,
and other phosphate esters. Plant growth and yield are dramatically reduced by P deciency because P is a component of high-energy compounds like ATP and is an essential component of the genetic material required for seed production. Phosphorus deciency symptoms occur primarily on seedling rice at the onset of tillering when rice begins to rapidly accumulate dry matter. Symptoms include severe stunting with plants having erect and dark green leaves. Phosphorus deciency reduces seedling height, tiller number, stem diameter, leaf size, and leaf duration. When P is decient, cell and leaf expansions are retarded more than chlorophyll formation. Thus, the chlorophyll content per unit leaf area increases, but the photosynthetic efciency per unit of chlorophyll decreases (Marschner, 1995). Phosphorus is not a constituent of chlorophyll; hence, the concentration of chlorophyll of P decient rice becomes comparatively high and the leaf color changes from green to dark green. If a P deciency persists, the older leaves may turn an orangish color and desiccate from the leaf tip back towards the base. Rice maturity can be delayed by as much as 10 12 days by P deciency (Fageria, 1980). Phosphorus deciency symptoms of many crops include a reddish or purple tint on leaves due to the accumulation of anthocyanins (Hewitt, 1963). However, the leaf purpling symptom has not been observed in P decient rice (Fageria and Barbosa Filho, 1994; Fageria and Gheyi, 1999). Phosphorus decient plants are more susceptible to some rice diseases. In Arkansas, excessive brown spot (Bipolaris oryzae) is commonly observed on nutritionally stressed rice. Phosphorus fertilization has also signicantly increased rice root growth, the number of panicles, and grain weight of rice grown on P decient soils (Fageria and Gheyi, 1999). When P is decient, rice does not respond to the application of N, K, or other fertilizers. Color photographs depicting typical P deciency symptoms of rice are available in various publications (Wallace, 1961; Mueller, 1974; Cheaney and Jennings, 1975; Ishizuka, 1978; Yoshida, 1981; Fageria, 1984; Bennett, 1993; Fageria and Barbosa Filho, 1994).
3. Critical Level in Plant and Uptake Phosphorus is a mobile element inside the plant; hence, the P concentration of individual leaves generally declines as leaf age increases. The top leaves have the highest P concentration and the bottom leaves have the lowest P concentration, especially when plant available P is limited (Westfall et al., 1973). Sims and Place (1968) reported that tissue P concentration varied less across plant development stages than N concentrations, which is generally true when P is not a growth-limiting factor. During early vegetative growth, the P concentration in rice tissue increases as P fertilizer application rates increase (Fig. 4). However, the difference in tissue P concentrations between P fertilizer rates diminishes as plant development progresses from the vegetative to reproductive growth stages.
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Figure 4 Phosphorus concentration in rice plant shoots at different growth stages in Brazil and Arkansas, USA.
Whole plant P concentration has been observed to either increase or decrease between the active tillering stage and panicle initiation depending on the soil pH, P fertilization, or both. On acid soils (pH 5.0 6.5) such as those represented by the data from Brazil (Fageria, unpublished data, 1997) in Fig. 4, tissue P concentrations tend to be high (2.5 4.5 g P kg21) during active tillering and decrease or remain constant as rice progresses into reproductive growth. On alkaline soils (pH . 7.0) as represented by data from Arkansas (Slaton, unpublished data, 2001) in Fig. 4, whole plant P concentrations are low (1.0 2.0 g P kg21) at the onset of tillering, increase until panicle initiation, and then are stable until owering. These described trends are presumably related to the availability of P after ooding as affected by P fertilization rate and the predominate forms of soil P. Nevertheless, tissue P concentrations remain nearly
constant from panicle initiation until owering on most soils. In studies examining a range of P fertilizer rates, differences in tissue P concentrations are seldom observed during this stage of reproductive growth, although plant P content may vary if different P fertilizer rates were applied. Because differences in tissue P concentration are not always evident, a critical P content at specic rice growth stages, rather than concentration, should perhaps be used to evaluate the P nutritional status of rice. After owering the lling rice grain becomes a strong sink for P and the straw P concentration declines. A wide variation exists among the critical tissue P concentrations for rice reported in the literature. Some of the variation can be attributed to differences among the production practices, cultivars, soils, and environments common to the various rice producing regions around the world. Other, more obvious, differences are due to the growth stage, plant part sampled, or the growth parameter used to establish the critical P concentration. In general, the critical tissue P concentrations for rice during vegetative growth range from 1.0 to 2.0 g P kg21. Yoshida (1981) reported that 2.0 g P kg21 in the Y-leaf (most recently matured leaf) was needed to realize the maximum tillering rate. When leaf blade P concentration was , 0.3 g P kg21 tillering did not occur. De Datta (1981) suggested that 1.0 g P kg21 in the rice leaf blades at active tillering was the critical concentration. Fageria et al. (1997a) reported that adequate P concentrations in rice shoots ranged from 2.5 to 4.5 g kg21 at 75 days of age (about panicle initiation). The decrease in P concentration during early plant development is probably related to the rapid increase in dry matter accumulation, which dilutes the P in the tissue until the P uptake rate can match dry matter accumulation. In general, it appears that Y-leaf or whole plant P concentrations during vegetative growth . 2.0 g P kg21 are sufcient for optimum rice growth and yield production, but even tissue P concentrations as low as 1.0 g P kg21 may be adequate to produce maximum yields provided that P availability and tissue P concentrations increase over time. The concentration of P in rice straw during reproductive growth or maturity is not commonly used to diagnose P deciencies since P typically limits early season vegetative growth. Slaton et al. (2001a) showed that ag-leaf P concentrations at the late-boot stage are generally above 2.0 g P kg21 for highyielding rice in Arkansas. Similarly, Dobermann et al. (1998) suggested that the rice ag-leaf P concentration should be . 1.7 g P kg21 for the production of 7 t ha21. In Asia, straw P concentrations , 0.6 g P kg21 at maturity indicate that P was decient (Dobermann et al., 1998). This critical concentration may be appropriate for rice grown in Asia, but appears too low for rice grown in the USA. Rough rice (grain hulls) P concentration averages about 3.0 g P kg21 (Nelson, 1980), but Dobermann et al. (1998) found that P concentration of rice grain produced in Asia ranged from 1.5 to 2.5 g P kg21. The P concentration of rice grain is relatively stable and is likely under genetic control, but varies
96
slightly among cultivars, which may partially explain the differences between grain P concentrations cited by Nelson (1980) and Dobermann et al. (1998). Batten et al. (2000) found that the P concentrations of brown rice of cultivars from Australia and other countries averaged 3.3 and 3.5 g P kg21, respectively. Moreover, they determined that brown rice of the various rice genotypes analyzed differed in their mineral composition when grown under the same environment and could often be grouped based on growth characteristics or their country of origin. Brown rice P concentration was negatively correlated with grain yield and harvest index, but positively correlated with the concentrations of some other elements in brown rice. These ndings are signicant because it shows the nutrient value of brown rice can be improved simply through breeding. Total P uptake by lowland rice during the growing season increases with plant development. The total, aboveground P uptake by a high-yielding rice crop may approach 60 kg P ha21, but more commonly ranges from 25 to 50 kg P ha21 with 60 75% of the plant total P contained in the panicles at maturity. Seasonal P uptake and dry matter accumulation tend to follow similar patterns. The accumulation of P is closely related to plant age. Fageria et al. (1997b) showed that plant age accounted for 70% of the variability in seasonal P accumulation. On P decient soils, total P uptake also increases as P fertilizer rate increases (Table V). In the straw, P accumulates with increasing plant age until owering. After owering, during the ripening period, straw P content decreases as P is translocated from the straw to the developing grain. A signicant portion of P is either translocated from the root system or absorbed between owering and
Table V as Phosphorus Accumulation in the Shoot and Grain of the Lowland Rice Cultivar Javae Inuenced by Plant Age and P Application Rates on a Brazialian Inceptisol (Fageria unpublished data, 1997) Days after sowing (growth stage) (kg P ha21) P Fertilizer rate (kg P ha21) 0 87 175 262 350 437 Average 18 (IT) 0.20 0.33 0.43 0.53 0.60 0.61 0.45 35 (AT) 0.6 1.1 1.5 2.1 2.3 1.7 1.5 63 (IP) 4.6 6.4 7.4 7.8 8.0 7.0 6.9 81 (B) 9.9 13.8 13.5 15.3 14.4 13.0 13.3 91 (F) 12.8 16.5 18.1 22.4 18.4 19.0 17.9 119 (PM) 4.5 4.6 6.5 8.7 6.4 7.5 6.3 119 (grain) 16.4 19.0 20.1 25.1 25.0 23.1 21.4
Total 20.1 23.6 26.6 33.7 31.4 30.6 27.8
IT, initiation of tillering; AT, active tillering; IP, initiation of panicle; B, booting; F, owering, PM, physiological maturity. Phosphorus rates were broadcast.
maturity to satisfy grain P requirements. About two-thirds of the total plant P at maturity can be accounted for in the aboveground biomass at owering (Table V). The straw P content at maturity is only about one-third of the total P content at owering and the grain content at maturity. The average P harvest index [Grain P/ (Straw Grain P)] generally ranges from 0.60 to 0.75. The harvest of rice grain removes a signicant portion of the total P taken up during the growing season. Thus, a major portion of P accumulated by the rice crop cannot be recycled for use by crops following rice in the rotation even though the rice straw is incorporated back into the soil.
4. Phosphorus Use Efciency Phosphorus use efciencies of rice are generally higher than use efciencies for K and can be higher than those of N, depending on the soil. Phosphorus use efciencies calculated based on results obtained in eld experiments conducted at the National Rice and Bean Research Center of EMBRAPA are presented in Table VI. All of the P use efciencies for this soil were greater compared to the same efciencies for N and K (Baligar and Fageria, 1997). Dobermann et al. (1998) reported the AE for P ranged from 0 to 114 kg grain kg21 P fertilizer applied. Singh et al. (2000) reported that APE in lowland rice varied from 235 to 316 kg grain kg21 P. Similarly, Witt et al. (1999) reported an APE value of 385 kg grain kg21 P when all production factors were at normal levels. The internal use efciency of P as reported by Dobermann et al. (1998) ranged from 282 to 724 kg grain kg21 total P uptake. Approximately 4 5 kg P is required to produce 1000 kg of rough rice grain (Fageria et al., 1997b). The efciency of P
Table VI Phosphorus Use Efciency by Lowland Rice Genotypes Agronomic efciency (Dkg grain kg21 P fertilizer added) 93 79 67 98 54 95 81 Physiological efciency (Dkg grain D kg21 P uptake) 360 533 363 407 484 491 440 Agro-physiological efciency (D kg grain D kg21 total P uptake) 158 226 234 187 127 252 197
Genotype Alianc a CAN 5751 CAN 6804 CAN 7238 CAN 7268 Metica1 Average
From Baligar and Fageria, 1997. Formulas used to calculate the different P use efciencies are given in section III.B.6
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use varies among rice genotypes. Genetic variability among rice genotypes has been reported by various authors (Ponnamperuma 1976a; Fageria, 1992; Fageria and Baligar, 1993; Baligar and Fageria, 1997, 1999; Fageria et al., 1997a; Fageria and Gheyi, 1999). 5. Phosphorus Fertilization Practices
Indiscriminate use of P fertilizer increases the cost of production, may reduce rice yields on some soils, and can contribute to water quality problems. The principal phosphate fertilizers commonly used to fertilize rice are the highly water-soluble fertilizers like single and triple-super phosphates, diammonium phosphate, and sometimes monoammonium phosphate. Soils used for lowland rice production commonly have low soil test P values, but may or may not respond to P fertilization (Shahandeh et al., 1994; Wilson et al., 1999). Routine soil test methods that extract plant-available nutrients may not provide a reliable estimate of the P available to lowland rice, but the use of calibrated soil test data is still the best criteria for making P fertilizer recommendations for most crops including rice. The anaerobic soil environment of ood-irrigated rice signicantly alters the availability of P and routine soil test methods may not adequately represent nutrient availability to rice. One of the most important factors affecting P fertilizer recommendations from calibration studies is the method used to extract soil P. A number of different extractants including Bray-1, Bray-2, Mehlich 1, Mehlich 3, and NaHCO3 (Olsens) have been used in different parts of the world to assess the status of plant-available soil P (Kamprath and Watson, 1980; Sharply et al., 1994). Many of these extractants tend to under- or over-estimate P availability to upland plants (Kamprath and Watson, 1980) and their ability to accurately predict the P fertilizer requirement of ood-irrigated rice are further compromised by the anaerobic soil conditions used for its production. In the United States, each of the primary rice producing states use a different extractant to estimate P fertilizer requirements of rice (Norman et al., 2003). The various extractants are used because no single extractant has shown a signicant advantage for making P recommendations on ood-irrigated rice, but the extractants have been calibrated for the soils and upland crops grown in each of the states. Sanyal and De Datta (1991) suggested that NaHCO3 (Olsen P) is perhaps the best routine method for predicting rice response to P as it is better correlated with the extraction of Fe-P. Fageria et al. (1997b) evaluated the Mehlich 1 soil P test to predict lowland rice response to P fertilization on a Brazilian Inceptisol. Based on relative grain yield, ranges of Mehlich 1 soil test P were categorized as very low (0 2.6 mg P kg21), low (2.6 8.8 mg P kg21), medium (8.8 13.0 mg P kg21), or high (. 13.0 mg P kg21). The amounts of broadcast P needed to increase soil P
concentration at very low, low, medium, and high soil classication categories were 444, 292, 220, and 80 P ha21, respectively. A medium soil test P was required for the production of . 95% relative yield. The rate of banded fertilizer P required to produce maximum yield was 66 kg P ha21 for the very low and low soil categories, 44 kg P ha21 for the medium category, and 22 kg P ha21 for the high category. Fageria et al. (1991) characterized the chemical properties of lowland rice soils of Brazil and reported a 65-fold range in Mehlich 1 extractable P concentrations in the 0- to 20-cm soil depth among 23 different municipalities. Similarly, Cochrane et al. (1984) reported great variation in Bray 2 extractable P of lowland soils of tropical America. Annual crop yield responses to P fertilization were expected on 40% of the soil samples that contained low (, 3 mg P kg21) to medium (3 7 mg P kg21) concentrations of extractable P. Fageria (1980) also reported P fertilization signicantly increased the yield of lowland rice grown in the Goias State of Brazil. Maximum grain yields were obtained when P was broadcast applied at 175 kg P ha21 as triple superphosphate. Dobermann et al. (2000) reported that at IRRI, 26 kg P ha21 is normally applied to obtain maximum yield of ooded rice. When P fertilizer is recommended in the USA, rates of 10 40 kg P ha21 are normally sufcient to produce maximum rice yields (Norman et al., 2003). Soils capable of xing large quantities of P required 97 (Fageria, 1980) to 175 kg P ha21 (Fageria et al., 1997b) to produce optimum yields when P fertilizer was broadcast applied, but only 22 44 kg P ha21 when P fertilizer was banded. Use of the most efcient P application method to reduce the optimum P rate is critical to offset the cost of P fertilizer and increase P fertilizer use efciency. The P fertilizer rates cited by Fageria et al. (1997b) required to produce maximum yields in Brazil are relatively high and suggest that low soil P content and high P xation capacity may be associated with the high P requirement in Brazilian acidic lowland soils. The time and placement of fertilizer P is critical for optimum uptake, especially on P decient soils. Patrick et al. (1974) showed that P placed with the rice seed during drilling was superior to broadcast application 2 weeks after seeding. Rice yield declined as the time of P application was delayed. They also generalized that broadcast preplant-incorporated P application would be equally effective as P drilled with the seed. However, broadcast application of P at the 5-leaf stage increased tissue P concentration, P uptake, and grain yield more than P broadcast applied to the soil surface at seeding on an alkaline soil in Arkansas (McGee et al., 2002) suggesting that xation potentially reduces P availability on some soils in a very short time. The prevention of early season P deciency is critical for the production of high yields. When P is decient, rice yield response to P fertilizer declines as the time of P fertilization is delayed (Patrick et al., 1974; Slaton et al., 1998).
100
D. POTASSIUM
Rice does not generally respond to K fertilization to the degree noted for either N or P. Many soils used for the production of continuous rice or rice wheat rotations can be cropped for extended periods without needing supplemental K to maintain crop production (Dobermann et al., 1996b). Rice grown in rotation with legumes, like soybean, may require annual K inputs due to the greater K requirement of the legume crop. Although annual K fertilization may not be required, the aboveground K content of rice is equal to or greater than the plant N content and greater than all other essential nutrients. Direct K fertilization of rice has produced grain yield increases ranging from 0 to 47% with yield responses of 0 10% being the norm (Dobermann et al., 1996b). Rice is highly efcient in scavenging plant available soil K because of its brous root system and the increase in K availability after ooding. However, on some soils, K deciency of rice may occur if rice and rotation crops are grown without regular applications of K fertilizer to replace the K removed by the harvested crops. Prior to the early 1990s, K deciencies of rice were rare in the rice producing areas of the USA. However, K deciency is now recognized as an annual problem on many soils as rice and rotation crop yields have increased, soils have been mined of K, and production practices have changed (Slaton et al., 1995; Williams and Smith, 2001). Many of the soils used for rice production throughout the world have low cation exchange capacities or are highly weathered which makes them poor reservoirs of plant available K. In highly weathered soils, the total soil K content may be quite low because of K decient parent materials and the climate. The high rainfall and warm temperatures common to the tropical rice growing areas have hastened the release and leaching of soil K over time (Tisdale et al., 1985). In Arkansas, approximately 20% of the soil samples analyzed for rice or irrigated-soybean, the primary crop rotated with rice, have soil K concentrations , 200 kg Mehlich 3 K ha21 (DeLong et al., 2001). The chemistry of soil and fertilizer reactions involving K is less complicated than the chemical and biological reactions that occur when N and P fertilizers are added to the soil. Potassium is not complexed into soil organic matter, susceptible to gaseous losses, or subject to precipitate into forms that are not plant available. Essentially all the soil K is associated with the soil mineral fraction (Foth and Ellis, 1988). The availability of soil K is inuenced by soil K concentration, soil texture, soil pH, cation exchange capacity, temperature, soil moisture, soil aeration, yield level, and root growth patterns. Unlike, N, P, and Zn, relatively few studies have been published regarding efcient K fertilization practices, diagnosis of K deciency, and correction of K deciency. Since De Datta and Mikkelsen (1985) reviewed K nutrition of rice, research has begun to increase our knowledge of the K nutritional requirements of rice.
NUTRIENT MANAGEMENT FOR IMPROVING LOWLAND RICE101
1.
Chemistry in the Soil
The soil minerals that are considered the most important K sources in the soil are the primary aluminosilicates that include K feldspars and micas and secondary aluminosilicates like illite. Potassium is released by the weathering of these minerals. The rate of weathering or K release depends on the mineral properties and predominate environmental conditions (Tisdale et al., 1985). Soil K can be divided into four fundamental categories including: (1) 0.1 0.2% as soil solution K, (2) 1 2% as exchangeable K, (3) 1 10% as nonexchangeable K, and (4) 90 98% as mineral K (Barber, 1995; Brady and Weil, 1996). Potassium ions move from one category to another whenever the removal or addition of K disturbs the equilibrium between these soil K pools. Equilibration between the soil solution and exchangeable K pools is rapid and usually complete within hours. However, the equilibration time between the nonexchangeable and exchangeable K is much slower, requiring days or even months. The conversion of K from the mineral form via weathering is extremely slow and varies among the soil K minerals. The weathering process is extremely slow and has little signicance in supplying plants with K during a single season (Barber, 1995). Most of the K used by plants during a single season comes from the soil solution and exchangeable K pools and is supplemented by K released from the nonexchangeable pool. The quantity of nonexchangeable K in soils depends on clay content and type of clay minerals (Sparks and Huang, 1985). Potassium is considered relatively immobile in the soil and moves primarily by diffusion in the soil plant system, especially for upland crops. Teo et al. (1995) found that diffusion, mass ow, and contact exchange accounted for , 57.8, 42, and , 0.3%, respectively, of K uptake by ooded rice. Although considered immobile, a signicant amount of K can be lost via leaching on some soils following displacement from the exchange complex after ooding. Leaching is a signicant problem in the humid tropical regions having acid soils with low cation exchange capacities. Liming an acid soil to raise its pH can reduce leaching losses of K because of the complementary ion effect and increasing the soil CEC (Brady and Weil, 1996). Signicant vertical K movement, via leaching, from the 0 20 cm soil depth to lower depths (20 40, 40 60 and 60 80 cm) has been reported after harvesting only four lowland rice crops on a Brazilian Inceptisol (Fageria et al., 1990c). The soil solution concentration of K increases after ooding. Soil reduction increases the soil solution concentrations of Fe, Mn and other soil cations, which displace K from the cation exchange sites into the soil solution. The result is an increase of K in the soil solution, where K can either be absorbed by rice plants or, on permeable soils, leached to depths below the rice root system (Patrick et al., 1986; Wells et al., 1993). The duration of increased availability of K has not been adequately characterized across an array of soils, but is believed to be relatively short lived. Preliminary data from Arkansas suggests that soil solution K
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concentrations of a DeWitt silt loam (ne, smectitic, thermic, Typic Albaqualfs) cropped to rice are highest within days after ooding and rapidly decline for 4 5 weeks, due presumably to plant uptake, until an equilibrium is reached and maintained until the ood is drained for harvest (Slaton, unpublished data, 2001). The initial increase in soil solution is considered an advantage for plant uptake, but also a disadvantage if the K is leached. Sandy soils high in organic matter and reducible Fe and Mn generally have higher solution K concentrations than soils with higher clay contents (Yoshida, 1981). The leaching loss of signicant amounts of soil K could result in K deciency later in the growing season if adequate K is not absorbed during vegetative growth or K is not supplemented shortly before the onset of reproductive growth.
2.
Functions and Deciency Symptoms
Potassium deciency symptoms initially appear on the lower, oldest rice leaves because K is highly mobile in the plant. The onset of K deciency of rice is difcult to diagnose because the only initial difference between K sufcient and decient rice is the color of the lower leaves. This deciency symptom can easily be confused for N deciency. Potassium deciency symptoms include stunted plants with little or no reduction in tillering, droopy and dark green upper leaves, and chlorosis of the interveinal areas and margins of the lower leaves starting at the leaf tip. Leaf tips will eventually die and turn brown with the progression of severe K deciency. The droopy leaves associated with K deciency are not always noted on K decient rice because Na may substitute for K in osmotic regulation functions. Potassium deciency reduces grain size and weight resulting in a direct yield loss. In the USA, K deciency symptoms are seldom observed during vegetative growth. More commonly, K deciency symptoms are observed during early reproductive growth beginning at panicle initiation. Potassium decient rice commonly has high levels of several diseases that infest the leaves, stems, and panicles (Slaton et al., 1995). As K deciency progresses, rice usually develops severe disease infestation due to the plants reduced ability to resist infestation. Diseases that are normally insignicant problems, such as brown leaf spot and stem rot (Sclerotium oryzae), may become severe, in addition to common diseases such as rice blast (Pyricularia grisea). The lodging associated with K decient rice may be related to the increased incidence and severity of stem diseases. Potassium is known to play an important role in the lignication of vascular bundles, a factor that contributes to the higher susceptibility to lodging and disease of K decient plants. The yield loss of K decient rice may actually be a combination of losses from insufcient K nutrition and losses caused directly by diseases that infest K decient rice. Limited research suggests that yields of rice low or decient in K do not respond
to K fertilizer applications made after the panicle differentiation growth stage (Slaton et al., 2001b). Thus, when K fertilizer is required for maximum yield production it must be applied during vegetative growth. Research has not been conducted to determine if a portion of the yield loss attributed to K deciency can be prevented or recovered by controlling or suppressing the diseases with timely fungicide applications. Additional research efforts are needed to verify whether application of K fertilizer to K decient rice during the reproductive growth stage is benecial. Color photographs of K deciency symptoms of rice are available in various publications (Wallace, 1961; Mueller, 1974; Cheaney and Jennings, 1975; Ishizuka, 1978; Yoshida, 1981; Fageria, 1984; Bennett, 1993; Fageria and Barbosa Filho, 1994; Wilson et al., 2001).
3. Critical Level in Plant and Uptake The concentration and content of nutrients in plants are important parameters used to evaluate the nutritional status of a crop. Knowledge of tissue nutrient concentrations at specic crop growth stages allow for the diagnosis and correction of nutritional disorders. The K nutritional status of plants is easily determined by routine analysis of plant tissues. However, the critical K concentrations of rice are not particularly well dened. Potassium has not historically been a major yield-limiting nutrient in many rice-producing areas and research efforts have been focused on other nutrients like N, P, and Zn. Thus, a somewhat limited database concerning K nutrition of rice is available in the literature. As grain yield increases the demand for plant nutrients, particularly K, also increases (Fageria et al., 1997a). The K concentration of whole rice plants decreases with plant age (Sims and Place, 1968). The critical nutrient concentration range required for optimum yields also decreases with plant age. Tanaka et al. (1977) compared the physiological response of 17 seedling crop species, including rice, to K nutrition. Rice was characterized as having a high capacity to absorb and deplete K in the growth medium. Fageria et al. (1997a) reported that tissue concentrations ranging from 15 to 40 g K kg21 in whole, aboveground plant samples at 75 days of age (about panicle initiation) were adequate for rice. During the vegetative growth phase, tillering stops when the K concentration in the leaf blade is , 5.0 g K kg21 (Yoshida, 1981). Kiuchi and Ishizaka (1961) reported that mature leaves should contain 20 g K kg21 at the booting stage to maximize the number of grains per panicle and decrease spikelet sterility. Fageria (1986) found K concentrations of 17 g K kg21 in the rice straw and 2.6 g K kg21 in the rough rice grain were sufcient at maturity. Straw K concentrations , 10 g K kg21 at maturity certainly indicate a K deciency (De Datta, 1981). Increasing the K concentration of rice stems and leaves is easily
104
done through the application of increasing rates of K fertilizer. Although luxury uptake of K may occur in rice vegetation, the K concentration of rice seed remains relatively constant regardless of K fertilization. The K concentration of rice seed ranges from 2.5 to 3.0 g K kg21, which is roughly equal to the grain P concentration (Fageria, 1986; Dobermann et al., 1998; Batten et al., 2000). Dobermann et al. (1996b) showed that K concentration of rice grain was stable and varied by less than 0.3 g K kg21 among K fertilizer treatments. Differences in K nutrition or availability are exhibited by a wide range of K concentrations in the rice straw, by reduced grain yields (grain K content), or both. Rice absorbs the majority of its K during the vegetative and early reproductive growth stages. Hirata (1995) reported that 75% of the total K uptake at maturity is absorbed prior to the booting stage and almost no absorption occurs between owering and maturity. While it is generally agreed that the majority of K uptake occurs before owering, some K is certainly absorbed during the ripening period. A major portion of the K absorbed before anthesis remains in the stems and leaves (Hirata, 1995). Approximately 80 90% of the aboveground K content of rice is found in the leaves and stems at maturity (Ishizuka and Tanaka, 1952a; Dobermann et al., 1996a; Wilson et al., 2001). The percentage of total K removed in harvested grain declines when luxury consumption occurs (Fageria, 1986). The grain concentrations of P and K are nearly equal in rice seeds, but, in contrast to P, a comparatively small amount of the total plant K content is translocated and stored in the grain. A single lowland rice crop may contain between 200 and 300 kg K ha21 at maturity (De Datta and Mikkelsen, 1985). If the rice straw is not physically removed from the eld, the majority of K is recycled back into the soil. If the rice straw is removed K fertilization practices must be altered to prevent from depleting the soil K. Rice hulls, which are separated from the grain during the milling process, comprise about 20% (by weight) of the harvested rough rice and have near equal K concentrations (, 2.8 g K kg21) as the brown rice. Most research suggests that approximately 40 kg K (straw plus grain content) is required to produce 1 t of rice grain (Fageria and Baligar, 1996; Fageria et al., 1997b).
4.
Potassium Use Efciency
Nutrient absorption and utilization capacity of a crop variety are important aspects for the improvement of crop yield and the reduction of production costs. Dobermann and Fairhurst (2000) suggested that the recovery efciency of applied K fertilizer is about 50%, but the uptake efciency is usually lower when all K is basal applied or higher when K fertilizer is topdressed in two or more split applications. Baligar and Fageria (1997) found that the recovery efciency of K fertilizer ranged from 37 to 58% and varied among lowland
rice genotypes (Table VII). The average fertilizer recovery for K is similar to N (39%, Table III) and typically higher than for P (, 10%, De Datta, 1981) although higher P fertilizer recovery efciencies have been reported (Baligar et al., 2001). Rice yield responses to K fertilization are generally lower than observed for N and P, especially on responsive soils. As such, the AE is much lower for K compared with N and P. Dobermann et al. (1998) found the AE for K in ve Asian countries ranged from 0 to 26 kg grain kg21 K fertilizer applied; however, AE values as high as 64 kg grain kg21 K fertilizer applied were reported by Baligar et al. (2001). In comparison, the AE for P ranged from 0 to 114 kg grain kg21 P fertilizer applied (Dobermann et al., 1998). The AE is dependent on the K fertility status of the soil and encompasses a wide range of values across the rice producing regions of the world. The internal use efciency of K as reported by Dobermann et al. (1998) ranged from 41 to 89 kg grain kg21 total K uptake and is less variable among rice producing regions because it reects the relative grain production per unit of total K uptake rather than per unit of fertilizer K absorbed (Dobermann et al., 1998). The K uptake requirements for rice yielding between 4 and 8 t ha21 ranged from 17 to 30 kg K t21 grain produced (Dobermann et al., 1996b). Rice genotypes also differ in their K use efciencies (Baligar et al., 2001). Fageria (unpublished data, 2001) found the APE among 14 rice genotypes ranged from 3.1 to 37.7 mg grain mg21 fertilizer K uptake. This indicates that some rice genotypes have higher K requirements than others and the possibility of developing rice cultivars that are adapted to soils with low K availability exists.
Table VII Potassium Use Efciency Among Lowland Rice Genotypes Agronomic efciency (Dkg grain kg21 K fertilizer added) 76 64 54 80 44 78 66 Physiological efciency (Dkg grain Dkg21 K uptake) 89 119 89 100 101 89 98 Agro-physiological efciency (Dkg grain Dkg21 total K uptake) 39 51 58 48 26 37 43 Recovery efciency (% fertilizer uptake) 81 58 51 75 73 61 67
Genotype Alianc a CAN 5751 CAN 6804 CAN 7238 CAN 7268 Metica 1 Average
From Baligar and Fageria, 1997. Formulas used to calculate the listed efciencies are given in the section III.B.6.
106
5.
Potassium Fertilization Practices
The importance of K fertilization is increasing for lowland rice due to the higher nutrient requirements of high-yielding modern cultivars, increased crop removal of K by intensive agricultural production, and the role of K in the control of diseases. Published data concerning rice response to K fertilization for lowland rice is scarce compared to that of other macronutrients. Dobermann et al. (1996b) reasoned that infrequent, signicant yield rice yield responses to K fertilization were due to high seasonal K inputs (7 60 kg K ha21 yr21) in irrigation waters, the release of nonexchangeable K, other nutritional limitations, or combinations of these factors. Fertilizer recommendations for immobile nutrients, like K, are made based on soil test calibration studies, which need to be conducted on a routine basis. Dobermann et al. (1996b) summarized critical soil concentrations of 1N NH4OAc extractable K from around the world and found the range varied from 0.08 to 0.41 cmol K kg21 soil. In the USA, K fertilizer is usually recommended for rice when exchangeable soil K is , 60 100 mg K kg21, regardless of the soil texture or extractant. For example, in California, K fertilizer is recommended when NH4OAc extractable K is , 60 mg K kg21 (Williams and Smith, 2001). Fageria et al. (1990a) found that annual K fertilizer applications signicantly increased rice yields during the rst and subsequent rice crops seeded on the same plots on an Inceptisol in central Brazil. Rice did not respond to K fertilization when soil test concentrations were . 50 mg K dm23 (Mehlich 1 extractable K; Fageria, 1999b). Dobermann et al. (1996b) found that mixed-bed exchange resins incubated for 2 weeks under ooded soil conditions were superior to K extracted by 1N NH4OAc for prediction of K uptake by rice. They reasoned that resin adsorbed K accounted for K diffusion rates, concentration of other cations that inuence K uptake by rice, and the K fertilization history of soils that resulted in accumulation or depletion of rapid and slow released pools of soil K after ooding. Dobermann et al. (2000) reported that at IRRI 50 kg K ha21 is normally applied in eld experiments to obtain maximum yields of ooded rice and is representative of K fertilizer rates used to fertilize rice in other parts of the world. The timing of K fertilizer application should consider two criteria: (1) the cost of application and (2) maximizing fertilizer use efciency by the crop. Sometimes, K is applied in a band at the time of sowing to increase its availability to seedling rice. In the USA and many other rice producing regions of the world, K fertilizer is broadcast applied immediately before seeding, after seeding, or split into multiple applications. In the humid tropical soils with low cation exchange capacity and clay content the possibility of K loss via leaching exists and K fertilizer is commonly broadcast applied as a topdressing, along with N. Fageria (1991) reported that lowland rice yields were higher when the total K fertilizer requirement was applied in split topdressed applications. Noguchi and Sugawara (1976), Su (1976), and Santos et al. (1999), also reported the benets
of split applications of K. Potassium application rates applied as topdressing depend on the initial soil K concentration with higher K rates required as the initial soil K concentration decreases. Preliminary data reported by Slaton et al. (2001b) suggests that a single application of K fertilizer applied preplant, preood (5-leaf stage), or at the panicle initiation stage were sufcient to maximize grain yields on a Calloway silt loam (Fine-silty, mixed, active, thermic Glossaquic Fragiudalfs) in Arkansas. However, K fertilizer applied during the boot stage did not increase yields above the untreated control. In general, a major portion, and sometimes all, of the K fertilizer should be applied at or near the time of seeding. A smaller portion of the total K fertilizer requirement should be topdressed, along with N, to reduce the cost of application, on soils where leaching losses of K are of concern. Potassium chloride is the most common source of K for most annual crops, including rice. Potassium chloride is highly effective and has the lowest cost per unit of K. Potassium sulfate is equally effective as a K source and furnishes S, but is also more expensive than KCl. Field experiments have shown little or no difference in rice response to KCl, K2SO4, and KNO3 fertilizers in Arkansas (Wilson et al., 1996). 6. Use of Efcient Genotypes The identication and selection of nutrient efcient cultivars is considered one of the most cost effective approaches for improving crop production in resourcepoor environments. Very little research has been conducted on nding genotypes that exhibit high nutrient use efciencies for K or other nutrients in the highly mechanized agricultural producing areas. Increased fertilizer prices, depletion of fertilizer resources, and the cultivation of marginal lands are excellent reasons for nding crop genotypes with superior nutrient utilization efciencies. Genetic differences in K uptake and utilization efciency among cultivars within crop species, including rice, have been reported in several studies (Glass et al., 1981; Siddiqi and Glass, 1981; Clark, 1990; Clark and Duncan, 1991; Fageria et al., 1997a). Plants have different uptake potentials for nonexchangeable K due to different root morphologies (Mengel, 1985).
E. CALCIUM, MAGNESIUM,
AND
SULFUR
Calcium, Mg, and S are often referred to as the secondary elements. These macronutrients are required in relatively large amounts for normal crop growth. Rice tissue macronutrient concentrations generally follow the order, from high to low, of N K . Ca . P . Mg . S. Calcium is usually the predominant soil cation and is present in relatively large amounts especially on alkaline soils.
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Further, Ca is a nontoxic mineral nutrient, even in high concentrations, and is very effective in detoxifying high concentrations of other mineral elements in plants (Marschner, 1995). Calcium and Mg are much more stable in the soil than S. Highly weathered Oxisols and Ultisols may be low in Ca, Mg, and plant available S due to excessive leaching. Sandy soils may also contain low levels of Ca, Mg, and S increasing the likelihood of crop deciencies of these nutrients. However, rice seldom suffers from Ca and Mg deciencies. In comparison, S deciency has been reported from nearly all rice producing regions of the world including Indonesia, Brazil, India, Bangladesh, Thailand, and the USA (De Datta, 1981; Wells et al., 1993). Yamaguchi (1997) reported that symptoms associated with S deciency of rice often occur in irrigated Vertisols of the lower Volta in Ghana, Africa. Sulfur deciency was a possible cause of these symptoms because farmers applied urea and high-analysis NPK fertilizers that contained little S. Blair et al. (1978) suggested that the low S content of most tropical soils was the primary cause of S deciency. Sulfur deciency of rice has increased for numerous reasons including: (1) increased crop removal of S via increased crop yields; (2) use of fertilizers lacking S; (3) the reduced industrial emissions of S lowering the input of atmospheric S; (4) reduction in soil organic matter; (5) leaching and weathering processes; (6) erosion; and (7) crop management practices (De Datta, 1981). 1. Chemistry in the Soil Calcium and Mg are alkaline earth cations. Calcium is the dominant exchangeable cation in most soils. Calcium and Mg concentrations in the soil solution are in equilibrium with the exchangeable forms of these cations. The degree of Ca and Mg saturation of the cation exchange sites, complimentary cations, the nature of the bonding with exchange sites, and the concentration of anions in the solution all interact to determine the Ca and Mg soil solution concentrations (Barber, 1995). Exchangeable Ca and Mg usually account for more than 60% of the exchangeable cations on soils at pH 5.5 or higher. Exchangeable Al3, H, K, and Na occupy the majority of the remaining 2 exchange sites (Barber, 1995). Calcium (Ca2), Mg2, and SO2 4 ions are the main forms absorbed by plants and mass ow is the principal mechanism by which these nutrients move to plant roots. The availability and absorption of 2 Ca2, Mg2, and SO2 4 are governed by the soil pH, ion concentrations in the soil solution, cation exchange capacity, organic matter content, type of soil colloid, the ratio of these ions to other cations or anions, and many plant factors. More than 90% of the total S in the A horizon of soils exists in the organic form. The N:S ratio in surface soils is relatively constant and averages about 10:1.3 (Foth and Ellis, 1988). Plant available S in the rice root zone is quickly depleted in many soils by plant uptake, SO4 leaching, and the reduction of SO4 to
sulde (S22). Consequently, the soil SO4 concentration declines after ooding, but may be accompanied by the accumulation of S22, which can be toxic to plants and may also be lost from the soil as H2S gas. Thus, the availability of soil S decreases as soil reduction proceeds. The rate of SO4 reduction in submerged soils depends on a number of soil properties. In neutral and alkaline soils, SO4 21 2 concentrations as high as 15,000 mg SO2 may be reduced to zero within 6 4 kg weeks of submergence (Ponnamperuma, 1972). The reduction of SO4 begins at an Eh of 2 0.15 to 2 0.2 V at pH 6.5 7.0 (Takai, 1978). Calcium and Mg concentrations in the soil solution are altered by the chemical changes associated with ooding, but their ionic forms are not affected. In contrast, the ionic forms of S undergo marked changes following ooding (Patrick and Mikkelsen, 1971). Flooded soils frequently become sufciently 2 reduced from a restricted oxygen supply and microbial activity to reduce SO2 4 to 22 3 2 2 22 S . Since Fe reduction to Fe precedes SO4 reduction, Fe is present in the soil solution by the time S22 is produced. The formation of insoluble FeS may prevent the formation of H2S and protect microorganisms and aquatic plants from the toxic effects of H2S gas (Patrick and Reddy, 1978). 2. Functions and Deciency Symptoms
Calcium accelerates the translocation of photosynthetic products in rice plants (Kawasaki, 1995). Kawasaki (1995) reported that Ca stimulated the absorption of P and K under certain ion concentration ranges in nutrient solutions. Among the various physiological functions performed by Ca in higher plants, the most important role is believed to be the maintenance of structure and the function of biomembranes. Injury due to heavy metals is reduced by Ca because it lowers heavy metal concentrations in rice plants (Kawasaki, 1995). Calcium is also a promoter of normal root growth and development. In contrast to Mg2, which is a strong activator of enzymes, Ca2 increases the activity of only a few enzymes. Hanson (1984) and Kirkby and Pilbeam (1984) presented comprehensive reviews on the functions of Ca as a plant nutrient. It is generally recognized that the most important physiological roles of Mg is as a constituent of chlorophyll and an activator of numerous enzymes. A balance between Ca and Mg concentrations within the plant is necessary for the maintenance of normal metabolic processes. A high proportion of the total plant Mg is involved in the regulation of cellular pH and the cation anion balance. Magnesium also functions as a bridging element for the aggregation of ribosome subunits, which is a process necessary for protein synthesis. Sulfur is an essential nutrient for all plants and animals because it is a constituent of essential amino acids (e.g., cysteine and methionine), several coenzymes (e.g., biotin, coenzyme A, thiamin pyrophosphate and lipoic acid), thioredoxins, and sulpholipids (Zhao et al., 1997). There are many other
110
S-containing compounds in plants that are not essential, but may be involved in defense mechanisms against pests and pathogens or contribute to the special taste and odor of some plants (Bennett and Wallsgrove, 1994). Apart from the effects on yield, the S nutrition of a crop often has a strong inuence on food quality because of its essential role in the synthesis of amino acids, proteins, and some secondary metabolites (Zhao et al., 1997). A sufcient S supply to crops is also important for the nutritional quality of legumes and the processing quality of cereal grains (Randall and Wrigley, 1986). Calcium deciency in plants is typically expressed as cell death due to the breakdown in critical functions such as selective membrane permeability and dysfunction of cellular signaling mechanisms (OBrien and Ferguson, 1997). In Ca decient rice the growing point becomes white and upper leaves roll and curl. Magnesium deciency of rice is characterized as pale green plants with droopy and wavy leaves. Magnesium is relatively mobile in the plant resulting in an interveinal chlorosis of the older leaves at the onset of deciency. The interveinal chlorosis gives the lower leaves an orangish-yellow color. Tillering and plant height of Mg decient rice are almost normal. The yellowing of the outer leaf edge of Mg decient plants is a characteristic difference between K and Mg deciency symptoms (Fageria and Barbosa Filho, 1994). Sulfur deciency symptoms are very similar to those described for N. However, S has limited mobility in the plant and produces a relatively uniform chlorosis of the plant. The general location of the chlorosis can be used to distinguish between N and S deciencies, especially during the early development of symptoms. Sulfur deciency is initially expressed as a chlorosis of the younger leaves while N deciency results in chlorotic older leaves. Additionally, prolonged N deciency results in a premature necrosis of the older leaves, which is not characteristic of S (Wells et al., 1993). Sulfur decient seedlings are yellow to pale green. Sulfur deciency largely affects the growth of leaf blades as the reduction in the dry weight of leaf blades is larger than in stems and roots. With S deciency, the chlorophyll content of leaves decreases and lowers the photosynthetic rate (Suzuki, 1995). In rice, severe S deciency may reduce the number of panicles, panicle length, and the number of spikelets per panicle. In Arkansas, late-season S deciency has recently been observed on a number of elds with sandy loam, sandy clay, and clay soil textures (Slaton et al., 2001a). These late-season symptoms generally appear on the top two or three leaves as an interveinal chlorosis that begins near the leaf tips and proceeds towards the leaf base shortly before the panicles exert from the boot. 3. Critical Level in Plant and Uptake
Rice plants generally have lower concentrations of Ca and Mg than other crops, especially dicots (Kawasaki, 1995). Calcium and Mg concentrations in rice
tend to decrease during vegetative growth and then stabilize, near the time of panicle initiation, for the remainder of the season (Table VIII). Calcium concentration in the shoot is typically higher than that of Mg. However, Mg concentration is generally higher in the rice grain than Ca presumably because it is more mobile. Fageria et al. (1997a) reported that adequate concentration ranges of Ca and Mg were 2.5 4.0 g Ca kg21 and 1.7 3.0 g Mg kg21, respectively, in the shoots of rice at 100 days of growth (about booting). De Datta (1981) reported the critical concentrations of 1.5 g Ca kg21 and 1.0 g Mg kg21 in rice straw at maturity. Knowledge of nutrient accumulation by a crop during its growth cycle is important to understand the nutrient requirements of the crop and know how much of a nutrient is taken up and removed from the soil. Such information is useful in maintaining the soil fertility at an adequate level for essential elements. Data listed in Table IX shows the seasonal accumulation of Ca and Mg by lowland rice at several different growth stages. Calcium accumulation was higher than that of Mg in the straw, but the opposite was true for the grain. In this study, lowland rice (straw grain) accumulated 6 kg Ca and 4 kg Mg t21 of grain produced, but harvested grain removed nearly double the amount Mg (Table IX). The critical S concentration in rice tissue, like that of N, varies with the stage of plant development and part of the plant that is sampled. Wells et al. (1993) reported that the critical concentration of S varies from approximately 2.5 g S kg21 at tillering to 1.0 g S kg21 at heading. Yoshida (1981) reported that the critical S concentrations in straw needed for maximum dry weight production
Table VIII Whole Plant Concentrations of Ca and Mg in the Shoots and Grain of the Flood-Irrigated Rice from Brazil (Fageria, unpublished 2001) and Arkansas (Slaton, unpublished data, 2001) at Different Growth Stages Brazila DAEc 22 35 71 97 112 140 140 Calcium (g Ca kg21) 4.0 4.0 3.2 3.0 2.9 3.3 0.8 Magnesium (g Mg kg21) 2.7 2.3 1.8 1.9 1.8 1.8 1.2 DAEc 55 81 96 111 145 146 Arkansasb Calcium (g Ca kg21) 8.3 5.8 5.5 5.1 5.3 0.24d Magnesium (g Mg kg21) 3.0 3.0 3.0 2.8 2.6 1.1d
Growth Stage Beginning tillering Active tillering Panicle initiation Boot stage Flowering Mature-straw Grain
a b
The rice cultivar was Metica 1 and was grown on an Inceptisol. In Arkansas, the rice cultivar was Wells and was grown on a slighty calcareous Calloway silt loam. c DAE, days after emergence. d Harvested grain. Whole panicle concentration at maturity was 0.9 g Ca kg21 and 1.7 g g kg21.
112 N. K. FAGERIA, N. A. SLATON AND V. C. BALIGAR

Table IX Whole Plant Contents of Ca and Mg in the Shoots and Grain of the Flood-Irrigated Rice from Brazil (Fageria, unpublished data, 2001) and Arkansas (Slaton, unpublished data, 2001) at Different Growth Stages Brazila Growth stage DAEc Beginning tillering Active tillering Panicle initiation Boot stage Flowering Mature-straw Grain Total uptake/t grain Grain content, % of total
a
Arkansasb Magnesium (kg Mg ha21) 0.8 2.6 10.1 19.5 24.1 16.3 7.7 3.8 32.1 DAEc 55 81 96 111 145 146 Calcium (kg Ca ha21) 4.8 24.6 48.0 60.3 52.4 1.9d 6.7 3.5
21
Calcium (kg Ca ha21) 1.4 4.6 18.4 30.6 37.3 31.3 4.8 5.7 13.3
Magnesium (kg Mg ha21) 1.8 12.7 26.4 33.1 25.7 8.6d 4.2 25.1
22 35 71 97 112 140 140
The rice cultivar was Metica 1 and was grown on an Inceptisol. The total, aboveground dry matter produced was 15,647 kg ha with a grain yield of 6323 kg ha21. b In Arkansas, the rice cultivar was Wells and was grown on a slightly calcareous Calloway silt loam. The total, aboveground dry matter produced was 17,640 kg ha21 with a grain yield of 8085 kg ha21. c DAE, days after emergence. d Ca and Mg concentrations in harvested rough rice grain.
varied from 1.6 g S kg21 at tillering to 0.7 g S kg21 at maturity. The critical N:S ratio in straw for maximum biomass production varies from 23 at active tillering to 13 at maturity. Fageria et al. (1997a) reported that adequate concentrations of S in the uppermost mature leaves at tillering were 2.0 6.0 g S kg21. Suzuki (1995) reported 1.0 g S kg21 as a critical level in the rice shoot at tillering and 0.55 g S kg21 in rough rice grains. Wang (1976) concluded that the critical concentration of S in rice straw should be 0.5 g S kg21 for optimum grain yield. Slaton et al. (2001a) observed late-season S deciency symptoms when rice ag leaves, immediately before panicle emergence from the boot, contained , 1.5 g S kg21. Rice grain S concentrations vary between 0.34 g S kg21 for S decient plants to 1.6 g S kg21 from plants that had no response to S application (De Datta, 1981). Wang et al. (1976) determined S uptake in the straw and grain of lowland rice grown in the Amazon Basin in the State of Para, Brazil (Table X). Grain and straw S contents increased as S and N fertilizer rates increased. Additionally, at low rates of S fertilization, grain S content was greater than straw S content, but straw and grain S contents were nearly equal at high rates of S fertilization. Wang (1976) reported that lowland rice grain yields of 5 7 t ha21 removed between 5 and 9 kg S ha21. The rate of S removal by lowland rice was affected by the cultivar, S application rate, and N fertilization. In Arkansas, rice total S uptake at maturity generally averages about 25 kg S ha21 with crop removal by harvested grain representing about 30% of total plant uptake (Wilson et al., 2001).
4.
Rate and Source of Application
Calcium and Mg deciency can be corrected with the application of dolomitic lime. The appropriate application rate should be based on lime recommendations, which are discussed in the liming section. If only the soil Ca level is low, gypsum (CaSO4) or CaCO3 can be applied to correct Ca deciency. Zia et al. (1997) reported that irrigated rice yield and N use efciency were improved when urea N was applied in combination with gypsum. Wang (1976) reported that at least 10 kg S ha21 is required from fertilizer for rice production on Brazilian lowland rice soils. These soils can tolerate very high levels of S (as much as 1000 kg S ha21 in eld and 2000 kg S ha21 in pot experiments) without reducing grain yields. Either ammonium sulfate or single superphosphate were good sources of S. Under pot culture, 10, 20, and 40 mg S kg21 applied once to the soil supported rice production for 1, 2, and 3 consecutive rice crops, respectively. Under eld conditions, 27 kg S ha21 applied once supported two crops (Wang, 1976). Immobilization was considered the major factor responsible for the reduced availability of residual S applied to the previous crops. In the USA, a portion of the total N requirement is sometimes
114 N. K. FAGERIA, N. A. SLATON AND V. C. BALIGAR
Table X Sulfur Uptake (kg ha21) in Shoot and Grain of Lowland Rice (Average of Two Cultivars) Under Five Different S Rates and Two N rates (After Wang et al., 1976) Shoot S Rate kg S ha21 Aboveground plant content (kg S ha21) 0 25 50 100 Mean 60 kg N ha21 120 kg N ha21 60 kg N ha21 Grain 120 kg N ha21 60 kg N ha21 Total 120 kg N ha21
0.89 2.93 3.41 4.12 2.84
0.99 3.82 4.45 4.72 3.50
1.57 3.30 3.37 3.50 2.94
1.92 4.37 3.81 3.80 3.48
2.46 6.23 6.78 7.62 5.78
2.91 8.19 8.26 8.52 6.98
supplied by application of ammonium sulfate, which also supplies adequate S when needed. Sulfur is primarily needed on permeable sandy soils or on highly reduced clay soils that are continuously cropped to rice. Irrigation water frequently contains adequate amounts of SO4 to supply seasonal crop requirements. Yamaguchi (1997) reported that S deciency in lowland rice could be corrected by the application of ammonium sulfate. Yamaguchi (1997) also reported that a mixture of ammonium sulfate (24% S) and urea increased rice dry matter production when the proportion of ammonium sulfate represented more than 25% of the total N application. According to De Datta (1981), S is generally supplied to rice as a component of other fertilizers such as ammonium sulfate (24% S), single superphosphate (14% S), potassium sulfate (16 18% S), or agricultural gypsum (15 18% S). Elemental S may also be used as a source of S provided an adequate time interval is allowed for the oxidation of S into a plant available form following its application. Sulfur, like N, is subject to many chemical and biological reactions that inuence its availability when added to the soil. Sulfur fertilizer applications should be timed based on the initial soil S status and the soil properties that inuence S availability during the course of the growing season. Routine soil testing for S is seldom used as a guideline. Soil properties (i.e., drainage, texture, and oxidation status) and eld histories are used by growers as a means of estimating the need for S fertilization. When the availability of S is initially low, SO4 containing fertilizers should be applied at seeding or by the 5-leaf stage when rapid plant growth and tillering begin. The application of SO4 containing fertilizers may also be necessary during the reproductive growth phase (i.e., panicle initiation or early boot stage) to prevent late-season S deciency on highly permeable or reduced soils.
F. MICRONUTRIENTS
Micronutrients are also called minor or trace elements. Their concentrations in plant tissues are present in small amounts relative to that of macronutrients. The essential micronutrients are Zn, Cu, B, Fe, Mn, Mo, and Cl. Accumulation of these micronutrients by plants generally follows the order of Cl . Mn . Fe . Zn . B . Cu . Mo. This order may change among plant species and growth conditions, but is generally correct for lowland rice. Micronutrient deciencies in crop plants are increasing because of (i) increased micronutrient demands from intensive cropping practices and adoption of high yielding cultivars which may have higher micronutrient demand, (ii) enhanced production of crops on marginal soils that contain low levels of available nutrients, (iii) increased use of high analysis fertilizers with low amounts of micronutrient contamination, (iv) decreased use of animal manures,
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composts, and crop residues, (v) cropping soils that are naturally low in micronutrient reserves, and (vi) involvement of natural and anthropogenic factors that limit adequate plant availability and create element imbalances (Fageria et al., 2002). Deciencies of B, Cu, Fe, Mn, and Zn have been reported in ooded rice. Zinc is, by far, the most common micronutrient deciency encountered throughout the worlds rice producing regions. In the USA, and possibly other rice growing regions of the world, Zn deciency is far more common than all the other essential elements except N. In addition to Fe deciency, Fe toxicity is a signicant problem in some parts of the world. Deciencies of B, Cu, and Mn are not common, but have been observed in production elds or greenhouse studies. Molybdenum deciencies of rice have not been reported and although Cl is essential for higher plants, its deciency has not been reported in rice (Obata, 1995). Chloride toxicity in the form of salt injury is a more common problem to some rice producing areas; however, Cl nutrition of rice is not discussed in this review. This review will discuss the management practices and nutrient requirements of four essential micronutrients in regards to lowland rice production. A more detailed discussion of micronutrient chemistry in the soil and functions in plant nutrition is available in reviews by Hodgson (1963), Hodgson et al. (1966), Romheld and Marschner (1991), Mortvedt (1994), Fageria (1999a), and Fageria et al. (2002). 1. Zinc
Zinc deciency has been reported in various parts of the world for a large number of annual crops including rice (Cakmak et al., 1998; Mandal et al., 2000; Fageria, 2001). A global study by FAO, showed that about 30% of the cultivated soils of the world are Zn decient (Sillanpaa, 1982). Additionally, about 50% of the soils used worldwide for cereal production contain low levels of plantavailable Zn (Graham et al., 1992; Welch, 1993). De Datta (1981) reported that Zn deciency is the second most serious nutritional disorder limiting the yield of lowland rice in the Philippines. Zinc deciency in crop plants reduces not only grain yield, but also the nutritional quality of the grain. Consumption of large quantities of cereal-based foods with low Zn concentrations, poor bioavailability of Zn, or both is believed to be a major factor in the widespread occurrence of Zn deciency in humans (Welch, 1993). In Brazil, Zn deciency has been reported in upland as well as lowland rice (Fageria and Barbosa Filho, 1994; Fageria, 2001) and is related to low concentrations of Zn in the highly weathered soils used for rice production and aggravated by high soil pH due to excessive lime application (Fageria and Baligar, 1993; Fageria and Gheyi, 1999). Chemistry in the soil. The availability of Zn to plants or its concentration in the soil solution is regulated by sorption desorption reactions at the surface of soil
colloids (Swift and McLaren, 1991). Desorption controls the amount and release rate of Zn into the soil solution for plant uptake. Desorption of Zn into the soil solution is controlled by the strength that Zn is adsorbed onto the surface of soil colloids. Other forms of Zn are also associated with organic matter, carbonates and oxide minerals, and Zn in primary and secondary minerals. Thus, the availability of Zn is inuenced by a number of soil characteristics including soil pH; organic matter content; CaCO3 content; cation-exchange capacity; clay content and mineralogy; and the quantity and types of Fe, Al, and Mn oxides (Harter, 1991; Hazra and Mandal, 1996; Singh et al., 1997). After ooded soils are drained, they contain relatively large amounts of amorphous Fe and Mn oxides that have large surface areas and a greater adsorption capacity as compared with their crystalline forms (Sah and Mikkelsen, 1986b; Quang and Dufey, 1995). Zinc uptake by rice depends not only on the concentration of Zn in the soil solution, but also on other factors, particularly the concentrations of Fe2 and Mn2 present in the soil solution. High concentrations of Fe2 and Mn2 in the soil solution antagonize Zn absorption (Sajwan and Lindsay, 1986; Mandal et al., 2000). High soil pH and the presence of free CaCO3 decrease the availability of Zn in soils. The solubility of soil Zn is highly pH dependent and decreases 100-fold for each unit increase in pH (Tisdale et al., 1985). The uptake, translocation, metabolism, and plant use of Zn is inhibited by high P availability or high rates of P fertilizer applications (Lindsay, 1979). Unlike the redox elements, Fe and Mn, the concentration of Zn in the soil solution generally decreases with time after ooding; however, Zn concentrations may increase briey immediately after ooding (Mikkelsen and Kuo, 1976; Gilmour, 1977a). A decrease in soil solution Zn concentration may be due to precipitation of ZnFe2O4 from the increased Fe solubility after ooding (Sajwan and Lindsay, 1986) or precipitation of ZnS under highly reduced soil conditions (Kittrick, 1976). Plant uptake of Zn depends not only upon the plant species, cultivar within species, and plant age, but also upon the predominate forms of Zn in the soil (i.e., amount of Zn associated with water soluble and exchangeable Zn fractions). Major factors affecting the availability of soil Zn include the soil pH, total soil Zn, Zn fertilizer source, soil organic matter content, and soil texture (Chlopecka and Adriano, 1996). Of these, soil pH extends the greatest inuence over Zn availability in most soils. Zinc deciency is most likely to occur on coarsetextured soils with high pH and low soil Zn, soils disturbed by land leveling, and highly eroded soils (Westfall et al., 1971). Functions and deciency symptoms. Zinc is a cofactor for several enzymes that are involved with N metabolism (e.g., glutamate dehydrogenase) and anaerobic metabolism (e.g., alcohol dehydrogenase). The reduction of acetaldehyde to ethanol in anaerobic metabolism requires alcohol dehydrogenase. The alcohol dehydrogenase activity of seedling rice roots increases dramatically after
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ooding and remains high for several weeks compared to rice seedlings that are not ooded (Pedrazzini and McKee, 1984). When Zn is decient, the activity of alcohol dehydrogenase is depressed, anaerobic root metabolism decreases, and the ability of the seedling rice to withstand anaerobic soil conditions is reduced (Moore and Patrick, 1988). This is one reason why Zn deciency symptoms are more dramatic after ooding rather than before ooding. When Zn deciency is diagnosed, draining the ood is commonly recommended to aid in plant recovery (Wilson et al., 2001). Removal of the ood allows seedlings to resume aerobic respiration as oxygen is reintroduced into the soil. The activity of glutamate dehydrogenase was not affected by Zn fertilization in studies conducted by Moore and Patrick (1988). Rice is considered susceptible to Zn deciency. The symptoms associated with Zn deciency of rice are well documented. Zinc, like the other micronutrients, is not very mobile within the plant; thus deciency symptoms are rst observed in the youngest leaves. Zinc deciency most commonly affects seedling rice plants, but if the deciency is mild and not corrected symptoms can also affect plants in the reproductive growth phase. In the early stages of Zn deciency, the youngest leaves usually become chlorotic, especially at the leaf base. As Zn deciency progresses, the midribs and base of older leaves may also turn yellow or pale green with brown blotches and streaks appearing on the lower leaves (Yoshida, 1981). Brown spots usually develop near the tip of the leaf blade as yellowing begins. Leaf collars may also be stacked as internode elongation is inhibited (Wilson et al., 2001). Zinc deciency tends to be more severe where high rates of N and P are applied (Mueller, 1974). Zinc decient rice plants do not respond to N fertilization (Cheaney and Jennings, 1975). Adequate Zn levels in the soil increase tillering and, consequently, the number of panicles per unit area of lowland rice (Fageria, 2001). Application of high rates of P fertilizer is known to aggravate Zn deciency too. The major reasons for P induced Zn deciency are believed to be the formation of Zn phosphate in soil solutions and/or an inhibitory effect of the excessive P on the metabolic functions of the Zn within the plant (Shimada, 1995). Zinc deciency symptoms of rice plants with color photographs are shown by Mueller (1974), Cheaney and Jennings (1975), Yoshida (1981), Fageria (1984), Wells et al. (1993), and Fageria and Barbosa Filho (1994). In the direct seeded, delayed ood management system used in Arkansas, Zn deciency symptoms are visible within 2 or 3 days after the ood is established on seedling rice and the severity of Zn deciency increases with ood depth (Wilson et al., 2001). When Zn deciency is severe, the symptoms are also visible before ooding. When Zn decient rice is ooded and severe Zn deciency symptoms are expressed seedling death may occur if the ood is not removed. For this reason, Zn deciency can result in complete crop failure. A mild Zn deciency may not be expressed in highly visible symptoms, but rather is characterized by slow growth. In such cases, plants may grow out of the Zn deciency or the more common symptoms described previously will be
expressed if the seedling Zn requirement is not satised. Mild Zn deciencies may be induced by cool temperatures and frequently disappear with warmer temperatures. Rice yield losses due to Zn deciency generally range from 10 to 60% in the untreated controls of research plots (Slaton et al., 2002). However, very little yield loss may occur if Zn deciency is recognized quickly and the appropriate corrective actions are taken. Critical level in plant and uptake. Zinc concentrations of plants typically range from 30 to 100 mg Zn kg21, depending upon species (Shimada, 1995). Zinc deciency of rice occurs primarily on seedling and tillering rice, hence, most studies initiated to evaluate critical tissue Zn concentrations have emphasized this growth stage. Zinc deciency of seedling rice is likely when leaf and/or whole plant concentrations are , 15 mg Zn kg21 (Forno et al., 1975; Adriano, 1986). During the vegetative growth stages the plant part sampled is not critical for rice. Although Zn is considered immobile in the plant, whole seedlings or individual leaves have similar Zn concentrations (Gilmour, 1977b). Fageria et al. (1997a) reported the Zn sufciency range in rice shoots at tillering was 20 150 mg Zn kg21. Yoshida et al. (1973) developed plant tissue analysis criteria for classifying the Zn nutritional status of rice. In their system whole seedling Zn concentrations , 10, 10 15, 15 20, and . 20 mg Zn kg21 are considered decient, probably decient, low, and sufcient, respectively. Research from all parts of the world agree that seedling Zn concentrations , 15 20 mg Zn kg21 are low or decient and require Zn fertilization for optimum rice growth. The tissue concentration of other elements can also be useful in diagnosing Zn deciency. Zinc decient rice tends to accumulate other divalent cations at the expense of monovalent cations. Thus, the concentrations of Ca, Cu, Fe, Mg, and Mn tend to be higher in Zn decient rice, but tissue concentrations of K and N are lower suggesting their uptake is inhibited in some way (Moore and Patrick, 1988). The tissue concentration of Zn in the rice plant uctuates during the growing season. Whole plant tissue Zn concentration is generally highest after ooding, decreases during tillering, and then, depending on the cultivar, may increase or remain stable through ripening (Gilmour, 1977a; Wells, 1980). The decline in tissue Zn concentration during tillering indicates that the rate of aboveground dry matter accumulation exceeds that of Zn uptake by the developing root system. The accumulation of Zn in the aboveground portion of ood-irrigated rice is relatively slow during vegetative growth, reaches a maximum during late vegetative to early reproductive growth, and then declines after anthesis (Gilmour, 1977b). The rate of maximum Zn uptake and dry matter accumulation occurs simultaneously. The rate of nutrient uptake is likely related to root growth and development. Slaton et al. (1990) found that the maximum rice root length was reached by the early boot stage, but the maximum root growth rate occurred
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by panicle initiation, which corresponds to the same growth stage that Gilmour (1977b) found maximum Zn uptake. The Zn concentration of rice at which toxicity occurs is not well documented. Marschner (1995) reported that the critical toxic concentration of Zn in leaves of crop plants is . 400 500 mg Zn kg21. Wells et al. (1993) indicated that rice is very tolerant to Zn, with toxicity levels estimated at more than 1500 mg Zn kg21. Fageria (2000a) reported a 10% reduction in rice shoot weight (critical toxic level) when the Zn concentration in 42-day old plants was 673 mg Zn kg21. Rattan and Shukla (1984) showed that application of excessive Zn fertilizer decreased rice dry matter and estimated that tissue Zn concentrations at owering . 190 mg Zn kg21 were toxic. The availability of Zn can be reduced or stabilized in the soil by the application of amendments such as lime, P, natural or synthetic zeolities, apatities, glauconite, iron oxide containing materials, and alkaline biosolids (Chlopecka and Adriano, 1996, 1997). Zinc fertilization practices. Routine soil testing is a valuable tool that can be used to assess the potential for Zn deciency in crops. Sims and Johnson (1991) reported that the critical soil Zn concentration range for most crops was between 0.5 to 2.0 mg Zn kg21 for DTPA and 0.5 3.0 mg Zn kg21 for Mehlich 1. Most research indicates that the critical soil test Zn concentrations for rice fall within the ranges suggested by Sims and Johnson (1991). Fageria (1989) reported that 1.0 mg Zn kg21 of soil extracted by the Mehlich 1 method was the critical concentration for lowland rice. Critical DTPA extractable soil Zn concentrations of 0.8 mg Zn kg21 has been reported for Indian soils for lowland rice (Tiwari and Dwivedi, 1994), whereas, 0.7 mg Zn kg21 (Sedberry et al., 1978) and 0.5 mg Zn kg21 (Hill et al., 1992) have been suggested for rice in the USA. Sedbery et al. (1980) and Wells (1980) both indicated that soil pH of silt loam soils was the best predictor of rice response to Zn fertilization. However, their research was conducted on soils that had not previously received applications of Zn fertilizer, were uniformly low in Zn, and micronutrients were not commonly measured in routine soil analysis. Thus, for a number of years Zn fertilizer recommendations were based exclusively on soil texture and soil pH, which triggered the recommendation to use Zn fertilizer on nearly every rice crop grown in the rotation on alkaline soils. In Arkansas, Zn fertilizer recommendations for ooded rice are now based on the soil pH, texture, and Mehlich 3 extractable Zn (Wilson et al., 2001). Zinc fertilizer is recommended for rice grown on silt and sandy loam soils having a pH . 6.0 and Mehlich 3 extractable Zn , 3.5 mg Zn kg21. Zinc deciencies are seldom observed on undisturbed clay soils in the USA. Precision land leveling often exposes Zn decient subsoils and Zn deciency is occasionally observed on leveled soils of all textures. The most common commercially manufactured granular Zn fertilizers are Zn sulfates, oxides, oxysulfates, lignosulfonates, and a number of organic chelated materials like ZnEDTA and ZnHEDTA. Excellent reviews of the manufacturing
and properties of Zn fertilizers are given by Tisdale et al. (1985), Foth and Ellis (1988), and Martens and Westermann (1991). Application of these Zn fertilizers to rice is performed in a variety of methods depending on the production system. Most commonly, relatively high rates of inorganic Zn fertilizers are applied to the soil before seeding. Application of 5 7 kg Zn ha21 as Zn sulfate was found to correct Zn deciency in lowland as well as upland rice grown on Inceptisols and Oxisols in Brazil (Fageria and Barbosa Filho, 1994). Slightly higher rates of 11 kg Zn ha21 are typically recommended for soil application in the USA. In the waterseeded system practiced in California, a surface, broadcast application of either ZnSO4 or Zn lignosulfonate is recommended because the roots of rice seedlings are positioned at or near the soil water interface (Wells et al., 1993). Highly water soluble ZnSO4 is generally regarded as the best Zn fertilizer used to correct Zn deciencies. Liscano et al. (2001) showed that the water solubility of inorganic Zn fertilizers was highly correlated to Zn uptake by seedling rice in greenhouse studies. They suggested a minimum of 40 50% of a Zn fertilizers total Zn content should be water-soluble to optimize Zn uptake. Amrani et al. (1999) and Gangloff et al. (2002) reported similar results for corn. In general, the water solubility of Zn sulfates and lignosulfonate sources is high and the water solubility of Zn oxides and Zn oxysulfate sources is low to moderate. However, in most cases, the Zn application rate is more critical than the water-soluble Zn content of the fertilizer, but research data clearly shows that tissue Zn concentration and total Zn uptake generally increase as water soluble Zn in a fertilizer increases. The use of the water-soluble Zn criteria for selecting a Zn fertilizer becomes more important as the severity of Zn deciency for the immediate crop increases. The recommended rates of soil applied Zn are about 20 times higher than the total crop uptake of Zn, but are required to obtain adequate distribution of Zn fertilizer granules. The primary advantage of soil applied Zn over other Zn fertilization methods that use much lower Zn application rates is the residual benet. A single Zn fertilizer application should provide adequate Zn for several years before additional Zn fertilizer is needed to optimize grain yields. The mobility of Zn in the soil following fertilization differs among Zn sources and inuences Zn uptake. When applied to the soil surface, water-soluble ZnSO4 showed much greater vertical, downward movement than the relatively waterinsoluble ZnO (Giordano and Mordvedt, 1972). The vertical mobility of Zn supplied from ZnSO4 and Zn lignosulfonate fertilizer sources are similar with both having greater mobility than ZnO and less mobility than ZnEDTA (Mikkelsen and Brandon, 1975). The water solubility of the Zn fertilizer source, the time allowed for vertical movement of the Zn, and the relative placement of the Zn fertilizer are all important aspects to consider in Zn fertilization practices. Granular Zn fertilizers do not need to be mechanically incorporated. Surface applications of Zn before seedling emergence are equally effective as preplant incorporated Zn (Giordano and Mordvedt, 1972; Slaton et al., 2001c). However, granular Zn applications should not be made immediately before ooding
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because the Zn is not positioned for immediate uptake. Generally, less than 5% of the broadcast soil applied Zn fertilizer is taken up by ood-irrigated rice (Giordano and Mordvedt, 1972). The highly water-soluble Zn sulfates and chelated Zn sources may also be sprayed in solutions to the soil or directly onto the seedling rice foliage before ooding. Compared with granular Zn applications, the application of Zn solutions has the advantage of uniform distribution. The chelated Zn fertilizer sources are generally more expensive per unit of Zn and are applied at relatively low rates (, 1 2 kg Zn ha21) so that little residual Zn is available for future crops. Foliar Zn applications are usually employed only when the growing crop shows deciency symptoms (De Datta, 1981) or they allow for more exible crop management in regards to production costs and application times (Wilson et al., 2001). In transplanted rice, dipping seedling roots in a 1% ZnO suspension has prevented Zn deciency (Yoshida, 1981). Abilay and De Datta (1978) recommended coating pregerminated rice seeds with ZnO prior to seeding followed by a foliar Zn application 5 7 days before panicle initiation. Slaton et al. (2001) showed that Zn application directly to rice seed at concentrations between 2.2 and 5.7 g Zn kg21 seed was a low-cost alternative to the standard recommendation of 11 kg Zn ha21 as a preplant incorporated inorganic Zn fertilizer for dry-seeded rice. Earlier research also showed that the application of low rates of Zn to rice seeds or dipping the roots of transplanted rice in a Zn solution were effective alternatives to broadcast applications of Zn fertilizer (Rush, 1972; Giordano and Mordvedt, 1973; Mengel et al., 1976; Haghighat and Thompson, 1982). Zinc is also present in manures and other organic amendments that, when readily available and soil applied at adequate rates, can supply crops with sufcient Zn (Ye and Yang, 1997). Lowering the pH of alkaline or calcareous soils, by application of acidforming fertilizers/amendments like elemental S, can improve Zn availability and uptake by rice (Slaton, 1998). The effectiveness of acidifying the soil is often limited by the economics and the practicality of soil pH reduction. Some soils contain very high quantities of CaCO3 or are highly buffered and require very high rates of acidic amendments to reduce pH. Additionally, the reduction of soil pH may be only temporary as the soil pH may gradually increase over time and return to near its initial value. The literature shows that a number of Zn application methods, times, sources, and rates are highly effective means of supplying Zn to rice in various production systems. Routine soil testing used in conjunction with the application of suitable Zn fertilizers at the proper rates and times is the best method to ensure that Zn nutrition is not a yield-limiting factor for rice production. The most efcient method of Zn fertilization is often dictated by the cultural production system, soil conditions, economics, the availability of Zn fertilizers, or a combination of several of these factors.
Use of efcient genotypes. Crop species differ markedly in their ability to adapt to or tolerate Zn decient soils (Graham, 1983, 1984; Graham and Rengel, 1993; Rengel, 1997). Among the cereal and grain crops, rice, sorghum, and corn are classied as sensitive to Zn deciency (Clark, 1990). The physiological mechanisms responsible for crop tolerance to Zn deciencies are not clearly understood (Graham and Rengel, 1993; Rengel and Graham 1996; Cakmak et al., 1998; Huang et al., 1996). Graham et al. (1997) reported signicant differences in the Zn concentration of rice grain among genotypes. These authors also reported that several tests had been conducted to examine the effect of soil climatic factors on the Zn content of grain. In rice, the lled grain trait was reasonably stable across the range of environments in which rice was grown, as well as over seasons, planting dates, and soil fertility. Hence, it is possible to select Zn efcient genotypes for production in different agro-ecological regions. Numerous other researchers have also established that differences exist among rice genotypes to Zn deciency (Ponnamperuma, 1976a; IRRI, 1977; Mahadevappa et al., 1981; Bowen, 1986, 1987; Fageria, 2001). Rice genotypes showed differences in Zn absorption that were not attributed to differences in root surface area (Bowen, 1986, 1987). Rice genotypes showed marked differences in Vmax (maximum ion uptake rate) and Km (Michaelis Menten constant, equal to the substrate ion concentration giving half the maximal rate of uptake) values. Efcient genotypes increase Zn translocation from the roots to the shoot and regulate Ca, Cu, Fe, Mg, and P transport in order to maintain balanced nutrient ratios with respect to Zn (Cayton et al., 1985). Resistance to Zn deciency appeared to be controlled polygenically in rice and is thought to be a dominant trait (Mahadevappa et al., 1981). Zinc deciency has been a common mineral nutrient problem in rice and genotypes have been screened extensively for this disorder at the International Rice Research Institute (Clark, 1990).
2.
Boron
Boron deciency has been reported in at least 80 countries and 132 crop species. It is estimated that about 15 million hectares are annually treated with B fertilizers (Shorrocks, 1997). Plant species vary in B requirement with dicotyledons generally requiring 3 4 times more B than monocotyledons (Bennett, 1993). A number of soil properties inuence B availability to plants and are reviewed by Fageria et al. (2002). Coarse-textured, low organic matter soils located in humid regions are the most prone to B deciency. The application of lime to acid soils can also induce B deciency because of increased B adsorption. Boron deciencies are not common to rice, but several environmental (i.e., high rainfall), soil (i.e., low organic matter, texture, and pH), and rice production (i.e., ood-irrigation) factors common to many rice growing regions of the world hint
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that B could very likely be a yield-limiting nutrient. Our lack of understanding of the B nutritional requirements, seasonal patterns of B uptake, and B partitioning within the rice plant demonstrates the need for increased research efforts for this micronutrient. Chemistry in the soil. The soil B pools can be categorized into the mineral, adsorbed, and soil solution fractions. From the plant nutrition standpoint, the soil solution fraction is of utmost importance. Boron is absorbed as H3BO3 (or B(OH)3) and exists in the soil solution as an undissociated molecule. In agricultural soils with a pH range of 5 9, undissociated H3BO3 is the predominant species in the soil solution. The transport mechanisms of B from the soil solution to plant roots are mass ow and diffusion with mass ow as the dominant mechanism contributing to plant uptake of B. Like most other micronutrients, the availability of B decreases as soil pH increases. The reduction in B availability from increasing soil pH by liming is caused by B adsorption by iron and aluminum hydroxides. Boron is adsorbed to the surface of these precipitated Fe and Al hydroxides. Adsorption of B is very pH dependent. Maximum adsorption by Al(OH)3 and Fe(OH)3 occurs in the soil pH range of 7 9 and corresponds to the soil pH range of lowest B availability (Tisdale et al., 1985). Organic matter also adsorbs B and acts as a reservoir to replenish soil solution B upon crop removal or loss via leaching. Replacement by other anions or mineralization of the organic matter releases B (Foth and Ellis, 1988). Clay contents also inuence B adsorption. Barber (1995) reported that B adsorption by ne-textured soils is 2 3 times greater than by coarsetextured soils. Despite this general knowledge, very little is known about the chemistry of B in ooded soils. The concentration of B in the soil solution is believed to remain more or less constant following soil submergence (Ponnamperuma, 1975). In ooded soils, with pH buffered around neutrality, H3BO3 is the dominant species in the soil solution. As with upland soils, the adsorption of B on Fe and Al oxides (Sims and Bingham, 1968) seems to be an important mechanism in governing B solubility in ooded soils (Patrick and Reddy, 1978). Functions and deciency symptoms. A thorough review of the literature suggests that B deciency symptoms of rice have not been documented in the eld in any rice-growing region of the world. However, deciency symptoms have been produced in nutrient solutions and induced in greenhouse studies. Boron is relatively immobile in plant tissues and deciency symptoms rst appear in the youngest growth. The tips of emerging leaves become white and rolled in B decient plants, which is similar to a symptom described for Ca deciency. The growing points may die in the case of severe B deciency. Obata (1995) noted that B deciency also retarded root elongation of plants. The B requirement for vegetative growth of plants, especially grasses, is very low, but the need for B
increases for seed production (Marschner, 1995). This is one possible reason why B deciency symptoms on rice leaves have not been documented in commercial rice elds. Boron deciency of rice may be expressed solely in the form of reduced grain yields from oret sterility and may be mistakenly blamed on poor environmental conditions during anthesis. This hypothesis is just speculation since it has yet to be proven in replicated eld trials. However, there is some preliminary evidence that supports this theory. Okuda et al. (1961) observed that the panicles of B decient rice plants failed to exert from the boot. Likewise, Dunn (1978) noted that B deciency produced similar symptoms as those associated with the physiological disorder straighthead that was induced by arsenic toxicity in greenhouse studies. Very little research has been conducted to verify or refute B deciency as a possible cause of straighthead. Further evidence supporting this theory is provided by the essential functions of B in plant growth. Boron plays important roles in cell development and elongation, protein synthesis, carbohydrate metabolism, pollen tube formation, and pollen viability (Bennett, 1993; Marschner, 1995). In the United States, interest in the B nutrition of rice has recently been stimulated by signicant yield increases from direct B fertilization of rice in Missouri (Dunn and Jones, 2002). Color photographs of B deciency symptoms of rice were published by Ishizuka (1978) and Fageria and Barbosa Filho (1994). Boron toxicity is also a major concern for rice since the soil and plant tissue B sufciency ranges, between decient and toxic concentrations, are quite narrow. The rice tissue concentration associated with B toxicity is 100 mg B kg21 in the Y-leaf of tillering rice, 35 mg B kg21 in the shoots at panicle initiation, and 100 mg B kg21 in the straw at maturity (Dobermann and Fairhurst, 2000). The plant part and the time of sampling are critical when using tissue analysis to diagnose B toxicity because B tends to accumulate in the leaf tips and may be leached from plant tissue by rain. Likewise, Dobermann and Fairhurst (2000) cited hot-water soluble soil B concentrations . 5 mg B kg21 as potentially toxic. Obata (1995) suggested a slightly higher hot-water soluble soil B concentration of $ 10 mg B kg21 was associated with B toxicity of rice. Application of high rates of B fertilizer may also induce B toxicity on soils that have low soil B concentrations (Fageria, 2000b). Toxicity symptoms include scorching of the tips and margins of older leaves and result in reduced dry matter and grain yield production. Critical level in plant and uptake. Boron contents vary widely among plant species. Rice, together with wheat and barley, has a lower requirement for B than do nongramineous crops (Obata, 1995). Boron requirement is higher during the reproductive growth stages than during vegetative growth due its important function in grain formation. Due to the lack of published research on B nutrition of rice, very little data is available on the critical tissue B concentrations required for the production of maximum rice yields. Dobermann and Fairhurst (2000)
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suggested that the optimum B concentration range for the Y-leaf of tillering rice was 6 15 mg B kg21. Fageria et al. (1997a) suggested a critical concentration of 8 mg B kg21 in rice straw at maturity. Yu and Bell (1998) reported 18.5 mg B kg21 in rice leaves and 8.9 mg B kg21 in rice stems were associated with maximum rice yield production. They also reported that B deciency in rice occurred when B concentrations in the top mature leaves was , 7.3 mg B kg21. In a review of the literature, Yu and Bell (1998) concluded that the sufcient B concentration range of rice varied from 5 to 67 mg B kg21 depending on the plant age and the part analyzed. Data in Table XI suggests that the B concentration of whole rice plants is relatively constant during the growing season with total B uptake increasing as plant age increased. Depending on the location, 22 34% of total aboveground B content was partitioned in the grain. Boron fertilization practices. Soil analysis is sometimes used as a basis for identifying potentially B decient soils and has been used to make fertilizer recommendations for some crops (Martens and Westermann, 1991; Sims and Johnson, 1991). However, B deciency of rice is yet to be recognized as a serious yield-limiting factor and insight to critical soil test B concentrations for rice have not been developed. Research conducted with other crops suggests that if positive rice yield responses to B fertilization are found fertilizer recommendations can potentially be based on routine soil analysis.
Table XI Whole Plant B Concentrations and Contents of Shoots and Grain of Flood-Irrigated Rice from Brazil (Fageria, unpublished data, 2001) and Arkansas (Slaton, unpublished data, 2001) at Different Growth Stages Brazila DAEc 22 35 71 97 112 140 140 Concentration (mg B kg21) 7.8 7.5 7.1 6.7 6.9 7.3 5.3 Content (g B ha21) 2.8 8.6 41.1 70.6 92.5 69.4 35.0 DAEc Arkansasb Concentration (mg B kg21) Content (g B ha21)
Growth stage Beginning tillering Active tillering Panicle initiation Boot stage Flowering Mature-straw Grain
55 81 96 111 145 146
7.4 5.0 4.9 5.3 5.2 1.9d
11 21 42 63 51 15d
a The rice cultivar was Metica 1 and was grown on an Inceptisol. Average yield (at maturity) was 9423 kg ha21 straw weight and 6389 kg ha21 grain yield. b In Arkansas, the rice cultivar was Wells and was grown on a slightly calcareous Calloway silt loam. Average yield (at maturity) was 9555 kg ha21 straw weight and 8085 kg ha21 grain yield. c DAE, Days after emergence. d Harvested grain. Whole panicle concentration at maturity was 3.8 g B kg21.
Boron extracted with the hot-water procedure is generally considered the standard extraction procedure to evaluate crop response to B. Sensitive crops are likely to respond to B fertilization if the level of hot water-soluble B is less than 0.5 mg B kg21 soil (Sims and Johnson, 1991). A particular problem with B is the narrow margin between deciency and toxicity. Sensitive crops can be affected by B toxicity at soil levels over 5.0 mg B kg21, so B fertilizer must be used with caution. Sims and Johnson (1991) reported critical soil B levels of 0.1 2.0 mg B kg21, depending on yield level, soil type, pH, and organic matter content. 3. Iron
Rice is considered very susceptible to Fe deciency, especially when grown under upland conditions. Rice yield losses caused by Fe deciency measured in research plots range from 10 to 50% (Snyder and Jones, 1988, 1991). Ferrous Fe (Fe2) is preferentially absorbed by plant root systems and is generally present in high concentrations in reduced soils (Marschner, 1995). Thus, Fe deciency does not commonly occur in ooded rice due to the increase in Fe availability associated with the anaerobic soil conditions used for its production. However, Fe deciency occurs primarily on seedling rice before ooding in some rice growing regions of the world (Yoshida, 1981; Snyder and Jones, 1988). Mori et al. (1991) suggested that seedling rice is highly susceptible to Fe deciency because rice roots produce comparatively low amounts of iron-chelating phytosiderophores compared to other grass species. Soil conditions that limit Fe availability coupled with plant limitations for obtaining Fe are the primary reasons why Fe deciency occurs. In many rice-growing regions of the world, Fe toxicity rather than Fe deciency is the more common problem and is associated with inherent soil properties (Yoshida, 1981; Fageria and Rabelo, 1987; Olaleye et al., 2001). Iron toxicity has been reported as a signicant problem in rice growing areas of southeast Asia, Africa, and South Africa that occurs primarily on acid sulfate soils (Ottow et al., 1983). Iron toxicity is believed to be caused by excessive Fe in the soil solution or induced by deciencies of other nutrients. The direct toxicity is dened as excessive Fe absorption by the plant resulting from high soil solution concentration of Fe (Howler, 1973). The indirect toxicity has been blamed on low soil fertility. Specically, low soil concentrations of Ca, Mg, K, and P have been cited as the common factors among soils expressing Fe toxicity symptoms (Benckiser et al., 1984). The name Fe toxicity is somewhat misleading in the sense that, in many cases, the Fe toxicity is actually an effect rather than a cause. Howler (1973) suggested that rice roots become coated with Fe oxide which reduces the plants ability to absorb sufcient quantities of other plant nutrients that are already present in low concentrations. The production of H2S, FeS, or both in highly reduced ooded soils may contribute to Fe toxicity. Hydrogen sulde and
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FeS reduce the oxidizing capacity of the rice root system, thereby increasing the susceptibility of the rice plant to Fe toxicity (Inada, 1966b; Tanaka et al., 1968). Application of (NH4)2SO4 fertilizer has been noted to increase the incidence of Fe toxicity (Inada, 1966a). Acidic soil conditions are likely to produce either type of Fe toxicity. Severe Fe toxicity can cause signicant rice yield reductions (Genon et al., 1994). In general, soil solution Fe concentrations of ooded soils are very high because of the anaerobic soil conditions, but rice is well adapted to this ooded environment and normally is able to regulate Fe uptake. Chemistry in the soil. Iron is a major constituent of most soils. Iron minerals commonly found in soils include goethite (FeOOH), hematite (Fe2O3), pyrite (FeS), siderite (FeCO3) and magnetite (Fe3O4). Therefore, Fe availability to plants is affected by the soil properties like soil pH and redox that inuence the solubility of Fe containing minerals. The Fe concentration in the soil solution decreases with an increase in soil pH. For each unit increase in soil pH there is a 1000-fold decrease in the solubility of Fe3 and a 100-fold decrease in the solubility of Fe2 (Tisdale et al., 1985). The amount of Fe2 increases rapidly at redox potentials below 200 mV. Ponnamperuma (1976b) found that the soil Fe2 concentration increased to a peak ranging from 0.1 to 600 mg kg21 for several soils shortly after submergence and then declined. Diffusion and mass ow are believed to be the two mechanisms responsible for the movement of Fe from soil to the root surface. The chemistry of ooded soils is dominated more by Fe than by any other redox element. The major reason for this dominance is the large amount of soil Fe that can undergo reduction, which usually exceeds the total amount of other redox elements by a factor of 10 or more (Patrick and Reddy, 1978). Under submerged soil conditions, Fe3 is reduced to Fe2 by respiring microorganisms. Although variable in composition, ferric oxyhydroxides in aerated soils can be represented by the formula Fe(OH)3, which can undergo reduction (Patrick and Reddy, 1978): FeOH3 3H e2 $ Fe2 3H2 O Although there is general agreement that the reduction of ferric compounds occur as a result of the respiration of facultative anaerobic bacteria, it has not been demonstrated conclusively that the reduction is brought about by enzymatic transfer of electrons directly to Fe3, or that the reduction is an indirect chemical reaction between bacterial metabolites and Fe3. In either case, it is likely that complexing of Fe with organic chelates plays an important role in making the Fe solution more reactive (Patrick and Reddy, 1978). Functions and symptoms of Fe deciency and toxicity. The concentration of Fe in rice tissue is generally higher than all other micronutrients, except Mn, and
reects the need for sufcient Fe nutrition for normal growth and seed production. The functions of Fe in plant nutrition and growth processes are thoroughly reviewed by Marschner (1995). Some of the essential functions of Fe in plant growth are clearly exhibited by the deciency symptoms expressed by rice and other plants. Iron deciency is most common to upland rice production systems and seldom occurs in lowland rice after ooding. However, many lowland rice production systems are dry-seeded and later ooded at the onset of tillering; thus the period between seeding and ooding is the time when Fe deciency is most likely to occur in lowland rice. Iron is not highly mobile within the plant and the youngest leaves are the rst to show Fe deciency symptoms. At the onset of Fe deciency, symptoms begin as an interveinal chlorosis of the youngest leaves giving plants a striped appearance. Further progression of the Fe deciency gives seedlings a uniform pale yellow to bleached white appearance (Snyder and Jones, 1988). These symptoms have been noted to occur within 1 week after emergence on Histosols used for rice production in the Florida Everglades. Iron deciency reduces seedling dry matter production, leaf chlorophyll content, panicle number per unit area, and grain yield (Snyder and Jones, 1988). Plant tissue analysis used to determine the total tissue Fe concentration has proven to be of little value for diagnosing Fe deciency of many plants, and rice is no exception (Mengel et al., 1984). The total Fe concentration of rice seedlings showing Fe deciency symptoms is generally equal to that of seedlings without symptoms (Snyder and Jones, 1988). Routine soil analysis is also of little value for predicting Fe decient soils, so diagnosis of Fe deciency from visual symptoms or eld history is generally relied upon. Color photographs of rice plants with Fe deciency are available in publications by Ishizuka (1978), Yoshida (1981), Fageria (1984), Fageria and Barbosa Filho (1994), and Dobermann and Fairhurst (2000). Iron toxicity causes the older leaves to turn a yellow orange color starting at the leaf tip with symptoms proceeding towards the leaf base. The tips of the lower leaves will eventually desiccate and give the plants a scorched appearance. Rice roots commonly have a black coating of FeS, which can also be used to help diagnose Fe toxicity. Toxic levels of Fe can induce deciencies of K, P, Cu, and Zn in rice (Fageria and Rabelo, 1987; Baruah and Nath, 1997). The degree of leaf bronzing has been suggested to be a good measure of the severity of Fe toxicity in ooded rice (IRRI, 1965). Critical level in plant and uptake. Iron has a relatively wide sufciency concentration range in plant tissue between the proposed critical concentrations for Fe deciency and toxicity. At the tillering stage, the sufciency range of Fe concentrations in the leaf blades ranges from 70 to 300 mg Fe kg21 (Wells et al., 1993). Iron deciency or toxicity occurs at concentrations below or above this sufciency range. Fageria et al. (1997a) also reported a similar Fe sufciency concentration range, but noted the concentration varied depending on the plant
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age and part analyzed. Dobermann and Fairhurst (2000) suggested the sufcient Fe concentration range of the Y-leaf was 75 150 mg Fe kg21 during vegetative growth. The sufcient Fe concentration of the whole shoots was somewhat lower at 60 100 mg Fe kg21. Gilmour (1977a) found that whole plant and bottom leaf Fe concentrations were signicantly greater than that for the Y-leaf up to 69 days after seeding. Similar to other nonmobile elements, the concentration of Fe usually increases as leaf age increases. The use of tissue Fe concentrations is useful only to the extent of establishing whether the total Fe concentration falls within the suggested sufciency range. The most promise for diagnosis of Fe deciency by tissue analysis is perhaps by analyzing for active Fe. Iron toxicity is believed to occur when leaf blade total Fe concentrations exceed 300 mg Fe kg21 (Tanaka et al., 1966). Dobermann and Fairhurst (2000) indicated this critical concentration was specically for the Y-leaf. The importance of leaf age, the plant part sampled, and sample cleanliness cannot be overemphasized for the diagnosis of Fe toxicity. The presence of toxic concentrations of Fe may simply be the result of another nutrient deciency. For example, Zn decient whole aboveground rice seedlings commonly contain Fe concentrations . 300 mg Fe kg21. However, clean Y-leaf tissue is seldom above this threshold. Whole plant samples of nutritionally healthy plant samples may easily exceed 300 mg Fe kg21 from contamination of Fe precipitates on the rice stems unless the tissues are thoroughly washed in a mild acid solution before drying and analysis. Representative Fe concentrations and total plant contents of lowland rice grown on an Inceptisol of central Brazil and an Alsol in Arkansas are listed in Table XII. The concentration of Fe in the whole-aboveground tissue is rather high during early tillering and then decreases during the growing season. Rice grain generally contains about 30 mg Fe kg21 so only a fraction of the total plant Fe content is actually removed in the harvested grain. Iron fertilization practices. Soil analysis is not a highly effective means of identifying Fe decient soils. Sims and Johnson (1991) reported that for most crops the critical soil Fe concentration range was 2.5 5.0 mg kg21 of DTPA extractable Fe, but is also inuenced by soil pH. For example, Fe deciency of upland rice occurs on Brazilian Oxisols with Mehlich 1 extractable Fe of more than 50 mg kg21 when the soil pH is raised above 6.0 by liming (Fageria, 2000c). As a general rule, Fe deciency is caused by soil chemical properties that affect Fe availability and not by low Fe content. Organic soils may be the exception to this rule. Snyder and Jones (1988) showed the soil ash content (i.e., weight) following combustion of soil at 5508C was not a reliable indicator of seedling chlorosis on Histosols in the Florida Everglades. However, the total iron content of the soil ash was consistently lower from eld areas that exhibited chlorosis. Further, they observed the color of the ash ranged from white to yellow to a yellow-red color. Soils with white-colored ash were associated with eld areas

Table XII Seasoncal Whole Plant Fe Concentration and Content in Rice Straw and Grain of FloodIrrigated Rice Grown in Arkansas (Slaton, unpublished data, 2001) and Brazil (Fageria, unpublished data, 2001) Brazila DAEc 22 35 71 97 112 140 140 Concentration (mg Fe kg21) 419 374 175 155 179 448 81 Content (g Fe ha21) 134 411 1045 1403 2202 3499 265 DAEc 55 81 96 111 145 146 Arkansasb Concentration (mg Fe kg21) 371 152 198 171 169 27d Content (g Fe ha21) 216 638 1535 1421 1653 219d
Growth stage Beginning tillering Active tillering Panicle initiation Boot stage Flowering Mature-straw Grain
a b
The rice cultivar was Metica 1 and was grown on an Inceptisol. The rice cultivar was Wells and was grown on a slighty calcareous Calloway silt loam. c DAE, Days after emergence. d Harvested grain. Whole panicle Fe concentration at maturity was 99 mg Fe kg21.
that produced chlorotic rice and appeared to be a highly reliable means of predicting Fe deciency on these organic soils. Examination of the ash color following soil combustion was highly correlated with rice response to Fe fertilization on these organic soils. The Fe deciency problems common to the organic soils used to produce rice in Florida initiated research to investigate the most efcient methods of Fe fertilization. Water-soluble Fe fertilizers that are broadcast applied to the soil are rapidly converted to insoluble forms that have limited effectiveness unless applied at very high rates (Martens and Westermann, 1991). Snyder and Jones (1988) found that application of 20 30 kg Fe ha21, as water-soluble Fe (i.e., FeSO4H2O and FeSO47H2O), with the seed was highly effective at preventing seedling chlorosis and increasing rice yield. Foliar application of Fe solutions varied in their effectiveness to prevent Fe chlorosis (Snyder and Jones, 1991). Foliar applications of chelated Fe (1 kg Fe ha21) made 1 week after emergence were equivalent to Fe drilled with the seed. Greenhouse studies also suggested that water-soluble FeSO4 was equal to chelated Fe when applied in a timely manner and with the proper water source. However, when foliar Fe was applied at 2 and 3 weeks after emergence, signicant yield losses occurred. Thus, under severe Fe deciency, preventative measures, before or at the time of seeding, are preferred over corrective measures that are initiated after deciency symptoms are expressed. Reducing soil pH to increase Fe availability has proven to be an effective means of ameliorating Fe deciencies of other crops (Martens and
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Westermann, 1991). However, soil acidication has not been thoroughly evaluated on Fe decient soils used for rice production to develop best management practices. The majority of published research regarding Fe fertilization of rice has been conducted on organic soils in Florida. We assume the same fertilization rates and methods proven effective on organic soils would be equally effective on Fe decient mineral soils. Fertilization practices used to ameliorate Fe toxicity include periodic surface drainage to reduce soil solution Fe concentrations, liming to increase soil pH, and the routine application of nutrients to maintain soil fertility. The Fe-excluding ability of rice plants is lowered by deciencies of P, K, Ca, and Mg (Obata, 1995). In particular, K deciency readily induces Fe toxicity. Use of tolerant rice cultivars is another alternative of producing rice on Fe toxic soils (Fageria et al., 1990b). Rice cultivars also show different susceptibilities to Fe toxicity (Gunawarkena et al., 1982; Fageria and Rabelo, 1987; Wu et al., 1997). 4. Manganese
Manganese deciencies and toxicities have been documented in some rice growing regions, but are not common in most soils used for rice production. Nutritional disorders related to Mn occur less frequently than those associated with Fe. Rice grown on Oxisols and Histosols are most likely to experience disorders related to Mn nutrition. Manganese deciency may cause yield losses that approach 100% from seedling mortality, but yield losses are usually less than 30% (Snyder et al., 1990). Chemistry in the soil. Similar to Fe, Mn is widely distributed in soils, but largely in the form of high-valency oxides, which are unavailable to plants. Manganese oxides are the most common manganese minerals in soil and include pyrolusite (MnO2), manganite (MnOOH), and hausmannite (Mn3O4). Soil manganese exists in three oxidation states including Mn2, Mn3, and Mn4 with Mn2 being the primary form absorbed by plants. The predominant Mn oxidation states in most soils are Mn2 and Mn4, with much more as Mn4 than Mn2 in aerated soils (Barber, 1995). Small amounts of Mn2 are present in soils as exchangeable ions and as organic complexes and these constitute the source for plant uptake. It has been reported that 80 90% of the Mn in the soil solution is complexed with organic matter (Foth and Ellis, 1988). The concentration of Mn2 in the soil is affected by soil pH and oxidation reduction reactions. Its solubility decreases 100-fold for each unit increase in pH (Barber, 1995; Tisdale et al., 1985). In submerged soils Mn4 is reduced to Mn2 due to oxygen depletion. The Mn2 concentration in the soil solution increases when redox potential (Eh ) values decrease. The reduction of Mn4 occurs after 3 NO2 reduction. Patrick and Reddy (1978) classied 3 reduction and before Fe
soils based on redox potential as aerated or well-drained soils, 700 to 500 mV; moderately reduced, 400 to 200 mV; reduced, 100 to 2 100 mV; and highly reduced, 2 100 to 2 300 mV. Under reduced soil conditions, rice uptake of Mn increases. The reduction of Mn can be either chemical or microbiological, although microbiological reduction is likely to predominate in ooded rice soils that are at about pH 5.5 6.0 (Patrick and Reddy, 1978). The reduction of Mn can be explained from the reduction equation as: MnO2 4H 2e2 $ Mn2 2H2 O The adequate Mn concentration for rice growth in water culture experiments has been reported as 0.1 0.5 mg L21 (Shimada, 1995) and concentrations higher than 10 mg L21 may be toxic (Yoshida, 1981). Functions and symptoms of Mn deciency and toxicity. Manganese is immobile in the plant and Mn deciency symptoms appear initially in the younger leaves. Manganese decient plants are chlorotic and develop an irregular yellow mottling between the leaf veins. In the case of Mn toxicity, yellow spots generally develop between leaf veins, extend to the interveinal areas, and eventually turn brown as the toxicity develops (Shimada, 1995). In rice, Mn toxicity is rarely observed because rice roots can effectively exclude Mn preventing its uptake (Tanaka et al., 1975) and the rice plant is relatively tolerant to high Mn concentrations. Excessive uptake of Mn by rice is generally suppressed antagonistically by the coexistence of high concentrations of Fe2. More than 60% of the Mn contained in higher plant leaves is found in the chloroplast, and, along with Fe and Cu, performs vital roles in the electron transport system (Obata, 1995). Manganese also functions as a cofactor to activate enzymes such as dehydrogenases and hydrolyses in glycolysis system and citric acid cycle and RNA polymerases in the chloroplasts. The protease enzyme contained in rice seeds is activated by Mn (Horiguchi and Kitagishi, 1976). The most well-known and extensively studied function of Mn in green plants is its involvement in photosynthetic O2 evolution (Marschner, 1995). Critical level in plant and uptake. Manganese deciency in rice occurs when the Mn concentration in the plant tissue is less than 20 mg Mn kg21 (Wells et al., 1993). The critical tissue concentration of Mn in most plants ranges from 10 to 20 mg Mn kg21 in mature leaves, and is surprisingly consistent regardless of the plant species, cultivar, or the prevailing environmental conditions (Marschner, 1995). Fageria et al. (1997a) reported whole plant Mn concentrations of 30 600 mg Mn kg21 at tillering were sufcient. Rice can tolerate tissue levels of more than 2500 mg Mn kg21 without adverse effects on either growth or grain yield (Wells et al., 1993). Cheng and Quellette (1971) reported a critical, toxic tissue concentration for rice of 7000 mg
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Mn kg21. Yoshida (1981) reported that in many cases, a high Mn content in rice tissues is frequently associated with high yields, possibly indicating that high Mn content in the soil is associated with various favorable soil conditions. Manganese fertilization practices. Soil analysis can be a useful guide for Mn fertilization of annual crops. Sims and Johnson (1991) reported the critical level of Mehlich 1 extractable Mn of 5 mg Mn kg21 at soil pH 6.0. These authors also reported the critical soil Mn concentration range for most crops of 1 5 mg Mn kg21 by the DTPA extraction method. Both inorganic and organic Mn fertilizers are used for correcting Mn deciencies. The use of MnSO4 is so common that it is usually included as the standard in research to determine the efcacy of other Mn sources (Martens and Westermann, 1991). Manganese deciency is usually corrected by foliar application of Mn or by banding Mn with an acid-forming fertilizer. If it is broadcast, a high rate is required. Martens and Westermann (1991) reported broadcast rates varied from 6 to 45 kg Mn ha21 and band application rates varied from 3 to 22 kg Mn ha21 for annual crops. If Mn deciency occurs, a foliar application of MnSO4 or MnCl2 can be effectively adopted as control measures. The deciency often occurs in neutral or alkaline soils. Application of acidic fertilizers can decrease soil pH and improve Mn uptake by plants as long as the acidic soil amendments are applied far enough in advance to reduce the soil pH. Manganese deciencies of rice occur on high-pH organic soils used for rice production in the Florida Everglades Agricultural Area (Snyder et al., 1990). Manganese drilled with the rice seed at planting was the most effective method of Mn fertilization on these organic soils. Soaking the rice seed in a Mn solution before seeding and foliar sprays failed to signicantly increase rice growth in greenhouse studies. Flooding the soil in advance of seeding to allow for soil reduction to increase the Mn concentration in the soil solution produced mixed results in studies conducted by Snyder et al. (1990), but could possibly be used as a cultural practice in waterseeded or transplanted systems to help alleviate Mn deciency if Mn fertilizer is not readily available. The Mn status of the main crop also signicantly affected the yield of the ratoon crop. Snyder et al. (1990) recommended drilling 15 kg Mn ha21 as a water-soluble Mn source with rice seed as the most economical and effective means of preventing Mn deciency of rice.
IV. CONCLUSIONS
Rice is the staple food of approximately one-half of the worlds population. Total rice production will need to increase to help feed a large proportion of the worlds increasing population. Increasing rice yields, especially in lowland
production systems, per unit of land area will be essential in order to meet this challenge. Forty years ago, when the Green Revolution started, persuading rice farmers to use modern varieties and accompanying fertilizer inputs was easy because the results, in terms of yield increases, were often spectacular and economically attractive. Further improvements in rice productivity, however, are likely to be much more incremental and knowledge-based. Further yield increases will mostly result from the positive interactions and integrated management of various agronomic inputs including nutrients, pest control, improved cultivars, and water management. Our knowledge of nutrient management and behavior under the ooded soil conditions used to produce lowland rice has progressed rapidly over the past several decades. Many nutrient management issues still require signicant improvement and will require a great of deal of research. Our ability to predict the need for supplemental fertilization of ood-irrigated rice, via routine plant and soil analysis, needs to be further rened to optimize production and nutrient use efciency. Flooded rice soils are characterized by the absence of oxygen. Oxygen diffusing into a ooded soil may be consumed by microbial respiration where it is used as an electron acceptor in the chemical oxidation of reduced Fe2 and Mn2, the biological oxidation of NH4 and C, and the oxidation of suldes. Upon depletion of oxygen, oxidized compounds are reduced by anaerobic organisms in the soil. Nitrate, NO2, the higher oxides of Mn, hydrated ferric oxide and SO4 will be reduced if an energy source is available to the microorganisms. The principal reduction reactions in ooded rice soils in sequence are O2/H2O, NO2 3 /N2, 2 Mn(IV, III)/Mn(II), Fe(III)/Fe(II), SO2 4 /H2S, and CO2/CH4. Flooding the soil has a signicant effect on the behavior of several essential plant nutrients and on the growth and yield of rice. Some nutrients are increased in availability to the crop, whereas others are subjected to greater xation or loss from the soil as a result of ooding. Changes in plant nutrient availability resulting from ooding are due to biological oxidation reduction processes brought into play by the depletion of oxygen from the ooded soil. Flooding the soil results in the potential loss of N through leaching and denitrication. When managed properly a high level of N use efciency can be obtained in oodirrigated rice systems. However, if N inputs are mismanaged N use efciency can be very low. Saturation of the soil with water increases the availability of soil P to rice. This has usually been attributed to the reduction of ferric phosphates to the more soluble ferrous phosphates, and to the hydrolysis of P compounds. The increase in soil pH of acidic soils as a result of submergence is also considered to aid in the solution of Fe and Al phosphates. Potassium is less affected by ooding than are N and P. Reduced soil conditions results in the displacement of exchangeable K from the exchange complex into the soil solution. This process makes K more available for uptake by rice, but may also enhance the potential for K leaching on some soils. The availability of some nutrients, such as Ca and Mg, are changed
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only to a limited extent by the biochemical changes associated with ooding. Other nutrients, such as S, Fe, and Mn undergo marked changes following waterlogging. The solubility, and presumably the availability, of Fe and Mn should be increased as a result of the increased solubility of sparingly soluble compounds, whereas B, Cu, and Zn availability may decrease due to adsorption as soil pH increases in acid soils. Reduction of Fe3 may increase the Fe2 concentration in the soil solution to the extent that toxicity can occur. Modern production agriculture requires efcient, sustainable, and environmentally sound management practices. Under these situations, increasing rice yield per unit area through the use of appropriate nutrient management practices has become an essential component of modern rice production technology. Adoption of proper nutrient management strategies that include the use of appropriate nutrient sources, rates, and application times used in conjunction with high-yielding rice cultivars bred for high-nutrient efciency may reduce production costs and improve rice yields. Development of efcient nutrient management recommendations for lowland rice production systems must integrate our basic knowledge of soil physical properties, nutrient cycles, biochemical transformation processes, and rice growth and nutrient uptake under ooded soil conditions. It is clear that the irrigation water must be used to manipulate these processes to produce nutrient management systems that are agronomically and environmentally efcient. Many of these basic relationships are well dened and already recommended in many rice-producing regions, but require some renement while others still require signicant improvement. Our challenge is to continue to incorporate new and emerging technologies into practical management recommendations and effectively demonstrate their value so that growers readily adopt them as routine.
ACKNOWLEDGMENTS
The authors are grateful to Dr C.D. Foy, USDA-ARS, Beltsville, for peer review and giving useful comments on the manuscript.
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NUTRIENT MANAGEMENT FOR IMPROVING LOWLAND RICE PRODUCTIVITY AND SUSTAINABILITY

N.K. Fageria,a N.A. Slatonb and V.C. Baligarc

NUTRIENT MANAGEMENT FOR IMPROVING LOWLAND RICE 65

II. SOILS USED FOR RICE PRODUCTION

N. K. FAGERIA, N. A. SLATON AND V. C. BALIGAR

NUTRIENT MANAGEMENT FOR IMPROVING LOWLAND RICE 67

III. MANAGEMENT PRACTICES TO IMPROVE NUTRIENT USE EFFICIENCY

N. K. FAGERIA, N. A. SLATON AND V. C. BALIGAR

NUTRIENT MANAGEMENT FOR IMPROVING LOWLAND RICE 69

N. K. FAGERIA, N. A. SLATON AND V. C. BALIGAR

NUTRIENT MANAGEMENT FOR IMPROVING LOWLAND RICE 71

N. K. FAGERIA, N. A. SLATON AND V. C. BALIGAR

NUTRIENT MANAGEMENT FOR IMPROVING LOWLAND RICE 73

Functions and Deciency Symptoms

N. K. FAGERIA, N. A. SLATON AND V. C. BALIGAR

Seasonal N Uptake and Partitioning

NUTRIENT MANAGEMENT FOR IMPROVING LOWLAND RICE 75

N. K. FAGERIA, N. A. SLATON AND V. C. BALIGAR

NUTRIENT MANAGEMENT FOR IMPROVING LOWLAND RICE 77

112 (F) 8 7 9 8 9 9 10 10 0.72*

140 (PM) 5 6 6 6 6 6 7 7 0.75**

97 (B) 50 62 82 88 91 113 137 130 0.95**

112 (F) 57 66 95 89 122 113 135 130 0.92**

140 (PM) 28 37 40 41 48 52 59 66 0.98**

N. K. FAGERIA, N. A. SLATON AND V. C. BALIGAR

NUTRIENT MANAGEMENT FOR IMPROVING LOWLAND RICE 79

N. K. FAGERIA, N. A. SLATON AND V. C. BALIGAR

NUTRIENT MANAGEMENT FOR IMPROVING LOWLAND RICE 81

Root 35a 35a 26a 32a 11 6

Unaccounted for N % of applied 15 N fertilizer 11 25 33 26 29 35 33 46 21 44 48 46 60 25 16

N. K. FAGERIA, N. A. SLATON AND V. C. BALIGAR

NUTRIENT MANAGEMENT FOR IMPROVING LOWLAND RICE 83

N. K. FAGERIA, N. A. SLATON AND V. C. BALIGAR

NUTRIENT MANAGEMENT FOR IMPROVING LOWLAND RICE 85

N Fertilizer rate (kg N ha21) 30 60 90 120 150 180 210 Average R

Agrophysiological efciency (Dkg grain Dkg21 N uptake) 72 73 75 66 57 51 46 63 0.87**

Apparent recovery efciency (%) 49 50 37 38 34 33 32 39 0.82**

*,**Signicant at the 5 and 1% probability levels, respectively.

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Total 20.1 23.6 26.6 33.7 31.4 30.6 27.8

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Chemistry in the Soil

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Functions and Deciency Symptoms

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Potassium Use Efciency

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