Nitrogen Use Efficiency

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Breeding for Nitrogen Use Efficiency in Cereals

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Indian Agricultural Research Institute, New Delhi Breeding For Nitrogen Use Efficiency in Cereals Rumesh Ranjan Division of Genetics IARI, New- Delhi-12. Welcome

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Indian Agricultural Research Institute, New Delhi Content I ntroduction a) Cereals b) Production of Cereals in India c) Use of Nitrogenous Fertilizer d) NUE e) Pathway of N from Rhizosphere to seed Breeding approaches for NUE a) Conventional Breeding Approaches b) Molecular Approaches c) Transgenic Approaches Patent for NUE Conclusion

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Indian Agricultural Research Institute, New Delhi Introduction: Cereals are the world’s most important sources of food - direct human consumption and as inputs to livestock production. > 70 % world populations staple food are cereals. FAO Cereal supply and Demand, 2015

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Indian Agricultural Research Institute, New Delhi India is the world's second largest producer of Rice, Wheat and other cereals. Huge demand for cereals in the global market create an excellent environment for the export of Indian cereal products. India's export of cereals (2014-15): Rice- INR. 58279.80 crore (including Basmati and Non Basmati ) wheat- INR 4991.8 crore Exports (2014-15) : Rice- 64.4 % Other Cereals + Wheat- 35.60% Major Importing Countries- Iran, UAE, Bangladesh and Indonesia. Economics Importance of Cereals- India APEDA, 2015

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Story of gloom to glory India pathetically depended on US food aid during 1960s (8-10 MT annual import). Paul Ehrlich predicted in his 60’s bestseller The Population Bomb that "hundreds of millions" would starve to death in the 1970s and 1980s. (Richard , 2003). Then came 14 gms of dwarf wheat on request from USDA that ushered wheat revolution, followed by rice revolution resulting in ‘GREEN REVOLUTION’. Increased nitrogen (N) fertilization , Irrigation, Plant protection, Cultural practices along with dwarf varieties was an important factor in the increases in yield achieved during the 20th century. Indian Agricultural Research Institute, New Delhi

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Indian Agricultural Research Institute, New Delhi Trends in Cereals production in India Production X5 Productivity X3 Area X0.25 FAO, 2012

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Indian Agricultural Research Institute, New Delhi Why Nitrogenous Fertilizer Nitrogen is 4 th most Essential element of Plant after C, H, O. Bio Chemical function of N: - major component of chlorophyll. - Amino acid - Enzyme - Energy transfer Compound ( ATP) - Nucleic acid ( DNA, RNA) Crop Nutrition.com

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Indian Agricultural Research Institute, New Delhi Global Fertilizer Trends Source: Potash Corp Haber–Bosch process (N fertilizer production)Requires approximately 1% of the world’s annual energy supply. Cost of production of N fertilizer increases .

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Indian Agricultural Research Institute, New Delhi Global Fertilizer Use by Crops Fertilizer use by region Cereals- 1/2 of world fertilizer consumption. Fruits, vegetables and oilseeds - 25 percent Remaining fertilizer use is split among crops such as cotton and sugar. Nearly 2/3 rd of fertilizer used by Developing countries. China & India- 42% Potash Corp, 2010

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Life Cycle of Nitrogen Most cereal crops take up roughly 40% of applied N, the remaining 60% is lost to the environment . Approximately, 19% of the applied N -- denitrification or bacterial uptake, 20% is leached into groundwater, 21% is volatilized into N2O, NH3 . Bouwman et al. , 2002 Indian Agricultural Research Institute, New Delhi

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Nitrogen Use Efficiency (NUE) The efficiency of uptake and the efficiency with which the N absorbed is utilized to produce grain, are the two primary components of N use efficiency (Moll et al., 1982). NUE is estimated at about 33%, with NUE’s of 42 and 29% in developed and developing countries ( Raun and Johnson 1999). There is scope to improve the NUE within adapted germplasm by up to 20% ( Foulkes et al ., 2009). As a Plant breeder must produce varieties that absorb N more efficiently and use it more efficiently to produce biomass. Indian Agricultural Research Institute, New Delhi

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Indian Agricultural Research Institute, New Delhi Definitions used to describe nutrient use efficiency in plants Definition Formula N itrogen use efficiency (NUE) NUE = (Total biomass (or grain wt. ) / (N supply) Uptake efficiency ( UpE ) UpE = (plant (or shoot) N content) / (N supply) Utilization efficiency (UtE) UtE = (grain mass) / (plant (or shoot) N content) Good et al., 2004

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Indian Agricultural Research Institute, New Delhi Nitrogen use by plants involves two main steps: Uptake Utilization Utilization can be further compartmentalized Assimilation Translocation⁄ remobilization ( Masclaux-Daubresse et al ., 2010). Fate of Nitrogen Use

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Indian Agricultural Research Institute, New Delhi Pathway of nitrogen from rhizosphere to plants www. studyblue.com

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Indian Agricultural Research Institute, New Delhi N-Uptake Plant roots have uptake systems for both NO3- and NH4+ with different affinities. Plants adapted to low pH , flooded wet land - ammonium Plants adapted to higher pH and more aerobic soils – nitrate ( Maathuis , 2009). N03 uptake at the root level - Two nitrate transport systems have been shown to coexist in plants and to act co- ordinately to take up nitrate from the soil solution. NRT1 gene family mediates the root low-affinity transport system(LATS) NRT 2- High-affinity transport system (HATS). ( Tsay et al., 2007). NH4 uptake occur by AMT gene , with 6 genes belonging to the same family of ammonium transporters were found in Arabidopsis ( Gazzarini et al., 1999).

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Indian Agricultural Research Institute, New Delhi N-Uptake and Assimilation Masclaux-Daubresse C, et al., 2010

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Indian Agricultural Research Institute, New Delhi McAllister et al, 2012. Nitrogen uptake, assimilation and remobilization in roots, leaves and seeds.

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Indian Agricultural Research Institute, New Delhi Nitrogen management in various crops Growth stage in context to N- uptake and Utilization The relative contribution (%) of N remobilization and post flowering N uptake in different crops Hirel B. et al 2007

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Indian Agricultural Research Institute, New Delhi Approaches to Improve NUE Conventional Breeding Approches Molecular and Genetic Engineering Approches Quantitative Trait Loci Transgenes for NUE

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Indian Agricultural Research Institute, New Delhi Conventional Breeding Approaches Genetic variability for morpho -physiological traits related to nitrogen use efficiency (NUE) and yield . Ortiz Monasterio et al. 1997, revealed at low N rates, uptake efficiency is dominant compared to utilization efficiency, whereas utilization efficiency is relatively more important than uptake efficiency at high N rates. Similarly, studies on nitrogen use efficiency (NUE) influencing traits on wheat genotypes revealed significant genetic variation and differences in some morphological characteristics (Le Gouis et al. , 2000 ; Gorny et al. , 2011; Miranzadeh et al., 2011).

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Indian Agricultural Research Institute, New Delhi Author and Year Material Used Findings Chandna et al. (2012) Ten genotypes of bread wheat Grown in complete nutrient solution with nitrogen-insufficient, nitrogen-moderate and nitrogen-sufficient nitrogen (N) levels. Genotypes, VL829 was identified as the most N-efficient, while HUW234 the most N-inefficient. . Khalilzade et al. (2012) 42 bread wheat landraces. Under two varied N fertilization levels (0, 200 kg N ha -1 ) in the field conditions. High variation between the genotypes for the NUE components, Landraces G19, G26, G27 and G41 were the best genotypes for NUE characters and high yield potential. Khalilzade et al. (2013) 20 Bread wheat cultivars High levels of variation were observed among genotypes. Cultivars Girmizigul-1, Azamatli-95, Chamran and Koohdasht produced the highest grain yield under zero nitrogen application.

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Indian Agricultural Research Institute, New Delhi Trend in Grain yield and NUE Modern cultivars are developed under conditions of high soil fertility in an effort to increase productivity. 0.55 t/ ha/ decade 0.21 t /ha/decade Grain yield and b. NUE : In 39 experimental trials with wheat cultivars evaluated. Sylvester-Bradley and Kindred (2009)

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Indian Agricultural Research Institute, New Delhi Conflict of gene action Additive and non-additive genetic effects are responsible for the genetic control of NUE and GY for maize grown in soils with high N availability. Under Low N, observed that the additive genetic effects were more important for NUE in maize than the non-additive effects. Brasília, 2008 Under high N-fertilization, the efficiency components were inherited in a manner favourable for wheat selection (preponderance of additive effects ). Euphytica (2011) 177:191–206

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Indian Agricultural Research Institute, New Delhi Quantitative Trait Loci (QTL) Previous studies have identified QTL controlling NUE and some of their component traits ( Agrama et al. 1999; Bertin and Gallais 2001; Hirel et al. 2001; Gallais and Hirel 2004) QTLs for N-uptake and N utilization efficiency QTLs for glutamine synthetase (GS) activity QTLs for glutamate dehydrogenase (GDH) Quantitative trait loci (QTL) mapping provides the best understanding of the genetic control and inheritance of NUE and can be used to indicate the best selection strategy.

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Main QTL studies on NUE in plants Indian Agricultural Research Institute, New Delhi

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Indian Agricultural Research Institute, New Delhi Arche X Récital Tolerant to Low N Susceptible 241 DHL Already studied by Laperche et al. (2006; 2007) Three experiments were carried out in 2004, 2006 and 2007 at Estrées -Mons INRA experimental station (Somme, Northern France).

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Indian Agricultural Research Institute, New Delhi Number of QTL detected for glutamine synthetase (GS) and glutamate dehydrogenase (NADHGDH) activities. QTL for NADH-GDH activity were detected on 18 chromosomal regions located on 13 chromosomes: 1A, 1D, 2A, 2B, 2D, 3B, 4B, 5A, 5B, 5D, 6A, 7A and 7B. QTL for GS activity were detected on 15 chromosomal regions spread on 12 chromosomes: 1D, 2A, 2B, 2D, 3B, 3D, 4A, 4B, 5A, 5B, 5D and 7A. Among these regions common to the two studies, three of them co-localized with known major genes controlling photoperiod sensitivity (Ppd-D1 on 2D), plant height(Rht-B1 on 4B) and awnedness (B1 on 5A).

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Indian Agricultural Research Institute, New Delhi Meta-analysis is an important tool for integrating information from multiple quantitative trait loci (QTLs) studies. Pooling of results from several studies allows greater statistical power for QTL detection and more precise estimation of their genetic effects. Chinese Spring x SQ1 ( Habash et al., 2007) Hanxuan 10 x Lumai 14 (An et al., 2006) and Arche x Recital ( Laperche et al., 2007)

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Indian Agricultural Research Institute, New Delhi

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Indian Agricultural Research Institute, New Delhi Genetic engineering for improving NUE With the aim of improving NUE, researchers have used various promoters (mainly CaMV 35S) to manipulate the expression of many candidate genes involved in N uptake and metabolism. Many transgenic approaches based on overexpressing candidate genes to improve NUE have also been used.

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Indian Agricultural Research Institute, New Delhi Gene Gene Source (Gene/promoter) Engineered Plant NUE Improvement (Percent) Grown in the Field? Glutamine synthetase (GS) Bean/rice Wheat 10 No Glutamine synthetase (GS) Corn/plant virus Corn 30 No Glutamate synthase (GOGAT) Rice/rice Rice 80 No Asparagine synthetase (AS) Arabidopsis/plant virus Arabidopsis 21 No Glutamate dehydrogenase E. coli/plant virus Tobacco 10 Yes Dof1 Corn/plant virus Arabidopsis 30 No Alanine aminotransferase (ALA) Barley/canola Canola 40 Yes Alanine aminotransferase (ALA) Barley/rice Rice 31–54 Yes Genes Used to Improve NUE through Genetic Engineering Chandra H. McAllister et al., 2012

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Indian Agricultural Research Institute, New Delhi Some genes were selected to test their functionality in NUE by a transgenic approach. One such example with positive effects on NUE is an early nodulin gene OsENOD93-1 . This OsENOD93-1 gene responded significantly to both, High N and Low N. Transgenic rice plants over-expressing the OsENOD93-1 gene had increased shoot dry biomass and seed yield. This OsENOD93-1 gene was expressed at high levels in roots of wild-type (WT) plants, and its protein product was localized in mitochondria. Transgenic plants accumulated higher concentrations of total amino acids and total N in roots. A higher concentration of amino acids in xylem sap was detected in transgenic plants, especially under N stress .

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Indian Agricultural Research Institute, New Delhi wild-type (WT) Oryza sativa japonica cv. Donjin plants. Transgenic plants, ‘ Kaybonnet ’ rice ( O. sativa L.) . The constructs for over-expressing NUE candidate genes were made using a ubiquitin promoter. Agrobacterium mediated transformation was performed. Table- Yield and yield attributes in transgenic rice plants over-expressing OsENOD93-1 and wild type (WT) at different nitrogen levels.

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Indian Agricultural Research Institute, New Delhi Conclusion Transgenic plants over-expressing this OsENOD93-1 gene had higher seed yield and yield-contributing factors than WT plant. In leguminous plants, the early nodulin ( ENOD ) genes are expressed at early stages of nodule development and may mediate rhizobial infection and/or nodule organogenesis. The OsENOD93-1 gene was expressed at high levels in roots in WT plants, and its protein product was localized in mitochondria. Over-expression of this OsENOD93-1 gene promoted the accumulation of amino acids in roots and in xylem sap. How this contributed to the increase in final seed yield and in yield-contributing traits in the transgenic plants is yet to be determined. The role of this mitochondrial OsENOD93-1 gene is still under investigation

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Indian Agricultural Research Institute, New Delhi Some Patents Methods of increasing nitrogen utilization efficiency in monocot plants through genetic modification to increase the levels of alanine aminotransferase expression and plants produced. In particular, methods for increasing the biomass and yield of transgenic monocot plants grown under nitrogen limiting conditions compared to non-transgenic plants are described. In this Way, monocot plant may produced a desirable yield while reducing the high level N application.

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Indian Agricultural Research Institute, New Delhi A method employing the glutamate decarboxylase gene to increase the nitrogen uptake and nitrogen use effciency of plants has been demonstrates.

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Indian Agricultural Research Institute, New Delhi “We hope that our collaboration with Arcadia on NUE rice will help farmers achieve better yields and improve their livelihoods,” Raju Barwale , Managing Director, Mahyco said. Rice is the world’s second-largest crop, grown on 160 million hectares annually. India has the world’s largest rice growing area, with more than 44 million hectares grown annually. Beyond NUE rice, Arcadia and Mahyco are also working on a number of other technologies and crops to increase farm productivity and reduce overall environmental impact in the region. “The NUE technology is an important step toward improving food security and the environmental footprint of agriculture in Southeast Asia,” Eric Rey, President and CEO, Arcadia said.

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Indian Agricultural Research Institute, New Delhi Conclusion Plant NUE is the integration of NUpE and NUtE , and is governed by multiple interacting genetic and environmental factors. Improving NUE either through genetic engineering or marker assisted breeding is still at the stage of proof of concept. Useful gene-based markers to breeders for the production of genetically modified plants or for marker assisted selection (MAS). Linking fundamental and strategic research on the one hand, with applied plant breeding on the other, to develop new genotypes with high NUE for sustainable agriculture. The road ahead for NUE crops appears bumpy , the necessity in creating crops that require decreased N fertilizer levels is the need and has been recognized in the call for a ‘ Second Green Revolution’ .

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Indian Agricultural Research Institute, New Delhi Thanks Fertilizers cannot check this growth in population but can help the world and its nations meet their increasing food, feed and fibre demands and in sustaining the humanity. Thus in developing countries reeling under population pressure the sustainable agriculture and efficient fertilizer use must go hand-in-hand for a better tomorrow. ( Borlaug , 15 th World Congress of Soil Science)

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Indian Agricultural Research Institute, New Delhi Schematic routes of N uptake from the rhizosphere Guohua Xu etal , 2012

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Indian Agricultural Research Institute, New Delhi Pathway for Chlorophyll Synthesis

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