Maqaalku waa qayb ka mid ah mowduuca cilmi-baarista "Tiknoolajiyada bioremediation-ka ee horumarsan iyo hababka dib-u-warshadaynta isku-dhafka dabiiciga ah ee synthetic organic (SOC)". Eeg dhammaan 14-ka maqaal
Hydrocarbons-ka udgoonka polycyclic-ga ee miisaanka yar (PAHs) sida naphthalene iyo naphthalenes-ka la beddelay (methylnaphthalene, naphthoic acid, 1-naphthyl-N-methylcarbamate, iwm.) si weyn ayaa loogu isticmaalaa warshado kala duwan waana sun genotoxic ah, isbeddel hidde-side ah iyo/ama kansar u leh noolaha. Isku-darkaan dabiiciga ah ee macmalka ah (SOCs) ama xenobiotics-ka waxaa loo arkaa inay yihiin wasakheeyayaasha mudnaanta leh waxayna khatar weyn ku yihiin deegaanka adduunka iyo caafimaadka dadweynaha. Xoogga hawlaha aadanaha (tusaale ahaan gaaska dhuxusha, sifeynta saliidda, qiiqa baabuurta iyo codsiyada beeraha) waxay go'aamisaa isku-darka, qaddarka iyo gaadiidka isku-dhafkan meel walba yaal iyo kuwa joogtada ah. Marka lagu daro hababka daaweynta/ka saarista jirka iyo kiimikada, teknoolojiyada cagaaran iyo kuwa deegaanka u fiican sida bioremediation, kuwaas oo adeegsada noolaha yaryar ee awood u leh inay si buuxda u burburiyaan POC-yada ama u beddelaan alaabada aan sunta ahayn, ayaa soo baxay beddel ammaan ah, kharash-ool ah oo rajo leh. Noocyo kala duwan oo bakteeriya ah oo ka tirsan phyla Proteobacteria (Pseudomonas, Pseudomonas, Comamonas, Burkholderia, iyo Neosphingobacterium), Firmicutes (Bacillus iyo Paenibacillus), iyo Actinobacteria (Rhodococcus iyo Arthrobacter) oo ku jira microbiota ciidda ayaa muujiyay awoodda ay u leeyihiin inay hoos u dhigaan isku-dhafka kala duwan ee dabiiciga ah. Daraasadaha dheef-shiid kiimikaadka, genomics, iyo falanqaynta metagenomic waxay naga caawinayaan inaan fahanno kakanaanta catabolic iyo kala duwanaanshaha ku jira qaababkan nolosha fudud, kuwaas oo si dheeraad ah loogu isticmaali karo burburka noolaha ee hufan. Jiritaanka muddada dheer ee PAHs waxay keentay soo ifbaxa noocyada cusub ee burburka iyada oo loo marayo wareejinta hidda-wadaha toosan iyadoo la adeegsanayo walxaha hidde-sideyaasha sida plasmids, transposons, bacteriophages, jasiiradaha genomic, iyo walxaha isku-dhafan ee isku-dhafan. Bayoolajiyada nidaamyada iyo injineernimada hidde-sideyaasha ee go'doominta gaarka ah ama bulshooyinka moodeelka (consortia) waxay awood u siin kartaa dib-u-habeyn dhammaystiran, degdeg ah oo hufan oo PAH-yadan ah iyada oo loo marayo saameyn isku-dhafan. Dib-u-eegistan, waxaan diiradda saareynaa waddooyinka kala duwan ee dheef-shiid kiimikaadka iyo kala duwanaanshaha, halabuurka hidde-sidaha iyo kala duwanaanshaha, iyo jawaabaha/habaynta unugyada ee naphthalene iyo bakteeriyada naphthalene-ka ee beddelka ah. Tani waxay bixin doontaa macluumaad deegaan oo loogu talagalay codsiga goobta iyo hagaajinta cadaadiska si loo helo daaweyn wax ku ool ah oo ku saabsan biomediation-ka.
Horumar degdeg ah oo warshadaha (petrochemicals, beeraha, daawooyinka, midabaynta dharka, qurxinta, iwm.) ayaa gacan ka geystay barwaaqo dhaqaale oo caalami ah iyo heerarka nolosha oo la hagaajiyay. Horumarkan baaxadda leh wuxuu keenay soo saarista tiro badan oo iskudhisyo dabiici ah oo macmal ah (SOCs), kuwaas oo loo isticmaalo in lagu soo saaro alaabooyin kala duwan. Iskudhisyadan shisheeye ama SOCs waxaa ka mid ah hydrocarbons aromatic polycyclic (PAHs), sunta cayayaanka, sunta cayayaanka, balaastikada, rinjiyeyaasha, daawooyinka, organophosphates, kuwa dabka celiya, dareerayaasha dabiiciga ah ee isbeddela, iwm. Waxaa lagu sii daayaa jawiga, nidaamyada deegaanka biyaha iyo dhulka halkaas oo ay ku leeyihiin saameyn dhinacyo badan leh, taasoo keenta saameyn waxyeello leh oo ku timaadda noocyada kala duwan ee noolaha iyada oo loo marayo isbeddelka sifooyinka fiisikokemikaalka iyo qaab-dhismeedka bulshada (Petrie et al., 2015; Bernhardt et al., 2017; Sarkar et al., 2020). Wasakhowga udgoonka badan ayaa saameyn xooggan oo burbur leh ku leh deegaanno badan oo aan isbeddelin/meelo kala duwan oo noolaha ah (tusaale ahaan xeebaha shacaabka, barafka Arctic/Antarctic, harooyinka buuraha dhaadheer, dhoobada badda qoto dheer, iwm.) (Jones 2010; Beyer et al. 2020; Nordborg et al. 2020). Daraasado juqraafiyeed oo dhowaan la sameeyay ayaa muujiyay in dhigista walxaha dabiiciga ah ee macmal ah (tusaale ahaan wasakhowga udgoonka) iyo waxyaabaha ka soo baxa dusha sare ee dhismayaasha macmalka ah (deegaanka la dhisay) (tusaale goobaha dhaxalka dhaqanka iyo taallooyinka laga sameeyay granite, dhagaxa, qoryaha iyo birta) ay dardar gelinayaan burburkooda (Gadd 2017; Liu et al. 2018). Hawlaha aadanuhu waxay sii xoojin karaan oo ay ka sii dari karaan burburka bayoolojiga ee taallooyinka iyo dhismayaasha iyada oo loo marayo wasakhowga hawada iyo isbeddelka cimilada (Liu et al. 2020). Wasakhowgan dabiiciga ah waxay la falgalaan uumiga biyaha ee jawiga waxayna ku degaan qaab-dhismeedka, taasoo keenta burbur jireed iyo kiimikadeed oo walxaha ah. Bayooloji-xumida waxaa si weyn loogu aqoonsadaa isbeddello aan loo baahnayn oo ku yimaada muuqaalka iyo sifooyinka walxaha ay sababaan noolaha nool ee saameeya ilaalintooda (Pochon iyo Jaton, 1967). Ficilka microbial dheeraad ah (dheef-shiid kiimikaad) ee isku-dhafkan ayaa yareyn kara hufnaanta qaab-dhismeedka, waxtarka ilaalinta iyo qiimaha dhaqanka (Gadd, 2017; Liu et al., 2018). Dhanka kale, xaaladaha qaarkood, la qabsiga microbial-ka iyo ka jawaab celinta qaab-dhismeedkan ayaa la ogaaday inay faa'iido leedahay maadaama ay sameeyaan biofilms iyo qolof kale oo ilaalin ah oo yareeya heerka qudhunka/burburka (Martino, 2016). Sidaa darteed, horumarinta istaraatiijiyado ilaalin waara oo wax ku ool ah oo loogu talagalay taallooyinka dhagaxa, birta iyo alwaaxa waxay u baahan tahay faham qoto dheer oo ku saabsan hababka muhiimka ah ee ku lug leh habkan. Marka la barbardhigo hababka dabiiciga ah (geedi socodka juqraafiyeed, dabka kaynta, qaraxyada foolkaanaha, falcelinta dhirta iyo bakteeriyada), hawlaha aadanaha waxay keenaan sii deynta tiro badan oo ah hydrocarbons-ka udgoonka polycyclic (PAHs) iyo kaarboon kale oo organic ah (OC) oo ku jira nidaamyada deegaanka. PAH-yo badan oo loo isticmaalo beeraha (cayayaanka iyo sunta cayayaanka sida DDT, atrazine, carbaryl, pentachlorophenol, iwm.), warshadaha (saliid cayriin ah, wasakhda saliidda/qashinka, balaastikada laga helo batroolka, PCB-yada, balaastikada, saabuunta, jeermiska dila, sunta dila, udgoonka, udgoonka iyo ilaaliyayaasha), alaabada daryeelka shakhsi ahaaneed (daawada qorraxda, jeermiska dila, sunta cayayaanka iyo musk-yada polycyclic) iyo rasaasta (qaraxyada sida 2,4,6-TNT) waa xenobiotics suurtagal ah oo saameyn kara caafimaadka meeraha (Srogi, 2007; Vamsee-Krishna iyo Phale, 2008; Petrie et al., 2015). Liiskan waxaa loo ballaarin karaa si loogu daro isku-dhafka laga helo batroolka (saliidaha shidaalka, saliidaha, asfaltenes), bioplastics-ka miisaanka sare leh ee molecular, iyo dareere ionic ah (Amde et al., 2015). Shaxda 1 waxay liis gareysaa wasakheeyayaasha udgoonka kala duwan iyo codsiyadooda warshado kala duwan. Sannadihii ugu dambeeyay, qiiqa aadanaha ee isku-dhafka dabiiciga ah ee isbeddela, iyo sidoo kale kaarboon laba ogsaydh iyo gaasaska kale ee aqalka dhirta lagu koriyo, ayaa bilaabay inay kordhaan (Dvorak et al., 2017). Si kastaba ha ahaatee, saameynta aadanaha ayaa si weyn uga badan kuwa dabiiciga ah. Intaa waxaa dheer, waxaan ogaanay in tiro ka mid ah SOCs ay ku sii jiraan deegaanno badan oo deegaan ah waxaana loo aqoonsaday inay yihiin wasakhowga soo baxaya oo leh saameyn xun oo ku leh noolaha (Jaantuska 1). Hay'adaha deegaanka sida Hay'adda Ilaalinta Deegaanka ee Mareykanka (USEPA) waxay ku dareen wasakhowgan badan liiskooda mudnaanta leh sababtoo ah sifooyinkooda sunta ah, sunta ah, isbeddelka unugyada, iyo kansarka. Sidaa darteed, waxaa loo baahan yahay xeerar adag oo qashin-qubka ah iyo istaraatiijiyado wax ku ool ah oo loogu talagalay daaweynta/ka saarista nidaamyada deegaanka ee wasakhaysan. Habab kala duwan oo daaweynta jirka iyo kiimikada ah sida pyrolysis, daaweynta kulaylka oksaydhka, hawo-mareenka hawada, buuxinta qashinka, gubista, iwm. waa kuwo aan waxtar lahayn oo qaali ah waxayna dhaliyaan daxal, sun ah oo ay adag tahay in la daweeyo waxyaabaha ka soo baxa. Iyadoo ay sii kordhayso wacyigelinta deegaanka ee caalamiga ah, noolaha yaryar ee awood u leh inay hoos u dhigaan wasakhowgan iyo waxyaabaha ay ka soo saaraan (sida halogenated, nitro, alkyl iyo/ama methyl) ayaa soo jiidanaya dareen sii kordhaya (Fennell et al., 2004; Haritash iyo Kaushik, 2009; Phale et al., 2020; Sarkar et al., 2020; Schwanemann et al., 2020). Isticmaalka noolaha yaryar ee asalka ah ee kaligood ama dhaqamada isku dhafan (gumeysiyada) si looga saaro wasakhowga udgoonka leh waxay leedahay faa'iidooyin marka la eego badbaadada deegaanka, kharashka, waxtarka, waxtarka, iyo waaritaanka. Cilmi-baarayaashu waxay sidoo kale sahaminayaan isku-darka hababka microbial-ka iyo hababka redox-ka elektaroonigga ah, gaar ahaan nidaamyada bioelectrochemical (BES), oo ah tignoolajiyad rajo leh oo loogu talagalay daaweynta/ka saarista wasakhowga (Huang et al., 2011). Tiknoolajiyadda BES waxay soo jiidatay dareen sii kordhaya sababtoo ah waxtarkeeda sare, kharash yar, badbaadada deegaanka, hawlgalka heerkulka qolka, agabka bayoolojiga la jaanqaadaya, iyo awoodda lagu soo ceshan karo wax soo saarka qiimaha leh (tusaale ahaan, korontada, shidaalka, iyo kiimikooyinka) (Pant et al., 2012; Nazari et al., 2020). Soo ifbixidda taxanaha hidde-sideyaasha ee wax-soo-saarka sare leh iyo aaladaha/hababka omics waxay bixisay macluumaad badan oo cusub oo ku saabsan nidaaminta hidde-sidaha, proteomics, iyo fluxomics ee falcelinta microorganisms-ka kala duwan ee burburiya. Isku-darka qalabkan iyo bayoolajiyada nidaamyada ayaa sii xoojisay fahamkeenna xulashada iyo hagaajinta waddooyinka catabolic ee bartilmaameedka ah ee microorganisms (tusaale ahaan, naqshadeynta dheef-shiid kiimikaadka) si loo gaaro burburka biograderation oo hufan oo wax ku ool ah. Si loo qaabeeyo istaraatiijiyado bioremediation oo wax ku ool ah iyadoo la adeegsanayo microorganisms-ka musharixiinta ah ee ku habboon, waxaan u baahanahay inaan fahanno awoodda biochemical, kala duwanaanshaha dheef-shiid kiimikaadka, halabuurka hidde-sidaha, iyo cilmiga deegaanka (autoecology/synecology) ee microorganisms-ka.
Jaantuska 1. Ilaha iyo waddooyinka PAH-yada molecular-ka hooseeya iyada oo loo marayo deegaanno kala duwan iyo arrimo kala duwan oo saameeya biota. Khadadka laalaaban waxay matalaan isdhexgalka ka dhexeeya walxaha deegaanka.
Dib-u-eegistan, waxaan isku daynay inaan soo koobno ​​xogta ku saabsan burburka PAH-yada fudud sida naphthalene iyo naphthalenes-ka la beddelay ee ay sameeyeen noocyo kala duwan oo bakteeriya ah oo daboolaya waddooyinka dheef-shiid kiimikaadka iyo kala duwanaanshaha, enzymes-ka ku lug leh burburka, halabuurka/cutubka hidde-sidaha iyo kala duwanaanshaha, jawaabaha unugyada iyo dhinacyada kala duwan ee hagaajinta biomediation-ka. Fahmidda heerarka bayoolaji iyo molecular-ka ayaa kaa caawin doonta aqoonsashada noocyada martida ku habboon iyo injineerintooda hidde-sidaha ee dheeraadka ah si loo helo daaweyn wax ku ool ah oo ku saabsan wasakhowga mudnaanta leh. Tani waxay kaa caawin doontaa horumarinta istaraatiijiyado lagu aasaasayo isku-xidhka bakteeriyada goobta gaarka u ah si loo helo daaweyn biomediation oo wax ku ool ah.
Joogitaanka tiro badan oo iskudhisyo udgoon oo sun ah oo khatar ah (ku qancinaya xeerka Huckel 4n + 2π elektaroono, n = 1, 2, 3, ...) waxay khatar weyn ku tahay warbaahinta deegaanka ee kala duwan sida hawada, ciidda, wasakhda, iyo dusha sare iyo biyaha dhulka hoostiisa (Puglisi et al., 2007). Iskudhisyadani waxay leeyihiin giraangiraha benzene ee hal-hal ah (monocyclic) ama giraangiraha benzene ee badan (polycyclic) oo ku habaysan qaab toosan, xagal ama kooxeed waxayna muujiyaan xasillooni (xasilloon/xasilloonnimo) deegaanka sababtoo ah tamar sare oo taban iyo firfircooni la'aan (aan firfircoonayn), taas oo lagu sharxi karo cabsidooda biyaha iyo xaaladdooda oo yaraatay. Marka giraanta udgoonka lagu beddelo kooxaha methyl (-CH3), carboxyl (-COOH), hydroxyl (-OH), ama sulfonate (-HSO3), waxay noqotaa mid deggan, waxay leedahay xiriir xooggan oo u dhexeeya macromolecules, waana mid ku urursan nidaamyada bayoolojiga (Seo et al., 2009; Phale et al., 2020). Qaar ka mid ah hydrocarbons-ka udgoonka polycyclic ee miisaanka yar (LMWAHs), sida naphthalene iyo waxyaabaha ka soo baxa [methylnaphthalene, naphthoic acid, naphthalenesulfonate, iyo 1-naphthyl N-methylcarbamate (carbaryl)], ayaa lagu daray liiska sunta dabiiciga ah ee mudnaanta leh ee Hay'adda Ilaalinta Deegaanka ee Mareykanka iyagoo ah sunta hidde-sideyaasha, isbeddelka, iyo/ama kansarka keena (Cerniglia, 1984). Sii deynta fasalkan NM-PAHs ee deegaanka waxay keeni kartaa isku-uruurinta walxahan dhammaan heerarka silsiladda cuntada, taasoo saameyn ku yeelanaysa caafimaadka nidaamyada deegaanka (Binkova et al., 2000; Srogi, 2007; Quinn et al., 2009).
Ilaha iyo waddooyinka PAHs ee loo maro biota waxay ugu horreyn ku yimaadaan socdaalka iyo isdhexgalka qaybaha kala duwan ee deegaanka sida carrada, biyaha dhulka hoostiisa, biyaha dusha sare, dalagyada iyo jawiga (Arey iyo Atkinson, 2003). Jaantuska 1aad wuxuu muujinayaa isdhexgalka iyo qaybinta PAH-yada miisaanka molecular-ka ee kala duwan ee nidaamyada deegaanka iyo dariiqyadooda u soo-gaadhista biota/aadanaha. PAH-yada waxaa lagu shubaa dusha sare natiijada wasakhowga hawada iyo iyada oo loo marayo socdaalka (qulqulka) qiiqa baabuurta, gaasaska qiiqa warshadaha (gaasaynta dhuxusha, gubashada iyo wax soo saarka kookaha) iyo dhigista. Hawlaha warshadaha sida soo saarista dharka macmal ah, midabaynta iyo rinjiga; ilaalinta alwaaxa; farsamaynta caagga; hawlaha wax soo saarka sibidhka; wax soo saarka cayayaanka; iyo codsiyada beeraha waa ilo muhiim ah oo PAH-yada ah ee nidaamyada dhulka iyo biyaha (Bamforth iyo Singleton, 2005; Wick et al., 2011). Daraasaduhu waxay muujiyeen in carrada ku taal magaalooyinka iyo magaalooyinka, meelaha u dhow waddooyinka waaweyn, iyo magaalooyinka waaweyn ay u nugul yihiin hydrocarbons-ka udgoon ee polycyclic (PAHs) sababtoo ah qiiqa ka imanaya warshadaha korontada, kuleylinta guryaha, culeyska gaadiidka hawada iyo waddooyinka, iyo hawlaha dhismaha (Suman et al., 2016). (2008) waxay muujiyeen in PAH-yada ku jira carrada u dhow waddooyinka ee New Orleans, Louisiana, USA ay ahaayeen kuwo aad u sarreeya ilaa 7189 μg/kg, halka meelaha bannaan, ay ahaayeen oo keliya 2404 μg/kg. Sidoo kale, heerarka PAH ee u sarreeya ilaa 300 μg/kg ayaa laga soo sheegay meelaha u dhow goobaha gaaska dhuxusha ee dhowr magaalo oo Mareykan ah (Kanaly iyo Harayama, 2000; Bamforth iyo Singleton, 2005). Carrada ka timid magaalooyin kala duwan oo Hindiya ah sida Delhi (Sharma et al., 2008), Agra (Dubey et al., 2014), Mumbai (Kulkarni iyo Venkataraman, 2000) iyo Visakhapatnam (Kulkarni et al., 2014) ayaa la sheegay inay ku jiraan xaddi badan oo PAH ah. Isku-dhafka udgoonka ayaa si fudud loogu nuugaa walxaha ciidda, walxaha dabiiciga ah iyo macdanta dhoobada, sidaas darteedna waxay noqdaan meelo waaweyn oo kaarboon ah oo ku jira nidaamyada deegaanka (Srogi, 2007; Peng et al., 2008). Ilaha ugu waaweyn ee PAHs ee nidaamyada deegaanka biyaha waa roobab (roobab qoyan/qalalan iyo uumiga biyaha), qulqulka magaalooyinka, dheecaanka biyaha wasakhda ah, dib u buuxinta biyaha dhulka hoostiisa iwm. (Srogi, 2007). Waxaa lagu qiyaasaa in qiyaastii 80% PAHs ee nidaamyada deegaanka badda ay ka yimaadaan roobabka, faleebada, iyo daadinta qashinka (Motelay-Massei et al., 2006; Srogi, 2007). Xoogagga sare ee PAH-yada ee ku jira biyaha dusha sare ama daadashada ka timaadda goobaha qashinka adag ayaa ugu dambeyntii ku daadanaya biyaha dhulka hoostiisa, taasoo khatar weyn ku ah caafimaadka dadweynaha maadaama in ka badan 70% dadka ku nool Koonfurta iyo Koonfur-bari Aasiya ay cabbaan biyaha dhulka hoostiisa (Duttagupta et al., 2019). Daraasad dhowaan ay sameeyeen Duttagupta et al. (2020) oo ka mid ah falanqaynta wabiga (32) iyo biyaha dhulka hoostiisa (235) ee ka socota Galbeedka Bengal, Hindiya, ayaa lagu ogaaday in qiyaastii 53% dadka deggan magaalooyinka iyo 44% dadka deggan miyiga (wadarta 20 milyan oo qof oo deggan) ay la kulmi karaan naphthalene (4.9–10.6 μg/L) iyo waxyaabaha ka soo baxa. Qaababka kala duwan ee isticmaalka dhulka iyo soo saarista biyaha dhulka hoostiisa ayaa loo arkaa inay yihiin arrimaha ugu muhiimsan ee xakameynaya gaadiidka toosan (ujeedka) ee PAH-yada miisaanka molecular-ka hooseeya ee ku jira dhulka hoostiisa. Biyaha daadanaya ee beeraha, biyaha wasakhda ah ee degmada iyo warshadaha, iyo qashinka adag/daadinta qashinka adag ayaa la ogaaday inay saameeyeen PAH-yada ku jira wabiyada iyo wasakhda dhulka hoostiisa. Roobabka hawada ayaa sii xumeeya wasakhda PAH. Xoogagga sare ee PAHs iyo waxyaabaha ka soo baxa alkyl (51 guud ahaan) ayaa laga soo sheegay webiyada/biyo-mareennada adduunka oo dhan, sida Webiga Fraser, Webiga Louan, Webiga Denso, Webiga Missouri, Webiga Anacostia, Webiga Ebro, iyo Webiga Delaware (Yunker et al., 2002; Motelay-Massei et al., 2006; Li et al., 2010; Amoako et al., 2011; Kim et al., 2018). Haraadiga dooxada webiga Ganges, naphthalene iyo phenanthrene ayaa la ogaaday inay yihiin kuwa ugu muhiimsan (laga helay 70% muunadaha) (Duttagupta et al., 2019). Intaa waxaa dheer, daraasaduhu waxay muujiyeen in koloriin ku daridda biyaha la cabbo ay horseedi karto sameynta PAHs sun ah oo oksijiin ku jira oo koloriin ku jira (Manoli iyo Samara, 1999). PAH-yadu waxay ku ururaan badarka, miraha iyo khudaarta natiijada ka dhalata nuugista dhirta carrada wasakhaysan, biyaha dhulka hoostiisa iyo roobka (Fismes et al., 2002). Nooleyaal badan oo biyaha ku jira sida kalluunka, mussels, clams iyo shrimp ayaa ku sumoobay PAHs iyada oo loo marayo isticmaalka cuntada wasakhaysan iyo biyaha badda, iyo sidoo kale iyada oo loo marayo unugyada iyo maqaarka (Mackay iyo Fraser, 2000). Hababka karinta/farsamaynta sida dubista, dubista, sigaarka, shiilinta, qalajinta, dubista iyo karinta dhuxusha ayaa sidoo kale horseedi kara xaddi badan oo PAHs ah oo ku jira cuntada. Tani waxay inta badan ku xiran tahay doorashada walxaha sigaarka lagu cabo, maaddada hydrocarbon-ka phenolic/aromatic, habka karinta, nooca kuleyliyaha, qoyaanka, sahayda oksijiinta iyo heerkulka gubashada (Guillén et al., 2000; Gomes et al., 2013). Hydrocarbons-ka udgoon ee polycyclic (PAHs) ayaa sidoo kale laga helay caanaha iyadoo ay ku jiraan xaddi kala duwan (0.75–2.1 mg/L) (Girelli et al., 2014). Ururinta PAH-yadan cuntada waxay sidoo kale ku xiran tahay sifooyinka fiisikokemikaalka cuntada, halka saameyntooda sunta ah ay la xiriirto hawlaha jirka, dhaqdhaqaaqa dheef-shiid kiimikaadka, nuugista, qaybinta iyo qaybinta jirka (Mechini et al., 2011).
Sunta iyo saamaynta waxyeelada leh ee polycyclic aromatic hydrocarbons (PAHs) ayaa la yaqaanay muddo dheer (Cherniglia, 1984). Hydrocarbons-ka aromatic polycyclic miisaanka hooseeya (LMW-PAHs) (laba ilaa saddex giraan) waxay si isku mid ah ugu xidhmi karaan macromolecules kala duwan sida DNA, RNA iyo borotiinno waana kansar keena (Santarelli et al., 2008). Sababtoo ah dabeecadooda hydrophobic, waxaa kala sooca xuubka dufanka. Aadanaha, cytochrome P450 monooxygenases waxay oxidiyaan PAHs ilaa epoksidyada, kuwaas oo qaarkood aad u falceliyaan (tusaale ahaan, baediol epoxide) waxayna horseedi karaan isbeddelka unugyada caadiga ah una beddelaan kuwo halis ah (Marston et al., 2001). Intaa waxaa dheer, wax soo saarka isbeddelka ee PAHs sida quinones, phenols, epoksidyada, diols, iwm. ayaa ka sun badan isku-dhafka waalidka. Qaar ka mid ah PAH-yada iyo dhexdhexaadintooda dheef-shiid kiimikaadka waxay saameyn karaan hoormoonada iyo enzymes-ka kala duwan ee dheef-shiid kiimikaadka, taasoo si xun u saameynaysa koritaanka, habdhiska dhexe ee neerfaha, habdhiska taranka iyo difaaca jirka (Swetha iyo Phale, 2005; Vamsee-Krishna et al., 2006; Oostingh et al., 2008). Soo-gaadhista muddada-gaaban ee PAH-yada miisaanka yar ee molecular-ka ayaa la soo sheegay inay sababaan shaqada sambabada oo xumaata iyo xinjirowga dhiigga ee neefta qaba iyo inay kordhiyaan halista kansarka maqaarka, sambabada, kaadiheysta iyo caloosha iyo mindhicirka (Olsson et al., 2010; Diggs et al., 2011). Daraasadaha xayawaanku waxay sidoo kale muujiyeen in soo-gaadhista PAH ay yeelan karto saameyn xun oo ku saabsan shaqada taranka iyo horumarka waxayna sababi kartaa cataracts, dhaawac kelyaha iyo beerka ah, iyo cagaarshow. Badeecooyin kala duwan oo PAH ah oo isbeddel ah sida diols, epoxides, quinones iyo xagjirrada xorta ah (cations) ayaa la muujiyay inay sameeyaan DNA-da. Duubabka xasilloon ayaa la muujiyay inay beddelaan mishiinnada taranka DNA-da, halka duubyada aan degganayn ay ka saari karaan DNA-da (badanaa adenine mararka qaarna guanine); labaduba waxay abuuri karaan khaladaad horseeda isbeddello (Schweigert et al. 2001). Intaa waxaa dheer, quinones (benzo-/pan-) waxay abuuri karaan noocyada oksijiinta ee falcelinta leh (ROS), taasoo keenta dhaawac dilaa ah oo soo gaara DNA-da iyo macromolecules-ka kale, taasoo saameynaysa shaqada/nololaanshaha unugyada (Ewa iyo Danuta 2017). Soo-gaadhista joogtada ah ee heerarka hoose ee pyrene, biphenyl iyo naphthalene ayaa la sheegay inay sababaan kansar xayawaanka tijaabada ah (Diggs et al. 2012). Sababtoo ah suntooda dilaaga ah, nadiifinta/ka saarista PAH-yadan meelaha ay saameeyeen/wasakhaysan waa mudnaan.
Habab kala duwan oo jireed iyo kiimiko ah ayaa loo isticmaalay in lagu saaro PAH-yada goobaha/deegaanka wasakhaysan. Habab sida gubista, ka saarista koloriin, oksaydhka UV, hagaajinta, iyo soo saarista dareeraha waxay leeyihiin khasaarooyin badan, oo ay ku jiraan sameynta wax soo saarka sunta ah, kakanaanta habka, arrimaha badbaadada iyo sharciyeynta, hufnaan hooseeya, iyo kharash sare. Si kastaba ha ahaatee, bayoolajiyeynta microbial (oo loo yaqaan bioremediation) waa hab kale oo rajo leh oo ku lug leh isticmaalka noolaha yaryar qaab dhaqameedyo saafi ah ama deegaanno. Marka la barbardhigo hababka jireed iyo kiimikada, habkani waa mid deegaan ahaan u wanaagsan, aan duullaan ahayn, kharash-ool ah, oo waara. Bioremediation waxaa lagu samayn karaa goobta ay dhibaatadu saameysey (in situ) ama goob si gaar ah loo diyaariyey (ex situ) sidaas darteedna waxaa loo arkaa hab dib-u-habayn waara oo ka badan hababka dhaqameed ee jireed iyo kiimikada (Juhasz iyo Naidu, 2000; Andreoni iyo Gianfreda, 2007; Megharaj et al., 2011; Phale et al., 2020; Sarkar et al., 2020).
Fahmidda tallaabooyinka dheef-shiid kiimikaadka microbial ee ku lug leh burburka wasakhowga udgoonku waxay leedahay saameyn weyn oo cilmiyeed iyo dhaqaale oo ku saabsan waaritaanka deegaanka iyo deegaanka. Qiyaastii 2.1×1018 garaam oo kaarboon ah (C) ayaa lagu kaydiyaa wasakhda iyo isku-dhafka dabiiciga ah (tusaale ahaan, saliidda, gaaska dabiiciga ah, iyo dhuxusha, tusaale ahaan, shidaalka fosil) adduunka oo dhan, taasoo gacan weyn ka geysaneysa wareegga kaarboonka adduunka. Si kastaba ha ahaatee, warshadaynta degdegga ah, soo saarista shidaalka fosil, iyo hawlaha aadanaha ayaa baabi'inaya kaydadka kaarboonka lithospheric-ka ah, iyagoo sii deynaya qiyaastii 5.5×1015 g oo kaarboon dabiici ah (sida wasakhowga) jawiga sannad kasta (Gonzalez-Gaya et al., 2019). Inta badan kaarboonkan dabiiciga ah wuxuu galaa nidaamyada dhulka iyo badda iyada oo loo marayo faleebo, gaadiid, iyo biyo-bax. Intaa waxaa dheer, wasakhowga cusub ee isku-dhafka ah ee laga helo shidaalka fosil, sida balaastikada, balaastikada iyo xasiliyeyaasha balaastikada (phthalates iyo isomers-kooda), waxay si xun u wasakheeyaan nidaamyada badda, carrada iyo biyaha iyo bayoolajiyadooda, taasoo sii xumeyneysa khataraha cimilada adduunka. Noocyo kala duwan oo ah microplastics, nanoplastics, jajabyo balaastig ah iyo walxo monomer sun ah oo laga soo qaatay polyethylene terephthalate (PET) ayaa ku ururay Badweynta Baasifigga ee u dhaxaysa Waqooyiga Ameerika iyo Koonfur-bari Aasiya, iyagoo sameeyay "Baloogga Qashinka Baasifigga Weyn", taasoo waxyeellaysa nolosha badda (Newell et al., 2020). Daraasado cilmiyeed ayaa xaqiijiyay inaysan suurtagal ahayn in laga saaro wasakhowga/qashinkaas habab jireed ama kiimiko ah. Xaaladdan, noolaha ugu waxtarka badan waa kuwa awood u leh inay si oksaydh ah u dheefshiido wasakhowga galay kaarboon laba ogsaydh, tamar kiimiko iyo waxyaabo kale oo aan sun ahayn oo ugu dambeyntii gala hababka kale ee wareegga nafaqada (H, O, N, S, P, Fe, iwm.). Sidaa darteed, fahamka deegaanka microbial ecophysiology ee macdanta wasakhowga udgoon iyo xakamaynta deegaanka ayaa muhiim u ah qiimeynta wareegga kaarboonka microbial, miisaaniyadda kaarboonka saafiga ah iyo khataraha cimilada mustaqbalka. Iyadoo la tixgelinayo baahida degdegga ah ee looga saarayo isku-dhafka noocaas ah deegaanka, warshado kala duwan oo deegaanka ah oo diiradda saaraya teknoolojiyada nadiifka ah ayaa soo baxay. Haddii kale, qiimaynta kiimikada qashinka warshadaha/qashinka ee ku urursan nidaamyada deegaanka (tusaale ahaan habka qashinka ilaa hantida) waxaa loo arkaa mid ka mid ah tiirarka dhaqaalaha wareega iyo yoolalka horumarinta waara (Close et al., 2012). Sidaa darteed, fahamka dhinacyada dheef-shiid kiimikaadka, ensaymyada iyo hidde-sidaha ee musharixiintan burburka suurtagalka ah ayaa aad muhiim ugu ah ka saarista wax ku oolka ah iyo hagaajinta bayoolajiyada wasakhda udgoonka leh.
Wasakhowga badan ee udgoonka leh, waxaan si gaar ah u fiirineynaa PAH-yada miisaanka yar ee molecular-ka sida naphthalene iyo naphthalenes-ka la beddelay. Isku-dhafkan waa qaybo muhiim ah oo ka mid ah shidaalka laga helo batroolka, midabyada dharka, alaabada macaamiisha, sunta cayayaanka (mothballs iyo cayayaan-celiyeyaasha), balaastikada iyo tannins sidaas darteedna waa kuwo ku baahsan nidaamyada deegaanka badan (Preuss et al., 2003). Warbixinnadii ugu dambeeyay waxay iftiiminayaan uruurinta naphthalene ee ku jirta sediments-ka biyaha, biyaha dhulka hoostiisa iyo carrada dhulka hoostiisa, aagagga vadose iyo sariiraha webiyada, taasoo soo jeedinaysa ururinta bioaccumulation-keeda ee deegaanka (Duttagupta et al., 2019, 2020). Shaxda 2 waxay soo koobaysaa sifooyinka fiisikokemikaalka, codsiyada iyo saameynta caafimaad ee naphthalene iyo waxyaabaha laga soo saaro. Marka la barbardhigo PAH-yada kale ee miisaanka sare leh ee molecular-ka, naphthalene iyo waxyaabaha laga soo saaro waa kuwo aan hydrophobic ahayn, biyo-milmi kara oo si ballaaran ugu baahsan nidaamyada deegaanka, sidaa darteed waxaa badanaa loo isticmaalaa sidii substrate moodel ah si loo barto dheef-shiid kiimikaadka, hidde-sideyaasha iyo kala duwanaanshaha dheef-shiid kiimikaadka ee PAH-yada. Tiro badan oo noolaha yaryar ah ayaa awood u leh inay dheefshiido naphthalene iyo waxyaabaha ka soo baxa, macluumaad dhammaystiranna waxaa laga heli karaa waddooyinkooda dheef-shiid kiimikaadka, enzymes-ka iyo sifooyinka sharciyeynta (Mallick et al., 2011; Phale et al., 2019, 2020). Intaa waxaa dheer, naphthalene iyo waxyaabaha ka soo baxa waxaa loo qoondeeyay inay yihiin isku-dhisyo tijaabo ah oo loogu talagalay qiimaynta wasakhowga deegaanka sababtoo ah tiro badan iyo helitaankooda bayoolojiga. Hay'adda Ilaalinta Deegaanka ee Mareykanka waxay ku qiyaastay in heerarka celceliska naphthalene ay yihiin 5.19 μg mitir cubic ah halkii qiiqa sigaarka, gaar ahaan gubasho aan dhammaystirnayn, iyo 7.8 ilaa 46 μg oo ka timaadda qiiqa dhinaca, halka soo-gaadhista creosote iyo naphthalene ay ka badan tahay 100 ilaa 10,000 oo jeer (Preuss et al. 2003). Gaar ahaan naphthalene waxaa la ogaaday inay leedahay sunta neefsashada ee noocyada, gobolka, iyo jinsiga gaarka ah iyo kansarka. Iyada oo lagu saleynayo daraasadaha xayawaanka, Hay'adda Caalamiga ah ee Cilmi-baarista Kansarka (IARC) waxay u aqoonsatay naphthalene "suurtogalnimada kansarka aadanaha ee suurtagalka ah" (Kooxda 2B)1. Soo-gaadhista naphthalenes-ka la beddelay, oo inta badan lagu neefsado ama lagu maamulo afka (afka), waxay sababtaa dhaawac unugyada sambabada waxayna kordhisaa dhacdooyinka burooyinka sambabada ee jiirka iyo jiirka (Barnaamijka Qaranka ee Sunta 2). Saamaynta degdegga ah waxaa ka mid ah lallabbo, matag, xanuun caloosha ah, shuban, madax-xanuun, jahawareer, dhidid badan, qandho, tachycardia, iwm. Dhanka kale, carbaryl-ka cayayaanka ee carbamate-ka ballaaran (1-naphthyl N-methylcarbamate) ayaa la soo sheegay inay sun u tahay dadka aan laf-dhabarta lahayn ee biyaha ku nool, amfibiyaanka, shinnida malabka iyo aadanaha waxaana la muujiyay inay joojiso acetylcholinesterase taasoo keenta curyaannimo (Smulders et al., 2003; Bulen iyo Distel, 2011). Sidaa darteed, fahamka hababka burburka microbial, nidaaminta hidde-sidaha, falgallada enzymatic iyo unugyada ayaa muhiim u ah horumarinta istaraatiijiyadaha bioremediation ee deegaannada wasakhaysan.
Shaxda 2. Macluumaad faahfaahsan oo ku saabsan sifooyinka fiisikokemikaalka, isticmaalka, hababka aqoonsiga iyo cudurrada la xiriira ee naphthalene iyo waxyaabaha ka soo baxa.
Meelaha wasakhaysan, wasakhowga udgoonka ee hydrophobic iyo lipophilic waxay sababi karaan saameyn unugyo kala duwan oo ku yimaada microbiome-ka deegaanka (bulshada), sida isbeddellada ku yimaada dareeraha xuubka, soo-gaadhista xuubka, bararka lakabka laba-jibbaaran ee lipid, carqaladaynta wareejinta tamarta (silsiladda gaadiidka elektaroonigga ah/xoogga ujeeddada proton), iyo dhaqdhaqaaqa borotiinnada la xiriira xuubka (Sikkema et al., 1995). Intaa waxaa dheer, qaar ka mid ah dhexdhexaadiyeyaasha milma sida catechols iyo quinones waxay abuuraan noocyada oksijiinta ee falcelinta leh (ROS) waxayna sameeyaan isku-darka DNA-da iyo borotiinnada (Penning et al., 1999). Sidaa darteed, tirada badan ee isku-dhafka noocaas ah ee ku jira nidaamyada deegaanka waxay cadaadis kala doorasho ah saartaa bulshooyinka microbial-ka si ay u noqdaan kuwo wax ku ool ah oo wax ku ool ah heerarka kala duwan ee jir ahaaneed, oo ay ku jiraan qaadashada/gaadiidka, isbeddelka unugyada, isku-darka/isticmaalka, iyo kala-qaybinta.
Baaritaan lagu sameeyay Mashruuca Xogta Ribosomal-II (RDP-II) ayaa lagu ogaaday in wadarta guud ee 926 nooc oo bakteeriya ah laga soocay dhaqamada warbaahinta ama kobcinta ee lagu wasakheeyay naphthalene ama waxyaabaha ka soo baxa. Kooxda Proteobacteria waxay lahayd tirada ugu badan ee wakiillada (n = 755), waxaana ku xigay Firmicutes (52), Bacteroidetes (43), Actinobacteria (39), Tenericutes (10), iyo bakteeriyada aan la kala saarin (8) (Jaantuska 2). Wakiillada γ-Proteobacteria (Pseudomonadales iyo Xanthomonadales) ayaa xukumayay dhammaan kooxaha Gram-negative oo leh heer sare oo G+C ah (54%), halka Clostridiales iyo Bacillales (30%) ay ahaayeen kooxo Gram-positive ah oo leh heer hoose oo G+C ah. Pseudomonas (tirada ugu badan, 338 nooc) ayaa la soo sheegay inay awood u leeyihiin inay baabi'iyaan naphthalene iyo waxyaabaha ka soo baxa methyl-ka ee ku jira nidaamyada kala duwan ee wasakhaysan (dhuxusha, batroolka, saliidda cayriin, dhoobada, daadashada saliidda, biyaha wasakhaysan, qashinka dabiiciga ah iyo meelaha qashinka lagu shubo) iyo sidoo kale nidaamyada deegaanka ee aan dhammaanayn (ciidda, webiyada, qashinka iyo biyaha dhulka hoostiisa) (Jaantuska 2). Intaa waxaa dheer, daraasadaha kobcinta iyo falanqaynta metagenomic ee qaar ka mid ah gobolladan ayaa shaaca ka qaaday in noocyada Legionella iyo Clostridium ee aan la dhaqmin ay yeelan karaan awood burburin, taasoo muujinaysa baahida loo qabo in la beero bakteeriyadan si loo barto waddooyin cusub iyo kala duwanaanshaha dheef-shiid kiimikaadka.
Jaantuska 2. Kala duwanaanshaha taxonomic iyo qaybinta deegaanka ee wakiilada bakteeriyada ee deegaannada ay ku sumoobeen naphthalene iyo naphthalene derivatives.
Ka mid ah noolaha kala duwan ee urta leh ee burburiya haydarojiin, intooda badan waxay awood u leeyihiin inay naphthalene ka dhigaan isha kaliya ee kaarboonka iyo tamarta. Taxanaha dhacdooyinka ku lug leh dheef-shiid kiimikaadka naphthalene ayaa lagu sharraxay Pseudomonas sp. (noocyada: NCIB 9816-4, G7, AK-5, PMD-1 iyo CSV86), Pseudomonas stutzeri AN10, Pseudomonas fluorescens PC20 iyo noocyo kale (ND6 iyo AS1) (Mahajan et al., 1994; Resnick et al., 1996; Annweiler et al., 2000; Basu et al., 2003; Dennis iyo Zylstra, 2004; Sota et al., 2006; Dheef-shiid kiimikaadka waxaa bilaabay dioxygenase [naphthalene dioxygenase (NDO), giraan hydroxylating dioxygenase ah] kaas oo kiciya oksaydhka mid ka mid ah giraanta udgoonka ee naphthalene iyadoo la adeegsanayo oksijiin molecular ah oo ah substrate kale, iyadoo naphthalene u beddelaysa cis-naphthalenediol (Jaantuska 3). Cis-dihydrodiol waxaa loo beddelaa 1,2-dihydroxynaphthalene oo ay samaysay dehydrogenase. Dioxygenase-ka kala-goynta giraanta, 1,2-dihydroxynaphthalene dioxygenase (12DHNDO), wuxuu 1,2-dihydroxynaphthalene u beddelaa 2-hydroxychromene-2-carboxylic acid. Isomerization-ka enzymatic cis-trans wuxuu soo saaraa trans-o-hydroxybenzylidenepyruvate, kaas oo ay kala-gooyso hydratase aldolase una beddelo salicylic aldehyde iyo pyruvate. Pyruvate-ka aashitada dabiiciga ah wuxuu ahaa isku-darka ugu horreeya ee C3 ee laga soo qaatay qalfoofka kaarboonka naphthalene wuxuuna u jiheeyay marinka kaarboonka dhexe. Intaa waxaa dheer, salicylaldehyde dehydrogenase-ka ku tiirsan NAD+ wuxuu salicylaldehyde u beddelaa salicylic acid. Dheef-shiid kiimikaadka marxaladdan waxaa loo yaqaan "waddada sare" ee naphthalene-ka. Waddadani aad bay ugu badan tahay inta badan bakteeriyada naphthalene-ka hoos u dhigta. Si kastaba ha ahaatee, waxaa jira dhawr ka reeban; tusaale ahaan, Bacillus hamburgii 2 ee heerkulka, Naafaynta naphthalene waxaa bilaaba naphthalene 2,3-dioxygenase si ay u samayso 2,3-dihydroxynaphthalene (Annweiler et al., 2000).
Jaantuska 3. Waddooyinka naphthalene, methylnaphthalene, naphthoic acid, iyo carbaryl oo burbura. Tirooyinka wareegsan waxay matalaan ensaymada mas'uulka ka ah beddelka isku xigxiga ee naphthalene iyo waxyaabaha ka soo baxa alaabada xiga. 1 - naphthalene dioxygenase (NDO); 2, cis-dihydrodiol dehydrogenase; 3, 1,2-dihydroxynaphthalene dioxygenase; 4, 2-hydroxychromene-2-carboxylic acid isomerase; 5, trans-O-hydroxybenzylidenepyruvate hydratase aldolase; 6, salicylaldehyde dehydrogenase; 7, salicylate 1-hydroxylase; 8, catechol 2,3-dioxygenase (C23DO); 9, 2-hydroxymuconate semialdehyde dehydrogenase; 10, 2-oxopent-4-enoate hydratase; 11, 4-hydroxy-2-oxopentanoate aldolase; 12, acetaldehyde dehydrogenase; 13, catechol-1,2-dioxygenase (C12DO); 14, muconate cycloisomerase; 15, muconolactone delta-isomerase; 16, β-ketoadipatenollactone hydrolase; 17, β-ketoadipate succinyl-CoA transferase; 18, β-ketoadipate-CoA thiolase; 19, succinyl-CoA: acetyl-CoA succinyltransferase; 20, salicylate 5-hydroxylase; 21 - gentisate 1,2-dioxygenase (GDO); 22, maleylpyruvate isomerase; 23, fumarylpyruvate hydrolase; 24, methylnaphthalene hydroxylase (NDO); 25, hydroxymethylnaphthalene dehydrogenase; 26, naphthaldehyde dehydrogenase; 27, 3-formylsalicylic acid oxidase; 28, hydroxyisophthalate decarboxylase; 29, carbaryl hydrolase (CH); 30, 1-naphthol-2-hydroxylase.
Iyada oo ku xidhan noolaha iyo qaab-dhismeedka hidde-sideyaasha, asiidhka salicylic ee ka dhasha ayaa si dheeraad ah loogu dheefshiidaa iyada oo loo marayo marinka catechol iyadoo la adeegsanayo salicylate 1-hydroxylase (S1H) ama iyada oo loo marayo marinka gentisate iyadoo la adeegsanayo salicylate 5-hydroxylase (S5H) (Jaantuska 3). Maadaama asiidhka salicylic uu yahay dhexdhexaadka ugu weyn ee dheef-shiid kiimikaadka naphthalene (marinka sare), tallaabooyinka laga bilaabo asiidhka salicylic ilaa dhexdhexaadka TCA waxaa badanaa loogu yeeraa marinka hoose, hiddo-sidayaashuna waxay u habaysan yihiin hal operon. Waa wax caadi ah in la arko in hiddo-sidayaasha ku jira marinka sare ee operon (nah) iyo marinka hoose ee operon (sal) ay nidaamiyaan arrimo caadi ah oo nidaamiya; tusaale ahaan, NahR iyo asiidhka salicylic waxay u dhaqmaan sidii kiciyeyaal, taasoo u oggolaanaysa labada operon inay si buuxda u dheefshiidaan naphthalene (Phale et al., 2019, 2020).
Intaa waxaa dheer, catechol waxaa si wareeg ah loogu kala gooyaa 2-hydroxymuconate semialdehyde iyada oo loo marayo marin-haweedka meta by catechol 2,3-dioxygenase (C23DO) (Yen et al., 1988) ka dibna waxaa lagu sii shubaa 2-hydroxymuconate semialdehyde hydrolase si loo sameeyo 2-hydroxypent-2,4-dienoic acid. 2-hydroxypent-2,4-dienoate waxaa markaa loo beddelaa pyruvate iyo acetaldehyde iyadoo la adeegsanayo hydratase (2-oxopent-4-enoate hydratase) iyo aldolase (4-hydroxy-2-oxopentanoate aldolase) ka dibna wuxuu galaa marin-haweedka dhexe ee kaarboonka (Jaantuska 3). Haddii kale, catechol waxaa si wareeg ah loogu kala gooyaa cis,cis-muconate iyada oo loo marayo marin-haweedka ortho by catechol 1,2-oxygenase (C12DO). Muconate cycloisomerase, muconolactone isomerase, iyo β-ketoadipate-nollactone hydrolase waxay cis,cis-muconate u beddelaan 3-oxoadipate, kaas oo gala marin-haweedka kaarboonka dhexe iyada oo loo marayo succinyl-CoA iyo acetyl-CoA (Nozaki et al., 1968) (Jaantuska 3).
Jidka gentisate (2,5-dihydroxybenzoate), giraanta aromatic-ka waxaa kala gooya gentisate 1,2-dioxygenase (GDO) si ay u samayso maleylpyruvate. Badeecadan si toos ah ayaa loo hydrolyze-ka karaa si ay u noqoto pyruvate oo malate, ama waxaa lagu kala gooyn karaa si ay u sameyso fumarylpyruvate, kaas oo markaa lagu hydrolyze-ka karo si uu u noqdo pyruvate oo uu u noqdo fumarate (Larkin iyo Day, 1986). Doorashada dariiqa kale waxaa lagu arkay bakteeriyada Gram-negative iyo Gram-positive labadaba heerarka biochemical iyo hidde-sideyaasha (Morawski et al., 1997; Whyte et al., 1997). Bakteeriyada Gram-negative (Pseudomonas) waxay doorbidaan inay isticmaalaan salicylic acid, kaas oo ah kiciyaha dheef-shiid kiimikaadka naphthalene, isagoo ka dhigaya catechol iyadoo la adeegsanayo salicylate 1-hydroxylase (Gibson iyo Subramanian, 1984). Dhanka kale, bakteeriyada Gram-positive (Rhodococcus), salicylate 5-hydroxylase waxay asiidhka salicylic u beddeshaa asiidhka gentisic, halka salicylic acid uusan saameyn ku yeelan qoraalka hiddo-wadaha naphthalene (Grund et al., 1992) (Jaantuska 3).
Waxaa la soo sheegay in noocyada sida Pseudomonas CSV86, Oceanobacterium NCE312, Marinhomonas naphthotrophicus, Sphingomonas paucimobilis 2322, Vibrio cyclotrophus, Pseudomonas fluorescens LP6a, Pseudomonas iyo noocyada Mycobacterium ay burburin karaan monomethylnaphthalene ama dimethylnaphthalene (Dean-Raymond iyo Bartha, 1975; Cane iyo Williams, 1982; Mahajan et al., 1994; Dutta et al., 1998; Hedlund et al., 1999). Kuwaas waxaa ka mid ah, waddada burburka 1-methylnaphthalene iyo 2-methylnaphthalene ee Pseudomonas sp. CSV86 si cad ayaa loo bartay heerarka bayoolaji iyo enzymatic (Mahajan et al., 1994). 1-Methylnaphthalene waxaa lagu dheefshiidaa laba waddo. Marka hore, giraanta aromatic-ka ah waxaa lagu shubaa hydroxylated (giraanta aan la beddelin ee methylnaphthalene) si ay u samayso cis-1,2-dihydroxy-1,2-dihydro-8-methylnaphthalene, taas oo si dheeraad ah loogu oksaydhiyo methyl salicylate iyo methylcatechol, ka dibna waxay gashaa marin-haweedka dhexe ee kaarboonka ka dib kala-goynta giraanta (Jaantuska 3). Jidkan waxaa loogu yeeraa "marin-haweedka isha kaarboonka". "Wadada ka-saarista sunta" ee labaad, kooxda methyl-ka waxaa hydroxylated kara NDO si ay u samayso 1-hydroxymethylnaphthalene, kaas oo si dheeraad ah loogu oksaydhiyo 1-naphthoic acid oo loo soo saaro habka dhaqanka oo ah badeecad aan dhammaad lahayn. Daraasaduhu waxay muujiyeen in nooca CSV86 uusan awoodin inuu ku koro aashitada 1- iyo 2-naphthoic oo ah isha kaliya ee kaarboonka iyo tamarta, taasoo xaqiijinaysa marin-haweedka ka-saarista sunta (Mahajan et al., 1994; Basu et al., 2003). 2-methylnaphthalene, kooxda methyl-ka waxay maraan hydroxylation iyadoo la adeegsanayo hydroxylase si ay u sameeyaan 2-hydroxymethylnaphthalene. Intaa waxaa dheer, giraanta aan la beddelin ee giraanta naphthalene waxay maraan hydroxylation giraanta si ay u sameeyaan dihydrodiol, kaas oo lagu oksaydhiyo 4-hydroxymethylcatechol taxane ah oo falgallo ensaym-ka ah oo galaya marinka kaarboonka dhexe iyada oo loo marayo marinka kala-goynta meta-ring. Sidoo kale, S. paucimobilis 2322 ayaa la sheegay inuu isticmaalo NDO si uu u isticmaalo hydroxylate 2-methylnaphthalene, kaas oo si dheeraad ah loogu oksaydhiyo si uu u sameeyo methyl salicylate iyo methylcatechol (Dutta et al., 1998).
Asiidhyada Naphthoic (kuwa la beddelay/aan la beddelin) waa waxyaabaha ka soo baxa sunta/bayoolajiga ee la sameeyay intii lagu jiray burburka methylnaphthalene, phenanthrene iyo anthracene oo lagu sii daayo habka dhaqanka ee la isticmaalay. Waxaa la soo sheegay in carradu ay kala soocdo Stenotrophomonas maltophilia CSV89 waxay awood u leedahay inay dheefshiido 1-naphthoic acid sidii ilo kaarboon ah (Phale et al., 1995). Dheef-shiid kiimikaadku wuxuu ka bilaabmaa dihydroxylation ee giraanta udgoonka si loo sameeyo 1,2-dihydroxy-8-carboxynaphthalene. Diol-ka ka dhasha waxaa lagu oksaydhiyaa catechol iyada oo loo marayo 2-hydroxy-3-carboxybenzylidenepyruvate, 3-formylsalicylic acid, 2-hydroxyisophthalic acid iyo salicylic acid waxayna gashaa marinka kaarboonka dhexe iyada oo loo marayo marinka kala-goynta meta-ring (Jaantuska 3).
Carbaryl waa sunta cayayaanka ee naphthyl carbamate. Tan iyo Kacaankii Cagaaran ee Hindiya 1970-meeyadii, isticmaalka bacriminta kiimikada iyo sunta cayayaanka ayaa horseeday kororka qiiqa polycyclic aromatic hydrocarbon (PAH) ee ka imanaya ilo aan muhiim ahayn oo beeraha ah (Pingali, 2012; Duttagupta et al., 2020). Qiyaastii 55% (85,722,000 hektar) oo ah dhulka dalaga ee Hindiya waxaa lagu daaweeyaa sunta cayayaanka kiimikada. Shantii sano ee la soo dhaafay (2015-2020), waaxda beeraha Hindiya waxay isticmaashay celcelis ahaan 55,000 ilaa 60,000 oo tan oo sunta cayayaanka ah sannad kasta (Waaxda Iskaashatooyinka iyo Daryeelka Beeraleyda, Wasaaradda Beeraha, Dowladda Hindiya, Agoosto 2020). Bannaanka waqooyiga iyo bartamaha Gangetic (gobolada leh tirada ugu badan ee dadka iyo tirada dadka), isticmaalka sunta cayayaanka ee dalagyada waa mid baahsan, iyadoo sunta cayayaanka ay aad u badan tahay. Carbaryl (1-naphthyl-N-methylcarbamate) waa cayayaan dila oo si dhexdhexaad ah ilaa sun ah oo loo isticmaalo beeraha Hindiya celcelis ahaan 100-110 tan. Waxaa si caadi ah loogu iibiyaa magaca ganacsiga Sevin waxaana loo isticmaalaa in lagu xakameeyo cayayaanka (aphids, quraanjada dabka, boodada, caarada, caarada iyo cayayaanka kale ee bannaanka) ee saameeya noocyo kala duwan oo dalagyo ah (galley, soybean, suuf, miro iyo khudaar). Noocyo yaryar ah sida Pseudomonas (NCIB 12042, 12043, C4, C5, C6, C7, Pseudomonas putida XWY-1), Rhodococcus (NCIB 12038), Sphingobacterium spp. (CF06), Burkholderia (C3), Micrococcus iyo Arthrobacter ayaa sidoo kale loo isticmaali karaa in lagu xakameeyo cayayaanka kale. Waxaa la soo sheegay in RC100 uu burburin karo carbaryl (Larkin iyo Day, 1986; Chapalamadugu iyo Chaudhry, 1991; Hayatsu et al., 1999; Swetha iyo Phale, 2005; Trivedi et al., 2017). Waddada burburka ee carbaryl waxaa si ballaaran loogu bartay heerarka bayoolaji, enzymatic iyo hidde-sideyaasha ee go'doominta ciidda ee Pseudomonas sp. Noocyada C4, C5 iyo C6 (Swetha iyo Phale, 2005; Trivedi et al., 2016) (Jaantuska 3). Waddada dheef-shiid kiimikaadka waxay ka bilaabataa hydrolysis-ka xiriirka ester-ka ee carbaryl hydrolase (CH) si loo sameeyo 1-naphthol, methylamine iyo carbon dioxide. 1-naphthol waxaa markaa loo beddelaa 1,2-dihydroxynaphthalene iyadoo la adeegsanayo 1-naphthol hydroxylase (1-NH), kaas oo si dheeraad ah loogu dheefshiido marinnada kaarboonka dhexe iyada oo loo marayo salicylate iyo gentisate. Bakteeriyada qaar ee carbaryl-degrading ayaa la sheegay inay dheefshiido u beddesho salicylic acid iyada oo loo marayo kala-goynta giraanta catechol ortho (Larkin iyo Day, 1986; Chapalamadugu iyo Chaudhry, 1991). Waxaa xusid mudan, bakteeriyada naphthalene-degrading waxay inta badan dheefshiido salicylic acid iyada oo loo marayo catechol, halka bakteeriyada carbaryl-degrading ay doorbiddo inay dheefshiido salicylic acid iyada oo loo marayo marinnada gentisate.
Naphthalenesulfonic acid/disulfonic acid iyo naphthylaminesulfonic acid derivatives waxaa loo isticmaali karaa dhexdhexaad ahaan soo saarista midabada azo, wakiilada qoynta, kala firdhiyaha, iwm. In kasta oo iskudhisyadani ay sun yar ku leeyihiin aadanaha, qiimaynta sunta ayaa muujisay inay dilaa u yihiin kalluunka, daphnia iyo algae (Greim et al., 1994). Wakiillada hidde-sideyaasha Pseudomonas (noocyada A3, C22) ayaa la soo sheegay inay bilaabaan dheef-shiid kiimikaad iyagoo labanlaabaya hydroxylation ee giraanta udgoonka leh ee ka kooban kooxda sulfonic acid si ay u sameeyaan dihydrodiol, kaas oo loo beddelo 1,2-dihydroxynaphthalene iyadoo la kala saarayo kooxda sulfite (Brilon et al., 1981). 1,2-dihydroxynaphthalene ee ka dhashay waxaa lagu kala qaadaa marinnada naphthalene ee caadiga ah, tusaale ahaan, marinnada catechol ama gentisate (Jaantuska 4). Waxaa la muujiyay in aminonaphthalenesulfonic acid iyo hydroxynaphthalenesulfonic acid si buuxda loo burburin karo iyadoo la adeegsanayo isku-dhafka bakteeriyada isku dhafan oo leh marinnada catabolic ee dhammaystiran (Nortemann et al., 1986). Waxaa la muujiyay in hal xubin oo ka mid ah isbahaysiga uu ka saaro aminonaphthalenesulfonic acid ama hydroxynaphthalenesulfonic acid 1,2-dioxygenation, halka aminosalicylate ama hydroxysalicylate lagu sii daayo habka dhaqanka isagoo ah metabolite aan dhammaad lahayn ka dibna waxaa qaata xubnaha kale ee ururka. Naphthalenedisulfonic acid waa mid si yar u janjeera laakiin si liidata u burbura oo sidaas darteed waxaa lagu dheefshiidi karaa marinno kala duwan. Desulfurization-ka ugu horreeya wuxuu dhacaa inta lagu jiro dihydroxylation regioselective ee giraanta aromatic iyo kooxda sulfonic acid; Kala-baxa labaad wuxuu dhacaa inta lagu jiro hydroxylation of 5-sulfosalicylic acid by salicylic acid 5-hydroxylase si loo sameeyo gentisic acid, kaas oo gala marinka dhexe ee kaarboonka (Brilon et al., 1981) (Jaantuska 4). Ensaymada mas'uulka ka ah burburka naphthalene ayaa sidoo kale mas'uul ka ah dheef-shiid kiimikaadka naphthalene sulfonate (Brilon et al., 1981; Keck et al., 2006).
Jaantuska 4. Waddooyinka dheef-shiid kiimikaadka ee burburka naphthalene sulfonate. Tirooyinka gudaha goobooyinka waxay matalaan ensaymada mas'uulka ka ah dheef-shiid kiimikaadka naphthyl sulfonate, oo la mid ah/la mid ah ensaymada lagu sharraxay Jaantuska 3.
PAH-yada miisaanka yar ee molecular-ka (LMW-PAHs) waa kuwo la dhimi karo, hydrophobic ah oo aan si fiican u milmi karin, sidaas darteedna uma nugul yihiin burburka/burburka dabiiciga ah. Si kastaba ha ahaatee, noolaha aerobic-ga ah ayaa awood u leh inay oksaydheeyaan iyagoo nuugaya oksijiinta molecular (O2). Ensaymadani waxay inta badan ka tirsan yihiin fasalka oxidoreductases waxayna samayn karaan falcelin kala duwan sida hydroxylation ring aromatic (mono- ama dihydroxylation), dehydrogenation iyo kala-goynta giraanta aromatic. Badeecadaha laga helo falgalladani waxay ku jiraan xaalad oksaydheyn sare waxayna si fudud ugu milmaan marinnada kaarboonka dhexe (Phale et al., 2020). Ensaymada ku jira marinnada burburka ayaa la sheegay inay yihiin kuwo la kicin karo. Dhaqdhaqaaqa ensaymadani aad ayuu u hooseeyaa ama waa la dayaci karaa marka unugyada lagu koriyo ilo kaarboon fudud sida gulukoos ama asiidhyo dabiici ah. Shaxda 3 waxay soo koobaysaa ensaymada kala duwan (oxygenases, hydrolases, dehydrogenases, oxidases, iwm.) oo ku lug leh dheef-shiid kiimikaadka naphthalene iyo waxyaabaha ka soo baxa.
Shaxda 3. Astaamaha bayoolojiga ee ensaymada mas'uulka ka ah burburka naphthalene iyo waxyaabaha ka soo baxa.
Daraasadaha Raadiyaha (18O2) waxay muujiyeen in ku darista molecular O2 ee giraanta udgoonka leh ee oxygenases ay tahay tallaabada ugu muhiimsan ee lagu kicinayo burburka bio ee isku-dhafka ah (Hayaishi et al., 1955; Mason et al., 1955). Ku darista hal atamka oksijiinta (O) ee ka timaadda oksijiinta molecular (O2) ee ku jirta substrate-ka waxaa bilaabaya monooxygenases endogenous ama exogenous (oo sidoo kale loo yaqaan hydroxylases). Atam kale oo oksijiin ah ayaa loo yareeyaa biyo. Monooxygenases-ka dibadda ka yimaada waxay yareeyaan flavin iyadoo la adeegsanayo NADH ama NADPH, halka flavin-ka endomonooxygenases-ka uu yareeyo substrate-ka. Booska hydroxylation-ku wuxuu keenaa kala duwanaansho sameynta badeecada. Tusaale ahaan, salicylate 1-hydroxylase hydroxylates salicylic acid booska C1, taasoo sameysa catechol. Dhanka kale, salicylate 5-hydroxylase oo ah qaybo badan (oo ka kooban reductase, ferredoxin, iyo subunits oksijiinase) ayaa hydroxylates salicylic acid ka dhigta booska C5, taasoo sameysanaysa gentisic acid (Yamamoto et al., 1965).
Dioxygenases-ku waxay ku daraan laba atam oo O2 ah substrate-ka. Iyada oo ku xidhan badeecadaha la sameeyay, waxay u qaybsan yihiin dioxygenases ring hydroxylating iyo dioxygenases ring cleaving. Dioxygenases-ka ring hydroxylating waxay substrates aromatic u beddelaan cis-dihydrodiols (tusaale ahaan, naphthalene) waxayna ku baahsan yihiin bakteeriyada. Ilaa hadda, waxaa la muujiyay in noolaha ka kooban dioxygenases ring hydroxylating ay awood u leeyihiin inay ku koraan ilo kala duwan oo kaarboon ah oo udgoon, ensaymyadanna waxaa loo kala saaraa NDO (naphthalene), toluene dioxygenase (TDO, toluene), iyo biphenyl dioxygenase (BPDO, biphenyl). Labada NDO iyo BDO waxay kicin karaan hydroxylation-ka labanlaabka ah ee oksaydhka iyo silsiladda dhinaca ee hydrocarbons-ka aromatic-ka ee kala duwan (toluene, nitrotoluene, xylene, ethylbenzene, naphthalene, biphenyl, fluorene, indole, methylnaphthalene, naphthalenesulfonate, phenanthrene, anthracene, acetophenone, iwm.) (Boyd iyo Sheldrake, 1998; Phale et al., 2020). NDO waa nidaam qaybo badan leh oo ka kooban oxidoreductase, ferredoxin, iyo qayb firfircoon oo oksijiinase ah oo goobta ka kooban (Gibson iyo Subramanian, 1984; Resnick et al., 1996). Cutubka katalytic-ga ee NDO wuxuu ka kooban yahay qayb weyn oo α ah iyo qayb yar oo β ah oo lagu habeeyey habaynta α3β3. NDO waxay ka tirsan tahay qoys ballaaran oo oksijiin ah, α-subunit-keeduna wuxuu ka kooban yahay goob Rieske ah [2Fe-2S] iyo bir aan lahayn heme mononuclear ah, taas oo go'aamisa gaar ahaanshaha substrate-ka ee NDO (Parales et al., 1998). Caadi ahaan, hal wareeg oo katalytic ah, laba elektaroono ah oo ka yimaada dhimista pyridine nucleotide ayaa loo wareejiyaa ion-ka Fe(II) ee goobta firfircoon iyada oo loo marayo reductase, ferredoxin iyo goobta Rieske. Kuwa wax u dhigma ee yareeya waxay kiciyaan oksijiinta molecular, taas oo ah shuruud u ah dihydroxylation substrate (Ferraro et al., 2005). Ilaa hadda, dhowr NDO ah ayaa la nadiifiyay oo si faahfaahsan loogu sifeeyay noocyo kala duwan, xakamaynta hidde-sidaha ee waddooyinka ku lug leh burburka naphthalene si faahfaahsan ayaa loo bartay (Resnick et al., 1996; Parales et al., 1998; Karlsson et al., 2003). Dioxygenase-ka kala-goynta giraanta (ensaymyada endo- ama ortho-ring-cleaving iyo enzymes-ka exodiol- ama meta-ring-cleaving) waxay ku dhaqmaan isku-dhisyo aromatic ah oo hydroxylated. Tusaale ahaan, dioxygenase-ka kala-goynta ortho-ring-cleaving waa catechol-1,2-dioxygenase, halka dioxygenase-ka kala-goynta meta-ring-ka uu yahay catechol-2,3-dioxygenase (Kojima et al., 1961; Nozaki et al., 1968). Marka laga soo tago oksijiinnada kala duwan, waxaa sidoo kale jira dehydrogenases kala duwan oo mas'uul ka ah ka-goynta hydrogenos-ka aromatic dihydrodiols, alcohols iyo aldehydes iyo isticmaalka NAD+/NADP+ sida aqbalayaasha elektarooniga, kuwaas oo ah qaar ka mid ah enzymes-ka muhiimka ah ee ku lug leh dheef-shiid kiimikaadka (Gibson iyo Subramanian, 1984; Shaw iyo Harayama, 1990; Fahle et al., 2020).
Ensaymyada sida hydrolases (esterases, amidases) waa fasalka labaad ee muhiimka ah ee ensaymyada isticmaala biyaha si ay u kala gooyaan isku xidhka covalent iyo inay muujiyaan gaar ahaanshaha substrate-ka ballaaran. Carbaryl hydrolase iyo hydrolases kale waxaa loo arkaa inay yihiin qaybo ka mid ah periplasm (transmembrane) ee xubnaha bakteeriyada Gram-negative (Kamini et al., 2018). Carbaryl waxay leedahay isku xidhka amide iyo ester; sidaa darteed, waxaa lagu milmi karaa esterase ama amidase si loo sameeyo 1-naphthol. Carbaryl ee Rhizobium rhizobium nooca AC10023 iyo Arthrobacter nooca RC100 ayaa la sheegay inay u shaqeeyaan sidii esterase iyo amidase, siday u kala horreeyaan. Carbaryl ee nooca Arthrobacter RC100 sidoo kale wuxuu u shaqeeyaa sidii amidase. RC100 waxaa la muujiyay inuu milmiyo afar sunta cayayaanka ee fasalka N-methylcarbamate sida carbaryl, methomyl, mefenamic acid iyo XMC (Hayaatsu et al., 2001). Waxaa la soo sheegay in CH ee Pseudomonas sp. C5pp uu ku shaqeyn karo carbaryl (100% firfircoon) iyo 1-naphthyl acetate (36% firfircoon), laakiin uusan ku shaqeyn 1-naphthylacetamide, taasoo muujinaysa inay tahay esterase (Trivedi et al., 2016).
Daraasadaha bayoolojiga, qaababka nidaaminta ensaymka, iyo falanqaynta hidde-sidaha ayaa muujiyay in hiddo-sidayaasha burburka naphthalene ay ka kooban yihiin laba cutub oo nidaamin ah oo la kicin karo ama "operons": nah ("waddada kor u socota", oo u beddelaysa naphthalene asiidh salicylic) iyo sal ("waddada hoose", oo u beddelaysa salicylic asiidh waddada dhexe ee kaarboonka iyada oo loo marayo catechol). Salicylic acid iyo analogues-keedu waxay u dhaqmi karaan sidii kiciyeyaal (Shamsuzzaman iyo Barnsley, 1974). Iyadoo ay joogaan gulukoos ama asiidhyo dabiici ah, operon-ka waa la cadaadiyaa. Jaantuska 5 wuxuu muujinayaa abaabulka hidde-sidaha oo dhammaystiran ee burburka naphthalene (qaabka operon). Noocyo/noocyo badan oo magac leh oo hiddo-sidaha nah ah (ndo/pah/dox) ayaa la sharraxay waxaana la ogaaday inay leeyihiin isku-midnimo taxane sare ah (90%) dhammaan noocyada Pseudomonas (Abbasian et al., 2016). Hiddo-sidayaasha waddada kor u socota ee naphthalene guud ahaan waxaa loo habeeyay si isku mid ah sida lagu muujiyey Jaantuska 5A. Hidde kale, nahQ, ayaa sidoo kale la soo sheegay inuu ku lug lahaa dheef-shiid kiimikaadka naphthalene wuxuuna badanaa ku yaalay inta u dhaxaysa nahC iyo nahE, laakiin shaqadiisa dhabta ah weli lama shaacin. Sidoo kale, hidda-wadaha nahY, oo mas'uul ka ah kemotaxis-ka xasaasiyadda naphthalene, ayaa laga helay dhammaadka fog ee operon-ka nah ee xubnaha qaarkood. Ralstonia sp., hidda-wadaha U2 ee codeeya glutathione S-transferase (gsh) ayaa la ogaaday inuu ku yaallo inta u dhaxaysa nahAa iyo nahAb laakiin ma uusan saameyn sifooyinka isticmaalka naphthalene (Zylstra et al., 1997).
Jaantuska 5. Ururinta hidde-sidaha iyo kala duwanaanshaha la arkay intii lagu jiray burburka naphthalene ee noocyada bakteeriyada; (A) Waddada sare ee naphthalene, dheef-shiid kiimikaadka naphthalene ilaa salicylic acid; (B) Waddada hoose ee naphthalene, salicylic acid iyada oo loo marayo catechol ilaa marinnada kaarboonka dhexe; (C) salicylic acid iyada oo loo marayo gentisate ilaa marinnada kaarboonka dhexe.
"Wadada hoose" (sal operon) waxay caadi ahaan ka kooban tahay nahGTHINLMOKJ waxayna salicylate u beddeshaa pyruvate iyo acetaldehyde iyada oo loo marayo marinka kala-goynta catechol metaring. Hidde-sidaha nahG (oo ku lifaaqan salicylate hydroxylase) ayaa la ogaaday in lagu keydiyay dhammaadka u dhow operon (Sawirka 5B). Marka la barbardhigo noocyada kale ee naphthalene-degrading, P. putida CSV86 nah iyo sal operons waa kuwo isku xiran oo aad isugu dhow (qiyaastii 7.5 kb). Bakteeriyada Gram-negative qaarkood, sida Ralstonia sp. U2, Polaromonas naphthalenivorans CJ2, iyo P. putida AK5, naphthalene waxaa lagu dheefshiidaa sidii metabolite kaarboon dhexe iyada oo loo marayo marinka gentisate (qaabka sgp/nag operon). Cajaladda hidda-wadaha waxaa badanaa lagu matalaa qaabka nagAaGHabAcAdBFCQEDJI, halkaas oo nagR (ku-darsanaysa nidaamiyaha nooca LysR) uu ku yaal dhammaadka sare (Jaantuska 5C).
Carbaryl wuxuu galaa wareegga kaarboonka dhexe iyada oo loo marayo dheef-shiid kiimikaadka 1-naphthol, 1,2-dihydroxynaphthalene, salicylic acid, iyo gentisic acid (Jaantuska 3). Iyada oo lagu saleynayo daraasadaha hidde-sideyaasha iyo dheef-shiid kiimikaadka, waxaa la soo jeediyay in waddadan loo qaybiyo "kor u kaca" (beddelka carbaryl ilaa salicylic acid), "dhexe" (beddelka salicylic acid ilaa gentisic acid), iyo "hoos u dhaca" (beddelka aashitada gentisic una gudubka dhexdhexaadinta marinnada kaarboonka dhexe) (Singh et al., 2013). Falanqaynta hidde-sideyaasha ee C5pp (supercontig A, 76.3 kb) waxay shaaca ka qaadday in hidda-wadaha mcbACBDEF uu ku lug leeyahay beddelka carbaryl ilaa salicylic acid, oo ay ku xigto mcbIJKL beddelka salicylic acid ilaa gentisic acid, iyo mcbOQP beddelka gentisic acid ilaa dhexdhexaadiyeyaasha kaarboonka dhexe (fumarate iyo pyruvate, Trivedi et al., 2016) (Jaantuska 6).
Waxaa la soo sheegay in ensaymada ku lug leh burburka haydaro-biyoodka udgoonka leh (oo ay ku jiraan naphthalene iyo salicylic acid) ay kicin karaan isku-dhafka u dhigma oo ay xannibi karaan ilo kaarboon oo fudud sida gulukoos ama asiidhyo dabiici ah (Shingler, 2003; Phale et al., 2019, 2020). Waddooyinka kala duwan ee dheef-shiid kiimikaadka ee naphthalene iyo waxyaabaha ka soo baxa, sifooyinka sharciyeynta ee naphthalene iyo carbaryl ayaa ilaa xad la bartay. Naphthalene, hiddo-wadaha ku jira labada waddo ee kor iyo hoosba waxaa nidaamiya NahR, oo ah nidaamiye togan oo LysR-nooc ah. Waxaa looga baahan yahay kicinta hidda-wadaha nah iyadoo la adeegsanayo salicylic acid iyo muujinta heerka sare ee xigta (Yen iyo Gunsalus, 1982). Intaa waxaa dheer, daraasaduhu waxay muujiyeen in qodobka martida isku-dhafan (IHF) iyo XylR (xakameeyaha qoraalka ee ku tiirsan sigma 54) ay sidoo kale muhiim u yihiin kicinta qoraalka ee hidda-wadaha ee dheef-shiid kiimikaadka naphthalene (Ramos et al., 1997). Daraasaduhu waxay muujiyeen in ensaymada marinnada furitaanka ee catechol meta-ring, oo ah catechol 2,3-dioxygenase, ay kiciyaan marka ay joogaan naphthalene iyo/ama salicylic acid (Basu et al., 2006). Daraasaduhu waxay muujiyeen in ensaymada marinnada furitaanka ee catechol ortho-ring, oo ah catechol 1,2-dioxygenase, ay kiciyaan marka ay joogaan benzoic acid iyo cis,cis-muconate (Parsek et al., 1994; Tover et al., 2001).
Nooca C5pp, shan hiddo-side, mcbG, mcbH, mcbN, mcbR iyo mcbS, ayaa codeeya nidaamiyeyaasha ka tirsan qoyska LysR/TetR ee nidaamiyeyaasha qoraalka ee mas'uulka ka ah xakamaynta burburka carbaryl. Hiddo-sideyaasha isku midka ah ee mcbG ayaa la ogaaday inay si dhow ula xiriiraan nidaamiyaha nooca LysR PhnS (58% aqoonsiga amino acid) ee ku lug leh dheef-shiid kiimikaadka phenanthrene ee Burkholderia RP00725 (Trivedi et al., 2016). Hiddo-sideyaasha mcbH waxaa la ogaaday inay ku lug leeyihiin marin-haweedka dhexe (beddelka aashitada salicylic una beddela aashitada gentisic) waxayna ka tirsan yihiin nidaamiyaha qoraalka nooca LysR NagR/DntR/NahR ee Pseudomonas iyo Burkholderia. Xubnaha qoyskan ayaa la sheegay inay aqoonsadeen aashitada salicylic inay tahay molecule saameyn gaar ah u leh kicinta hiddo-sideyaasha burburka. Dhanka kale, saddex hiddo-side, mcbN, mcbR iyo mcbS, oo ka tirsan nidaamiyeyaasha qoraalka nooca LysR iyo TetR, ayaa lagu aqoonsaday marin-haweedka hoose (metabolites-ka marin-haweedka kaarboon-dhexe).
Prokaryotes-ka, hababka wareejinta hidda-wadaha ee toosan (helitaanka, is-weydaarsiga, ama wareejinta) iyada oo loo marayo plasmids-ka, transposon-ka, prophages-ka, jasiiradaha genomic-ga, iyo walxaha isku-xidhka isku-dhafan (ICE) ayaa ah sababaha ugu waaweyn ee balaastikada hidde-sideyaasha bakteeriyada, taasoo horseedaysa faa'iidada ama lumitaanka shaqooyinka/astaamaha gaarka ah. Waxay u oggolaanaysaa bakteeriyada inay si dhakhso leh ula qabsato xaaladaha deegaanka ee kala duwan, iyadoo siinaysa faa'iidooyin dheef-shiid kiimikaad oo suurtagal ah martida, sida burburka isku-dhafka udgoonka. Isbeddellada dheef-shiid kiimikaadka waxaa badanaa lagu gaaraa iyada oo la hagaajinayo operons-ka burburka, hababka nidaaminta, iyo gaar ahaan enzyme-ka, taas oo sahlaysa burburka noocyo badan oo isku-dhafan oo udgoon ah (Nojiri et al., 2004; Phale et al., 2019, 2020). Cajaladaha hidda-wadaha ee burburka naphthalene ayaa la ogaaday inay ku yaalliin walxo kala duwan oo guurguura sida plasmids-ka (isku-xidhka iyo kuwa aan isku-xidhka ahayn), transposons-ka, genome-yada, ICE-yada, iyo isku-darka noocyada bakteeriyada kala duwan (Jaantuska 5). Pseudomonas G7, operons-ka nah iyo sal ee plasmid NAH7 waxaa lagu qoraa isla jihada waxayna qayb ka yihiin transposon cilladaysan oo u baahan transposase Tn4653 si loogu abaabulo (Sota et al., 2006). Nooca Pseudomonas NCIB9816-4, hidda-wadaha waxaa laga helay pDTG1 plasmid isku-dhafan laba operons (qiyaastii 15 kb oo kala fog) kuwaas oo lagu qoray jihooyin iska soo horjeeda (Dennis iyo Zylstra, 2004). Nooca Pseudomonas putida AK5, plasmid-ka aan isku-xidhnayn ee pAK5 wuxuu codeeyaa enzyme-ka mas'uulka ka ah burburka naphthalene iyada oo loo marayo marinka gentisate (Izmalkova et al., 2013). Nooca Pseudomonas ee PMD-1, nah operon wuxuu ku yaal koromosoomka, halka sal operon uu ku yaal plasmid-ka isku-xidhka pMWD-1 (Zuniga et al., 1981). Si kastaba ha ahaatee, Pseudomonas stutzeri AN10, dhammaan hiddo-wadaha naphthalene ee burburay (nah iyo sal operons) waxay ku yaalliin koromosoomka waxaana loo maleynayaa in lagu qoro iyada oo loo marayo dhacdooyinka isbeddelka, dib-u-mideynta, iyo dib-u-habaynta (Bosch et al., 2000). Pseudomonas sp. CSV86, nah iyo sal operons waxay ku yaalliin hidde-side qaab ICE ah (ICECSV86). Qaab-dhismeedka waxaa ilaaliya tRNAGly oo ay ku xigto ku celcelin toos ah oo tilmaamaysa goobaha isku-dhafka/ku-xirnaanta (attR iyo attL) iyo isku-darka u eg phage oo ku yaal labada daraf ee tRNAGly, sidaas darteed qaab-dhismeed ahaan la mid ah curiyaha ICEclc (ICEclcB13 ee Pseudomonas knackmusii ee burburka chlorocatechol). Waxaa la soo sheegay in hiddo-sideyaasha ICE lagu wareejin karo isku-xidhnaan leh soo noqnoqosho wareejin aad u hooseeya (10-8), taasoo keentay in hantida burburka loo wareejiyo qaataha (Basu iyo Phale, 2008; Phale et al., 2019).
Inta badan hiddo-sidayaasha mas'uulka ka ah burburka carbaryl waxay ku yaalliin balaasmaha. Arthrobacter sp. RC100 wuxuu ka kooban yahay saddex balaasmaha (pRC1, pRC2 iyo pRC300) oo laba balaasmaha isku xiran, pRC1 iyo pRC2, ay ku dhejiyaan ensaymada u beddela carbaryl gentisate. Dhanka kale, ensaymada ku lug leh beddelka gentisate una beddela metabolites-ka kaarboonka dhexe waxay ku yaalliin koromosoomka (Hayaatsu et al., 1999). Bakteeriyada hidde-sidaha Rhizobium. AC100 oo la shiiday, oo loo isticmaalo beddelka carbaryl una beddela 1-naphthol, waxay ka kooban tahay plasmid pAC200, kaas oo sita hidda-wadaha cehA ee CH ku qeexaya qayb ka mid ah transposon-ka Tnceh oo ay ku hareeraysan yihiin taxane u eg walxaha gelinta (istA iyo istB) (Hashimoto et al., 2002). Nooca Sphingomonas CF06, hidda-wadaha burburka carbaryl waxaa la rumeysan yahay inuu ku jiro shan plasmid: pCF01, pCF02, pCF03, pCF04, iyo pCF05. Isku-dhafka DNA-da ee plasmid-yadan waa mid sarreeya, taasoo muujinaysa jiritaanka dhacdo nuqul hidde ah (Feng et al., 1997). Isku-dhafka carbaryl-degrading symbiont oo ka kooban laba nooc oo Pseudomonas ah, nooca 50581 wuxuu ka kooban yahay plasmid isku-dhafan pCD1 (50 kb) oo qeexaya hidda-wadaha carbaryl hydrolase ee mcd, halka plasmid isku-dhafan ee nooca 50552 uu qeexayo enzyme 1-naphthol-degrading (Chapalamadugu iyo Chaudhry, 1991). Nooca Achromobacter WM111, hidda-wadaha mcd furadan hydrolase wuxuu ku yaal plasmid 100 kb (pPDL11). Hidde-sidahan waxaa la muujiyay inuu ku jiro plasmids kala duwan (100, 105, 115 ama 124 kb) bakteeriyada kala duwan ee ka kala yimid gobollo juqraafiyeed oo kala duwan (Parekh et al., 1995). Pseudomonas sp. C5pp, dhammaan hiddo-sidayaasha mas'uulka ka ah burburka carbaryl waxay ku yaalliin hidde-sidayaasha oo ku fidsan 76.3 kb oo taxane ah (Trivedi et al., 2016). Falanqaynta hidda-sidayaasha (6.15 Mb) waxay muujisay jiritaanka 42 MGE iyo 36 GEIs, kuwaas oo 17 MGEs ay ku yaalliin supercontig A (76.3 kb) oo leh celcelis ahaan G+C aan sinnayn (54–60 mol%), taasoo soo jeedinaysa dhacdooyin wareejin hidda-sidayaal toosan oo suurtagal ah (Trivedi et al., 2016). P. putida XWY-1 waxay muujinaysaa habayn la mid ah hiddo-sidayaasha burburiya carbaryl, laakiin hiddo-sidayaashani waxay ku yaalliin plasmid (Zhu et al., 2019).
Marka laga soo tago waxtarka dheef-shiid kiimikaadka heerarka bayoolaji iyo hidde-sideyaasha, noolaha yaryar waxay sidoo kale muujiyaan sifooyin ama jawaabo kale sida kiimikada kiimikada, sifooyinka wax ka beddelka dusha sare ee unugyada, qaybinta qaybaha, isticmaalka doorbidka, wax soo saarka bayoosurfactant, iwm., kuwaas oo ka caawiya inay si hufan u dheefshiido wasakhowga udgoonka ee deegaannada wasakhaysan (Jaantuska 7).
Jaantuska 7. Istaraatiijiyado kala duwan oo ka jawaab celinta unugyada ee bakteeriyada udgoon ee ku habboon ee burburisa haydaro-biyoodka si loogu helo burburka noolaha ee wax ku oolka ah ee isku-dhafka wasakhowga shisheeye.
Jawaabaha kiimikada waxaa loo arkaa inay yihiin arrimo kor u qaada burburka wasakhowga dabiiciga ah ee nidaamyada deegaanka ee si kala duwan u wasakhaysan. (2002) waxay muujiyeen in kiimikada kiimikada ee Pseudomonas sp. G7 ilaa naphthalene ay kordhisay heerka burburka naphthalene ee nidaamyada biyaha. Nooca duurjoogta ah ee G7 ayaa si aad uga dhakhso badan u burburiyay naphthalene marka loo eego nooca mutant-ka ee aan lahayn chemotaxis. Borotiinka NahY (538 amino acids oo leh topology xuubka) ayaa la ogaaday in lala qoray hiddo-wadaha marinnada metacleave ee ku yaal plasmid-ka NAH7, iyo sida kuwa loo yaqaan chemotaxis transducers, borotiinkani wuxuu u muuqdaa inuu u shaqeeyo sidii chemoreceptor si loo burburiyo naphthalene (Grimm iyo Harwood 1997). Daraasad kale oo ay sameeyeen Hansel et al. (2009) waxay muujiyeen in borotiinku uu yahay kemotactic, laakiin heerka burburkiisu uu sarreeyo. (2011) waxay muujiyeen jawaab celin kiimiko ah oo Pseudomonas (P. putida) u ah naphthalene gaseous, halkaas oo faafinta wejiga gaaska ay keentay qulqul joogto ah oo naphthalene ah oo unugyada ah, taas oo xakamaysay jawaabta kiimikada ee unugyada. Cilmi-baarayaashu waxay ka faa'iideysteen dhaqankan kiimikada si ay u farsameeyaan microbes kuwaas oo kor u qaadi doona heerka burburka. Daraasaduhu waxay muujiyeen in waddooyinka dareemayaasha kiimikada ay sidoo kale nidaamiyaan hawlaha kale ee unugyada sida qaybinta unugyada, nidaaminta wareegga unugyada, iyo sameynta biofilm, taasoo gacan ka geysaneysa xakamaynta heerka burburka. Si kastaba ha ahaatee, ka faa'iideysiga hantidan (chemotaxis) si loo helo burbur hufan waxaa caqabad ku ah dhowr caqabadood. Caqabadaha ugu waaweyn waa: (a) soo-dhoweeyayaasha paralogous ee kala duwan waxay aqoonsadaan isku-dhafka/ligands isku mid ah; (b) jiritaanka soo-dhoweeyayaasha kale, tusaale ahaan, tropism-ka firfircoon; (c) kala duwanaansho taxane ah oo muhiim ah oo ku jira qaybaha dareenka ee qoyska soo-dhoweeyaha isku midka ah; iyo (d) la'aanta macluumaadka ku saabsan borotiinnada dareemayaasha bakteeriyada ee waaweyn (Ortega et al., 2017; Martin-Mora et al., 2018). Mararka qaar, burburka bayoolajiyada ee haydaro-biyoodka udgoonku wuxuu soo saaraa metabolites/dhexdhexaadin badan, kuwaas oo laga yaabo inay yihiin kuwo kiimiko ah oo loogu talagalay hal koox oo bakteeriya ah laakiin u adkaysta kuwa kale, taasoo sii adkeynaysa habka. Si loo aqoonsado isdhexgalka ligands (hydrocarbons-ka udgoonka leh) iyo soo-dhoweeyayaasha kiimikada, waxaan dhisnay borotiinno dareemayaal isku-dhafan (PcaY, McfR, iyo NahY) annagoo isku darayna qaybaha dareemayaasha iyo calaamadaynta ee Pseudomonas putida iyo Escherichia coli, kuwaas oo bartilmaameedsada soo-dhoweeyayaasha asiidhyada udgoonka leh, dhexdhexaadiyeyaasha TCA, iyo naphthalene, siday u kala horreeyaan (Luu et al., 2019).
Iyada oo ay saameyn ku leedahay naphthalene iyo hydrocarbons-ka kale ee aromatic-ka ah (PAHs), qaab-dhismeedka xuubka bakteeriyada iyo hufnaanta noolaha yaryar ayaa isbeddello muhiim ah sameeya. Daraasaduhu waxay muujiyeen in naphthalene ay farageliso isdhexgalka silsiladda acyl iyada oo loo marayo isdhexgalka hydrophobic, taasoo kordhinaysa bararka iyo dareeraha xuubka (Sikkema et al., 1995). Si looga hortago saameyntan waxyeelada leh, bakteeriyadu waxay nidaamisaa dareeraha xuubka iyadoo beddeleysa saamiga iyo isku-dhafka asiidhka dufanka leh ee u dhexeeya asiidhada dufanka leh ee silsiladda laanta ah ee iso/anteiso iyo isomerizing asiidhyada dufanka leh ee cis-unsaturated ee isomers-ka trans-isomers ee u dhigma (Heipieper iyo de Bont, 1994). Pseudomonas stutzeri oo lagu beeray daaweynta naphthalene, saamiga asiidhka dufanka leh ee buuxa ilaa aan buuxin ayaa ka kordhay 1.1 ilaa 2.1, halka Pseudomonas JS150 saamigani uu ka kordhay 7.5 ilaa 12.0 (Mrozik et al., 2004). Markii lagu koray naphthalene, unugyada Achromobacter KAs 3–5 waxay muujiyeen isku-darka unugyada agagaarka kiristaalo naphthalene iyo hoos u dhac ku yimid dallacaadda dusha sare ee unugyada (laga bilaabo -22.5 ilaa -2.5 mV) oo ay weheliso isku-darka cytoplasmic iyo vacuolization, taasoo muujinaysa isbeddellada qaab-dhismeedka unugyada iyo sifooyinka dusha sare ee unugyada (Mohapatra et al., 2019). Inkasta oo isbeddellada unugyada/dusha sare ay si toos ah ula xiriiraan qaadashada wanaagsan ee wasakhowga udgoonka, istaraatiijiyadaha injineernimada bio ee khuseeya si fiican looma hagaajin. Wax ka qabashada qaabka unugyada si dhif ah ayaa loo isticmaalay si loo wanaajiyo hababka bayoolojiga (Volke iyo Nikel, 2018). Tirtiridda hiddo-wadaha saameeya kala-qaybinta unugyada waxay keentaa isbeddello ku yimaada qaab-dhismeedka unugyada. Tirtiridda hiddo-wadaha saameeya kala-qaybinta unugyada waxay keentaa isbeddello ku yimaada qaab-dhismeedka unugyada. Bacillus subtilis, borotiinka septum-ka unugyada SepF ayaa la muujiyay inuu ku lug leeyahay sameynta septum waxaana loo baahan yahay tallaabooyinka xiga ee kala-qaybinta unugyada, laakiin maaha hiddo-wade muhiim ah. Tirtiridda hiddo-sidayaasha codeeya peptide glycan hydrolases ee Bacillus subtilis waxay keentay fidinta unugyada, kordhinta heerka koritaanka gaarka ah, iyo hagaajinta awoodda wax soo saarka ensaymka (Cui et al., 2018).
Kala qaybsanaanta wadada burburka carbaryl ayaa la soo jeediyay si loo gaaro burbur hufan oo noocyada Pseudomonas C5pp iyo C7 ah (Kamini et al., 2018). Waxaa la soo jeediyay in carbaryl loo qaado booska periplasmic iyada oo loo marayo septum-ka xuubka dibadda iyo/ama iyada oo loo marayo porins-ka faafin kara. CH waa ensaym periplasmic ah oo kiciya hydrolysis-ka carbaryl ilaa 1-naphthol, kaas oo ah mid aad u deggan, hydrophobic badan oo sun badan. CH waxay ku taal periplasm waxayna leedahay xiriir hoose oo carbaryl ah, sidaas darteed waxay xakameysaa sameynta 1-naphthol, taasoo ka hortageysa ururinta unugyada iyo yaraynta sunta unugyada (Kamini et al., 2018). 1-naphthol ee ka dhasha waxaa loo qaadaa cytoplasm-ka xuubka gudaha iyadoo la kala qaybinayo iyo/ama faafin, ka dibna waxaa loo hydroxylase gareeyaa 1,2-dihydroxynaphthalene iyadoo la adeegsanayo enzyme-ka aadka u sarreeya 1NH si loogu sameeyo dheef-shiid kiimikaad dheeraad ah wadada kaarboonka dhexe.
In kasta oo noolaha yaryar ay leeyihiin awoodo hidde iyo dheef-shiid kiimikaad si ay u burburiyaan ilaha kaarboonka xenobiotic, qaab-dhismeedka kala-sareynta ee isticmaalkooda (tusaale ahaan, isticmaalka doorbidka ah ee ilaha kaarboonka fudud marka loo eego kuwa adag) waa caqabad weyn oo ku timaadda burburka noolaha. Joogitaanka iyo isticmaalka ilaha kaarboonka fudud waxay hoos u dhigtaa hiddo-sidayaasha codeeya ensaymada kuwaas oo burburiya ilaha kaarboonka ee adag/aan la doorbidin sida PAHs. Tusaale si fiican loo bartay ayaa ah marka gulukooska iyo lactose si wadajir ah loogu quudiyo Escherichia coli, gulukooska waxaa loo isticmaalaa si ka hufan lactose (Jacob iyo Monod, 1965). Pseudomonas waxaa la soo sheegay inay burburiyaan noocyo kala duwan oo PAHs ah iyo isku-darka xenobiotic sida ilaha kaarboonka. Kala-sareynta isticmaalka ilaha kaarboonka ee Pseudomonas waa asiidhyo dabiici ah > glucose > isku-darka aromatic (Hylemon iyo Phibbs, 1972; Collier et al., 1996). Si kastaba ha ahaatee, waxaa jira wax ka reeban. Waxaa xiiso leh, Pseudomonas sp. CSV86 waxay muujisaa qaab-dhismeed gaar ah oo kala sarreyn ah kaas oo si gaar ah u isticmaala hydrocarbons-ka udgoonka leh (benzoic acid, naphthalene, iwm.) halkii laga isticmaali lahaa gulukooska oo si wadajir ah u dheefshiida hydrocarbons-ka udgoonka leh asiidhyada dabiiciga ah (Basu et al., 2006). Bakteeriyadan, hiddo-sidayaasha burburka iyo gaadiidka hydrocarbons-ka udgoonka ah hoos looma sameeyo xitaa marka ay joogaan ilo kaarboon labaad sida gulukooska ama asiidhyada dabiiciga ah. Markii lagu koray gulukooska iyo hydrocarbons-ka udgoonka leh, waxaa la arkay in hiddo-sidayaasha gaadiidka gulukooska iyo dheef-shiid kiimikaadka la dhimay, hydrocarbons-ka udgoonka leh ayaa la isticmaalay marxaladda koowaad ee log, gulukooskana waxaa loo isticmaalay marxaladda labaad ee log (Basu et al., 2006; Choudhary et al., 2017). Dhanka kale, joogitaanka asiidhyada dabiiciga ah ma saameynin muujinta dheef-shiid kiimikaadka hydrocarbon-ka udgoonka leh, sidaa darteed bakteeriyadan waxaa la filayaa inay noqoto nooc musharrax u ah daraasadaha burburka bayoolojiga (Phale et al., 2020).
Waa la ogyahay in isbeddelka bayoolojiga ee hydrocarbon uu sababi karo walbahaar oksaydh iyo kor u kaca ensaymada antioxidant-ka ee ku jira noolaha yaryar. Bayoolojiga aan waxtarka lahayn ee naphthalene labadaba unugyada marxaladda taagan iyo joogitaanka iskudhisyada sunta ah waxay horseedaa sameynta noocyada oksijiinta ee falcelinta leh (ROS) (Kang et al. 2006). Maadaama ensaymada naphthalene-degrading ay ka kooban yihiin kooxo bir-sulfur ah, cadaadiska oksaydhka hoostiisa, birta ku jirta heme iyo borotiinnada birta-sulfur ayaa la oksaydhin doonaa, taasoo horseedaysa joojinta borotiinka. Ferredoxin-NADP+ reductase (Fpr), oo ay weheliso superoxide dismutase (SOD), waxay dhexdhexaadisaa falcelinta redox ee la rogi karo ee u dhaxaysa NADP+/NADPH iyo laba molecule oo ferredoxin ah ama flavodoxin ah, sidaas darteedna waxay baabi'inaysaa ROS waxayna soo celinaysaa xarunta birta-sulfur-ka iyadoo la raacayo cadaadiska oksaydhka (Li et al. 2006). Waxaa la soo sheegay in Fpr iyo SodA (SOD) labadaba ee Pseudomonas ay sababi karto walbahaarka oksaydhka, iyo kororka hawlaha SOD iyo catalase ayaa lagu arkay afar nooc oo Pseudomonas ah (O1, W1, As1, iyo G1) inta lagu jiro koritaanka xaaladaha lagu daray naphthalene (Kang et al., 2006). Daraasaduhu waxay muujiyeen in ku darista antioxidants sida ascorbic acid ama iron ferrous (Fe2+) ay kordhin karto heerka koritaanka naphthalene. Markii Rhodococcus erythropolis uu ku koray bartamaha naphthalene, qoraalka hiddo-sideyaasha cytochrome P450 ee la xiriira walbahaarka oksaydhka oo ay ku jiraan sodA (Fe/Mn superoxide dismutase), sodC (Cu/Zn superoxide dismutase), iyo recA ayaa la kordhiyay (Sazykin et al., 2019). Falanqaynta proteomic-ka ee isbarbardhigga ah ee unugyada Pseudomonas ee lagu beeray naphthalene waxay muujisay in kor u qaadista borotiinnada kala duwan ee la xiriira jawaabta walbahaarka oksaydhka ay tahay istaraatiijiyad la qabsiga walbahaarka (Herbst et al., 2013).
Noolaha yaryar ayaa la soo sheegay inay soo saaraan noolaha noolaha iyadoo la adeegsanayo ilaha kaarboonka ee hydrophobic. Sunta-saareyaashan waa isku-dhisyo firfircoon oo dusha sare ah oo samayn kara isku-darka biyaha-saliidda ama biyaha-hawada. Tani waxay kor u qaadaysaa milmida beenta ah waxayna sahlaysaa nuugista haydaro-biyoodka udgoonka leh, taasoo keenta burburka noolaha oo hufan (Rahman et al., 2002). Sababo la xiriira sifooyinkan, noolaha noolaha waxaa si weyn loogu isticmaalaa warshado kala duwan. Ku darista sunta-biyoodka kiimikada ama noolaha noolaha ee dhaqamada bakteeriyada waxay kor u qaadi kartaa hufnaanta iyo heerka burburka haydaro-biyoodka. Qaybaha sunta-biyoodka, rhamnolipids-ka ay soo saarto Pseudomonas aeruginosa si ballaaran ayaa loo bartay oo loo sifeeyay (Hisatsuka et al., 1971; Rahman et al., 2002). Intaa waxaa dheer, noocyada kale ee bayosurfactants waxaa ka mid ah lipopeptides (mucins ka yimid Pseudomonas fluorescens), emulsifier 378 (ka yimid Pseudomonas fluorescens) (Rosenberg iyo Ron, 1999), lipids trehalose disaccharide ka yimid Rhodococcus (Ramdahl, 1985), lichenin ka yimid Bacillus (Saraswathy iyo Hallberg, 2002), iyo surfactant ka yimid Bacillus subtilis (Siegmund iyo Wagner, 1991) iyo Bacillus amyloliquefaciens (Zhi et al., 2017). Surfactants-kan xooggan ayaa la muujiyay inay yareeyaan xiisadda dusha sare laga bilaabo 72 dynes/cm ilaa wax ka yar 30 dynes/cm, taasoo u oggolaanaysa nuugista hydrocarbon oo wanaagsan. Waxaa la soo sheegay in Pseudomonas, Bacillus, Rhodococcus, Burkholderia iyo noocyada kale ee bakteeriyada ay soo saari karaan noocyo kala duwan oo ah biosurfactants ku salaysan rhamnolipid iyo glycolipid marka lagu beero warbaahinta naphthalene iyo methylnaphthalene (Kanga et al., 1997; Puntus et al., 2005). Pseudomonas maltophilia CSV89 waxay soo saari kartaa biosurfactant-ka ka baxsan unugyada Biosur-Pm marka lagu beero iskudhisyo udgoon sida naphthoic acid (Phale et al., 1995). Kinetics-ka sameynta Biosur-Pm waxay muujisay in isku-darka uu yahay geedi socod korriin iyo pH ku xiran. Waxaa la ogaaday in xaddiga Biosur-Pm ee ay soo saaraan unugyada pH-ga dhexdhexaadka ah uu ka sarreeyay kan pH 8.5. Unugyada lagu beero pH 8.5 waxay ahaayeen kuwo hydrophobic badan waxayna lahaayeen xiriir sare oo u dhexeeya iskudhisyo udgoon iyo aliphatic marka loo eego unugyada lagu beero pH 7.0. Rhodococcus spp. N6, saamiga kaarboonka sare ilaa nitrogen (C:N) iyo xaddidaadda birta ayaa ah xaaladaha ugu habboon ee soo saarista bayosurfactants-ka ka baxsan jirka (Mutalik et al., 2008). Waxaa la sameeyay isku day lagu horumarinayo bayosurfactants-ka (surfactants) iyadoo la hagaajinayo noocyada iyo halsano. Si kastaba ha ahaatee, titer-ka surfactant-ka ee ku jira habka dhaqanka waa mid hooseeya (1.0 g/L), taas oo caqabad ku ah wax soo saarka baaxadda weyn (Jiao et al., 2017; Wu et al., 2019). Sidaa darteed, hababka injineernimada hidde-sideyaasha ayaa loo isticmaalay in lagu hagaajiyo bayosurthesis-keeda. Si kastaba ha ahaatee, wax ka beddelka injineerinteeda way adag tahay sababtoo ah cabbirka weyn ee operon-ka (qiyaastii 25 kb) iyo nidaaminta bayosurthetic-ga ee adag ee nidaamka dareenka quorum (Jiao et al., 2017; Wu et al., 2019). Tiro isbeddello ah oo ku saabsan injineernimada hidde-sidaha ayaa lagu sameeyay bakteeriyada Bacillus, oo inta badan loogu talagalay in lagu kordhiyo wax soo saarka surfactin iyadoo la beddelayo promoter-ka (srfA operon), iyadoo si xad dhaaf ah loo muujiyo borotiinka dhoofinta surfactin YerP iyo arrimaha sharciyeynta ComX iyo PhrC (Jiao et al., 2017). Si kastaba ha ahaatee, hababkan injineernimada hidde-sidaha waxay gaareen hal ama dhowr isbeddel hidde-sidaha ah oo keliya mana aysan gaarin wax soo saar ganacsi. Sidaa darteed, daraasad dheeraad ah oo ku saabsan hababka hagaajinta aqoonta ku salaysan ayaa lagama maarmaan ah.
Daraasadaha PAH ee burburka bayoolojiga waxaa inta badan lagu sameeyaa xaaladaha caadiga ah ee shaybaarka. Si kastaba ha ahaatee, goobaha wasakhaysan ama deegaannada wasakhaysan, arrimo badan oo aan dabiici ahayn iyo bayooloji (heerkulka, pH, oksijiin, helitaanka nafaqada, helitaanka bayoolojiga ee substrate-ka, xenobiotics kale, joojinta badeecada dhammaadka, iwm.) ayaa la muujiyay inay beddelaan oo saameeyaan awoodda burburka ee noolaha yaryar.
Heerkulku wuxuu saameyn weyn ku leeyahay burburka PAH. Marka heerkulku kordho, fiirsashada oksijiinta ee milmi karta ayaa hoos u dhacda, taas oo saameysa dheef-shiid kiimikaadka noolaha aerobic-ga, maadaama ay u baahan yihiin oksijiin molecular oo ah mid ka mid ah substrate-yada oksijiinta ee fuliya falgallada hydroxylation ama ring cleavage. Badanaa waxaa la ogaadaa in heerkulka sare uu u beddelo PAH-yada waalidka ah iskudhisyo sun ah, taasoo joojinaysa burburka noolaha (Muller et al., 1998).
Waxaa la ogaaday in goobo badan oo wasakhaysan oo PAH ah ay leeyihiin qiimayaal pH oo aad u daran, sida goobaha wasakhaysan ee daadinta aashitada (pH 1–4) iyo goobaha gaaska dabiiciga ah/dhaxalka ee lagu wasakheeyay alkaline leachate (pH 8–12). Xaaladahani waxay si xun u saameyn karaan habka burburka bayoolojiga. Sidaa darteed, ka hor inta aan la isticmaalin noolaha noolaha ah ee daaweynta bayoolojiga, waxaa lagu talinayaa in la hagaajiyo pH iyadoo lagu darayo kiimikooyin ku habboon (oo leh awood dhexdhexaad ah ilaa mid aad u hooseeya oo yareynaysa oksaydhka) sida ammonium sulfate ama ammonium nitrate carrada alkaline ama xaddididda kalsiyum carbonate ama magnesium carbonate meelaha aashitada leh (Bowlen et al. 1995; Gupta iyo Sar 2020).
Sahayda oksijiinta ee aagga ay dhibaatadu saameysey waa qodobka xaddidaya heerka wasakhowga PAH. Sababtoo ah xaaladaha redox ee deegaanka, hababka hagaajinta bioredition-ka ee goobta waxay badanaa u baahan yihiin soo bandhigid oksijiin ah oo ka timaadda ilo dibadda ah (qolitaanka, nuugista hawada, iyo ku darista kiimikada) (Pardieck et al., 1992). Odenkranz et al. (1996) waxay muujiyeen in ku darista magnesium peroxide (isku-darka sii daaya oksijiinta) ee biyo-mareenka wasakhaysan ay si wax ku ool ah u hagaajin karto isku-dhafka BTEX. Daraasad kale ayaa baartay burburka in goobta ee phenol iyo BTEX ee biyo-mareenka wasakhaysan iyadoo la durayo sodium nitrate iyo dhisidda ceelasha soo saarista si loo gaaro daaweyn bioredition oo wax ku ool ah (Bewley iyo Webb, 2001).