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'''Kriptohromi''' (od [[grčki jezik|grčkog]] κρυπτός χρώμα – ''kriptos'' + ''hroma'' = "skrivena boja") su klasa [[flavoprotein]]a pronađenih u [[biljke|biljkama]] i [[životinja]]ma koje su osjetljivi na [[Vidljivi spektar|plavo svjetlo]]. Uključeni su u [[cirkadijski ritam|cirkadijanske ritmove]] i [[magnetorecepcija|osjećanje magnetnih polja]] kod brojnih [[vrsta]]. Naziv ''kriptohrom'' je predložen kao ''[[portmanteau]]'' koji kombinuje ''[[hromatin|hromatsku]]'' prirodu [[fotoreceptor]]skih proteina i ''[[kriptosjemenjače|kriptogamnih]]'' organizama na kojima su provedena mnoga istraživanja plavog [[svjetlost]]i.<ref>{{Cite journal|last=Gressel|first=J.|date=1979|title=Blue Light Photoreception|journal=Photochemistry and Photobiology|language=en|volume=30|issue=6|pages=749–754|doi=10.1111/j.1751-1097.1979.tb07209.x|s2cid=98643540|issn=1751-1097}}</ref><ref>{{cite journal | vauthors = Yang Z, Liu B, Su J, Liao J, Lin C, Oka Y | title = Cryptochromes Orchestrate Transcription Regulation of Diverse Blue Light Responses in Plants | journal = Photochemistry and Photobiology | volume = 93 | issue = 1 | pages = 112–127 | date = |
'''Kriptohromi''' (od [[grčki jezik|grčkog]] κρυπτός χρώμα – ''kriptos'' + ''hroma'' = "skrivena boja") su klasa [[flavoprotein]]a pronađenih u [[biljke|biljkama]] i [[životinja]]ma koje su osjetljivi na [[Vidljivi spektar|plavo svjetlo]]. Uključeni su u [[cirkadijski ritam|cirkadijanske ritmove]] i [[magnetorecepcija|osjećanje magnetnih polja]] kod brojnih [[vrsta]]. Naziv ''kriptohrom'' je predložen kao ''[[portmanteau]]'' koji kombinuje ''[[hromatin|hromatsku]]'' prirodu [[fotoreceptor]]skih proteina i ''[[kriptosjemenjače|kriptogamnih]]'' organizama na kojima su provedena mnoga istraživanja plavog [[svjetlost]]i.<ref>{{Cite journal|last=Gressel|first=J.|date=1979|title=Blue Light Photoreception|journal=Photochemistry and Photobiology|language=en|volume=30|issue=6|pages=749–754|doi=10.1111/j.1751-1097.1979.tb07209.x|s2cid=98643540|issn=1751-1097}}</ref><ref>{{cite journal | vauthors = Yang Z, Liu B, Su J, Liao J, Lin C, Oka Y | title = Cryptochromes Orchestrate Transcription Regulation of Diverse Blue Light Responses in Plants | journal = Photochemistry and Photobiology | volume = 93 | issue = 1 | pages = 112–127 | date = januar 2017 | pmid = 27861972 | doi = 10.1111/php.12663 | pmc = 6167254 }}</ref> |
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Dva [[gen]]a "Cry1" i "Cry2" [[kodon|kodiraju]] dva kriptohromna [[protein]]a, CRY1 i CRY2.<ref>{{cite journal | vauthors = van der Spek PJ, Kobayashi K, Bootsma D, Takao M, Eker AP, Yasui A | title = Cloning, tissue expression, and mapping of a human photolyase homolog with similarity to plant blue-light receptors | journal = Genomics | volume = 37 | issue = 2 | pages = 177–182 | date = |
Dva [[gen]]a "Cry1" i "Cry2" [[kodon|kodiraju]] dva kriptohromna [[protein]]a, CRY1 i CRY2.<ref>{{cite journal | vauthors = van der Spek PJ, Kobayashi K, Bootsma D, Takao M, Eker AP, Yasui A | title = Cloning, tissue expression, and mapping of a human photolyase homolog with similarity to plant blue-light receptors | journal = Genomics | volume = 37 | issue = 2 | pages = 177–182 | date = oktobar 1996 | pmid = 8921389 | doi = 10.1006/geno.1996.0539 | hdl-access = free | hdl = 1765/55742 | url = http://repub.eur.nl/pub/55742 }}</ref> Kod [[insekti|insekata]] i [[biljke|biljaka]], CRY1 reguliše [[cirkadijski sat]] na način ovisan o svjetlosti, dok kod [[sisar]]a, CRY1 i CRY2 djeluju kao svjetlosno-nezavisni inhibitori [[CLOCK]]-[[ARNTL|BMAL1]], komponente cirkadijskog sata. U biljkama, fotorecepcija plavog svjetla može se koristiti za signaliziranje razvojnih signala. Osim [[hlorofil]]a, kriptohromi su jedini proteini za koje se zna da formiraju fotoindukovane [[mehanizam radikalnih parova|radikalne parove]] ''[[in vivo]]''.<ref>{{cite journal | vauthors = Hore PJ, Mouritsen H | title = The Radical-Pair Mechanism of Magnetoreception | journal = Annual Review of Biophysics | volume = 45 | issue = 1 | pages = 299–344 | date = juli 2016 | pmid = 27216936 | doi = 10.1146/annurev-biophys-032116-094545 | s2cid = 7099782 | url = https://ora.ox.ac.uk/objects/uuid:c1e3c8ca-98b3-4e9d-8efd-0b9ad9b965eb }}</ref> Čini se da one omogućavaju nekim životinjama da otkriju [[magnetno polje|magnetna polja]]. |
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Kriptohromi su bili u fokusu nekoliko dosadašnjih napora u [[optogenetika|optogenetici]]. Koristeći [[transfekcija|transfekciju]], početne studije na [[kvasci|kvascu]] su iskoristile potencijal [[heterodimerizacija]] Cry2 da kontroliše ćelijske procese, uključujući [[ekspresija gena|ekspresiju gena]], pomoću svetlosti. |
Kriptohromi su bili u fokusu nekoliko dosadašnjih napora u [[optogenetika|optogenetici]]. Koristeći [[transfekcija|transfekciju]], početne studije na [[kvasci|kvascu]] su iskoristile potencijal [[heterodimerizacija]] Cry2 da kontroliše ćelijske procese, uključujući [[ekspresija gena|ekspresiju gena]], pomoću svetlosti. |
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Nova hipoteza predlaže da u biljnim kriptohromima transdukcija svjetlosnog signala u hemijski signal koji mogu osjetiti partnerske molekule može biti pokrenuta fotoinduciranim negativnim nabojem unutar proteina - na kofaktor FAD ili na susjednu [[aspartat|asparaginsku kiselinu]]. Ovaj negativni naboj bi elektrostatički odbio molekulu [[Adenozin-trifosfat|ATP]] vezanu za proteine, a time i protein C-terminalni domen, koji pokriva [[Adenozin trifosfat|ATP]] vezni džep prije apsorpcije [[foton]]a. Rezultirajuća promjena u konformaciji proteina mogla bi dovesti do [[fosforilacije]] prethodno nedostupnih fosforilacijskih mjesta na [[C-terminal]]u i dati fosforilirani segment bi tada mogao osloboditi [[transkripcijski faktor]] HY5, takmičeći se za isto [[mjesto vezivanja]] na negativnom regulatoru fotomorfogeneze [[COP1]] . |
Nova hipoteza predlaže da u biljnim kriptohromima transdukcija svjetlosnog signala u hemijski signal koji mogu osjetiti partnerske molekule može biti pokrenuta fotoinduciranim negativnim nabojem unutar proteina - na kofaktor FAD ili na susjednu [[aspartat|asparaginsku kiselinu]]. Ovaj negativni naboj bi elektrostatički odbio molekulu [[Adenozin-trifosfat|ATP]] vezanu za proteine, a time i protein C-terminalni domen, koji pokriva [[Adenozin trifosfat|ATP]] vezni džep prije apsorpcije [[foton]]a. Rezultirajuća promjena u konformaciji proteina mogla bi dovesti do [[fosforilacije]] prethodno nedostupnih fosforilacijskih mjesta na [[C-terminal]]u i dati fosforilirani segment bi tada mogao osloboditi [[transkripcijski faktor]] HY5, takmičeći se za isto [[mjesto vezivanja]] na negativnom regulatoru fotomorfogeneze [[COP1]] . |
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Kod ''[[Drosophila]]'' može funkcionirati drugačiji mehanizam. Pravo osnovno stanje kofaktora flavina u ''Drosophila'' CRY se još uvijek raspravlja, a neki modeli ukazuju da je FAD u oksidiranom obliku, dok drugi podržavaju model u kojem flavinski kofaktor postoji u [[anion]]skom [[radikal (hemija)|radikalnom]] obliku, {{chem|FAD|-}}•. Nedavno je u istraživanjimaa primiječeno da se oksidirani FAD lahko redukuje svjetlom u {{chem|FAD|-}}•. Štaviše, mutacije da blokirana fotoredukcija nije imala efekta na razgradnju CRY izazvanu svjetlom, dok su mutacije koje su promijenile stabilnost {{chem|FAD|-}}• uništile funkciju CRY fotoreceptora.<ref name="Song_2007">{{cite journal | vauthors = Song SH, Oztürk N, Denaro TR, Arat NO, Kao YT, Zhu H, Zhong D, Reppert SM, Sancar A | display-authors = 6 | title = Formation and function of flavin anion radical in cryptochrome 1 blue-light photoreceptor of monarch butterfly | journal = The Journal of Biological Chemistry | volume = 282 | issue = 24 | pages = 17608–17612 | date = |
Kod ''[[Drosophila]]'' može funkcionirati drugačiji mehanizam. Pravo osnovno stanje kofaktora flavina u ''Drosophila'' CRY se još uvijek raspravlja, a neki modeli ukazuju da je FAD u oksidiranom obliku, dok drugi podržavaju model u kojem flavinski kofaktor postoji u [[anion]]skom [[radikal (hemija)|radikalnom]] obliku, {{chem|FAD|-}}•. Nedavno je u istraživanjimaa primiječeno da se oksidirani FAD lahko redukuje svjetlom u {{chem|FAD|-}}•. Štaviše, mutacije da blokirana fotoredukcija nije imala efekta na razgradnju CRY izazvanu svjetlom, dok su mutacije koje su promijenile stabilnost {{chem|FAD|-}}• uništile funkciju CRY fotoreceptora.<ref name="Song_2007">{{cite journal | vauthors = Song SH, Oztürk N, Denaro TR, Arat NO, Kao YT, Zhu H, Zhong D, Reppert SM, Sancar A | display-authors = 6 | title = Formation and function of flavin anion radical in cryptochrome 1 blue-light photoreceptor of monarch butterfly | journal = The Journal of Biological Chemistry | volume = 282 | issue = 24 | pages = 17608–17612 | date = juni 2007 | pmid = 17459876 | doi = 10.1074/jbc.M702874200 | doi-access = free }}</ref><ref name="Oztürk_2011">{{cite journal | vauthors = Ozturk N, Selby CP, Annayev Y, Zhong D, Sancar A | title = Reaction mechanism of Drosophila cryptochrome | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 108 | issue = 2 | pages = 516–521 | date = januar 2011 | pmid = 21187431 | pmc = 3021015 | doi = 10.1073/pnas.1017093108 | doi-access = free | bibcode = 2011PNAS..108..516O }}</ref> Ova zapažanja pružaju podršku za osnovno stanje {{chem|FAD|-}}•. Istraživači su također nedavno predložili model u kojem se {{chem|FAD|-}} pobuđuje u svoje [[dvostruko stanje|dubletno]] ili kvartetno stanje apsorpcijom fotona, što zatim dovodi do konformacijske promjene u CRY proteinu .<ref name="Oztürk_2011"/> |
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Također, prstenaste oči larve [[sunđeri|demosunđera]] ''Amphimedon queenslandica'' izražavaju kriptohrom osetljiv na plavo svetlo (Aq-Cry2), koji bi mogao da posreduje [[fototaksija|fototaksiju]]. Nasuprot tome, [[oči]] većine životinja koriste fotoosjetljive [[opsin]]e eksprimirane u [[fotoreceptor]]skim ćelijama, koje prenose informacije o svjetlosti iz okoline u [[nervni sistem]]. Međutim, ''A. queenslandica'' nema nervni sistem, kao i druge [[spužve]]. A također nema [[opsin]]ski [[gen]] u svom potpuno sekvenciranom [[genom]]u, uprkos tome što ima mnogo drugih [[G-protein-spregnuti receptor|G-protein-spregnutih receptora]] (GPCR). Prema tome, jedinstvene oči spužve mora da su razvile drugačiji mehanizam za detekciju svjetlosti i posredovanje fototaksije, vjerovatno pomoću kriptohroma ili drugih proteina. |
Također, prstenaste oči larve [[sunđeri|demosunđera]] ''Amphimedon queenslandica'' izražavaju kriptohrom osetljiv na plavo svetlo (Aq-Cry2), koji bi mogao da posreduje [[fototaksija|fototaksiju]]. Nasuprot tome, [[oči]] većine životinja koriste fotoosjetljive [[opsin]]e eksprimirane u [[fotoreceptor]]skim ćelijama, koje prenose informacije o svjetlosti iz okoline u [[nervni sistem]]. Međutim, ''A. queenslandica'' nema nervni sistem, kao i druge [[spužve]]. A također nema [[opsin]]ski [[gen]] u svom potpuno sekvenciranom [[genom]]u, uprkos tome što ima mnogo drugih [[G-protein-spregnuti receptor|G-protein-spregnutih receptora]] (GPCR). Prema tome, jedinstvene oči spužve mora da su razvile drugačiji mehanizam za detekciju svjetlosti i posredovanje fototaksije, vjerovatno pomoću kriptohroma ili drugih proteina. |
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=== Funkcija šarenice === |
=== Funkcija šarenice === |
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[[Šarenica]] [[kokoš]]ijih [[embrion]]a osjeća svjetlo kratkih talasa preko kriptohroma, a ne opsina.<ref>{{cite journal | vauthors = Tu DC, Batten ML, Palczewski K, Van Gelder RN | title = Nonvisual photoreception in the chick iris | journal = Science | volume = 306 | issue = 5693 | pages = 129–131 | date = |
[[Šarenica]] [[kokoš]]ijih [[embrion]]a osjeća svjetlo kratkih talasa preko kriptohroma, a ne opsina.<ref>{{cite journal | vauthors = Tu DC, Batten ML, Palczewski K, Van Gelder RN | title = Nonvisual photoreception in the chick iris | journal = Science | volume = 306 | issue = 5693 | pages = 129–131 | date = oktobar 2004 | pmid = 15459395 | doi = 10.1126/science.1101484 | bibcode = 2004Sci...306..129T | s2cid = 26821205 }}</ref> |
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=== [[Cirkadijski ritam]] === |
=== [[Cirkadijski ritam]] === |
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{{glavni|Magnetorecepcija}} |
{{glavni|Magnetorecepcija}} |
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Magnetorecepcija je čulo koje omogućava organizmu da detektuje magnetsko polje kako bi uočio pravac, nadmorsku visinu ili lokaciju. Eksperimentalni podaci sugeriraju da su kriptohromi u [[fotoreceptorskim ćelijama|fotoreceptorskim neuronima]] ptičjih očiju uključeni u magnetnu orijentaciju tokom [[migracija|migracije]]. Smatra se da su i kriptohromi uključeni u magnetnu orijentaciju tokom [[migracija ptica]] od suštinskog značaja za sposobnost ''[[Drosophila]]'' zavisne od svetlosti da oseti [[magnetna polja]]. Nekada je prijavljeno da magnetna polja utiču na kriptohrome i u ''[[Arabidopsis thaliana]]'': činilo se da na ponašanje utieču magnetna polja u prisustvu plave (ali ne crvene) svjetlosti<ref name="Ahmad_1993">{{cite journal | vauthors = Ahmad M, Cashmore AR | title = HY4 gene of A. thaliana encodes a protein with characteristics of a blue-light photoreceptor | journal = Nature | volume = 366 | issue = 6451 | pages = 162–166 | date = |
Magnetorecepcija je čulo koje omogućava organizmu da detektuje magnetsko polje kako bi uočio pravac, nadmorsku visinu ili lokaciju. Eksperimentalni podaci sugeriraju da su kriptohromi u [[fotoreceptorskim ćelijama|fotoreceptorskim neuronima]] ptičjih očiju uključeni u magnetnu orijentaciju tokom [[migracija|migracije]]. Smatra se da su i kriptohromi uključeni u magnetnu orijentaciju tokom [[migracija ptica]] od suštinskog značaja za sposobnost ''[[Drosophila]]'' zavisne od svetlosti da oseti [[magnetna polja]]. Nekada je prijavljeno da magnetna polja utiču na kriptohrome i u ''[[Arabidopsis thaliana]]'': činilo se da na ponašanje utieču magnetna polja u prisustvu plave (ali ne crvene) svjetlosti<ref name="Ahmad_1993">{{cite journal | vauthors = Ahmad M, Cashmore AR | title = HY4 gene of A. thaliana encodes a protein with characteristics of a blue-light photoreceptor | journal = Nature | volume = 366 | issue = 6451 | pages = 162–166 | date = novembar 1993 | pmid = 8232555 | doi = 10.1038/366162a0 | s2cid = 4256360 | bibcode = 1993Natur.366..162A }}</ref> |
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Ipak, kasnije se pokazalo da su ovi rezultati neponovljivi pod strogo kontroliranim uvjetima u drugom laboratorija, sugerirajući da biljni kriptohromi ne reaguju na magnetna polja. |
Ipak, kasnije se pokazalo da su ovi rezultati neponovljivi pod strogo kontroliranim uvjetima u drugom laboratorija, sugerirajući da biljni kriptohromi ne reaguju na magnetna polja. |
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[[slika:Quantum Magnetoreception in Birds.svg|thumb|center|upright=3.5| Mehanizam radikalnih parova je predložen za kvantnu magnetorecepciju kod ptica.<ref name="Hore Mouritsen 2022">{{cite journal |last1=Hore |first1=Peter J. |last2=Mouritsen |first2=Henrik |title=The Quantum Nature of Bird Migration |journal=Scientific American |date= |
[[slika:Quantum Magnetoreception in Birds.svg|thumb|center|upright=3.5| Mehanizam radikalnih parova je predložen za kvantnu magnetorecepciju kod ptica.<ref name="Hore Mouritsen 2022">{{cite journal |last1=Hore |first1=Peter J. |last2=Mouritsen |first2=Henrik |title=The Quantum Nature of Bird Migration |journal=Scientific American |date=april 2022 |pages=24–29}}</ref>]] |
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Kriptohrom formira par [[radikal (hemija)|radikal]] sa koreliranim [[spin (fizika)|spinom]] kada je izložen plavoj svjetlosti.<ref name="Biskup_2009">{{cite journal | vauthors = Biskup T, Schleicher E, Okafuji A, Link G, Hitomi K, Getzoff ED, Weber S | title = Direct observation of a photoinduced radical pair in a cryptochrome blue-light photoreceptor | journal = Angewandte Chemie | volume = 48 | issue = 2 | pages = 404–407 | year = 2009 | pmid = 19058271 | pmc = 4329312 | doi = 10.1002/anie.200803102}}</ref> |
Kriptohrom formira par [[radikal (hemija)|radikal]] sa koreliranim [[spin (fizika)|spinom]] kada je izložen plavoj svjetlosti.<ref name="Biskup_2009">{{cite journal | vauthors = Biskup T, Schleicher E, Okafuji A, Link G, Hitomi K, Getzoff ED, Weber S | title = Direct observation of a photoinduced radical pair in a cryptochrome blue-light photoreceptor | journal = Angewandte Chemie | volume = 48 | issue = 2 | pages = 404–407 | year = 2009 | pmid = 19058271 | pmc = 4329312 | doi = 10.1002/anie.200803102}}</ref> |
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Parovi radikala se također mogu generirati reoksidacijom [[kofaktor (biohemija)|kofaktora]] flavina u mraku nezavisnom od svjetla molekulskim kisikom kroz formiranje parova radikala koreliranih sa FADH-superoksidom. Pretpostavlja se da magnetorecepcija funkcionišu kroz uticaj okolnog magnetnog polja na [[korelacija|korelaciju]] (paralelnu ili antiparalelnu) ovih radikala, što utiče na životni vijek aktiviranog oblika kriptohroma. Aktivacija kriptohroma može uticati na osetljivost na svjetlost [[neuron]]a [[retina|mrežnjače]]l, sa ukupnim rezultatom da životinja može da oseti magnetno polje.<ref name="Chandler_2011">{{cite web | url = http://www.ks.uiuc.edu/Research/cryptochrome/ | title = Cryptochrome and Magnetic Sensing | vauthors = Chandler D, Ilia Solov'yov I, Schulten K | publisher = Beckman Institute for Advanced Science and Technology, University of Illinois Urbana–Champaign | access-date = |
Parovi radikala se također mogu generirati reoksidacijom [[kofaktor (biohemija)|kofaktora]] flavina u mraku nezavisnom od svjetla molekulskim kisikom kroz formiranje parova radikala koreliranih sa FADH-superoksidom. Pretpostavlja se da magnetorecepcija funkcionišu kroz uticaj okolnog magnetnog polja na [[korelacija|korelaciju]] (paralelnu ili antiparalelnu) ovih radikala, što utiče na životni vijek aktiviranog oblika kriptohroma. Aktivacija kriptohroma može uticati na osetljivost na svjetlost [[neuron]]a [[retina|mrežnjače]]l, sa ukupnim rezultatom da životinja može da oseti magnetno polje.<ref name="Chandler_2011">{{cite web | url = http://www.ks.uiuc.edu/Research/cryptochrome/ | title = Cryptochrome and Magnetic Sensing | vauthors = Chandler D, Ilia Solov'yov I, Schulten K | publisher = Beckman Institute for Advanced Science and Technology, University of Illinois Urbana–Champaign | access-date = 14. 4. 2011 }}</ref> |
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Životinjski kriptohromi i blisko srodne životinje (6-4 ) fotolijaze sadrže duži lanac [[triptofan]]a za prijenos elektrona od ostalih proteina natporodice kriptohrom-[[fotolijaza]] (triptofanska tetrada umjesto trijade). Što je duže lanac dovodi do boljeg razdvajanja i preko 1000 puta dužeg vijeka trajanja fotoinduciranih parova radikala flavin-[[triptofan]] nego u proteinima sa trijadom triptofana. Odsustvo spin-selektivnih rekombinacija ovih radikalnih parova na vremenskim skalama od nanosekunde do mikrosekunde izgleda nekompatibilna sa sugestijom da je magnetorecepcija kriptohroma zasnovana na reakciji prednjeg svjetla. |
Životinjski kriptohromi i blisko srodne životinje (6-4 ) fotolijaze sadrže duži lanac [[triptofan]]a za prijenos elektrona od ostalih proteina natporodice kriptohrom-[[fotolijaza]] (triptofanska tetrada umjesto trijade). Što je duže lanac dovodi do boljeg razdvajanja i preko 1000 puta dužeg vijeka trajanja fotoinduciranih parova radikala flavin-[[triptofan]] nego u proteinima sa trijadom triptofana. Odsustvo spin-selektivnih rekombinacija ovih radikalnih parova na vremenskim skalama od nanosekunde do mikrosekunde izgleda nekompatibilna sa sugestijom da je magnetorecepcija kriptohroma zasnovana na reakciji prednjeg svjetla. |
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==Reference== |
==Reference== |
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<ref name="Ahmad_2007">{{cite journal | vauthors = Ahmad M, Galland P, Ritz T, Wiltschko R, Wiltschko W | title = Magnetic intensity affects cryptochrome-dependent responses in Arabidopsis thaliana | journal = Planta | volume = 225 | issue = 3 | pages = 615–624 | date = |
<ref name="Ahmad_2007">{{cite journal | vauthors = Ahmad M, Galland P, Ritz T, Wiltschko R, Wiltschko W | title = Magnetic intensity affects cryptochrome-dependent responses in Arabidopsis thaliana | journal = Planta | volume = 225 | issue = 3 | pages = 615–624 | date = februar 2007 | pmid = 16955271 | doi = 10.1007/s00425-006-0383-0 | s2cid = 96263}} |
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*{{cite press release |date= |
*{{cite press release |date=7. 9. 2006 |title=The "sixth sense" of plants |website=Centre national de la recherche scientifique |url=http://www2.cnrs.fr/en/664.htm?debut=16 |archive-url=https://web.archive.org/web/20110716080827/http://www2.cnrs.fr/en/664.htm?debut=16 |archive-date=16. 7. 2011}}</ref> |
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<ref name="Berndt_2007">{{cite journal | vauthors = Berndt A, Kottke T, Breitkreuz H, Dvorsky R, Hennig S, Alexander M, Wolf E | title = A novel photoreaction mechanism for the circadian blue light photoreceptor Drosophila cryptochrome | journal = The Journal of Biological Chemistry | volume = 282 | issue = 17 | pages = 13011–13021 | date = |
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<ref name="Thompson_2004">{{cite book | vauthors = Thompson CL, Sancar A | editor = Lenci F, Horspool WM | title = CRC handbook of organic photochemistry and photobiology | publisher = CRC Press | location = Boca Raton | year = 2004 | pages = 1381–89 | isbn = 978-0-8493-1348-6 | chapter= Cryptochrome: Discovery of a Circadian Photopigment}}</ref> |
<ref name="Thompson_2004">{{cite book | vauthors = Thompson CL, Sancar A | editor = Lenci F, Horspool WM | title = CRC handbook of organic photochemistry and photobiology | publisher = CRC Press | location = Boca Raton | year = 2004 | pages = 1381–89 | isbn = 978-0-8493-1348-6 | chapter= Cryptochrome: Discovery of a Circadian Photopigment}}</ref> |
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<ref name="Todo_1996">{{cite journal | vauthors = Todo T, Ryo H, Yamamoto K, Toh H, Inui T, Ayaki H, Nomura T, Ikenaga M | display-authors = 6 | title = Similarity among the Drosophila (6-4)photolyase, a human photolyase homolog, and the DNA photolyase-blue-light photoreceptor family | journal = Science | volume = 272 | issue = 5258 | pages = 109–112 | date = |
<ref name="Todo_1996">{{cite journal | vauthors = Todo T, Ryo H, Yamamoto K, Toh H, Inui T, Ayaki H, Nomura T, Ikenaga M | display-authors = 6 | title = Similarity among the Drosophila (6-4)photolyase, a human photolyase homolog, and the DNA photolyase-blue-light photoreceptor family | journal = Science | volume = 272 | issue = 5258 | pages = 109–112 | date = april 1996 | pmid = 8600518 | doi = 10.1126/science.272.5258.109 | s2cid = 23151554 | bibcode = 1996Sci...272..109T }}</ref> |
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<ref name="Ukai-Tadenuma_2011">{{cite journal | vauthors = Ukai-Tadenuma M, Yamada RG, Xu H, Ripperger JA, Liu AC, Ueda HR | title = Delay in feedback repression by cryptochrome 1 is required for circadian clock function | journal = Cell | volume = 144 | issue = 2 | pages = 268–281 | date = |
<ref name="Ukai-Tadenuma_2011">{{cite journal | vauthors = Ukai-Tadenuma M, Yamada RG, Xu H, Ripperger JA, Liu AC, Ueda HR | title = Delay in feedback repression by cryptochrome 1 is required for circadian clock function | journal = Cell | volume = 144 | issue = 2 | pages = 268–281 | date = januar 2011 | pmid = 21236481 | doi = 10.1016/j.cell.2010.12.019 | s2cid = 8159963 }}</ref> |
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<ref name="urlwww.rcsb.org">{{cite web | url = http://www.rcsb.org/pdb/images/1U3C_ram_m_500.pdf | title = MolProbity Ramachandran analysis,1U3C, model 1 | publisher = www.rcsb.org | access-date = |
<ref name="urlwww.rcsb.org">{{cite web | url = http://www.rcsb.org/pdb/images/1U3C_ram_m_500.pdf | title = MolProbity Ramachandran analysis,1U3C, model 1 | publisher = www.rcsb.org | access-date = 13. 4. 2011 | archive-url = https://web.archive.org/web/20121021035015/http://www.rcsb.org/pdb/images/1U3C_ram_m_500.pdf | archive-date = 21. 10. 2012 | url-status = dead }}</ref> |
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<!--<ref name="van_der_Spek_1996">{{cite journal | vauthors = van der Spek PJ, Kobayashi K, Bootsma D, Takao M, Eker AP, Yasui A | title = Cloning, tissue expression, and mapping of a human photolyase homolog with similarity to plant blue-light receptors | journal = Genomics | volume = 37 | issue = 2 | pages = 177–82 | date = |
<!--<ref name="van_der_Spek_1996">{{cite journal | vauthors = van der Spek PJ, Kobayashi K, Bootsma D, Takao M, Eker AP, Yasui A | title = Cloning, tissue expression, and mapping of a human photolyase homolog with similarity to plant blue-light receptors | journal = Genomics | volume = 37 | issue = 2 | pages = 177–82 | date = oktobar 1996 |pmid=8921389 |doi=10.1006/geno.1996.0539 |url=http://repub.eur.nl/pub/55742 |hdl=1765/55742 }}</ref>--> |
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<ref name="Vitaterna_1999">{{cite journal | vauthors = Vitaterna MH, Selby CP, Todo T, Niwa H, Thompson C, Fruechte EM, Hitomi K, Thresher RJ, Ishikawa T, Miyazaki J, Takahashi JS, Sancar A | display-authors = 6 | title = Differential regulation of mammalian period genes and circadian rhythmicity by cryptochromes 1 and 2 | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 96 | issue = 21 | pages = 12114–12119 | date = |
<ref name="Vitaterna_1999">{{cite journal | vauthors = Vitaterna MH, Selby CP, Todo T, Niwa H, Thompson C, Fruechte EM, Hitomi K, Thresher RJ, Ishikawa T, Miyazaki J, Takahashi JS, Sancar A | display-authors = 6 | title = Differential regulation of mammalian period genes and circadian rhythmicity by cryptochromes 1 and 2 | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 96 | issue = 21 | pages = 12114–12119 | date = oktobar 1999 | pmid = 10518585 | pmc = 18421 | doi = 10.1073/pnas.96.21.12114 | doi-access = free | bibcode = 1999PNAS...9612114V }}</ref> |
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<ref name="Weber_2005">{{cite journal | vauthors = Weber S | title = Light-driven enzymatic catalysis of DNA repair: a review of recent biophysical studies on photolyase | journal = Biochimica et Biophysica Acta (BBA) - Bioenergetics | volume = 1707 | issue = 1 | pages = 1–23 | date = |
<ref name="Weber_2005">{{cite journal | vauthors = Weber S | title = Light-driven enzymatic catalysis of DNA repair: a review of recent biophysical studies on photolyase | journal = Biochimica et Biophysica Acta (BBA) - Bioenergetics | volume = 1707 | issue = 1 | pages = 1–23 | date = februar 2005 | pmid = 15721603 | doi = 10.1016/j.bbabio.2004.02.010 | doi-access = free }}</ref> |
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<ref name="Zhu_2008a">{{cite journal | vauthors = Zhu H, Sauman I, Yuan Q, Casselman A, Emery-Le M, Emery P, Reppert SM | title = Cryptochromes define a novel circadian clock mechanism in monarch butterflies that may underlie sun compass navigation | journal = PLOS Biology | volume = 6 | issue = 1 | pages = e4 | date = |
<ref name="Zhu_2008a">{{cite journal | vauthors = Zhu H, Sauman I, Yuan Q, Casselman A, Emery-Le M, Emery P, Reppert SM | title = Cryptochromes define a novel circadian clock mechanism in monarch butterflies that may underlie sun compass navigation | journal = PLOS Biology | volume = 6 | issue = 1 | pages = e4 | date = januar 2008 | pmid = 18184036 | pmc = 2174970 | doi = 10.1371/journal.pbio.0060004 }}</ref> |
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<ref name="Zhu_2008b">{{cite journal | vauthors = Zhu H, Yuan Q, Briscoe AD, Froy O, Casselman A, Reppert SM | title = The two CRYs of the butterfly | journal = Current Biology | volume = 15 | issue = 23 | pages = R953–R954 | date = |
<ref name="Zhu_2008b">{{cite journal | vauthors = Zhu H, Yuan Q, Briscoe AD, Froy O, Casselman A, Reppert SM | title = The two CRYs of the butterfly | journal = Current Biology | volume = 15 | issue = 23 | pages = R953–R954 | date = decembar 2005 | pmid = 16332522 | doi = 10.1016/j.cub.2005.11.030 | s2cid = 2130485 | doi-access = free }}</ref> |
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== Vanjski linkovi == |
== Vanjski linkovi == |
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* {{MeshName|cryptochrome}} |
* {{MeshName|cryptochrome}} |
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* [http://www.umassmed.edu/neuroscience/faculty/reppert.cfm?start=0 Cryptochrome circadian clock in Monarch Butterflies] {{Webarchive|url=https://web.archive.org/web/20111121112812/http://www.umassmed.edu/neuroscience/faculty/reppert.cfm?start=0 |date= |
* [http://www.umassmed.edu/neuroscience/faculty/reppert.cfm?start=0 Cryptochrome circadian clock in Monarch Butterflies] {{Webarchive|url=https://web.archive.org/web/20111121112812/http://www.umassmed.edu/neuroscience/faculty/reppert.cfm?start=0 |date=21. 11. 2011 }}, by Steven M. Reppert, Department of Neurobiology, University of Massachusetts |
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* [http://www.ks.uiuc.edu/Research/cryptochrome/ Cryptochrome and Magnetic Sensing], ''Theoretical and Computational Biophysics Group'' at the University of Illinois at Urbana-Champaign |
* [http://www.ks.uiuc.edu/Research/cryptochrome/ Cryptochrome and Magnetic Sensing], ''Theoretical and Computational Biophysics Group'' at the University of Illinois at Urbana-Champaign |
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* [http://www.rcsb.org/pdb/explore.do?structureId=2IJG 2IJG] at the [[Protein Data Bank]]; 3-D structure of ''Arabidopsis'' cryptochrome 3, obtained by X-ray crystallography. |
* [http://www.rcsb.org/pdb/explore.do?structureId=2IJG 2IJG] at the [[Protein Data Bank]]; 3-D structure of ''Arabidopsis'' cryptochrome 3, obtained by X-ray crystallography. |
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{{Flavoproteini}} |
{{Flavoproteini}} |
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{{popravak DNK}} |
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[[Kategorija:Fiziologija]] |
[[Kategorija:Fiziologija]] |
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[[Kategorija:Biološki pigmenti]] |
[[Kategorija:Biološki pigmenti]] |
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Trenutna verzija na dan 29 januar 2023 u 22:30
| Kriptohrom-1 | |||||||
|---|---|---|---|---|---|---|---|
 Kristalografska struktura kriptohroma-1 | |||||||
| Identifikatori | |||||||
| Simbol | CRY1 | ||||||
| NCBI gen | 1407 | ||||||
| HGNC | 2384 | ||||||
| OMIM | 601933 | ||||||
| PDB | 5T5X | ||||||
| RefSeq | NP_004066 | ||||||
| UniProt | Q16526 | ||||||
| Ostali podaci | |||||||
| Lokus | Hrom. 12 q23.3 | ||||||
| |||||||
| Kriptohrom-2 | |||||||
|---|---|---|---|---|---|---|---|
 | |||||||
| Identifikatori | |||||||
| Simbol | CRY2 | ||||||
| NCBI gen | 1408 | ||||||
| HGNC | 2385 | ||||||
| OMIM | 603732 | ||||||
| PDB | 4MLP | ||||||
| RefSeq | NP_066940 | ||||||
| UniProt | Q49AN0 | ||||||
| Ostali podaci | |||||||
| Lokus | Hrom. 11 p11.2 | ||||||
| |||||||
Kriptohromi (od grčkog κρυπτός χρώμα – kriptos + hroma = "skrivena boja") su klasa flavoproteina pronađenih u biljkama i životinjama koje su osjetljivi na plavo svjetlo. Uključeni su u cirkadijanske ritmove i osjećanje magnetnih polja kod brojnih vrsta. Naziv kriptohrom je predložen kao portmanteau koji kombinuje hromatsku prirodu fotoreceptorskih proteina i kriptogamnih organizama na kojima su provedena mnoga istraživanja plavog svjetlosti.[1][2]
Dva gena "Cry1" i "Cry2" kodiraju dva kriptohromna proteina, CRY1 i CRY2.[3] Kod insekata i biljaka, CRY1 reguliše cirkadijski sat na način ovisan o svjetlosti, dok kod sisara, CRY1 i CRY2 djeluju kao svjetlosno-nezavisni inhibitori CLOCK-BMAL1, komponente cirkadijskog sata. U biljkama, fotorecepcija plavog svjetla može se koristiti za signaliziranje razvojnih signala. Osim hlorofila, kriptohromi su jedini proteini za koje se zna da formiraju fotoindukovane radikalne parove in vivo.[4] Čini se da one omogućavaju nekim životinjama da otkriju magnetna polja.
Kriptohromi su bili u fokusu nekoliko dosadašnjih napora u optogenetici. Koristeći transfekciju, početne studije na kvascu su iskoristile potencijal heterodimerizacija Cry2 da kontroliše ćelijske procese, uključujući ekspresiju gena, pomoću svetlosti.
Evolucijska historija i struktura
[uredi | uredi izvor]Kriptohromi (CRY1, CRY2) su evolucijski stari i visoko konzervirani proteini koji pripadaju natporodici flavoproteina koja postoji u svim carstvima života. Svi njeni članovi imaju karakteristike N-terminalnu homologijiju fotolijaze (PHR) domena. PHR domen se može vezati za flavin-adenin dinukleotidni (FAD) kofaktor i hromofor koji sakuplja svjetlost. Kriptohromi su izvedeni od i blisko srodne fotoliazama, koje su bakterijski enzimi koji se aktiviraju svjetlošću i koji su uključeni u popravak oštećenja DNK izazvanog UV zračenjem. Kod eukariota, kriptohromi više ne zadržavaju ovu originalnu enzimsku aktivnost. Struktura kriptohroma uključuje nabor koji je veoma sličan onom fotoliaze, sa jednim molekulom FAD-a koji je nekovalentno vezan za protein, Ovi proteini imaju promjenjive dužine i površine na C-terminalnom kraju, zbog promjena u genomu i izgledu koje su rezultat nedostatka enzima za popravak DNK. Ramachandranov plot pokazuje da je sekundarna struktura CRY1 proteina prvenstveno desnoruki alfa-heliks sa malo ili nimalo sternih preklapanja. Struktura CRY1 je skoro u potpunosti sastavljena od alfa heliksa, sa nekoliko petlji i nekoliko beta-listova. Molekul je raspoređen kao ortogonalni snop.
Funkcija
[uredi | uredi izvor]Fototropizam
[uredi | uredi izvor]Kod biljaka, kriptohromi posreduju fototropizam, ili usmjereni rast prema izvoru svjetlosti, kao odgovor na plavo svjetlo. Sada je poznato da ovaj odgovor ima svoj vlastiti skup fotoreceptora, fototropina.
Za razliku od fitohroma i fototropina, kriptohromi nisu kinaze. Njihov flavinski hromofor se redukuje pod uticajem svetlosti i transportuje u ćelijsko jedro, gde utiče na turgorski pritisak i uzrokuje naknadno izduživanje stabljike. Preciznije, „Cry2“ je odgovoran za kotiledone, posredovano plavom svjetlošću i širenje listova. Prekomjerna ekspresija Cry2 u transgenim biljkama povećava širenje kotiledona, stimulirano plavim svjetlom, što rezultira velikim brojem širokih listova i bez cvjetova, umjesto nekoliko primarnih listova s cvijetom. Dvostruka mutacija gubitka funkcije u genima Arabidopsis thaliana rnocvjetajućeg elf3) i Cry2 odgađa cvjetanje pod kontinuiranim svjetlom i pokazalo se da ga ubrzava tokom dugih i kratkih dana, što sugerira da Arabidopsis CRY2 može imati ulogu u ubrzavanju vremena cvjetanja tokom neprekidnog svetla.
Fotomorfogeneza
[uredi | uredi izvor]Receptori kriptohroma uzrokuju da biljke reaguju na plavo svjetlo putem fotomorfogeneze. Pomažu u kontroli razvoja sjemena i sadnica, kao i prelasku iz vegetativne u fazu cvjetanja. U Arabidopsis je pokazano da kriptohromi kontrolišu rast biljaka tokom suboptimalnih uslova plavog svetla.
Snimanje svjetla
[uredi | uredi izvor]Uprkos velikom broju istraživanja o ovoj temi, kriptohromska fotorecepcija i fototransdukcija kod Drosophila i Arabidopsis thaliana još uvijek su slabo shvaćene. Poznato je da kriptohromi posjeduju dva hromofora: pterin (u obliku 5,10-meteniltetrahidrofolne kisekine (MTHF)) i flavin (u obliku FAD-a). Oba mogu apsorbirati foton, a u arabidopsisu se čini da pterin apsorbuje na talasnoj dužini od 380 nm, a flavin na 450 nm. Prethodne studije su podržale model kojim se energija zarobljena pterinom prenosi na flavin. Pod ovim modelom fototransdukcije, FAD bi tada bio reduciran u FADH, koji vjerovatno posreduje u fosforilaciji određenog domena u kriptohromu. Ovo bi onda moglo pokrenuti lanac transdukcija signala, što bi moglo utjecati na regulaciju gena u ćelijskom jedru.
Nova hipoteza predlaže da u biljnim kriptohromima transdukcija svjetlosnog signala u hemijski signal koji mogu osjetiti partnerske molekule može biti pokrenuta fotoinduciranim negativnim nabojem unutar proteina - na kofaktor FAD ili na susjednu asparaginsku kiselinu. Ovaj negativni naboj bi elektrostatički odbio molekulu ATP vezanu za proteine, a time i protein C-terminalni domen, koji pokriva ATP vezni džep prije apsorpcije fotona. Rezultirajuća promjena u konformaciji proteina mogla bi dovesti do fosforilacije prethodno nedostupnih fosforilacijskih mjesta na C-terminalu i dati fosforilirani segment bi tada mogao osloboditi transkripcijski faktor HY5, takmičeći se za isto mjesto vezivanja na negativnom regulatoru fotomorfogeneze COP1 .
Kod Drosophila može funkcionirati drugačiji mehanizam. Pravo osnovno stanje kofaktora flavina u Drosophila CRY se još uvijek raspravlja, a neki modeli ukazuju da je FAD u oksidiranom obliku, dok drugi podržavaju model u kojem flavinski kofaktor postoji u anionskom radikalnom obliku, FAD−•. Nedavno je u istraživanjimaa primiječeno da se oksidirani FAD lahko redukuje svjetlom u FAD−•. Štaviše, mutacije da blokirana fotoredukcija nije imala efekta na razgradnju CRY izazvanu svjetlom, dok su mutacije koje su promijenile stabilnost FAD−• uništile funkciju CRY fotoreceptora.[5][6] Ova zapažanja pružaju podršku za osnovno stanje FAD−•. Istraživači su također nedavno predložili model u kojem se FAD− pobuđuje u svoje dubletno ili kvartetno stanje apsorpcijom fotona, što zatim dovodi do konformacijske promjene u CRY proteinu .[6]
Također, prstenaste oči larve demosunđera Amphimedon queenslandica izražavaju kriptohrom osetljiv na plavo svetlo (Aq-Cry2), koji bi mogao da posreduje fototaksiju. Nasuprot tome, oči većine životinja koriste fotoosjetljive opsine eksprimirane u fotoreceptorskim ćelijama, koje prenose informacije o svjetlosti iz okoline u nervni sistem. Međutim, A. queenslandica nema nervni sistem, kao i druge spužve. A također nema opsinski gen u svom potpuno sekvenciranom genomu, uprkos tome što ima mnogo drugih G-protein-spregnutih receptora (GPCR). Prema tome, jedinstvene oči spužve mora da su razvile drugačiji mehanizam za detekciju svjetlosti i posredovanje fototaksije, vjerovatno pomoću kriptohroma ili drugih proteina.
Funkcija šarenice
[uredi | uredi izvor]Šarenica kokošijih embriona osjeća svjetlo kratkih talasa preko kriptohroma, a ne opsina.[7]
Studije na životinjama i biljkama sugeriraju da kriptohromi imaju ključnu ulogu u stvaranju i održavanju cirkadijalnih ritmova. Slično tome, kriptohromi imaju važnu ulogu i u uvlačenju cirkadijskih ritmova u biljke. Kod Drosophila, kriptohrom (dCRY) djeluje kao fotoreceptor plave svjetlosti koji direktno modulira ulaz svjetlosti u cirkadijski sat, dok kod sisara kriptohromi (CRY1 i CRY2) djeluju kao transkripcijski represor unutar cirkadijsog sata. Neki insekti, uključujući reptira monarha, imaju sisarima i drozofilama slične verzije kriptohroma, pružajući dokaze za prastari mehanizam sata koji uključuje i ulogu senzora svetlosti i uloge transkripcione represije za kriptohrom.
Cry mutant je izmijenio cirkadijske ritmove, pokazujući da utiče na cirkadijski pejsmejker. Drosophila sa mutiranim Cry pokazuje malo ili nimalo ciklusa iRNK. Tačkasta mutacija u cryb, koja je potrebna za asocijaciju flavina u CRY proteinu, rezultira bez PER ili TIM cikliranja proteina ni u DD ni u LD. Osim toga, miševi kojima nedostaju "Cry1" ili "Cry2" geni pokazuju različito izmijenjene slobodne periode trčanja, ali su i dalje sposobni za fotopomjeranje. Međutim, miševi kojima nedostaju i Cry1 i Cry2 su aritmični u LD i DD i uvijek imaju visoke Per1 nivoe iRNK. Ovi rezultati sugeriraju da kriptohromi imaju fotoreceptivnu ulogu, kao i da djeluju kao negativni regulatori ekspresije gena Per kod miševa.
Magnetorecepcija
[uredi | uredi izvor]Magnetorecepcija je čulo koje omogućava organizmu da detektuje magnetsko polje kako bi uočio pravac, nadmorsku visinu ili lokaciju. Eksperimentalni podaci sugeriraju da su kriptohromi u fotoreceptorskim neuronima ptičjih očiju uključeni u magnetnu orijentaciju tokom migracije. Smatra se da su i kriptohromi uključeni u magnetnu orijentaciju tokom migracija ptica od suštinskog značaja za sposobnost Drosophila zavisne od svetlosti da oseti magnetna polja. Nekada je prijavljeno da magnetna polja utiču na kriptohrome i u Arabidopsis thaliana: činilo se da na ponašanje utieču magnetna polja u prisustvu plave (ali ne crvene) svjetlosti[8] Ipak, kasnije se pokazalo da su ovi rezultati neponovljivi pod strogo kontroliranim uvjetima u drugom laboratorija, sugerirajući da biljni kriptohromi ne reaguju na magnetna polja.
Kriptohrom formira par radikal sa koreliranim spinom kada je izložen plavoj svjetlosti.[10] Parovi radikala se također mogu generirati reoksidacijom kofaktora flavina u mraku nezavisnom od svjetla molekulskim kisikom kroz formiranje parova radikala koreliranih sa FADH-superoksidom. Pretpostavlja se da magnetorecepcija funkcionišu kroz uticaj okolnog magnetnog polja na korelaciju (paralelnu ili antiparalelnu) ovih radikala, što utiče na životni vijek aktiviranog oblika kriptohroma. Aktivacija kriptohroma može uticati na osetljivost na svjetlost neurona mrežnjačel, sa ukupnim rezultatom da životinja može da oseti magnetno polje.[11] Životinjski kriptohromi i blisko srodne životinje (6-4 ) fotolijaze sadrže duži lanac triptofana za prijenos elektrona od ostalih proteina natporodice kriptohrom-fotolijaza (triptofanska tetrada umjesto trijade). Što je duže lanac dovodi do boljeg razdvajanja i preko 1000 puta dužeg vijeka trajanja fotoinduciranih parova radikala flavin-triptofan nego u proteinima sa trijadom triptofana. Odsustvo spin-selektivnih rekombinacija ovih radikalnih parova na vremenskim skalama od nanosekunde do mikrosekunde izgleda nekompatibilna sa sugestijom da je magnetorecepcija kriptohroma zasnovana na reakciji prednjeg svjetla.
Reference
[uredi | uredi izvor]- ^ Gressel, J. (1979). "Blue Light Photoreception". Photochemistry and Photobiology (jezik: engleski). 30 (6): 749–754. doi:10.1111/j.1751-1097.1979.tb07209.x. ISSN 1751-1097. S2CID 98643540.
- ^ Yang Z, Liu B, Su J, Liao J, Lin C, Oka Y (januar 2017). "Cryptochromes Orchestrate Transcription Regulation of Diverse Blue Light Responses in Plants". Photochemistry and Photobiology. 93 (1): 112–127. doi:10.1111/php.12663. PMC 6167254. PMID 27861972.
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Vanjski linkovi
[uredi | uredi izvor]- cryptochrome na US National Library of Medicine Medical Subject Headings (MeSH)
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