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Conversion of molecular nitrogen into the biologically-accessible nitrogen compounds

Nitrogen fixation is a chemical procedure by which molecular nitrogen (N
2 ), with a strong triple covalent bail, in the air is converted into ammonia (NH
iii ) or related nitrogenous compounds, typically in soil or aquatic systems [1] but as well in manufacture. Atmospheric nitrogen is molecular dinitrogen, a relatively nonreactive molecule that is metabolically useless to all only a few microorganisms. Biological nitrogen fixation or diazotrophy is an important microbially mediated process that converts dinitrogen (Due north2) gas to ammonia (NH3) using the nitrogenase protein circuitous (Nif).[2] [3]

Nitrogen fixation is essential to life because fixed inorganic nitrogen compounds are required for the biosynthesis of all nitrogen-containing organic compounds, such as amino acids and proteins, nucleoside triphosphates and nucleic acids. As part of the nitrogen cycle, information technology is essential for agronomics and the industry of fertilizer. It is also, indirectly, relevant to the manufacture of all nitrogen chemical compounds, which includes some explosives, pharmaceuticals, and dyes.

Nitrogen fixation is carried out naturally in soil by microorganisms termed diazotrophs that include bacteria such as Azotobacter and archaea. Some nitrogen-fixing bacteria have symbiotic relationships with plant groups, specially legumes.[iv] Looser non-symbiotic relationships betwixt diazotrophs and plants are ofttimes referred to equally associative, as seen in nitrogen fixation on rice roots. Nitrogen fixation occurs between some termites and fungi.[5] It occurs naturally in the air by means of NOx production by lightning.[6] [seven]

All biological reactions involving the process of nitrogen fixation are catalysed by enzymes called nitrogenases.[eight] These enzymes contain iron, often with a second metal, usually molybdenum but sometimes vanadium.

History [edit]

Schematic representation of the nitrogen cycle. Abiotic nitrogen fixation has been omitted.

Biological nitrogen fixation was discovered by Jean-Baptiste Boussingault in 1838.[ix] Afterward, in 1880, the process by which it happens was discovered past German language agronomist Hermann Hellriegel and Hermann Wilfarth [de] [x] and was fully described by Dutch microbiologist Martinus Beijerinck.[11]

"The protracted investigations of the relation of plants to the acquisition of nitrogen begun by Saussure, Ville, Lawes and Gilbert and others culminated in the observe of symbiotic fixation by Hellriegel and Wilfarth in 1887."[12]

"Experiments by Bossingault in 1855 and Pugh, Gilbert & Lawes in 1887 had shown that nitrogen did non enter the found straight. The discovery of the function of nitrogen fixing leaner past Herman Hellriegel and Herman Wilfarth in 1886-eight would open a new era of soil science."[xiii]

In 1901 Beijerinck showed that azotobacter chroococcum was able to fix atmospheric nitrogen. This was the outset species of the azotobacter genus, so-named by him. It is also the first known diazotroph, species that use diatomic nitrogen as a pace in the complete nitrogen cycle.

Biological [edit]

Biological nitrogen fixation (BNF) occurs when atmospheric nitrogen is converted to ammonia by a nitrogenase enzyme.[1] The overall reaction for BNF is:

Northward two + sixteen ATP + 16 H 2 O + 8 e − + 8 H + ⟶ two NH 3 + H 2 + 16 ADP + 16 {\displaystyle {\ce {N2 + 16ATP + 16H2O + 8e- + 8H+ -> 2NH3 +H2 + 16ADP + sixteen}}} P i {\displaystyle {\text{P}}_{i}}

The process is coupled to the hydrolysis of 16 equivalents of ATP and is accompanied by the co-formation of one equivalent of H
2 .[14] The conversion of Northward
two into ammonia occurs at a metal cluster chosen FeMoco, an abbreviation for the iron-molybdenum cofactor. The mechanism proceeds via a series of protonation and reduction steps wherein the FeMoco agile site hydrogenates the N
two substrate.[15] In free-living diazotrophs, nitrogenase-generated ammonia is assimilated into glutamate through the glutamine synthetase/glutamate synthase pathway. The microbial nif genes required for nitrogen fixation are widely distributed in diverse environments.[16]

For example, decomposing wood, which generally has a low nitrogen content, has been shown to host a diazotrophic community.[17] [eighteen] The bacteria enrich the wood substrate with nitrogen through fixation, thus enabling deadwood decomposition by fungi.[19]

Nitrogenases are chop-chop degraded past oxygen. For this reason, many bacteria stop production of the enzyme in the presence of oxygen. Many nitrogen-fixing organisms be only in anaerobic weather, respiring to draw downwardly oxygen levels, or bounden the oxygen with a protein such as leghemoglobin.[ane]

Importance of nitrogen [edit]

Atmospheric nitrogen is inaccessible to most organisms,[twenty] because its triple covalent bond is very strong. Life takes up fixed nitrogen in various ways. Because cantlet acquisition, for every 100 atoms of carbon, roughly 2 to 20 atoms of nitrogen are assimilated. The atomic ratio of carbon (C) : nitrogen (N) : phosphorus (P) observed on average in planktonic biomass was originally described by Alfred Redfield.[21] The Redfield Ratio, the stoichiometric relationship betwixt C:North:P atoms, is 106:16:1.[21]

Nitrogenase [edit]

The protein circuitous nitrogenase is responsible for catalyzing the reduction of nitrogen gas (North2) to ammonia (NH3).[22] In Blue-green alga, this enzyme organisation is housed in a specialized cell called the heterocyst.[23] The production of the nitrogenase circuitous is genetically regulated, and the action of the protein complex is dependent on ambient oxygen concentrations, and intra- and extracellular concentrations of ammonia and oxidized nitrogen species (nitrate and nitrite).[24] [25] [26] Additionally, the combined concentrations of both ammonium and nitrate are thought to inhibit NFix, specifically when intracellular concentrations of 2-oxoglutarate (2-OG) exceed a critical threshold.[27] The specialized heterocyst cell is necessary for the performance of nitrogenase every bit a result of its sensitivity to ambient oxygen.[28]

Nitrogenase consist of two proteins, a catalytic fe-dependent poly peptide, commonly referred to as MoFe poly peptide and a reducing atomic number 26-only protein (Fe protein). There are iii different fe dependent proteins, molybdenum-dependent, vanadium-dependent, and iron-only, with all three nitrogenase protein variations containing an iron protein component. Molybdenum-dependent nitrogenase is the most ordinarily present nitrogenase.[22] The unlike types of nitrogenase can exist adamant past the specific iron protein component.[29] Nitrogenase is highly conserved. Gene expression through Dna sequencing can distinguish which protein circuitous is nowadays in the microorganism and potentially being express. Near oftentimes, the nifH factor is used to place the presence of molybdenum-dependent nitrogenase, followed by closely related nitrogenase reductases (component II) vnfH and anfH representing vanadium-dependent and iron-simply nitrogenase, respectively.[30] In studying the environmental and evolution of nitrogen-fixing leaner, the nifH gene is the biomarker most widely used.[31] nifH has 2 similar genes anfH and vnfH that also encode for the nitrogenase reductase component of the nitrogenase complex.[32]

Microorganisms [edit]

Diazotrophs are widespread within domain Bacteria including cyanobacteria (e.g. the highly significant Trichodesmium and Cyanothece), as well as dark-green sulfur bacteria, Azotobacteraceae, rhizobia and Frankia. Several obligately anaerobic bacteria fix nitrogen including many (but not all) Clostridium spp. Some archaea as well gear up nitrogen, including several methanogenic taxa, which are significant contributors to nitrogen fixation in oxygen-deficient soils.[33]

Cyanobacteria, commonly known as blue-dark-green algae, inhabit nearly all illuminated environments on Earth and play key roles in the carbon and nitrogen cycle of the biosphere. In general, blue-green alga tin use diverse inorganic and organic sources of combined nitrogen, such every bit nitrate, nitrite, ammonium, urea, or some amino acids. Several cyanobacteria strains are also capable of diazotrophic growth, an ability that may have been present in their final common ancestor in the Archean eon.[34] Nitrogen fixation non merely naturally occurs in soils but besides aquatic systems, including both freshwater and marine.[35] [36] Indeed, the corporeality of nitrogen fixed in the ocean is at least as much as that on land.[37] The colonial marine cyanobacterium Trichodesmium is thought to fix nitrogen on such a scale that it accounts for almost half of the nitrogen fixation in marine systems globally.[38] Marine surface lichens and non-photosynthetic bacteria belonging in Proteobacteria and Planctomycetes fixate meaning atmospheric nitrogen.[39] Species of nitrogen fixing cyanobacteria in fresh waters include: Aphanizomenon and Dolichospermum (previously Anabaena).[xl] Such species have specialized cells chosen heterocytes, in which nitrogen fixation occurs via the nitrogenase enzyme.[41] [42]

Root nodule symbioses [edit]

Legume family unit [edit]

Nodules are visible on this broad bean root

Plants that contribute to nitrogen fixation include those of the legume family—Fabaceae— with taxa such equally kudzu, clover, soybean, alfalfa, lupin, peanut and rooibos. They contain symbiotic rhizobia leaner within nodules in their root systems, producing nitrogen compounds that help the plant to grow and compete with other plants.[43] When the plant dies, the fixed nitrogen is released, making information technology available to other plants; this helps to fertilize the soil.[1] [44] The great majority of legumes have this clan, but a few genera (e.1000., Styphnolobium) practise not. In many traditional farming practices, fields are rotated through diverse types of crops, which normally include i consisting mainly or entirely of clover.[ citation needed ]

Fixation efficiency in soil is dependent on many factors, including the legume and air and soil conditions. For case, nitrogen fixation past red clover tin can range from 50 to 200 lb/acre (56 to 224 kg/ha).[45]

Non-leguminous [edit]

A sectioned alder tree root nodule

The ability to fix nitrogen in nodules is present in actinorhizal plants such as alder and bayberry, with the help of Frankia bacteria. They are plant in 25 genera in the orders Cucurbitales, Fagales and Rosales, which together with the Fabales form a nitrogen-fixing clade of eurosids. The power to fix nitrogen is not universally present in these families. For instance, of 122 Rosaceae genera, only iv ready nitrogen. Fabales were the get-go lineage to co-operative off this nitrogen-fixing clade; thus, the ability to prepare nitrogen may be plesiomorphic and after lost in most descendants of the original nitrogen-fixing institute; all the same, it may exist that the basic genetic and physiological requirements were present in an incipient state in the most recent common ancestors of all these plants, but simply evolved to full function in some of them.[46]

In addition, Trema (Parasponia), a tropical genus in the family Cannabaceae, is unusually able to collaborate with rhizobia and form nitrogen-fixing nodules.[47]

Non-legumious nodulating plants
Family Genera Species
Betulaceae
  • Alnus (alders)
 About or all species
Boraginaceae
  • Phacelia
  • Phacelia tanacetifolia
Cannabaceae
  • Trema (Parasponia)
  • Trema orientale
  • Trema lamarckiana
Casuarinaceae
  • Allocasuarina
  • Casuarina
  • Ceuthostoma
  • Gymnostoma
Coriariaceae
  • Coriaria
  • Coriaria arborea
  • Coriaria myrtifolia
Datiscaceae
  • Datisca
Elaeagnaceae
  • Elaeagnus (silverberries)
  • Hippophae (bounding main-buckthorns)
  • Shepherdia (buffaloberries)
Myricaceae
  • Comptonia (sweetfern)
  • Myrica (babyberries)
Posidoniaceae
  • Posidonia (seagrass)
Rhamnaceae
  • Ceanothus
  • Colletia
  • Discaria
  • Kentrothamnus
  • Retanilla
  • Talguenea
  • Trevoa
Rosaceae
  • Cercocarpus (mountain mahoganies)
  • Chamaebatia (mount miseries)
  • Dryas
  • Purshia/Cowania (bitterbrushes/cliffroses)

Cyanobiont [edit]

Some other plants alive in association with cyanobacteria (such every bit Nostoc) which fix nitrogen for them:

  • Some lichens such equally Lobaria and Peltigera
  • Musquito fern (Azolla species)
  • Cycads[48]
  • Gunnera
  • Blasia (liverwort)
  • Hornworts[49]

Industrial processes [edit]

Historical [edit]

A method for nitrogen fixation was first described by Henry Cavendish in 1784 using electric arcs reacting nitrogen and oxygen in air. This method was implemented in the Birkeland–Eyde process of 1903.[l] The fixation of nitrogen by lightning is a very like natural occurring process.

The possibility that atmospheric nitrogen reacts with certain chemicals was offset observed past Desfosses in 1828. He observed that mixtures of element of group i oxides and carbon react with nitrogen at loftier temperatures. With the use of barium carbonate as starting cloth, the kickoff commercial process became available in the 1860s, adult by Margueritte and Sourdeval. The resulting barium cyanide reacts with steam, yielding ammonia. In 1898 Frank and Caro developed what is known every bit the Frank–Caro process to set nitrogen in the class of calcium cyanamide. The process was eclipsed past the Haber process, which was discovered in 1909.[51] [52]

Haber process [edit]

Equipment for a study of nitrogen fixation by alpha rays (Fixed Nitrogen Research Laboratory, 1926)

The dominant industrial method for producing ammonia is the Haber procedure as well known as the Haber-Bosch process.[53] Fertilizer production is now the largest source of human-produced fixed nitrogen in the terrestrial ecosystem. Ammonia is a required precursor to fertilizers, explosives, and other products. The Haber process requires high pressures (around 200 atm) and high temperatures (at least 400 °C), which are routine conditions for industrial catalysis. This procedure uses natural gas as a hydrogen source and air as a nitrogen source. The ammonia production has resulted in an intensification of nitrogen fertilizer globally[54] and is credited with supporting the expansion of the homo population from around 2 billion in the early 20th century to roughly viii billion people now.[55]

Homogeneous catalysis [edit]

Much research has been conducted on the discovery of catalysts for nitrogen fixation, often with the goal of lowering energy requirements. All the same, such research has thus far failed to arroyo the efficiency and ease of the Haber process. Many compounds react with atmospheric nitrogen to requite dinitrogen complexes. The start dinitrogen complex to be reported was Ru(NH
three )
v (N
2 )ii+.[56] Some soluble complexes do catalyze nitrogen fixation.[57]

Lightning [edit]

Lightning heats the air around it breaking the bonds of Due north
ii starting the formation of nitrous acid.

Nitrogen can be fixed by lightning converting nitrogen gas (Due north
2 ) and oxygen gas (O
2 ) in the atmosphere into NOx (nitrogen oxides). The N
2 molecule is highly stable and nonreactive due to the triple bond between the nitrogen atoms.[58] Lightning produces plenty energy and heat to intermission this bond[58] allowing nitrogen atoms to react with oxygen, forming NO
x . These compounds cannot exist used by plants, only every bit this molecule cools, it reacts with oxygen to form NO
2 ,[59] which in plough reacts with h2o to produce HNO
2 (nitrous acid) or HNO
3 (nitric acid). When these acids seep into the soil, they make NO −
three (nitrate), which is of use to plants.[60] [58]

Run across also [edit]

  • Birkeland–Eyde process: an industrial fertiliser production procedure
  • George Washington Carver: an American botanist
  • Denitrification: an organic process of nitrogen release
  • Heterocyst
  • Nitrification: biological production of nitrogen
  • Nitrogen cycle: the menstruation and transformation of nitrogen through the surroundings
  • Nitrogen deficiency
  • Nitrogen fixation package for quantitative measurement of nitrogen fixation by plants
  • Nitrogenase: enzymes used past organisms to fix nitrogen
  • Ostwald process: a chemical procedure for making nitric acrid (HNO
    three     )
  • Push–pull technology: the utilise of both repellent and attractive organisms in agriculture
  • Carbon fixation

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External links [edit]

  • Hirsch AM (2009). "A Brief History of the Discovery of Nitrogen-fixing Organisms" (PDF). University of California, Los Angeles.
  • "Marine Nitrogen Fixation laboratory". University of Southern California.
  • "Travis P. Hignett Drove of Stock-still Nitrogen Research Laboratory Photographs // Scientific discipline History Institute Digital Collections". digital.sciencehistory.org . Retrieved 16 August 2019. Science History Institute Digital Collections (Photographs depicting numerous stages of the nitrogen fixation process and the various equipment and appliance used in the production of atmospheric nitrogen, including generators, compressors, filters, thermostats, and vacuum and boom furnaces).
  • "Proposed Process for the Fixation of Atmospheric Nitrogen", historical perspective, Scientific American, xiii July 1878, p. 21
  • A global ocean snapshot of nitrogen fixers past matching sequences to cells in the Tara Ocean

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Source: https://en.wikipedia.org/wiki/Nitrogen_fixation

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