Biological nitrogen fixation: A scientific perspective

The discoveries of Hellriegel and Wilfarth ended the period of controversy about the existence of biological N₂ fixation and launched a period featuring the agronomic application of the inoculation of legumes. Serious studies of the biochemistry of N₂ fixation started in the late 1920's, and defined some of the basic properties of the N₂-fixing system. Application of¹⁵N as a tracer gave definitive evidence for the role of ammonia as the key intermediate in biological N₂ fixation. It was demonstrated in the 1950's and 1960's that nitrogenase could reduce substrates other than N₂. With the achievement of consistent cell-free N₂ fixation it was possible to resolve the nitrogenase system into two proteins, electron donors, and ATP-hydrolyzing and regenerating systems. The sequence of electron transfer was established. Recently, studies of the genetics of the nitrogenase system have defined in detail how the system is assembled and controlled.     

Biological nitrogen fixation in mixed legume-cereal cropping systems

Cereal/legume intercropping increases dry matter production and grain yield more than their monocultures. When fertilizer N is limited, biological nitrogen fixation (BNF) is the major source of N in legume-cereal mixed cropping systems. The soil N use patterns of component crops depend on the N source and legume species. Nitrogen transfer from legume to cereal increases the cropping system's yield and efficiency of N use. The use of nitrate-tolerant legumes, whose BNF is thought to be little affected by application of combined N, may increase the quantity of N available for the cereal component. The distance between the cereal and legume root systems is important because N is transferred through the intermingling of root systems. Consequently, the most effective planting distance varies with type of legume and cereal. Mutual shading by component crops, especially the taller cereals, reduces BNF and yield of the associated legume. Light interception by the legume can be improved by selecting a suitable plant type and architecture. Planting pattern and population at which maximum yield is achieved also vary among component species and environments. Crops can be mixed in different proportions from additive to replacement or substitution mixtures. At an ideal population ratio a semi-additive mixture may produce higher gross returns.     

Fertilizers and biological nitrogen fixation as sources of plant nutrients: Perspectives for future agriculture

Biological nitrogen fixation (BNF) has an assured place in agriculture, mainly as a source of nitrogen for legumes. Legumes are currently grown mostly as a source of vegetable oil and as food for humans and animals, but not as nitrogen source.Other crops with BNF capability may be eventually be developed eventually. Such crops will also need mineral fertilizers to maintain a good status of soil nutrients, but their possible effects to the environment is also a concern. Fertilizers, however, will remain a necessary and sustainable input to agriculture to feed the present and increasing human population. It is not a case of whether BNF is better or worse than mineral fertilizers because both plays an important role in agriculture.     

Biological nitrogen fixation: Investments,expectations and actual contributions to agriculture

Inputs of biologically fixed N into agricultural systems may be derived from symbiotic relationships involving legumes and Rhizobium spp., partnerships between plants and Frankia spp. or cyanobacteria, or from non-symbiotic associations between free-living diazotrophs and plant roots. It is assumed that these N₂-fixing systems will satisfy a large portion of their own N requirements from atmospheric N₂, and that additional fixed N will be contributed to soil reserves for the benefit of other crops or forage species. This paper reviews the actual levels of N₂ fixation attained by legume and non-legume associations and assesses their role as a source of N in tropical and sub-tropical agriculture. We discuss factors influencing N₂ fixation and identify possible strategies for improving the amount of N₂ fixed.     

Biosynthesis and beneficial effects of microbial gibberellins on crops for sustainable agriculture

Soil microbes promote plant growth through several mechanisms such as secretion of chemical compounds including plant growth hormones. Among the phytohormones, auxins, ethylene, cytokinins, abscisic acid and gibberellins are the best understood compounds. Gibberellins were first isolated in 1935 from the fungus Gibberella fujikuroi and are synthesized by several soil microbes. The effect of gibberellins on plant growth and development has been studied, as has the biosynthesis pathways, enzymes, genes and their regulation. This review revisits the history of gibberellin research highlighting microbial gibberellins and their effects on plant health with an emphasis on the early discoveries and current advances that can find vital applications in agricultural practices.     

Sustainable agriculture and plant diseases: an epidemiological perspective

The potential for modern biology to identify new sources for genetical, chemical and biological control of plant disease is remarkably high. Successful implementation of these methods within globally and locally changing agricultural environments demands new approaches to durable control. This, in turn, requires fusion of population genetics and epidemiology at a range of scales from the field to the landscape and even to continental deployment of control measures. It also requires an understanding of economic and social constraints that influence the deployment of control. Here I propose an epidemiological framework to model invasion, persistence and variability of epidemics that encompasses a wide range of scales and topologies through which disease spreads. By considering how to map control methods onto epidemiological parameters and variables, some new approaches towards optimizing the efficiency of control at the landscape scale are introduced. Epidemiological strategies to minimize the risks of failure of chemical and genetical control are presented and some consequences of heterogeneous selection pressures in time and space on the persistence and evolutionary changes of the pathogen population are discussed. Finally, some approaches towards embedding epidemiological models for the deployment of control in an economically plausible framework are presented.     

Mycorrhiza and Phosphate Solubilizing Bacteria: Potential Bioagents for Sustainable Phosphorus Management in Agriculture

Phosphorus (P) is a critical nutrient that plays an essential role in improving soil fertility for optimum plant growth and productivity. It is one of the most deficient macro-nutrients in agricultural soils after nitrogen and is considered inadequate for plant growth and production. To P availability in soils, the farmers are applying huge amounts of synthetic P fertilizers that adversely affect the wider environment, groundwater, soil fertility and microbial population. Many beneficial microbes are known to release and supply soluble P for improving growth and yield of a variety of plants in a sustainable manner in P deficient soils. Thus, inoculation of these microbes, including arbuscular mycorrhizal fungi (AMF) and phosphate solubilizing bacteria (PSB) to soil to enhance crop production without harming the environment, is an alternative approach to chemical fertilizers. The combined role of AMF and PSB in P solubilization is not well understood and the application and mode of action of these microbial groups are often naive due to variation in the environment. Therefore, the current review article would develop a better understanding of the interactive role and mechanisms of AMF and PSB in improving P availability from both organic and inorganic sources in a sustainable crop production system. Finally, the current review would loop out further avenues for researchers interested to commercially produce effective AMF and PSB-based biofertilizers for sustainable management of phosphorus over a wide range of agricultural crops worldwide.     

Diazotrophic endophytes: progress and prospects for nitrogen fixation in monocots

Spatial variability of plant growth has been a major problem for plant physiologists, agronomists, agro-foresters and soil scientists comparing or modelling treatment effects on acid sandy soils in the Sahel. While aerial photographs from aeroplanes or satellite images may provide valuable information for the surveying of large areas, their use for individual small experiments or farmers' fields has been limited due to high costs, restricted availability, and unsatisfactory resolution. As a simple alternative, a commercially available Zeppelin-type balloon, dragged on a rope by a camel, was fitted with an standard 35 mm camera and a remote control system. Flight altitude varied from 20 m to 500 m above ground. A ground-based camera mounted vertically on a tripod was used to center the Zeppelin over the target area. The high-resolution true colour negatives and colour infrared slides obtained by this device were used to unravel the history of farmers' management strategies for maintaining soil productivity, to monitor treatment effects and crop growth variability in an on-station experiment, and to visualize light absorption by photosynthesis in crops and trees. Such non-destructively collected data may serve as quick but reliable references for ground measurements in a wide range of experiments with loosely-spaced crops, bushes, and trees. ei]B E Clothier     

Use of white clover as an alternative to nitrogen fertiliser for dairy pastures in nitrate vulnerable zones in the UK: productivity, environmental impact and economic considerations

Perennial ryegrass and perennial ryegrass/white clover permanent dairy pastures are compared with respect to productivity, environmental impact and financial costs in nitrate vulnerable zones (NVZ) in the UK. With appropriate management, and utilisation of recommended perennial ryegrass and white clover cultivars, white clover is likely to stabilise at around 20% of total dry matter production in a mixed pasture. Plant dry matter production and milk production from a perennial ryegrass/white clover pasture are likely to be similar to that from a perennial ryegrass pasture receiving 200 kg N ha⁻¹ annum⁻¹ and around 70% of that obtained with perennial ryegrass supplied with 350–400 kg N ha⁻¹ annum⁻¹. Nitrate, phosphorus and methane losses from the system and decreases in biodiversity relative to a grazed indigenous sward are likely to be similar for a perennial ryegrass/white clover pasture and a perennial ryegrass pasture receiving 200 kg N ha⁻¹ annum⁻¹: nitrate leachate from both systems is likely to comply with European legislation. Greenhouse gas emissions resulting from nitrogen (N) fertiliser production would be avoided with the perennial ryegrass/white clover pasture. Within NVZ stocking rate restrictions, white clover can provide the N required by a pasture at a lower financial cost than that incurred by the application of N fertiliser.     

Bradyrhizobium technology: a promising substitute for chemical nitrogen fertilizer in Bangladesh agriculture

Nitrogen is the most limiting element in Bangladesh soils and urea is the fertilizer commonly used for supplying it. Bradyrhizobium/Rhizobium inoculant was tried as a source of N nutrition for grain legumes in a number of field experiments. The inoculants markedly increased nodule number, nodule mass, shoot weight and yield of the crops compared to uninoculated control and urea-N treatments. For soybean (Glycine max), inoculation increased yield 113 percent over the control and 49 percent over the urea treament. For groundnut (Arachis hypogaea), the increases were 36 and 11 percent; for lentil (Lens culinaris), 30 and 13 percent; and for mungbean (Vigna radiata), 47 and 7 percent. The local inoculant strains were suitable for dependable inoculant production. The inoculant technology can be used as a promising and cheap substitute of urea for growing food legume crops in Bangladesh.     

Biological nitrogen fixation associated with sugar cane and rice: Contributions and prospects for improvement

¹⁵N isotope and N balance studies performed over the last few years have shown that several Brazilian varieties of sugarcane are capable of obtaining over 60% of their nitrogen (<150 kg N ha^-1 year^-1) from biological nitrogen fixation (BNF). This may be due to the fact that this crop in Brazil has been systematically bred for high yields with low fertilizer N inputs. In the case of wetland rice, N balance experiments performed both in the field and in pots suggest that 30 to 60 N ha^-1 crop^-1 may be obtained from plant-associated BNF and that different varieties have different capacities to obtain N from this source. ¹⁵N₂ incorporation studies have proved that wetland rice can obtain at least some N from BNF and acetylene reduction (AR) assays also indicate differences in N₂-fixing ability between different rice varieties. However in situ AR field estimates suggest plant-associated BNF inputs to be less than 8 kg N ha^-1 crop^-1. The problems associated with the use of the ¹⁵N dilution technique for BNF quantification are discussed and illustrated with data from a recent study performed at EMBRAPA-CNPAB. Although many species of diazotrophs have been isolated from the rhizosphere of both sugarcane and wetland rice, the recent discovery of endophytic N₂-fixing bacteria within roots, shoots and leaves of both crops suggests, at least in the case of sugarcane, that these bacteria may be the most important contributors to the observed BNF contributions. In sugarcane both Acetobacter diazotrophicus and Herbaspirillum spp. have been found within roots and aerial tissues and these microorganisms, unlike Azospirillum spp. and other rhizospheric diazotrophs, have been shown to survive poorly in soil. Herbaspirillum spp. are found in many graminaceous crops, including rice (in roots and aerial tissue), and are able to survive and pass from crop to crop in the seeds. The physiology, ecology and infection of plants by these endophytes are fully discussed in this paper. The sugarcane/endophytic diazotroph association is the first efficient N₂-fixing system to be discovered associated with any member of the gramineae. As yet the individual roles of the different diazotrophs in this system have not been elucidated and far more work on the physiology and anatomy of this system is required. However, the understanding gained in these studies should serve as a foundation for the improvement/development of similar N₂-fixing systems in wetland rice and other cereal crops.     

Potentials for win-win alliances among animal agriculture and forest products industries: Application of the principles of industrial ecology and sustainable development

Commercial forests in many parts of the world are deficient in nitrogen and phosphorus. These nutrient-deficient forests often exist in close proximity to large animal feeding operations, meat processing and other food, textile, or other biomass-processing plants, and municipal waste treatment facilities. Many of these facilities produce large surpluses of nitrogen,phosphorus, and organic matter as gaseous ammonia, urea, uric acid, phosphorus compounds,bacterial sludges, and partially treated municipal wastewaters. These co-existing and substantial nutrient deficiencies and surpluses offer ready-made opportunities for discovery, demonstration,and commercial development of science-based, technology-facilitated, environmentally sound,economically viable, and socially acceptable "win-win alliances" among these major industries based on the principles of industrial ecology and sustainable development. The major challenge is to discover practical means to capture the surplus nutrients and put them to work in forest stands from which value-added products can be produced and sold at a profit.     

Atmospheric nitrogen deposition in world biodiversity hotspots: the need for a greater global perspective in assessing N deposition impacts

Increased atmospheric nitrogen (N) deposition is known to reduce plant diversity in natural and semi‐natural ecosystems, yet our understanding of these impacts comes almost entirely from studies in northern Europe and North America. Currently, we lack an understanding of the threat of N deposition to biodiversity at the global scale. In particular, rates of N deposition within the newly defined 34 world biodiversity hotspots, to which 50% of the world's floristic diversity is restricted, has not been quantified previously. Using output from global chemistry transport models, here we provide the first estimates of recent (mid‐1990s) and future (2050) rates and distributions of N deposition within biodiversity hotspots. Our analysis shows that the average deposition rate across these areas was 50% greater than the global terrestrial average in the mid‐1990s and could more than double by 2050, with 33 of 34 hotspots receiving greater N deposition in 2050 compared with 1990. By this time, 17 hotspots could have between 10% and 100% of their area receiving greater than 15 kg N ha^?1 yr^?1, a rate exceeding critical loads set for many sensitive European ecosystems. Average deposition in four hotspots is predicted to be greater than 20 kg N ha^?1 yr^?1. This elevated N deposition within areas of high plant diversity and endemism may exacerbate significantly the global threat of N deposition to world floristic diversity. Overall, we highlight the need for a greater global approach to assessing the impacts of N deposition.     

Hemp (Cannabis sativa L.) leaf photosynthesis in relation to nitrogen content and temperature: implications for hemp as a bio‐economically sustainable crop

Hemp (Cannabis sativa L.) may be a suitable crop for the bio‐economy as it requires low inputs while producing a high and valuable biomass yield. With the aim of understanding the physiological basis of hemp's high resource‐use efficiency and yield potential, photosynthesis was analysed on leaves exposed to a range of nitrogen and temperature levels. Light‐saturated net photosynthesis rate (A_max) increased with an increase in leaf nitrogen up to 31.2 ± 1.9 μmol m^?2 s^?1 at 25 °C. The A_max initially increased with an increase in leaf temperature (T_L), levelled off at 25–35 °C and decreased when T_L became higher than 35 °C. Based on a C₃ leaf photosynthesis model, we estimated mesophyll conductance (g_m), efficiency of converting incident irradiance into linear electron transport under limiting light (κ_2LL), linear electron transport capacity (J_max), Rubisco carboxylation capacity (V_cmax), triose phosphate utilization capacity (T_p) and day respiration (R_d), using data obtained from gas exchange and chlorophyll fluorescence measurements at different leaf positions and various levels of incident irradiance, CO₂ and O₂. The effects of leaf nitrogen and temperature on photosynthesis parameters were consistent at different leaf positions and among different growth environments except for κ_2LL, which was higher for plants grown in the glasshouse than for those grown outdoors. Model analysis showed that compared with cotton and kenaf, hemp has higher photosynthetic capacity when leaf nitrogen is <2.0 g N m^?2. The high photosynthetic capacity measured in this study, especially at low nitrogen level, provides additional evidence that hemp can be grown as a sustainable bioenergy crop over a wide range of climatic and agronomic conditions.     

Potentials for win-win alliances among animal agriculture and forest products industries: Application of the principles of industrial ecology and sustainable development

Commercial forests in many parts of the world are deficient in nitrogen and phosphorus. These nutrient-deficient forests often exist in close proximity to large animal feeding operations, meat processing and other food, textile, or other biomass-processing plants, and municipal waste treatment facilities. Many of these facilities produce large surpluses of nitrogen, phosphorus, and organic matter as gaseous ammonia, urea, uric acid, phosphorus compounds, bacterial sludges, and partially treated municipal wastewaters. These co-existing and substantial nutrient deficiencies and surpluses offer ready-made opportunities for discovery, demonstration, and commercial development of science-based, technology-facilitated, environmentally sound, economically viable, and socially acceptable "win-win alliances" among these major industries based on the principles of industrial ecology and sustainable development. The major challenge is to discover practical means to capture the surplus nutrients and put them to work in forest stands from which value-added products can be produced and sold at a profit.     

A Simulation Tool for Industrial Ecology: Creating a Board Game

This article presents a board game that was developed for use as a simulation tool in teaching the basic concepts of industrial ecology (IE). The game, with the automobile industry as its theme, includes realistic numbers and displays a variety of IE principles. The objectives of the simulation, however, transcend the automobile industry and apply to other manufacturing industries. They include: pollution prevention, design for environment (in several forms, including design for disassembly), environmental management, and life-cycle assessment. The game has already been played by engineering and business professors, graduate students in environmental engineering, government representatives, and industry executives. A statistical analysis performed on pre- and post-game questionnaires indicates that the game is an effective teaching tool.     

Plants for nitrogen management in riparian zones: A proposed trait‐based framework to select effective species

Restoring plants to the riparian zone is regarded as management best practice in river restoration and has the potential to reduce the impact of nitrogen (N) pollution on aquatic organisms and improve water quality for human use. Plant characteristics and the interplay of hydrology and biogeochemistry control N retention in the riparian zone. The balance between processes such as denitrification and plant assimilation determines riparian N retention. Plant traits are likely to mediate these N removal processes through variations in root form, growth character, foliage production (quantity, quality and rate of return to the soil) and by altering conditions in the rhizosphere soil. Vegetation can slow N transfer via direct plant uptake of N (during periods of rapid vegetation growth) and changes induced to soil hydrology, nutrient cycling and microbial activity, principally denitrification. Few studies have focused on species‐dependent effects on N movement through soil and across boundaries. We propose a new framework, based on a literature review of plant traits with respect to N cycling, which can be used to select plant species with traits likely to maximise N removal during transport through the riparian zone. In the proposed framework, inter‐specific differences in traits known to influence N mobility: root form, growth rate, foliar characteristics and rhizosphere processes, are used to describe species’ potential impact on N removal. Plant trait data may be drawn from studies outside the riparian zone; for example forest ecology, horticulture or forestry research, and candidate species are scored to predict N removal efficiency. We apply the framework to New Zealand's native riparian plant assemblages to demonstrate the trait‐based approach. This framework can guide restoration management decisions and investment in riparian revegetation in a manner that is not restricted to geographically specific or well‐studied species.     

Olfactory receptors in Bactrocera species for sustainable fruit fly management: A review and future perspectives

Molecular studies on odorant receptors (ORs), odorant-binding proteins (OBPs) and the functioning of the receptor and pheromone signal transduction in fruit fly Bactrocera species have expanded exponentially during the past few decades. OBPs contribute to the sensing of the olfactory system (OS) via the transduction of odorants through the sensillum lymph. However, ORs, a family of G-protein-coupled receptors in Bactrocera and various other species, exhibit heightened responsiveness to multiple chemical odours such as hormones, sensory stimuli and neurotransmitters. The apparent mechanism involves a combinatorial code encompassing both peripheral and antennal lobe processing, facilitating the reception of sexual pheromones and environmental cues. The OS is specifically designed to recognize and process information from volatile chemical signals, and these chemical signals play an important function in various flies. Insects rely on these chemicals to navigate and comprehend their surroundings. A mature insect OS is composed of two pairs of sensillae-covered palps, antennae and two primary pairs of olfactory appendages on the anterior head. It has been shown that chemosensory gene families contribute in odour perception. These include various neuroreceptor families, such as OBPs, chemosensory proteins and sensory neuron membrane proteins. Additionally, there are three divergent chemoreceptors, namely ORs, ionotropic receptors and gustatory receptors. Methods based on systematic biology, molecular biology and bioinformatics tools have rapidly emerged to investigate the insect communication systems and provide new insights for the management of many agricultural pest. Several aromatic compounds, including semiochemicals and pheromones, have been employed to defend crops and animals from destructive fruit flies and other invasive and frugivorous species. To promote the expansion of the cropping system, the utilization of phytochemical lures can be convenient for sustainable agriculture production and enhance food security. Hence, this review examined the state of the art in chemical communication of insects with a focus on fruit fly pest species to identify OS and their semiochemical receptors, protein receptors and chemosensory receptors (CSRs), as well as their practical applications for biological control and integrated pest management are highlighted.     

Potential for Entomopathogenic Nematodes in Biological Control: A Meta-Analytical Synthesis and Insights from Trophic Cascade Theory

Entomopathogenic nematodes (EPN) are ubiquitous and generalized consumers of insects in soil food webs, occurring widely in natural and agricultural ecosystems on six continents. Augmentative releases of EPN have been used to enhance biological control of pests in agroecosystems. Pest managers strive to achieve a trophic cascade whereby natural-enemy effects permeate down through the food web to suppress host herbivores and increase crop production. Although trophic cascades have been studied in diverse aboveground arthropod-based systems, they are infrequently investigated in soil systems. Moreover, no overall quantitative assessment of the effectiveness of EPN in suppressing hosts with cascading benefits to plants has been made. Toward synthesizing the available but limited information on EPN and their ability to suppress prey and affect plant yield, we surveyed the literature and performed a meta-analysis of 35 published studies. Our analysis found that effect sizes for arthropod hosts as a result of EPN addition were consistently negative and indirect effects on plants were consistently positive. Results held across several different host metrics (abundance, fecundity and survival) and across measures of plant performance (biomass, growth, yield and survival). Moreover, the relationship between plant and host effect sizes was strikingly and significantly negative. That is, the positive impact on plant responses generally increased as the negative effect of EPN on hosts intensified, providing strong support for the mechanism of trophic cascades. We also review the ways in which EPN might interact antagonistically with each other and other predators and pathogens to adversely affect host suppression and dampen trophic cascades. We conclude that the food web implications of multiple-enemy interactions involving EPN are little studied, but, as management techniques that promote the long-term persistence of EPN are improved, antagonistic interactions are more likely to arise. We hope that the likely occurrence of antagonistic interactions in soil food webs should stimulate researchers to conduct field experiments explicitly designed to examine multiple-enemy interactions involving EPN and their cascading effects to hosts and plants.