Biology:9-cis-epoxycarotenoid dioxygenase

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Short description: Class of enzymes
9-cis-epoxycarotenoid dioxygenase
Corn 9-cis-epoxycarotenoid dioxygenase 1 PDB=3npe.png
Corn/maize 9-cis-epoxycarotenoid dioxygenase in complex with iron and molecular oxygen PDB: 3NPE​
Identifiers
EC number1.13.11.51
CAS number199877-10-6
Databases
IntEnzIntEnz view
BRENDABRENDA entry
ExPASyNiceZyme view
KEGGKEGG entry
MetaCycmetabolic pathway
PRIAMprofile
PDB structuresRCSB PDB PDBe PDBsum
Gene OntologyAmiGO / QuickGO

9-cis-epoxycarotenoid dioxygenase (EC 1.13.11.51, nine-cis-epoxycarotenoid dioxygenase, NCED, AtNCED3, PvNCED1, VP14) is an enzyme in the biosynthesis of abscisic acid (ABA),[1] with systematic name 9-cis-epoxycarotenoid 11,12-dioxygenase.[2][3][4][5][6] This enzyme catalyses the following chemical reaction

(1) a 9-cis-epoxycarotenoid + O2 [math]\displaystyle{ \rightleftharpoons }[/math] 2-cis,4-trans-xanthoxin + a 12'-apo-carotenal
(2) 9-cis-violaxanthin + O2 [math]\displaystyle{ \rightleftharpoons }[/math] 2-cis,4-trans-xanthoxin + (3S,5R,6S)-5,6-epoxy-3-hydroxy-5,6-dihydro-12'-apo-beta-caroten-12'-al
(3) 9'-cis-neoxanthin + O2 [math]\displaystyle{ \rightleftharpoons }[/math] 2-cis,4-trans-xanthoxin + (3S,5R,6R)-5,6-dihydroxy-6,7-didehydro-5,6-dihydro-12'-apo-beta-caroten-12'-al

9-cis-epoxycarotenoid dioxygenase contains iron(II).

Gene family

NCED belongs to a gene family called Carotenoid Cleavage Dioxygenases (CCD), which contains both CCD genes and NCED genes.[7] CCD is available in all plants (including algae), while NCED is currently only observed in land plants.[8] Please note some algae CCD genes have been incorrectly named NCED. There are usually multiple copies of NCED in a species.

Gene function

The enzyme catalyses the rate-limiting step of ABA biosynthesis.[1] Interestingly, though ABA is also produced in algae, NCED is currently only observed in land plants, suggesting ABA is produced in a different pathway in algae compare to land plants.[8]

Though first identified in maize/corn,[2][3] it is now quite extensively studied in the model plant Arabidopsis thaliana (Arabidopsis). The most studied NCED in Arabidopsis is the AtNCED3. Overexpression of the AtNCED3 gene improves the tolerance of transgenic plants to dehydration stress[6] as well as to salinity stress. Overexpression also leads to the overexpression of other genes induced by drought-stress. Transgenics containing AtNCED3 had greater root biomass, bigger pith size and higher level of photosynthesis.[9] It has been suggested that the AtNCED3 gene promoter contains G-box-like cis-acting elements which are responsible for dehydration-induced expression.[10]

Gene expression differs by plant organ. The gene may have a dual role in roots by either promoting or inhibiting the development of lateral roots.[11][12] AtNCED3 functions in seeds by regulating the seed establishment and abortion, maturation of the embryo, and seed dormancy. Maternal ABA functions in the early stage of zygote development, while embryonal AtNCED3 expresses later for ABA synthesis in case of dormancy. Expression of the gene mainly happens in the maternal tissues in the basal part of seeds or funiculus.[11]

AtNCED3 gene expression responds to drought-stress.[13][6] and salt-stress.[14][9] The signal caused by low moisture in the air is first induced by the stomata of leaves and transferred to other cells and tissues, which upregulate the expression of the AtNCED3 gene and ABA synthesis. Stomata closure limits various processes, including air exchange, water loss and O2 release. The AtNCED3 gene is active and expressed under these circumstances.[13][6] Accumulation of AtNCED3 mRNA and AtNCED3 protein was first found in the vascular parenchyma cells under drought stress, which suggested that plant drought tolerance relates to the development of plant vascular tissue.[13] Also, overexpression of AtNCED3 gene can improve plant salt tolerance.

Other NCED genes in Arabidopsis are less characterised. However, Tan et. al. (2003) [11] observed that other NCED is important in plants and seed development. For example, AtNCED2 and AtNCED9 genes are important in flower development, while AtNCED6 is important in seed dormancy and development. AtNCED5 is found to be important in seed dormancy, and it can interact with AtNCED3 for drought response.[15]

References

  1. ↑ 1.0 1.1 "The 9-cis-epoxycarotenoid cleavage reaction is the key regulatory step of abscisic acid biosynthesis in water-stressed bean". Proceedings of the National Academy of Sciences of the United States of America 96 (26): 15354–15361. December 1999. doi:10.1073/pnas.96.26.15354. PMID 10611388. Bibcode: 1999PNAS...9615354Q. 
  2. ↑ 2.0 2.1 "Specific oxidative cleavage of carotenoids by VP14 of maize". Science 276 (5320): 1872–1874. June 1997. doi:10.1126/science.276.5320.1872. PMID 9188535. 
  3. ↑ 3.0 3.1 "Genetic control of abscisic acid biosynthesis in maize". Proceedings of the National Academy of Sciences of the United States of America 94 (22): 12235–12240. October 1997. doi:10.1073/pnas.94.22.12235. PMID 9342392. Bibcode: 1997PNAS...9412235C. 
  4. ↑ "The 9-cis-epoxycarotenoid cleavage reaction is the key regulatory step of abscisic acid biosynthesis in water-stressed bean". Proceedings of the National Academy of Sciences of the United States of America 96 (26): 15354–15361. December 1999. doi:10.1073/pnas.96.26.15354. PMID 10611388. Bibcode: 1999PNAS...9615354Q. 
  5. ↑ "Ectopic expression of a tomato 9-cis-epoxycarotenoid dioxygenase gene causes over-production of abscisic acid". The Plant Journal 23 (3): 363–374. August 2000. doi:10.1046/j.1365-313x.2000.00789.x. PMID 10929129. 
  6. ↑ 6.0 6.1 6.2 6.3 "Regulation of drought tolerance by gene manipulation of 9-cis-epoxycarotenoid dioxygenase, a key enzyme in abscisic acid biosynthesis in Arabidopsis". The Plant Journal 27 (4): 325–333. August 2001. doi:10.1046/j.1365-313X.2002.01347.x. PMID 11532178. 
  7. ↑ "Enzymology of the carotenoid cleavage dioxygenases: reaction mechanisms, inhibition and biochemical roles". Archives of Biochemistry and Biophysics 544: 105–111. February 2014. doi:10.1016/j.abb.2013.10.005. PMID 24144525. 
  8. ↑ 8.0 8.1 "Surviving a Dry Future: Abscisic Acid (ABA)-Mediated Plant Mechanisms for Conserving Water under Low Humidity". Plants 6 (4): E54. November 2017. doi:10.3390/plants6040054. PMID 29113039. 
  9. ↑ 9.0 9.1 "Overexpression of AtSTO1 leads to improved salt tolerance in Populus tremula × P. alba". Transgenic Research 23 (5): 817–26. October 2014. doi:10.1007/s11248-014-9808-x. PMID 24929937. 
  10. ↑ "Characterization of the promoter region of an Arabidopsis gene for 9-cis-epoxycarotenoid dioxygenase involved in dehydration-inducible transcription". DNA Research 20 (4): 315–324. August 2013. doi:10.1093/dnares/dst012. PMID 23604098. 
  11. ↑ 11.0 11.1 11.2 "Molecular characterization of the Arabidopsis 9-cis epoxycarotenoid dioxygenase gene family". The Plant Journal 35 (1): 44–56. July 2003. doi:10.1046/j.1365-313x.2003.01786.x. PMID 12834401. 
  12. ↑ "Confirmation that abscisic acid accumulation is required for maize primary root elongation at low water potentials.". Journal of Experimental Botany 45 (Special Issue): 1743–1751. November 1994. doi:10.1093/jxb/45.Special_Issue.1743. 
  13. ↑ 13.0 13.1 13.2 "Drought induction of Arabidopsis 9-cis-epoxycarotenoid dioxygenase occurs in vascular parenchyma cells". Plant Physiology 147 (4): 1984–1993. August 2008. doi:10.1104/pp.108.116632. PMID 18550687. 
  14. ↑ "Both abscisic acid (ABA)-dependent and ABA-independent pathways govern the induction of NCED3, AAO3 and ABA1 in response to salt stress". Plant, Cell & Environment 29 (10): 2000–8. October 2006. doi:10.1111/j.1365-3040.2006.01576.x. PMID 16930325. 
  15. ↑ "Epoxycarotenoid cleavage by NCED5 fine-tunes ABA accumulation and affects seed dormancy and drought tolerance with other NCED family members". The Plant Journal 70 (3): 501–512. May 2012. doi:10.1111/j.1365-313x.2011.04887.x. PMID 22171989. 

External links



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