Biological nitrogen fixation and prospects for ecological intensification in cereal-based cropping systems.

The demand for nitrogen (N) for crop production increased rapidly from the middle of the twentieth century and is predicted to at least double by 2050 to satisfy the on-going improvements in productivity of major food crops such as wheat, rice and maize that underpin the staple diet of most of the world's population. The increased demand will need to be fulfilled by the two main sources of N supply - biological nitrogen (gas) (N2) fixation (BNF) and fertilizer N supplied through the Haber-Bosch processes. BNF provides many functional benefits for agroecosystems. It is a vital mechanism for replenishing the reservoirs of soil organic N and improving the availability of soil N to support crop growth while also assisting in efforts to lower negative environmental externalities than fertilizer N. In cereal-based cropping systems, legumes in symbiosis with rhizobia contribute the largest BNF input; however, diazotrophs involved in non-symbiotic associations with plants or present as free-living N2-fixers are ubiquitous and also provide an additional source of fixed N. This review presents the current knowledge of BNF by free-living, non-symbiotic and symbiotic diazotrophs in the global N cycle, examines global and regional estimates of contributions of BNF, and discusses possible strategies to enhance BNF for the prospective benefit of cereal N nutrition. We conclude by considering the challenges of introducing in planta BNF into cereals and reflect on the potential for BNF in both conventional and alternative crop management systems to encourage the ecological intensification of cereal and legume production.

Steady agronomic and genetic interventions are essential for sustaining productivity in intensive rice cropping.

Intensive systems with two or three rice (Oryza sativa L.) crops per year account for about 50% of the harvested area for irrigated rice in Asia. Any reduction in productivity or sustainability of these systems has serious implications for global food security. Rice yield trends in the world's longest-running long-term continuous cropping experiment (LTCCE) were evaluated to investigate consequences of intensive cropping and to draw lessons for sustaining production in Asia. Annual production was sustained at a steady level over the 50-y period in the LTCCE through continuous adjustment of management practices and regular cultivar replacement. Within each of the three annual cropping seasons (dry, early wet, and late wet), yield decline was observed during the first phase, from 1968 to 1990. Agronomic improvements in 1991 to 1995 helped to reverse this yield decline, but yield increases did not continue thereafter from 1996 to 2017. Regular genetic and agronomic improvements were sufficient to maintain yields at steady levels in dry and early wet seasons despite a reduction in the yield potential due to changing climate. Yield declines resumed in the late wet season. Slower growth in genetic gain after the first 20 y was associated with slower breeding cycle advancement as indicated by pedigree depth. Our findings demonstrate that through adjustment of management practices and regular cultivar replacement, it is possible to sustain a high level of annual production in irrigated systems under a changing climate. However, the system was unable to achieve further increases in yield required to keep pace with the growing global rice demand.

Can productivity and profitability be enhanced in intensively managed cereal systems while reducing the environmental footprint of production? Assessing sustainable intensification options in the breadbasket of India.

In the most productive area of the Indo-Gangetic Plains in Northwest India where high yields of rice and wheat are commonplace, a medium-term cropping system trial was conducted in Haryana State. The goal of the study was to identify integrated management options for further improving productivity and profitability while rationalizing resource use and reducing environmental externalities (i.e., "sustainable intensification", SI) by drawing on the principles of diversification, precision management, and conservation agriculture. Four scenarios were evaluated: Scenario 1 - "business-as-usual" [conventional puddled transplanted rice (PTR) followed by (fb) conventional-till wheat]; Scenario 2 - reduced tillage with opportunistic diversification and precision resource management [PTR fb zero-till (ZT) wheat fb ZT mungbean]; Scenario 3 - ZT for all crops with opportunistic diversification and precision resource management [ZT direct-seeded rice (ZT-DSR) fb ZT wheat fb ZT mungbean]; and Scenario 4 - ZT for all crops with strategic diversification and precision resource management [ZT maize fb ZT wheat fb ZT mungbean]. Results of this five-year study strongly suggest that, compared with business-as-usual practices, SI strategies that incorporate multi-objective yield, economic, and environmental criteria can be more productive when used in these production environments. For Scenarios 2, 3, and 4, system-level increases in productivity (10-17%) and profitability (24-50%) were observed while using less irrigation water (15-71% reduction) and energy (17-47% reduction), leading to 15-30% lower global warming potential (GWP), with the ranges reflecting the implications of specific innovations. Scenario 3, where early wheat sowing was combined with ZT along with no puddling during the rice phase, resulted in a 13% gain in wheat yield compared with Scenario 2. A similar gain in wheat yield was observed in Scenario 4 vis-à-vis Scenario 2. Compared to Scenario 1, wheat yields in Scenarios 3 and 4 were 15-17% higher, whereas, in Scenario 2, yield was either similar in normal years or higher in warmer years. During the rainy (kharif) season, ZT-DSR provided yields similar to or higher than those of PTR in the first three years and lower (11-30%) in Years 4 and 5, a result that provides a note of caution for interpreting technology performance through short-term trials or simply averaging results over several years. The resource use and economic and environmental advantages of DSR were more stable through time, including reductions in irrigation water (22-40%), production cost (11-17%), energy inputs (13-34%), and total GWP (14-32%). The integration of "best practices" in PTR in Scenario 2 resulted in reductions of 24% in irrigation water and 21% in GWP, with a positive impact on yield (0.9 t/ha) and profitability compared to conventional PTR, demonstrating the power of simple management changes to generate improved SI outcomes. When ZT maize was used as a diversification option instead of rice in Scenario 4, reductions in resource use jumped to 82-89% for irrigation water and 49-66% for energy inputs, with 13-40% lower GWP, similar or higher rice equivalent yield, and higher profitability (27-73%) in comparison to the rice-based scenarios. Despite these advantages, maize value chains are not robust in this part of India and public procurement is absent. Results do demonstrate that transformative opportunities exist to break the cycle of stagnating yields and inefficient resource use in the most productive cereal-based cropping systems of South Asia. However, these SI entry points need to be placed in the context of the major drivers of change in the region, including market conditions, risks, and declining labor availability, and matching with the needs and interests of different types of farmers.

Role of nitrogen fertilization in sustaining organic matter in cultivated soils.

Soil organic matter (SOM) is essential for sustaining food production and maintaining ecosystem services and is a vital resource base for storing C and N. The impact of long-term use of synthetic fertilizer N on SOM, however, has been questioned recently. Here we tested the hypothesis that long-term application of N results in a decrease in SOM. We used data from 135 studies of 114 long-term experiments located at 100 sites throughout the world over time scales of decades under a range of land-management and climate regimes to quantify changes in soil organic carbon (SOC) and soil organic nitrogen (SON). Published data of a total of 917 and 580 observations for SOC and SON, respectively, from control (unfertilized or zero N) and N-fertilized treatments (synthetic, organic, and combination) were analyzed using the SAS mixed model and by meta-analysis. Results demonstrate declines of 7 to 16% in SOC and 7 to 11% in SON with no N amendments. In soils receiving synthetic fertilizer N, the rate of SOM loss decreased. The time-fertilizer response ratio, which is based on changes in the paired comparisons, showed average increases of 8 and 12% for SOC and SON, respectively, following the application of synthetic fertilizer N. Addition of organic matter (i.e., manure) increased SOM, on average, by 37%. When cropping systems fluctuated between flooding and drying, SOM decreased more than in continuous dryland or flooded systems. Flooded rice ( L.) soils show net accumulations of SOC and SON. This work shows a general decline in SOM for all long-term sites, with and without synthetic fertilizer N. However, our analysis also demonstrates that in addition to its role in improving crop productivity, synthetic fertilizer N significantly reduces the rate at which SOM is declining in agricultural soils, worldwide.

Spatial and seasonal distribution of nitrate-N in groundwater beneath the rice-wheat cropping system of India: a geospatial analysis.

Increased use of nitrogenous fertilizers in the intensively cultivated rice (Oryza sativa)-wheat (Triticum aestivum) cropping system (covers a 13.5-ha m area in South Asia) has led to the concentration of nitrates (NO(3)-N) in the groundwater (GW) in Haryana State of India. Six districts from the freshwater zone were selected to identify factors affecting NO(3)-N enrichment in GW. Water and soil samples were collected from 1,580 locations and analyzed for their chemical properties. About 3% (26,796, and 10,588 ha) of the area was estimated to be under moderately high (7.5-10 mg l( -1)) and high (>10 mg l( -1)) risk categories, respectively. The results revealed that NO(3)-N was 10-50% higher during the pre-monsoon season than in the monsoon season. Nitrate-N decreased with the increase in aquifer depth (r (2) = 0.99). Spatial and proximity analyses using ArcGIS (9.2) revealed that (1) clay material in surface and sub-surface texture restricts N leaching, (2) piedmont and rolling plains act as an N sink, and (3) perennial rivers bring a dilution effect whereas seasonal rivers provide favorable conditions for NO(3) (-) enrichment. The study concludes that chemical N fertilizers applied in agro-ecosystems are not the sole factor determining the NO(3) in groundwater; rather, it is an integrated process governed by several other factors including physical and chemical properties of soils, proximity and type of river, and geomorphologic and geographical aspects. Therefore, future studies should adopt larger area (at least watershed scale) to understand the mechanistic pathways of NO(3) enrichment in groundwater and interactive role of the natural drainage system and surrounding physical features. In addition, the study also presents a conceptual framework to describe the process of nitrate formation and leaching in piedmont plains and its transportation to the mid-plain zone.

Metabolic engineering of rice with soybean isoflavone synthase for promoting nodulation gene expression in rhizobia.

Isoflavonoids are derived from a flavonone intermediate, naringenin, that is ubiquitously present in plants, and play a critical role in plant development and defence response. Isoflavonoids secreted by the legumes also play an important role in promoting the formation of nitrogen-fixing nodules by symbiotic rhizobia. In these plants, the key enzyme that redirects phenylpropanoid pathway intermediates from flavonoids to isoflavonoids is the cytochrome P450 mono-oxygenase, isoflavone synthase. In an effort to develop a rice variety possessing the ability to induce nodulation (nod) genes in rhizobia, the IFS gene from soybean was incorporated into rice (Oryza sativa L. cv. Murasaki R86) under the control of the 35S promoter. The presence of IFS in transgenic rice was confirmed by PCR and Southern blot analysis. Analyses of the 35S-IFS transgenic lines demonstrated that the expression of the IFS gene led to the production of the isoflavone genistein in rice tissues. These results showed that the soybean IFS gene-expressed enzyme is active in the R86 rice plant, and that the naringenin intermediate of the anthocyanin pathway is available as a substrate for the introduced foreign enzyme. The genistein produced in rice cells was present in a glycoside form, indicating that endogenous glycosyltransferases were capable of recognizing genistein as a substrate. Studies with rhizobia demonstrated that the expression of isoflavone synthase confers rice plants with the ability to produce flavonoids that are able to induce nod gene expression, albeit to varied degrees, in different rhizobia.

Infection and colonization of rice seedlings by the plant growth-promoting bacterium Herbaspirillum seropedicae Z67.

A beta-glucoronidase (GUS)-marked strain of Herbaspirillum seropedicae Z67 was inoculated onto rice seedling cvs. IR42 and IR72. Internal populations peaked at over 10(6) log CFU per gram of fresh weight by 5 to 7 days after inoculation (DAI) but declined to 10(3) to 10(4) log CFU per gram of fresh weight by 28 DAI. GUS staining was most intense on coleoptiles, lateral roots, and at the junctions of some of the main and lateral roots. Bacteria entered the roots via cracks at the points of lateral root emergence, with cv. IR72 appearing to be more aggressively infected than cv. IR42. H. seropedicae subsequently colonized the root intercellular spaces, aerenchyma, and cortical cells, with a few penetrating the stele to enter the vascular tissue. Xylem vessels in leaves and stems were extensively colonized at 2 DAI but, in later harvests (7 and 13 DAI), a host defense reaction was often observed. Dense colonies of H. seropedicae with some bacteria expressing nitrogenase Fe-protein were seen within leaf and stem epidermal cells, intercellular spaces, and substomatal cavities up until 28 DAI. Epiphytic bacteria were also seen. Both varieties showed nitrogenase activity but only with added C, and the dry weights of the inoculated plants were significantly increased. Only cv. IR42 showed a significant (approximately 30%) increase in N content above that of the uninoculated controls, and it also incorporated a significant amount of 15N2.