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{{enzyme


| Name = nitrogenase
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| EC_number = 1.18.6.1
 
| CAS_number = 9013-04-1
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| caption =
 
}}'''Nitrogenases''' ({{EC number|1.18.6.1}}{{EC number|1.19.6.1}}) are [[enzyme]]s used by some organisms to fix atmospheric [[nitrogen]] gas (N<sub>2</sub>). There is only one known family of enzymes that accomplishes this process. [[Dinitrogen]] is quite inert because of the strength of its N≡N [[triple bond]].
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[[File:Nitrogenase.png|thumb|Nitrogenase]]
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Whilst the equilibrium formation of ammonia from molecular hydrogen and nitrogen has an overall negative [[enthalpy of reaction]] (<math> \Delta H^{0} = -45.2 \ \mathrm{kJ} \, \mathrm{mol^{-1}} \; \mathrm{NH_3} </math>), the [[Activation energy|energy barrier]] to activation is very high (<math> E_A = 420 \ \mathrm{kJ} \, \mathrm{mol^{-1}} </math>) without the assistance of catalysis.<ref>Modak, J. M., 2002, Haber Process for Ammonia Synthesis, ''Resonance''. 7, 69-77.</ref>
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In addition to [[reducing agents]], such as [[dithionite]] ''[[in vitro]]'', or [[ferredoxin]] or [[flavodoxin]] ''[[in vivo]]'', the enzymatic reduction of dinitrogen to ammonia therefore also requires an input of chemical energy, released from the [[hydrolysis]] of [[Adenosine triphosphate|ATP]], to overcome the activation energy barrier. The [[enzyme]] is composed of the heterotetrameric MoFe protein that is transiently associated with the [[homodimeric]] Fe [[protein]]. Electrons for the reduction of nitrogen are supplied to nitrogenase when it associates with the reduced, nucleotide-bound homodimeric Fe protein. The heterocomplex undergoes cycles of [[cycle of association|association]] and disassociation to transfer one [[electron]], which is the [[rate-limiting step]] in nitrogen reduction {{Citation needed|date=June 2012}}. ATP supplies the energy to drive the transfer of electrons from the Fe protein to the MoFe protein. The reduction potential of each electron transferred to the MoFe protein is sufficient to break one of dinitrogen's chemical bonds, though it has not yet been shown that exactly three cycles are sufficient to convert one molecule of N<sub>2</sub> to ammonia. Nitrogenase ultimately bonds each atom of nitrogen to three hydrogen atoms to form [[ammonia]] (NH<sub>3</sub>), which is in turn bonded to [[glutamate]] to form [[glutamine]]. The nitrogenase reaction additionally produces molecular [[hydrogen]] as a side product.
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The exact mechanism of [[catalysis]] is unknown due to the difficulty in obtaining [[crystal]]s of nitrogen bound to nitrogenase. This is because the resting state of the MoFe protein does not bind nitrogen and also requires at least three electron transfers to perform [[catalysis]]. Nitrogenase is able to reduce acetylene, but is inhibited by carbon monoxide, which binds to the enzyme and thereby prevents binding of dinitrogen. Dinitrogen will prevent acetylene binding, but acetylene does not inhibit binding of dinitrogen and requires only one electron for reduction to [[ethylene]].<ref>Seefeldt LC, Dance IG, Dean DR. 2004. Substrate interactions with nitrogenase: Fe versus Mo. ''Biochemistry''. 43(6):1401-9.</ref>
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All nitrogenases have an iron- and sulfur-containing cofactor that includes a heterometal complex in the active site (e.g., [[FeMoCo]]). In most, this heterometal complex has a central [[molybdenum]] atom, though in some species it is replaced by a [[vanadium]] <ref>Robson, R. L.; Eady, R. R.; Richardson, T. H.; Miller, R. W.; Hawkins, M.; Postgate, J. R., 1986, The alternative nitrogenase of Azotobacter chroococcum is a vanadium enzyme, ''Nature (London)''. 322, 388-390.</ref> or [[iron]] atom.
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Due to the oxidative properties of [[oxygen]], most nitrogenases are irreversibly inhibited by [[dioxygen]], which degradatively oxidizes the Fe-S cofactors. This requires mechanisms for nitrogen fixers to protect nitrogenase from oxygen ''[[in vivo]]''. Despite this problem, many use oxygen as a terminal electron acceptor for respiration. One known exception is the nitrogenase of ''[[Streptomyces thermoautotrophicus]]'', which is unaffected by the presence of oxygen.<ref>{{pmid|9334244}}</ref> Although the ability of some nitrogen fixers such as [[Azotobacteraceae]] to employ an oxygen-labile nitrogenase under aerobic conditions has been attributed to a high [[metabolic rate]], allowing oxygen reduction at the [[cell membrane]], the effectiveness of such a mechanism has been questioned at oxygen concentrations above 70 µM (ambient concentration is 230 µM O<sub>2</sub>), as well as during additional nutrient limitations.<ref>Oelze J. 2000. Respiratory protection of nitrogenase in ''[[Azotobacter]]'' species: Is a widely-held hypothesis unequivocally supported by experimental evidence? ''FEMS Microbiol Rev''. 24(4):321&ndash;33.</ref>
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The reaction that this enzyme performs is:
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:<math> \mathrm{N_2 + 8 \ H^+ + 8 \ e^- + 16 \ ATP + 16 \ H_2O \longrightarrow 2 \ NH_3 + H_2 + 16 \ ADP + 16 \ P_i} </math>
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==Nonspecific reactions==
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In addition to performing the reaction N≡N → 2 NH<sub>3</sub>, nitrogenase is also capable of catalyzing the following reactions:<ref name=Burris1>{{cite journal | author = Rivera-Ortiz, José M., and Burris, Robert H. | title = Interactions among substrates and inhibitors of nitrogenase | journal = J Bacteriol | volume = 123 | issue = 2 | pages = 537&ndash;545 | year = 1975 | url = http://jb.asm.org/cgi/content/abstract/123/2/537 | doi = | pmid = 1150625 | pmc = 235759 }}</ref><ref name=Schrauzer>{{cite journal | author = G. N. Schrauzer | title = Nonenzymatic Simulation of Nitrogenase Reactions and the Mechanism of Biological Nitrogen Fixation | journal = Angewandte Chemie International Edition in English | volume = 14 | issue = 8 | pages = 514&ndash;522 | year = 2003 | url = http://www3.interscience.wiley.com/journal/106580477/abstract | doi = 10.1002/anie.197505141 | pmid = 810048 }}</ref>
:[[acetylene|HC≡CH]] → [[ethylene|H<sub>2</sub>C=CH<sub>2</sub>]]
:[[nitrous oxide|N<sup>&ndash;</sup>=N<sup>+</sup>=O]] → N<sub>2</sub> + H<sub>2</sub>O
:[[azide|N=N=N<sup>&ndash;</sup>]] → N<sub>2</sub> + NH<sub>3</sub>
:[[cyanide|{{chem|C≡N|-}}]] → [[methane|CH<sub>4</sub>]], NH<sub>3</sub>, [[ethane|H<sub>3</sub>C–CH<sub>3</sub>]], H<sub>2</sub>C=CH<sub>2</sub> (CH<sub>3</sub>NH<sub>2</sub>)
:N≡C–R → RCH<sub>3</sub> + NH<sub>3</sub>
:C≡N–R → CH<sub>4</sub>, H<sub>3</sub>C–CH<sub>3</sub>, H<sub>2</sub>C=CH<sub>2</sub>, [[propane|C<sub>3</sub>H<sub>8</sub>]], [[propylene|C<sub>3</sub>H<sub>6</sub>]], RNH<sub>2</sub>
:[[Carbonyl sulfide|O=C=S]] → [[carbon monoxide|CO]] + [[Hydrogen sulfide|H<sub>2</sub>S]] <ref name=Seefeldt1>{{cite journal | author = Lance C. Seefeldt, Madeline E. Rasche, Scott A. Ensign | title = Carbonyl sulfide and carbon dioxide as new substrates, and carbon disulfide as a new inhibitor, of nitrogenase | journal = Biochemistry | volume = 34 | issue = 16 | pages = 5382&ndash;5389 | year = 1995 | url = http://pubs.acs.org/doi/abs/10.1021/bi00016a009 | doi = 10.1021/bi00016a009 | pmid = 7727396 }}</ref><ref name="Seefeldt2" />
:[[carbon dioxide|O=C=O]] → CO + H<sub>2</sub>O <ref name="Seefeldt1" />
:[[thiocyanate|S=C=N<sup>&ndash;</sup>]] → H<sub>2</sub>S + HCN <ref name=Seefeldt2>{{cite journal | author = Madeline E. Rasche and Lance C. Seefeldt | title = Reduction of Thiocyanate, Cyanate, and Carbon Disulfide by Nitrogenase: Kinetic Characterization and EPR Spectroscopic Analysis | journal = Biochemistry | volume = 36 | issue = 28 | pages = 8574&ndash;8585 | year = 1997 | url = http://pubs.acs.org/doi/abs/10.1021/bi970217e | doi = 10.1021/bi970217e | pmid = 9214303 }}</ref>
:[[Cyanate|O=C=N<sup>&ndash;</sup>]] → H<sub>2</sub>O + [[Hydrogen cyanide|HCN]], CO + NH<sub>3</sub> <ref name="Seefeldt2" />


Furthermore, [[dihydrogen]] functions as a [[competitive inhibition|competitive inhibitor]],<ref name=Burris2>{{cite journal | author = Joseph H. Guth, Robert H. Burris | title = Inhibition of nitrogenase-catalyzed ammonia formation by hydrogen | journal = Biochemistry | volume = 22 | issue = 22 | pages = 5111&ndash;5122 | year = 1983 | url = http://pubs.acs.org/doi/abs/10.1021/bi00291a010 | doi = 10.1021/bi00291a010 | pmid = 6360203 }}</ref> [[carbon monoxide]] functions as a [[Non-competitive inhibition|non-competitive inhibitor]],<ref name="Burris1" /><ref name="Schrauzer" /> and [[carbon disulfide]] functions as a [[Enzyme inhibitor#Reversible inhibitors|rapid-equilibrium inhibitor]]<ref name="Seefeldt1" /> of nitrogenase.
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</ul>


[[Vanadium nitrogenase]]s have also been shown to catalzye the conversion of CO into [[alkanes]] through a reaction comparable to Fischer-Tropsch synthesis.
== New Balance 574  WilkesBarre ==


==Organisms that synthesize nitrogenase==
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* Free-living [[diazotroph]]s, e.g.
 
** [[Green sulfur bacteria]]
  <li>[http://enseignement-lsf.com/spip.php?article64#forum18071069 http://enseignement-lsf.com/spip.php?article64#forum18071069]</li>
** [[Cyanobacteria]] (with or without<ref>http://www.ncbi.nlm.nih.gov/pubmed/20532747</ref> differentiated [[heterocyst]]s)
 
** [[Azotobacteraceae]]
  <li>[http://www.peakrealestate.com.au/bbs/forum.php?mod=viewthread&tid=197477&fromuid=42305 http://www.peakrealestate.com.au/bbs/forum.php?mod=viewthread&tid=197477&fromuid=42305]</li>
* Symbiotic diazotrophs, e.g.
 
** [[Rhizobia]]
  <li>[http://slapfish.co.uk/forum/viewtopic.php?f=1&t=286082 http://slapfish.co.uk/forum/viewtopic.php?f=1&t=286082]</li>
** [[Frankia]]
 
Typically nitrogenases are coded by [[Nif gene]]s.
  <li>[http://www.ltc1688.com/thread-1487928-1-1.html http://www.ltc1688.com/thread-1487928-1-1.html]</li>
 
 
== Similarity to other proteins ==
  <li>[http://www.2sov.com/forum.php?mod=viewthread&tid=485536 http://www.2sov.com/forum.php?mod=viewthread&tid=485536]</li>
The three subunits of nitrogenase exhibit significant sequence similarity to three subunits of the light-independent version of [[protochlorophyllide reductase]] that performs the conversion of [[protochlorophyllide]] to [[chlorophyll]]. This protein is present in gymnosperms, algae, and photosynthetic bacteria but has been lost by angiosperms during evolution.<ref>{{pmid|8305874}}</ref>
 
 
</ul>
Separately, two of the nitrogenase subunits (NifD and NifD) have homologues in [[methanogen]]s that do not fix nitrogen e.g. ''[[Methanocaldococcus jannaschii]]''.<ref name=jannaschii>{{pmid|17660283}}</ref> Little is understood about the function of these "class IV" ''nif'' genes,<ref>{{pmid|14694078}}</ref> though they occur in many methanogens. In ''M. jannaschii'' they are known to interact with each other and are constitutively expressed.<ref name=jannaschii />
 
==See also==
* [[Nitrogen fixation]]
 
== References ==
* {{cite journal | author = Zumft WG, Mortenson LE | year = 1975 | title = The nitrogen-fixing complex of bacteria | journal = Biochim. Biophys. Acta. | volume = 416 | pages = 1&ndash;52 | pmid = 164247 | issue = 1 }}
{{reflist}}
 
{{Other oxidoreductases}}
 
[[Category:EC 1.18.6]]
[[Category:Iron-sulfur proteins]]
[[Category:Nitrogen metabolism]]

Latest revision as of 11:13, 13 January 2015

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