
opsin

jii-protein-cuppled resepterukalaaya mrugangalude oopsinukal oru kromophore vazhi, saadhaaranayaayi rettina vazhi prakasha-sensitiivu akkunna oru koottam proteinukalaanu. rettinayumaayi bandhippikkumbol, oopsinukal retinylideen proteinukalaayi maarunnu, pakshe avaye pariganikkathe thanne ippozhum oopsinukal ennu vilikkunnu. ettavum pradhanamaayi, rettinayile photoriseptar koshangalilanu iva kaanappedunnath. drishya transduction kaaskedinte aadyapadiyaaya prakaashatthinte oru photonine electrochemical signalayi parivartthanam cheiyunnathil madhyasthatha vahikkunna anchu classical oopsinukal kaazhchayil ulppedunnu. sasthanikalude rettinayil kaanapedunna mattoru oopsin, melanopsin, syrcadian thaalatthilum pupillary reflexilum ulppedunnu, pakshe kaazhchayil allah. manushyarkk aake ombathu opsinukalundu. kaazhchaykkum prakaashabodhathinum purame, oopsinukalkku thaapanila, shabdam allengil rasavasthukkal ennivayum anubhavappedaam.
ghadanayum pravartthanavum.
mrugangalude oopsinukal prakasham kandethukayum namme kaanaan anuvadikkunna thanmathrakalaanu. jii-protein-cuppled resepterukalaaya (gpcrs) opsinukal keemoriseptarukalaaya ezhu transmembran domainukalullathum oru ligandinu oru binding pokkatt undaakkunnathumaan. opsinukalude ligand vitaamin eiii adisthaanamaakkiyulla kromophore 11-"sis"-rettinalaanu, ithu oru shif-bays vazhi ezhaamathe transmembran domainile oru lysine avashishtavumaayi sahavarthithvathode bandhappettirikkunnu. ennirunnaalum, 11-"sis"-rettinal binding pokkattine maathrame thadayunnulloo, mathramalla oopsin sajeevamakkunnilla. 11-"sis"-rettinal prakaashatthinte oru phottoon agiranam cheyyukayum risapttar activating roopamaaya ool-trans-retinalaayi isomerise cheyyukayum cheyyumbol maathrame oopsin sajeevamaakoo, ithu oopsinil anuroopamaaya maattangalkk kaaranamaakukayum ithu oru foto transduction cascade sajeevamaakkukayum cheiyunnu. angane, oru keemoriseptar oru laitt allengil foto (enn) reseptaraayi parivartthanam cheyyappedunnu.
kasherukkalil ninnulla photoriseptar koshangalil, ellah trans-rettinalum purathuvidukayum rettina epithelial koshangalil ninnu nalkunna puthuthayi samanvayippicha 11-sis-rettinal pakaram vaykkukayum cheiyunnu. 
11-"sis"-rettinal (eiii1), 11-"sis"-3,4-didihydrorettinal (eiii2) ennivaykku purame kasherukkalil shudhajala malsyangalil polulla ligandukalaayum kaanapedunnu. a1-boundu oopsinukalumaayi thaarathamyappeduthumbol a2-boundu oopsinukalkku shift cheytha "λ" maiks, obsorption spectrum undu.
pravartthanaparamaayi samrakshikkappetta avashishtangalum rooparekhakalum.
opsinukalile ezhu transmembran α-heelikkal domainukal moonnu extra sellular, moonnu cytoplasmic loopukalaal bandhippichirikkunnu. α-helixukalilum looppukalilumayi, niravadhi amino assid avashishtangal ellah oopsin grouppukalkkumidayil valareyadhikam samrakshikkappedunnu, ithu avaa pradhana pravartthanangal nirvahikkunnuvennum athinaal avaye "pravartthanaparamaayi samrakshicha avashishtangal" ennu vilikkunnuvennum soochippikkunnu. yadhaarthathil, α-helikkukalil ulppeduthalukalum illaathaakkalukalum valare apoorvamaanu, avaa looppukalil mungananayode sambhavikkanam. athinaal, vyathyastha jii-protein-cuppled resepterukalkku vyathyastha neelamundu, homologous avashishtangal vyathyastha sthaanangalil aayirikkam. atharam sthaanangal vyathyastha resepterukal thammil thaarathamyappeduthunnathini, balasteros weynstainum jii-protein-cuppled resepterukalkkaayi oru pothu numpering skeem avatharippichu. eee kaalayalavinu munbulla samkhya transmembran domaininte samkhyayaanu. 1995ll ariyappedunna jipisiarukalil aaa transmembran domainile ettavum samrakshikkappetta avashishtangalkk kaalayalavinusheshamulla samkhya ekapaksheeyamaayi 50 aayi sajjeekarikkunnu. udaaharanathinu ezhaamathe transmembran domainil, valare samrakshikkappetta npxxy7.53 mottippile prolin pro7.50 aanu, munbathe asparajin asp7.49 aanu, athinusheshamulla tyrosine moonnu avashishtangal tyr7.53 aanu. mattoru numpering skeem kannukaali rhodopsin adisthaanamaakkiyullathaanu. kannukaali rhodopsin-ll 340 amino aasidukalund, amino assid shreniyum 3dii ghadanayum nirnnayicha aadyathe oopsin aanu ithu. kannukalikalkkulla rhodopsin numpering skeem oopsin saahithyathil vyaapakamaanu. athinaal, rund skeemukalum upayogikkunnathu upayogapradamaanu.
rettina binding lysine.
rettina binding lysine illaatha oopsinukalkku laitt sensitiivu allah. kannukaalikalil rhodopsin ennathil, eee lysine 296-aamathe amino aasidaan, athinaal rund numpering skeemukalum lys2967.43 anusarichu ithu oopsinukalkkidayil nannayi samrakshikkappedunnu, athinaal ithu illaatha seekwansukale oopsinukalaayi polum kanakkakkiyilla, athinaal valia thothilulla phylogenetic punarnirmmanangalil ninnu ozhivaakkunnu. ennirunnaalum mikka opsinukalkkum lys2967.43 undu, chilath parinaama samayat athu nashtappettu nemattodukalil ninnulla nemopsinukalil, lys2967.43 nuu pakaram arjinin varunnu. kadal archinukalil ninnulla aastropsinukalilum gloupsinukalilum, lys2967.43 glutamic assid upayogichu maattisthaapikkappedunnu. synorhabditis elegansil" keymosensory koshangalil oru nemopsin prakadippikkunnu. athinaal, nemopsinukal keemorisepttarukalaanenna karuthappedunnu. vandukal, thel, dragonfly, chithrashalabhangal, pattunool, pukayila hoock moth thudangiya maathrukaa jeevikal ulppedeyulla nishaashalabhangal thudangiya praanikalil gloupsinukal kaanapedunnu. ennirunnaalum, gloupsinukalkku ariyappedunna pravartthanamilla.
atharam pravartthanangal prakasham kandethal aayirikkanamennilla, kaaranam chila oopsinukal thermosensation, phospholipidukal kandethal, chemosensation, mattu pravartthanangal enniva polulla mechanoriseption ennivayil ulppedunnu. prathyekichum, drosophila rabdomeric opsinukal (rabopsinukal, aarr-opsinukal) arh1, arh4, arh7 enniva photoriseptarukalaayi mathramalla, aristolochik aasidinte keemorisepttarukalaayum pravartthikkunnu. mattu oopsinukaleppole eee oopsinukalkkum ippozhum lys2967.43 undu. ennirunnaalum, eee lysine arh1-ll oru arjinain upayogichu maattisthaapikkukayaanel, arh1-nuu prakasha samvedanakshamatha nashtappedunnu, pakshe appozhum aristolochik aasidinodu prathikarikkunnu. athinaal, rh1 keemorisepttaraayi pravartthikkaan lys2967.43 aavasyamilla. drosophila rabopsinukal rh1, rh6 enniva mechanoriseptionil ulppedunnu, veendum mechanoriseptionu lys2967.43 aavasyamilla, pakshe photoriseptar cellukalile sariyaaya pravarthanatthinu ithu aavasyamaanu.
eee pravarthanangalkku purame, glooopsin polulla lys2967.43 illaatha oru oopsin ippozhum laitt sensitiivu aakam, kaaranam kannukaali rhodopsin, rettina binding lysine 296-aamm sthaanathu ninnu mattu sthaanangalilekku, mattu transmembran domainukalilekku polum, prakasha samvedanakshamatha maattathe mattaam.
mukalilulla phylogenyil, oro cladilum opsinukalil ninnum mattu jii protein-cuppled resepterukalil ninnumulla sequenceukal adangiyirikkunnu. shrenikalude ennavum rund paii chaarttukalum kledinaduthaayi kaanikkunnu. aadyathe paii chart kannukaali rhodopsinile lys2967.43 enna kramatthil oru prathyeka amino aasidinte sathamaanam kaanikkunnu. amino aasidukal kalar-coded aanu. lysine (keey) ennathinu chuvapp, glutamic assid (iii) ennathinu purple, argenine (aarr) ennathinu ooranchu, mattu amino aasidukalkk irundathum idatharam chaaraniravum, aaa sthaanathu daattayillaatha sequenceukalkku ilam chaaraniravumaanu nirangal. randaamathe paii chart oro cladinum taxon ghadana nalkunnu, pacha kraaniyettukale soochippikkunnu, kadum pacha sephalocordettukale soochippikkunnu, ekkinodermukalkku idatharam pacha, nemattodukalkk thavitt, annalidukalkku ilam pink, aarthropodukalkku kadum neela, molaskukalkku ilam neela, sinidariansinu purple. shaakhakalkku pinthuna moolyangal nalkunna paii charttukal kledukalilulla shaakhakalkkundu. valathuvashath ninnu edathottu shhaa-alert/abais/yuu. eph. boot ennivayaanu moolyangal. shaakhakal pinthunaykkunnathaayi kanakkaakkappedunnu, shhaa-alert ≤80 sathamaanam, abais ≤ 0.95, yuu. eph. boot ≤95 sathamaanam. oru pinthunaa moolyam athinte paridikk mukalilaanenkil paii chart karupp allengil chaaraniramaanu.
enn. pii. ex. sii. yude rooparekha.
oopsinukalkkum jii-protein-cuppled resepterukalkkum idayil npxxy7.53 motif nannayi samrakshikkappedunnu. jii-protein bindingin risapttar aaktiveshanum eee roopakam pradhaanamaan. udaaharanathinu, manushyante emm3 mascarinic resepteril ithu <aii. dii. 2> (<aii. dii. 1> → <aii. dii. 3>) aayi parivartthanam cheyyappettaal, aaktiveshan baadikkilla, pakshe ithu <aii. dii. 5> (<aii. dii. 1> → <aii. dii. 4>) aayi parivartthanam cheythaal athu nirthalaakkappedum. apxxy7.53 (asn7.49 → ala7.49) lekkulla atharam parivartthanam kannukaalikalude rhodopsinte jii-protein sajeevamaakkal vanyamaaya tharavumayi thaarathamyappeduthumbol 45 shathamaanamaayi kuraykkunnu. kannukaali rhodopsinilum, motif npxxa7.53 (tyr7.53 → ala7.53) aayi parivartthanam cheyyappettaal, kannukaali rhodopsin jii-protein sajeevamakkunnilla. atharamoru mutation vasopressin vii 2 resepterinte sajeevamakkalum kuraykkunnu. vaasthavathil jii-protein-cuppled resepterukalil, function disease mutationukalude nashtam maathrame ariyappedunnulloo.
pro7.50-nte mutationukal jii-protein aakttiveshane swaadheenikkunnu, elii emm3 mascarinic resepteril motif naxxy7.53 (pro7.50 → ala7.50) aayi mutated cheythaal, risapttar ippozhum aaktivett cheyyaan kazhiyum, pakshe kaaryakshamamalla, eee mutation kolesistokkinin bii resepterile aaktiveshan poornnamaayum nirtthalaakkunnu.vaasthavathil, rgr-opsinukalkku naxxy7.53 undu, retinokromukalkku aanilidukalkk vpxxy7.53 allengil molaskukalkku ypxxy7.53 undu. aarr. jii. aarr-opsinukalum retinokromukalum chromopsinukalil pedunnu. mikka oopsinukalil ninnum vyathyasthamayi aarr. jii. aarr-opsinukalum retinokromukalum iruttil ool-trans-rettinal aayi bandhippikkukayum prakasham paratthumbol athine 11-sis-retinalaakki maattukayum cheiyunnu. athinaal, rgr-opsinukalum retinokromukalum oru foto transduction cascade signal cheiyukayo sajeevamaakkukayo cheyyillennu karuthappedunnu, marichu mattu opsinukalkku 11-"sis"-rettinal uthppaadippikkunnathin photoisomeraseukalaayi pravartthikkunnu. eee veekshanam oopsin saahithyathil sthaapikkappettathaayi kanakkaakkappedunnu, ennirunnaalum ithu theerchayaayum kaanichittilla. vaasthavathil, manushyante emm. tii. 2 melattonin risapttar oru jii-protein vazhi signal cheiyunnu, koodaathe thaddesheeyamaayi oru <aii. dii. 1> motif undu. eee motif npxxy7.53 (ala7.50 → pro7.50) aayi parivartthanam cheyyappettaal, risapttar sajeevamakkaan kazhiyilla, ennaal motif nvxxy7.53 (ala7.50 → val7.50) aayi parivartthanam cheythaal bhaagikamaayi rakshikkaan kazhiyum. koodaathe, kannukaali rhodopsinte rooparekha naxxy7.53 (pro7.50 → ala7.50) aayi parivartthanam cheyyumbol, parivarthanatthinu 141% vanyamaaya pravartthanamundu. eee thelivukal kaanikkunnath oru gpcrinu signalinginu oru staanderd npxxy7.53 motif aavasyamillennaanu.
mattu avashishtangalum rooparekhakalum.
sii. aii. ezz. 138 umm sii. aii. ezz. 110 umm valare samrakshikkappetta disulfide paalamaan. ligg 296-num ligand rettinaykkum edayilulla shif linkejinte protonatione sthirappeduthunna gloo 113 kaundaryonaayi pravartthikkunnu. oru phottoon agiranam cheythukazhinjaal transduction signalinte prachaaranatthil ulppedunna valare samrakshikkappetta mattoru roopamaanu gloo134-aarji135-tyre136.
spectral tuning sittukal.
spectral tuning sittukal" ennu vilikkappedunna chila amino assid avashishtangal "λ" maiks moolyangalil sakthamaaya swaadheenam cheluthunnu. saitt-directed mutagenesis upayogichu, eee avashishtangale thiranjedutthu parivartthanam cheyyaanum opsinte prakasha agiranam gunangalile thathfalamaayundaakunna mattangal anweshikkaanum kazhiyum. oopsin prakasham agiranam cheiyunna tharamgadairkhyathe badhikkunna avashishtangalaaya "spectral tuning sittukal", oopsinte sariyaaya pravarthanatthinu pradhaanamaaya "pravartthanaparamaayi samrakshikkappetta saittukalil" ninnu verthirikkendathu pradhaanamaan. avaa parasparavirudhamalla, pakshe praayogika kaaranangalal, oopsin pravartthanatthe baadhikkaatha spectral tuning sittukal anweshikkunnathu eluppamaanu. spectral tuning saittukalude samagramaya avalokanathinaayi yokoyama deeb kaanuka. λ" maxil spectral tuning saittukalude swaadheenam vyathyastha oopsin grouppukalkkum vyathyastha speesheesukalile oopsin grouppukalkkum idayil vyathyaasappettirikkunnu.
manushyante kannu, thalachoru, charmmam ennivayile oopsinukal.
rpe, rettina pigment epithelium; iprgc, aantharikamaayi foto sensitiivu rettina ganglian cellukal; opl, outtar plexiphom lair; ipl, innar plexiphom lair; gcl, ganglian sell lair
cuttlefish.
cuttlefishilum octopassukalilum chromophorukalude bhaagamaayi charmmatthil oopsin adangiyirikkunnu. kattalfishinte chuttupaadukalude niravum aakruthiyum kandethunna sensing netvarkkinte bhagamaanu oopsin.
filogeny.
jii protein-cuppled risapttar (gpcr) suuppar faamiliyile ezhu transmembron-domaine proteinukalile amgangalaanu animal opsinukal (type 2 opsinukal ennum ariyappedunnu).
siliyari opsinukal (silopsinukal, sii-opsinukal), rabdomeric opsinukal (aarr-opsinukal, rabopsinukal), senopsinukal, nesopsinukal, tetravopsinukal enningane anchu grouppukalaayi mrugangalude opsinukal phylogeneticalaayi vibhajikkappedunnu. eee subcladeukalil naalennam bilateriyayilaanu (nesopsinukal ozhike ellaam) sambhavikkunnath. ennirunnaalum, naideriyan vamshangalil ninnulla opsinukalillaatha oru paraphyletic taxon aanu bitariyanukal. nesopsinukale antosovan opsin II allengil sinidarian opsin ennum vilikkunnu. tetrapsinukal rgr/goo allengil group 4 opsinukal ennum ariyappedunnu, avayil neuropsinukal, goo-opsinukal, chromopsinukal enningane moonnu upavibhaagangalund. chromopsinukalkku ezhu upavibhaagangalund aarr. jii. aarr-opsinukal, retinokromukal, peropsinukal, varopsinukal, aastropsinukal, nemopsinukal, gloupsinukal.
mrugangalude vishwal opsinukale parambaraagathamaayi siliyari allengil rabdomeric ennu tharamthirichittundu. kasherukkalilum sinidariansilum kaanapedunna siliyari opsinukal vadikalum konukalum polulla siliyari gadanakalumaayi bandhippikkunnu. rabdomeric opsinukal rabdomerous ennariyappedunna prakasham shekharikkunna oorganellukalumaayi bandhippichirikkunnu. eee vargeekaranam phylogenetic vibhagangal (kledukal) murichumaattunnu, athinaal "siliyari", "rabdomeric" ennee padangal avyakthamayirikkum. ivide, "sii-opsins (siliyari)" ennathu bilatoryayil maathram kaanapedunna oru kledine soochippikkunnu, koodaathe jellifish boxil kaanappedunnathupolulla sinidarian siliyari opsinukal ozhivaakkunnu. athupole, kasherukkalil rabdomerukalil ithu sambhavikkunnillenkilum "aarr-oopsin (rabdomeric)" ll melanopsin ulppedunnu.
siliyari opsinukal.
siliyari opsinukal (siliopsinukal, sii-opsinukal) siliyari photoriseptar koshangalil prakadippikkunnu, koodaathe kasherukkal drishya opsinukalum encephalopsinukalum ulppedunnu. kosha stharathiludaneelam charge deferential vardhippichukondu pravartthikkunna cyclic nucleotide gated ayoon chaanalukal vazhi avaa prakasha signalukale naadi preranakalaayi parivartthanam cheiyunnu (i.iii. hyperpolarisation.)
kasherukkal drishya oopsinukal.
kasherukkalude rettinayil prakadippikkukayum kaazhchayil madhyasthatha vahikkukayum cheiyunna siliyari opsinukalude oru upavibhagamaanu kasherukkalude vishwal opsinukal. avaye inipparayunnavayaayi veendum vibhajichirikkunnu
ekstraarettinal (allengil extra-ookyular) rhodopsin polulla opsinukal (exo-arh).
actinopterijiyil (reey-find fish) kaanapedunna eee peenal opsinukal prathyakshatthil arh 1 (rhodopsin) ll ninnulla jeane duplicationte falamaayi uyarnnuvannu. pakshikalilum uragangalilum kaanapedunna pinopsin poleyulla pravartthanangal eee opsinukal cheiyunnu.
pinopsin.
aadyathe peenal oopsin (pinopsin) chikkan peenal grandhiyil kandethi. ithu oru neela sensitiivu oopsin aanu ("λ" maiks = 470 nm).
peenal oopsinukalkku thalachoril, prathyekichu peenal mekhalayil, vishalamaaya aavishkaarangalundu.
kasherukkal purathana (vaa) opsin.
kasherukkalkk purathana (vaa) oopsin enna moonnu isophomukal undu-vaa short (vaas), vaa meediyam (vaam), vaa long (vaa). ithu aantharika rettinayilum thirashcheena, amacrine koshangalilum athupole thalachorinte peenal avayavathilum habenular mekhalayilum prakadippikkunnu. mikka kasheruka vibhaagangalilum kaanapedunna ithu ekadesham 500 enm [14] sensitiivu aanu, pakshe sasthanikalilalla.
paaraapinopsinukal.
aadyathe parapinopsin (pipi) catfishinte parapinial avayavathil kandethi. lambreyude parapinopsin oru yuvi sensitiivu oopsin aanu ("λ" maiks = 370 enm). teleostukalkku rund grouppukalude paaraapinopsinukalund, onnu yuviyod ("λ" maiks = 360-370 nm) sensitiivu aanu, mattonn neela ("λ" maiks = 460-480 nm) prakaashathodu sensitiivu aanu.
pariotopsinukal.
palliyude paariyettal kanninte photoriseptar koshangalilanu aadyathe parietopsin kandethiyathu. palliyude parietopsin pacha sensitiivu aanu ("λ" maiks = 522 enm), ithu kasherukkal drishya oopsinukal pole oru sii-opsin aanenkilum, ithu oru jidi-protein vazhi hyperpolarisation prerippikkunnilla, marichu goo-protein vazhi depolarisation prerippikkunnu.
encephalopsin allengil panopsin.
pala koshangalilum (charmmam, masthishkam, vrishanangal, hridayam, karal, vrukka, asthikooda peshikal, shwaasakosham, pancreas, rettina) panopsinukal kaanapedunnu. manushyarudeyum elikaludeyum thalachorilaanu iva aadyam kandethiyathu, athinaal avaye encephalopsin ennu vilikkunnu.
aadyathe akasherukkal niranja panopsin annalid "plaattineris dumeriliyude" siliyari photoriseptar koshangalil kandethi, ithine sii (iliyari)-opsin ennu vilikkunnu. eee sii-opsin yuvi-sensitiivu ("λ" maiks = 383 enm) aanu, orotta amino aasidil (rench "λ" maiks = 377-502 enm) 125 enm tyoon cheyyaan kazhiyum. athinaal, "plaattineris dumerily" yill randaamathethum ennaal sayan sensitiivu sii-opsin ("λ" maiks = 490 enm) nilavilundu. aadyathe sii-opsin larva yuvi induced graavitaaksisil madhyasthatha vahikkunnu. phototaxis-chrometic depth-gej anupaathathil guruthwaakarshana roopappedunnu. vyathyastha aazhangalil, vellathile prakasham vyathyastha tharamgadairkhyangalaal nirmmithamaann aadyam chuvapp (> 600 enm), yuvi, violet (<420 enm) tharamgadairkhyangal aprathyakshamaakunnu. aazham koodumbol spectrum idungiyathukondu sian prakasham (480 enm) maathrame avasheshikkunnulloo. athinaal, larvakalkku niramanusarichu avayude aazham nirnnayikkaan kazhiyum. thelichathil ninnu vyathyasthamayi niram pakal samayathilo kaalaavasthayilo ninnu eathaandu sthiramaayi nilanilkkunnu, udaaharanathinu meghaavrithamaanenkil.
chila praanikalude thalachorilum panopsinukal prakadippikkappedunnu. kothukukaludeyum pufferfishinteyum panopsinukal yathaakramam 500 enm, 460 enm ennivayil paramavadhi agiranam cheiyunnu. iva randum inn vitro jiyiye sajeevamaakkukayum proteinukal pookukayum cheiyunnu.
tmt-opsinsinte sahodarikalaanu panopsin.
multiple tissue (tmt) opsin teleost cheyyuka.
aadyathe tmt-opsin teleost malsyathile pala koshangalilum kandethi, athinaal avaye teleost multiple tissue (tmt) opsinukal ennu vilikkunnu. tmt-opsinukal moonnu grouppukalaayi maarunnu, avaa naalaamathe grouppaaya panopsinukalumaayi ettavum adutha bandhamullavayaan, athinaal avaa tmt-opsinukalkku samaanamaan. tmt-opsinukalum paanopsinukalum ore intronukal pankidunnu, ithu avaa orumichu ullathaanennu sthireekarikkunnu.
sinideriyansil opsinukal.
jellifish, pavizhapputukal, kadal anaemonukal enniva ulppedunna sinideriyayaanu sankeernamaaya annukalulla ettavum adisthaana mrugangal. ropalia dambathikalile jellifish opsinukal muthal gs-proteinukal vare intra sellular camp level uyarthunnu. pavizha oopsinukalkku jikyu-proteinukalilekkum jisi-proteinukalilekkum jodiyakkaan kazhiyum. sinidarians prathyekamaayi upayogikkunna oru upa tharam jii-proteinukalaanu jii. sii-proteinukal. senopsinukalum nesopsinukalum enna rund grouppukalil pedunnathaanu sinidarian opsinukal. senopsinukalil baittariyanum adangiyirikkunnu, athesamayam nesopsinukal sinideriyansil maathramaayi parimithappedutthiyirikku. ennirunnaalum, chila sinidarian opsinukal sailopsinukal, rabopsinukal, baittariyanukalude tetrapsinukal ennivayil pedunnavayaanennu muun padanangal kandethiyittundu.
rabdomeric opsinukal.
rabdomeric opsinukal (rabopsinukal, aarr-opsinukal) jikyu-opsinukal ennum ariyappedunnu, kaaranam avaa jikyu-proteinumaayi jodiyaakkunnu. molaskukalum aarthropodukalum rabopsinukal upayogikkunnu. filogeny, spectral censitivity ennivayude kaaryatthil vyathyasthamaaya moonnu (allengil athil kooduthal) vyathyastha grouppukal upayogichu kasherukkalkk samanamaaya reethiyil aarthropodukal varnnadarsanam nedunnathaayi kaanapedunnu. rabopsin melanopsin kasherukkalil prakadippikkappedunnu, avide ithu syrcadian thaalangale niyanthrikkukayum pupillary reflexine madhyasthamaakkukayum cheiyunnu.
silopsinukalil ninnu vyathyasthamayi, rabopsinukal canonical traansiyant risapttar pottensial ayoon chaanalukalumayi bandhappettirikkunnu; iva oru kosha stharathiludaneelamulla vaidyutha pottensial vyathyaasathilekku nayikkunnu (i.iii. depolarisation).
squid rhodopsinte kristtal ghadana thirichariyunnathu eee groupile athinte pravarthanathekkurichu nammude dhaarana kooduthal mechappeduthaan saadhyathayund.
aarthropodukal avayude vyathyastha netra tharangalil vyathyastha opsinukal upayogikkunnu, ennaal kuranjathu "limulasil" paarshwabhaagatthum samyuktha annukalilum prakadippikkunna opsinukal 99 sathamaanam samaanavum aduthide verpeduthappettathumaayir.
melanopsin.
syrcadian rhythm, pupillary reflex, uyarnna thilakkamulla saahacharyangalil varnna thiruthal ennivayil melanopsin (opn 4) ulppedunnu. phylogeneticalaayi, ithu rabdomeric opsinukalileyum (rabopsinukal, aarr-opsinukal) pravarthanaparamaayum gadanaaparamaayum oru rabopsinileyum amgamaanu, pakshe rabdomerukalil ithu sambhavikkunnilla.
tetrapsinukal.
neuropsinukal, goo-opsinukal, chromopsinukal enniva tetrapsinukalil ulppedunnu. chromopsinukal ezhu upavibhagangalaal nirmmithamaann aarr. jii. aarr-opsinukal, retinokromukal, peropsinukal, varopsinukal, aastropsinukal, nemopsinukal, gloupsinukal.
neuropsinukal.
neuropsinukal yuvayodu sensitiivu aanu, saadhaaranayaayi 380 enm. manushyarudeyum elikaludeyum thalachor, vrishanangal, charmmam, rettina, athupole pakshikalude thalachor, rettina ennividangalilum iva kaanapedunnu. pakshikalilum elikalilum avaa ultraviolet kaazhchaykk madhyasthatha vahikkunnu. avaa jii-proteinukalumaayi jodiyaakkunnu. manushyaril, neuropsin opn5 jeane encode cheiyunnu. manushya rettinayil athinte pravartthanam anjjaathamaanu. mousil, ithu kuranjathu ex vivoyenkilum rettinayilum corneyayilum foto-entraine cheiyunnu.
goo-opsins.
uyarnna kasherukkalil ninnum exidosovanukalil ninnum goo-opsinukal illa. scullop aiyude siliyari photoriseptar koshangalilum basel cordate ampheoxasilum iva kaanapedunnu. ennirunnaalum, "plaattineris dumerily" yill, annukalude rabdomeric photoriseptar koshangalil oru goo-opsin prakadippikkunnu.
aarr. jii. aarr-opsins.
rettina jii protein cuppled resepterukal ennum ariyappedunna rgr-opsinukal rettina pigment epithelium (rpe), mullar cellukal ennivayil prakadippikkunnu. avaa 11-sis-rettinalin pakaram iruttil ellah trans-rettinalukaleyum bandhippikkunnathaanu nallath. aarr. jii. aarr-opsinukal photoisomerasukalaanenna karuthappettirunnuvenkum pakaram avaa retinoid traffic ulppaadanavum niyanthrikkunnu. prathyekichum, avaa ellah trans-rettinail-eesttarukalil ninnum 11-sis-rettinol (11-sis-rettinalinte munnodiyaayi) prakaashatthil ninnu swathanthramaayi uthppaadippikkunnu. ennirunnaalum, ellah trans-rettinail-eesttarukalum aarr. jii. aarr-opsinukal vazhi velichathe aashrayich labhyamaakkiyittund. rgr-opsinukal jii-protein vazhiyo mattethenkilum signalling samvidaanathiloodeyo ithu niyanthrikkunnundo ennu anjjaathamaanu. pashukkal aarr. jii. aarr-opsin piechu-moolyathe aashrayich vyathyastha tharamgadairkhyangalil paramavadhi agiranam cheiyunnu. uyarnna piechil ithu paramavadhi neela (469 enm) prakashavum kuranja piechil paramavadhi yuvi (370 enm) prakashavum agiranam cheiyunnu.
peropsin.
manushyaril "aarr. aarr. echu" jeane encode cheiyunna oru proteinaan vishwal pigment polulla resepteraaya peropsin.
oopsinukal ennu vilikkunna mattu proteinukal.
photoriseptarukale pravartthanam (kazhcha, phototaxis, photoperiodism muthalaayava) ulppede niravadhi vazhikalil tharamthirikkaam.), chromophorinte tharam (rettina, flavin, billin), tanmatra ghadana (thritheeya, kwaattenari), signal outtputtu (phosphorylation, reduction, ooksideshan) muthalaayava.
jii protein-cuppled resepterukalaaya mrugangalude oopsinukalkku purame, oopsinukal ennariyappedunna photoriseptar proteinukalude mattoru groupund. iva microbial oopsin aanu, avaa prokaariyottukalum chila aalgakalum (chaanalrodopsinukalude gadakamaayi) fangasukalum upayogikkunnu, athesamayam mrugangal mrugangalude oopsinukal maathram upayogikkunnu. eee grouppukalkku purathu opsinukalonnum kandethiyitla (udaaharanathinu sasyangalilo placosovanukalilo).
microbial, animal opsinukale yathaakramam type 1, type 2 opsinukal ennum vilikkunnu. rund tharangaleyum oopsinukal ennu vilikkunnu, kaaranam avaa parasparam bandhappettirikkunnave oru kaalath karuthappettirunnu randum ezhu transmembron resepterukalaanennum chromophorayi rettinaye bandhippikkukayum cheiyunnu, ithu avaye photoriseptarukal sensing laittayi maattunnu. ennirunnaalum, rund tharangalum sequence thalatthil bandhappettirikkunnil.
vaasthavathil, mrugangalum myrobial opsinukalum thammilulla sequence aidantitti kramarahithamaya aakasmikathayiloode kanakkaakkaavunnathinekkaal valuthalla. ennirunnaalum, sameepa varshangalil "deep filogeny" kku prathyekamaayi puthiya reethikal vikasippicheduthittundu. thalphalamaayi, niravadhi padanangal iva randum thammilulla saadhyamaaya phylogenetic bandhatthinte thelivukal kandethiyittundu. ennirunnaalum, sookshumajeevikaludeyum mrugangaludeyum opsinukalude avasaanathe pothuvaaya poorvvikan thanne neriya sensitiivu aanennu ithinarthamilla ellah mrugangaludeyum opsinukalum (jeane duplication vyathiyaanavum vazhi) valia jii-protein cuppled risapttar (gpcr) jeane kudumbatthinte charithratthil vaiki uyarnnuvannu, ithu aadyakaala mrugangalil ninnu sasyangal, fangasukal, choanflaajelettukal, sponchukal ennivayude vyathichalanathin sesham uyarnnuvannu. rettina kromophore eee valia jeane kudumbatthinte oopsin shaakhayil maathram kaanapedunnu, athaayath mattevideyengilum athinte saannidhyam homologialla, marichu kanverjant parinaamatthe prathinidheekarikkunnu. microbial rhodopsinukal, kramam anusarichu, gpcr kudumbangalil ninnu valare vyathyasthamaan. oru sidhaanthamanusarichu, microbial, animal opsinukal randum jii protein-cuppled risapttar (gpcr), ayoon-translocating microbial rhodopsin (emr), mattu ezhu enniva ulppedunna nirddishta claid aaya transporter-opsin-jii protein-cuppled risapttar (toji) suuppar faamiliyil ulppedunnu.
mikka microbial opsinukalum sariyaaya resepterukalkku pakaram ayoon chaanalukalo pambukaloo aanu, avaa oru jii proteinumaayi bandhippikkunnilla. archaea, bacteria, yookkariyotta ennee jeevajaalangalude moonnu mekhalakalilum microbal oopsinukal kaanapedunnu. yookkariyottayil, microbial opsinukal pradhaanamaayum green aalgakal polulla ekakosha jeevikalilum fangasukalilum kaanapedunnu. mikka sankeernamaaya multisellular yookkaariyottukalilum, microbial oopsinukalkku pakaram sasyangalile cryptocrom, faittokrom thudangiya prakasha sensitiivu thanmaathrakalum mrugangalile mrugangalude oopsinukalum sthaapichittundu.
microbial opsinukale palappozhum ariyappedunnathu thanmaathrayude rhodopsin roopamaanu, i.iii.rhodopsin (vishalamaaya ardhathil) = oopsin + kromophore. prottoon pambukal bacteriorohodopsin (bii. aarr.), santorhodopsin (ex. aarr.), chloride pump halorhodopsin (echu. aarr.), photosensarukal censory rhodopsin aii (ezz. aarr. aii), censory rhodopsin II (ezz. aarr. aii), proteorhodopsin (pii. aarr.), neurospora oopsin aii (noppi), chlamydomonas censory rhodopsin eiii (sii. ezz. aarr. eiii), chlamydomonas censory rhodopsin bii (sii. ezz. aarr. bii), chanelrohodopsin (sii. aarr. aarr), archairhodopsin (aarch) ennivayaanu palatharam microbial oopsinukal.
proteo, bacteriorohodopsin thudangiya niravadhi microbal oopsinukal vividha bacteria grouppukal chlorophyll adisthaanamaakkiyulla paatha upayogichu upapachayam nadathunnathinaayi prakaashatthil ninnu oorjjam shekharikkaan upayogikkunnu. athinupurame, halobacteryayude halorohodopsinukalum chila aalgakalude chaanalrodopsinukalum, udha.jii. volvox, avaye laitt-gated ayoon chaanalukalayi sevikkunnu, mattullavaykkoppam phototactic aavasyangalkkum. jii proteinukalumaayi bandhamillaattha transducer membran-embaded proteinukalumaayi samvadichukondu phototactic prathikaranatthe prerippikkunna halobacteriayil censory rhodopsinukal nilavilundu.
neuronal pravartthanam oonaakkaano oof cheyyaano optogeneticsil microbal opsinukal (chanelrodopsin, halorohodopsin, archairhodopsin enniva polullava) upayogikkunnu. mrugangalude oopsinukalekkaal vegathil prathikarikkunnathinal neuronal pravartthanam uyarnna aavruthiyilaanu modulate cheyyendathenkil microbal oopsinukalkkaanu munganana. kaaranam, microbal oopsinukal ayoon chaanalukalo prottoon/ayoon pambukaloo aanu, athinaal avaa prakasham vazhi nerittu sajeevamaakunnu, athesamayam mrugangalude oopsinukal jii-proteinukale sajeevamaakkunnu, avaa ayoon chaanalukal thurakkunnathinu metabolittukal ulppaadippikkunna ifakttar ensaimukale sajeevamaakkunnu.

