Iyi nkuru ni igice cy'insanganyamatsiko y'ubushakashatsi igira iti “Ikoranabuhanga rigezweho ryo gutunganya ibintu bikomoka ku binyabuzima n'uburyo bwo kongera gukoresha ibintu bikomoka ku bimera (SOC). Reba ingingo zose 14
Karubone zikozwe muri polycyclic aromatic hydrocarbons (PAHs) zifite uburemere buke bwa molekile nka naphthalene na naphthalene zisimbuwe (methylnaphthalene, aside naphthoic, 1-naphthyl-N-methylcarbamate, nibindi) zikoreshwa cyane mu nganda zitandukanye kandi zitera uburozi mu miterere y’ibinyabuzima, zihindura imiterere y’ibinyabuzima cyangwa zitera kanseri. Izi organic compounds (SOCs) cyangwa xenobiotics zifatwa nk'ibintu byangiza ibidukikije kandi bibangamira cyane ibidukikije ku isi no ku buzima rusange. Ubukana bw'ibikorwa bya muntu (urugero: gukwirakwiza imyuka y'amakara, gutunganya peteroli, imyuka ihumanya ikirere n'ikoreshwa mu buhinzi) bigena ubwinshi, iherezo n'ubwikorezi bw'izi chemicals zikunze kugaragara kandi zihoraho. Uretse uburyo bwo kuvura/gukuraho ibidukikije n'imiti, ikoranabuhanga rirengera ibidukikije nk'ubuvuzi bw'ibinyabuzima, bukoresha udukoko dushobora kwangiza burundu POCs cyangwa kuzihinduramo ibintu bitari uburozi, byagaragaye nk'uburyo bwizewe, buhendutse kandi butanga icyizere. Amoko atandukanye ya bagiteri ari mu bwoko bwa phyla Proteobacteria (Pseudomonas, Pseudomonas, Comamonas, Burkholderia, na Neosphingobacterium), Firmicutes (Bacillus na Paenibacillus), na Actinobacteria (Rhodococcus na Arthrobacter) mu butaka bw’ibinyabuzima byagaragaje ubushobozi bwo kwangiza ibintu bitandukanye by’ibinyabuzima. Inyigo za metabolike, genomics, n’isesengura rya metagenomike bidufasha gusobanukirwa imiterere y’imiterere n’ubusumbane bw’ibinyabuzima biri muri ubu bwoko bw’ibinyabuzima byoroshye, bishobora kurushaho gukoreshwa mu kwangiza ibinyabuzima neza. Kubaho igihe kirekire kwa PAH kwatumye habaho uburyo bushya bwo kwangirika binyuze mu kwimura gene mu buryo butambitse hakoreshejwe ibintu bya gene nka plasmids, transposons, bacteriophages, ibirwa bya genomike, n’ibintu bihuza ibinyabuzima. Ubuzima bw’ibinyabuzima n’ikoranabuhanga rya gene ry’ibintu byihariye cyangwa imiterere y’uturemangingo (consortia) bishobora gutuma habaho uburyo bwo kuvura ibinyabuzima by’ibimera byihariye, byihuse kandi neza binyuze mu ngaruka z’ubufatanye. Muri iri suzuma, twibanda ku nzira zitandukanye z’imikorere y’uturemangingo n’ubutandukanye, imiterere n’ubutandukanye bw’uturemangingo, hamwe n’uburyo uturemangingo twitabira/twihinduranya naphthalene hamwe na bagiteri zangiza naphthalene. Ibi bizatanga amakuru ku bidukikije kugira ngo bikoreshwe mu murima no kunoza uburyo ibintu bimera neza.
Iterambere ryihuse ry’inganda (ibikomoka kuri peteroli, ubuhinzi, imiti, amarangi y’imyenda, amavuta yo kwisiga, nibindi) ryagize uruhare mu iterambere ry’ubukungu bw’isi no kunoza imibereho. Iri terambere ryimbitse ryatumye habaho umubare munini w’ibinyabutabire by’umwimerere (SOCs), bikoreshwa mu gukora ibintu bitandukanye. Ibi binyabutabire by’amahanga cyangwa SOC birimo polycyclic aromatic hydrocarbons (PAHs), imiti yica udukoko, imiti yica ibyatsi, plasticizers, amarangi, imiti, organophosphates, ibibuza umuriro, ibintu bishongesha ibinyabuzima bihindagurika, nibindi. Bishyirwa mu kirere, mu mazi no ku isi aho bigira ingaruka nyinshi, bigatera ingaruka mbi ku binyabuzima bitandukanye binyuze mu guhindura imiterere ya fizikikemike n’imiterere y’umuryango (Petrie et al., 2015; Bernhardt et al., 2017; Sarkar et al., 2020). Imyuka myinshi ihumanya ikirere igira ingaruka zikomeye kandi zangiza ku rusobe rw'ibinyabuzima byinshi bidafite aho bihuriye/ahantu hakunze kugaragara urusobe rw'ibinyabuzima (urugero: amabuye ya korali, urubura rwa Arctique/Antarctica, ibiyaga birebire byo mu misozi miremire, ibisigazwa byo mu nyanja nini, nibindi) (Jones 2010; Beyer et al. 2020; Nordborg et al. 2020). Ubushakashatsi buherutse gukorwa ku binyabuzima bwagaragaje ko gushyira ibintu bikomoka ku binyabuzima (urugero: imyanda ihumanya ikirere) n'ibikomoka kuri byo ku buso bw'inyubako z'ubukorano (ibidukikije byubatswe) (urugero: ahantu h'umurage w'umuco n'inzibutso zikozwe mu mabuye ya granite, amabuye, ibiti n'ibyuma) byihutisha kwangirika kwabyo (Gadd 2017; Liu et al. 2018). Ibikorwa bya muntu bishobora kongera no kwangiza ibidukikije by'inyubako n'inyubako binyuze mu kwanduza ikirere n'imihindagurikire y'ikirere (Liu et al. 2020). Iyi myanda ihumanya ikirere ihura n'umwuka w'amazi mu kirere ikagwa ku nyubako, bigatuma ibintu byangirika mu buryo bw'umubiri no mu buryo bwa shimi. Kwangirika kw'ibinyabuzima bizwi cyane nk'impinduka zidakwiye mu miterere n'imiterere y'ibintu biterwa n'ibinyabuzima bigira ingaruka ku kubungabungwa kwabyo (Pochon na Jaton, 1967). Ingaruka nyinshi ku mikorobe (metabolism) z'ibi bintu zishobora kugabanya ubuziranenge bw'imiterere, imikorere myiza yo kubungabunga no kurengera ibidukikije n'agaciro k'umuco (Gadd, 2017; Liu et al., 2018). Ku rundi ruhande, mu bihe bimwe na bimwe, guhuza mikorobe no guhangana n'ibi bintu byagaragaye ko ari ingirakamaro kuko bikora biofilms n'ibindi bice by'umubiri birinda bigabanya umuvuduko wo kwangirika/kubora (Martino, 2016). Kubwibyo, guteza imbere ingamba zirambye zo kubungabunga amabuye, ibyuma n'ibiti bisaba gusobanukirwa neza inzira z'ingenzi zikoreshwa muri iyi nzira. Ugereranije n'imikorere karemano (imikorere y'ubutaka, inkongi z'umuriro mu mashyamba, iruka ry'ibirunga, ingaruka z'ibimera na bagiteri), ibikorwa by'abantu bituma habaho kurekurwa kwa hydrocarboni nyinshi za polycyclic aromatic (PAHs) n'izindi karuboni organic (OC) mu rusobe rw'ibinyabuzima. Ingano nyinshi za PAH zikoreshwa mu buhinzi (imiti yica udukoko n'imiti yica udukoko nka DDT, atrazine, carbaryl, pentachlorophenol, nibindi), inganda (amavuta mabi, imyanda y'amavuta, pulasitiki ikomoka kuri peteroli, PCB, pulasitiki, isabune, imiti yica udukoko, imiti ihumura neza n'imiti irinda ubushyuhe), ibikoresho byo kwita ku buzima bwite (imiti irinda izuba, imiti yica udukoko, imiti yica udukoko n'imizi ya polycyclic) n'ibisasu (ibisasu nka 2,4,6-TNT) ni ibinyabuzima bishobora kugira ingaruka ku buzima bw'isi (Srogi, 2007; Vamsee-Krishna na Phale, 2008; Petrie et al., 2015). Uru rutonde rushobora kwaguka rukubiyemo ibintu bikomoka kuri peteroli (amavuta ya lisansi, amavuta, asphaltenes), bioplastics ifite uburemere bwinshi, n'amazi ya ionic (Amde et al., 2015). Imbonerahamwe ya 1 igaragaza imyuka ihumanya ikirere itandukanye n'ikoreshwa ryayo mu nganda zitandukanye. Mu myaka ya vuba aha, imyuka ihumanya ikirere ituruka ku binyabutabire bihinduka, kimwe na dioxyde de carbone n'indi myuka ihumanya ikirere, yatangiye kwiyongera (Dvorak et al., 2017). Ariko, ingaruka zituruka ku binyabutabire ziruta cyane izisanzwe. Byongeye kandi, twasanze hari ubwoko bwinshi bwa SOC bukomeje kuba mu bidukikije byinshi kandi byagaragaye ko ari imyuka ihumanya ikirere igaragara igira ingaruka mbi ku binyabuzima (Ishusho ya 1). Inzego zishinzwe ibidukikije nka Leta Zunze Ubumwe za Amerika zishinzwe kurengera ibidukikije (USEPA) zashyize byinshi muri ibi binyabutabire ku rutonde rw'ibintu byangiza ibidukikije bitewe n'imiterere yabyo yangiza ibidukikije, uburozi bwa genoside, uburozi bwa mutagenic, na kanseri. Kubwibyo, hakenewe amabwiriza akomeye yo guta imyanda n'ingamba nziza zo gutunganya/gukura imyanda mu bidukikije byanduye. Uburyo butandukanye bwo kuvura ibidukikije n'imiti nka pyrolysis, oxidative thermal treatment, air emission, purchasing imyanda, incineration, nibindi ntacyo bumaze kandi buhenze kandi butanga ibintu byangiza, uburozi kandi bigoye kubitunganya. Kubera ko ubumenyi ku bidukikije bukomeje kwiyongera ku isi, udukoko dushobora kwangiza iyi myanda n'ibiyikomokaho (nk'udukoko twa halogenated, nitro, alkyl na/cyangwa methyl) turimo gukurura abantu benshi (Fennell et al., 2004; Haritash na Kaushik, 2009; Phale et al., 2020; Sarkar et al., 2020; Schwanemann et al., 2020). Gukoresha utu dukoko gakondo gusa cyangwa mu moko atandukanye (amakoloni) mu gukuraho imyanda ihumura bifite inyungu mu bijyanye n'umutekano w'ibidukikije, ikiguzi, imikorere myiza, imikorere myiza, no kuramba. Abashakashatsi barimo kandi gusuzuma uburyo bwo guhuza imikorobe n'uburyo bwa electrochemical redox, ari bwo sisitemu za bioelectrochemical (BES), nk'ikoranabuhanga ryiza ryo kuvura/gukuraho imyanda ihumanya (Huang et al., 2011). Ikoranabuhanga rya BES ryakuruye ibitekerezo byinshi bitewe n’ubushobozi bwaryo bwo hejuru, igiciro gito, umutekano w’ibidukikije, imikorere y’ubushyuhe bw’icyumba, ibikoresho bihuye n’ibinyabuzima, ndetse n’ubushobozi bwo kugarura umusaruro w’agaciro (urugero: amashanyarazi, lisansi, n’ibinyabutabire) (Pant et al., 2012; Nazari et al., 2020). Kuza kw’uburyo bwo gusesengura genome bufite ubushobozi bwo hejuru n’ibikoresho/uburyo bwa omics byatanze amakuru mashya menshi ku miterere y’uturemangingo, proteomics, na fluxomics y’imikorere y’uturemangingo dutandukanye twononekara. Guhuza ibi bikoresho n’ibinyabuzima by’imiti byarushijeho kunoza uburyo dusobanukirwa guhitamo no gutunganya inzira za katabolic zigamije muri utwo turemangingo (ni ukuvuga imiterere ya metabolic) kugira ngo tugere ku kubora neza kandi ku buryo bunoze. Kugira ngo dushyireho ingamba nziza zo kuvura bioremediation hakoreshejwe utwo turemangingo dukwiriye, tugomba gusobanukirwa ubushobozi bwa biochemical, ubwoko butandukanye bwa metabolic, imiterere ya genetic, na ecology (autoecology/synecology) ya utwo turemangingo.
Ishusho ya 1. Inkomoko n'inzira za PAH zifite molekile nkeya zinyura mu bidukikije bitandukanye n'ibintu bitandukanye bigira ingaruka ku binyabuzima. Imirongo ihanamye igaragaza imikoranire hagati y'ibintu bigize urusobe rw'ibinyabuzima.
Muri iri suzuma, twagerageje gusobanura muri make amakuru yerekeye kwangirika kwa PAH zoroshye nka naphthalene na naphthalene zasimbuwe na bagiteri zitandukanye zikubiyemo inzira za metabolisme n'ubutandukanye, enzymes zigira uruhare mu kwangirika, imiterere/ibirimo bya gene n'ubutandukanye, ibisubizo by'uturemangingo n'ibindi bintu bitandukanye byo kuvura indwara. Gusobanukirwa urwego rwa biochemical na molekile bizafasha mu kumenya ubwoko bukwiye bw'ibinyabuzima n'ubuhanga bwazo mu bijyanye no kuvura neza indwara nk'izo z'ingenzi. Ibi bizafasha mu gushyiraho ingamba zo gushyiraho itsinda ry'udukoko twihariye mu rwego rwo kuvura indwara.
Kuba hari umubare munini w'ibintu bihumura nabi kandi biteje akaga (bikurikije itegeko rya Huckel 4n + 2π electrons, n = 1, 2, 3, …) biteza akaga gakomeye ku bintu bitandukanye bidukikije nk'umwuka, ubutaka, imyanda, n'ubuso n'amazi yo munsi y'ubutaka (Puglisi et al., 2007). Ibi bintu bifite impeta imwe ya benzene (monocyclic) cyangwa impeta nyinshi za benzene (polycyclic) zitondetse mu buryo bugororotse, buto cyangwa bugizwe n'amatsinda kandi bigaragaza ituze (ituze/ihungabana) mu bidukikije bitewe n'ingufu nyinshi za resonance mbi n'ubudahangarwa (inertness), bishobora gusobanurwa no kudahuza amazi n'imiterere yabyo. Iyo impeta ihumura isigaye isimbuzwa na methyl (-CH3), carboxyl (-COOH), hydroxyl (-OH), cyangwa sulfonate (-HSO3), irushaho gukomera, ikagira uruhare rukomeye mu macromolecules, kandi ikomeza kwiyongera mu buryo bw'ibinyabuzima (Seo et al., 2009; Phale et al., 2020). Hari hydrocarbons nke za polycyclic aromatic hydrocarbons (LMWAHs) zifite uburemere buke bwa molekile, nka naphthalene n'ibiyikomokaho [methylnaphthalene, aside naphthoic, naphthalenesulfonate, na 1-naphthyl N-methylcarbamate (carbaryl)], zashyizwe ku rutonde rw'ibintu bihumanya ibidukikije by'ibanze n'Ikigo cya Amerika gishinzwe kurengera ibidukikije nk'ibihumanya ibidukikije, bihindura imiterere y'ibinyabuzima, na/cyangwa bitera kanseri (Cerniglia, 1984). Gusohora uru rwego rwa NM-PAH mu bidukikije bishobora gutuma ibi bintu byiyongera mu buryo bw’umubiri ku nzego zose z’uruhererekane rw’ibiribwa, bityo bikagira ingaruka ku buzima bw’urusobe rw’ibinyabuzima (Binkova et al., 2000; Srogi, 2007; Quinn et al., 2009).
Inkomoko n'inzira za PAH zijya kuri biota ahanini ni ukwimuka no gukorana hagati y'ibice bitandukanye by'urusobe rw'ibinyabuzima nk'ubutaka, amazi yo munsi y'ubutaka, amazi yo hejuru, ibihingwa n'ikirere (Arey na Atkinson, 2003). Ishusho ya 1 igaragaza imikoranire n'ikwirakwizwa rya PAH zitandukanye zifite uburemere buke mu rusobe rw'ibinyabuzima n'inzira zazo zijya kuri biota/abantu. PAH zibikwa ku buso bitewe n'ihumana ry'ikirere no mu kwimuka (gutemba) kw'ibyuka bihumanya ikirere, imyuka ihumanya ikirere mu nganda (gutwika no gukora coke) no kuzishyira mu bikorwa. Ibikorwa by'inganda nko gukora imyenda, amarangi n'irangi by'ubukorano; kubungabunga ibiti; gutunganya kabure; ibikorwa byo gukora sima; gukora imiti yica udukoko; no gukoresha mu buhinzi ni isoko nyamukuru ya PAH mu buryo bw'ubutaka no mu mazi (Bamforth na Singleton, 2005; Wick et al., 2011). Ubushakashatsi bwagaragaje ko ubutaka bwo mu nkengero z'umujyi n'imijyi, hafi y'imihanda minini, no mu mijyi minini bukunze kwibasirwa cyane n'ibyuka bihumanya ikirere (PAHs) bitewe n'imyuka iva mu nganda z'amashanyarazi, ubushyuhe bwo mu ngo, ikirere n'imihanda, ndetse n'ibikorwa by'ubwubatsi (Suman et al., 2016). (2008) bwagaragaje ko PAH mu butaka hafi y'imihanda muri New Orleans, Louisiana, muri Amerika zari 7189 μg/kg, mu gihe ahantu hafunguye, zari 2404 μg/kg gusa. Mu buryo nk'ubwo, urugero rwa PAH ruri hejuru ya 300 μg/kg rwagaragaye mu turere twegereye ahantu hakoreshwa gaze y'amakara mu mijyi myinshi ya Amerika (Kanaly na Harayama, 2000; Bamforth na Singleton, 2005). Ubutaka buturuka mu mijyi itandukanye yo mu Buhinde nka Delhi (Sharma et al., 2008), Agra (Dubey et al., 2014), Mumbai (Kulkarni na Venkataraman, 2000) na Visakhapatnam (Kulkarni et al., 2014) byavuzwe ko burimo ibipimo byinshi bya PAH. Ibinyabutabire bihumura byoroshye kwinjizwa ku duce tw’ubutaka, ibintu bikomoka ku bimera n’imyunyu ngugu y’ibumba, bityo bikaba ibintu bikomeye bishongera karuboni mu rusobe rw’ibinyabuzima (Srogi, 2007; Peng et al., 2008). Inkomoko nyamukuru ya PAH mu rusobe rw’ibinyabuzima byo mu mazi ni imvura (imvura itose/yumye n’umwuka w’amazi), amazi atemba mu mijyi, amazi yanduye, amazi yo munsi y’ubutaka n’ibindi (Srogi, 2007). Bivugwa ko hafi 80% bya PAH mu rusobe rw’ibinyabuzima byo mu mazi bikomoka ku imvura, gushonga kw’amazi, no gusohora imyanda (Motelay-Massei et al., 2006; Srogi, 2007). Ubwinshi bw'amazi ya PAH mu mazi yo hejuru cyangwa aturuka ahantu ho kumena imyanda ikomeye amaherezo agwa mu mazi yo munsi y'ubutaka, bigatera ikibazo gikomeye ku buzima bw'abaturage kuko abaturage barenga 70% bo muri Aziya y'Amajyepfo n'Amajyepfo y'Uburasirazuba banywa amazi yo munsi y'ubutaka (Duttagupta et al., 2019). Ubushakashatsi buherutse gukorwa na Duttagupta et al. (2020) ku migezi (32) n'amazi yo munsi y'ubutaka (235) bwakorewe muri West Bengal, mu Buhinde, bwagaragaje ko abaturage bo mu mijyi bagera kuri 53% na 44% by'abaturage bo mu cyaro (abaturage miliyoni 20 bose hamwe) bashobora guhura na naphthalene (4.9–10.6 μg/L) n'ibikomoka kuri yo. Imiterere itandukanye y'imikoreshereze y'ubutaka n'ubwiyongere bw'amazi yo munsi y'ubutaka bifatwa nk'ibintu by'ingenzi bigenzura ubwikorezi bw'amazi ya PAH afite uburemere buke mu mazi yo munsi y'ubutaka. Amazi yavuye mu buhinzi, amazi yavuye mu mihanda no mu nganda, ndetse n'imyanda yavuye mu myanda/imyanda byagaragaye ko bigizweho ingaruka na PAH mu bibaya by'imigezi n'ibisigazwa byo munsi y'ubutaka. Imvura igwa mu kirere yongera umwanda wa PAH. Ingano nini ya PAH n'ibikomoka kuri alkyl (51 muri rusange) byagaragaye mu migezi/amazi ku isi, nko mu ruzi rwa Fraser, uruzi rwa Louan, uruzi rwa Denso, uruzi rwa Missouri, uruzi rwa Anacostia, uruzi rwa Ebro, n'uruzi rwa Delaware (Yunker et al., 2002; Motelay-Massei et al., 2006; Li et al., 2010; Amoako et al., 2011; Kim et al., 2018). Mu myanda yo mu kibaya cy'uruzi rwa Ganges, naphthalene na phenanthrene byagaragaye ko ari byo bifite akamaro kanini (byabonetse muri 70% by'ibipimo) (Duttagupta et al., 2019). Byongeye kandi, ubushakashatsi bwagaragaje ko chlorine mu mazi yo kunywa ishobora gutuma habaho PAH zifite ogisijeni n'chlorine nyinshi (Manoli na Samara, 1999). PAH zirundanya mu binyampeke, imbuto n'imboga bitewe n'uko ibimera bitwarwa n'ubutaka bwanduye, amazi yo mu butaka n'imvura (Fismes et al., 2002). Udukoko twinshi two mu mazi nk'amafi, imineke, clams na shrimps byandujwe na PAHs binyuze mu kurya ibiryo byanduye n'amazi yo mu nyanja, ndetse no mu ruhu (Mackay na Fraser, 2000). Uburyo bwo guteka/gutunganya nko guteka, gukaranga, kunywesha itabi, gukaranga, kumisha, guteka no guteka amakara nabyo bishobora gutuma PAHs nyinshi mu biribwa ziboneka. Ibi biterwa ahanini n'amahitamo y'ibikoresho byo kunywa itabi, ingano ya phenolic/aromatic hydrocarbon, uburyo bwo guteka, ubwoko bw'ubushyuhe, ingano y'ubushuhe, ingano ya ogisijeni n'ubushyuhe bwo gutwika (Guillén et al., 2000; Gomes et al., 2013). Polycyclic aromatic hydrocarbons (PAHs) nazo zabonetse mu mata ku rugero rutandukanye (0.75–2.1 mg/L) (Girelli et al., 2014). Kwirundanya kw'izi PAH mu biribwa biterwa kandi n'imiterere ya fizikiki y'ibiryo, mu gihe ingaruka zabyo z'uburozi zifitanye isano n'imikorere y'umubiri, imikorere ya metabolike, kwinjiza, gukwirakwira no gukwirakwira k'umubiri (Mechini et al., 2011).
Uburozi n'ingaruka mbi za hydrocarboni aromatic za polycyclic (PAHs) bimaze igihe kinini bizwi (Cherniglia, 1984). Hydrocarboni aromatic za polycyclic zifite uburemere buke bwa molekile (LMW-PAHs) (impeta ebyiri kugeza kuri eshatu) zishobora gufatana ku macromolecules atandukanye nka ADN, RNA na poroteyine kandi zitera kanseri (Santarelli et al., 2008). Bitewe n'imiterere yazo yo kudakoresha amazi, zitandukanywa n'uturemangingo twa lipide. Mu bantu, cytochrome P450 monooxygenases zigabanya PAH zikajya kuri epoxides, zimwe muri zo zikaba zikora cyane (urugero, baediol epoxide) kandi zishobora gutuma uturemangingo dusanzwe tuba uturemangingo twa kanseri (Marston et al., 2001). Byongeye kandi, umusaruro wa PAH nka quinones, phenols, epoxides, diols, nibindi ni uburozi kurusha ibintu bikurura. Amwe mu ma PAH n'ibice byayo bya metabolisme bishobora kugira ingaruka ku misemburo na enzymes zitandukanye mu mikorere y'umubiri, bityo bigira ingaruka mbi ku mikurire, mu mitsi yo hagati, ku myororokere no ku budahangarwa bw'umubiri (Swetha na Phale, 2005; Vamsee-Krishna et al., 2006; Oostingh et al., 2008). Guhura n'udukoko duto twa PAHs byagaragaye ko bitera imikorere mibi y'ibihaha no kugabanuka kw'amaraso mu mubiri ku barwayi ba asima ndetse bikongera ibyago byo kurwara kanseri y'uruhu, ibihaha, uruhago n'igifu (Olsson et al., 2010; Diggs et al., 2011). Ubushakashatsi ku nyamaswa bwagaragaje kandi ko guhura n'udukoko twa PAH bishobora kugira ingaruka mbi ku mikorere y'imyororokere no ku mikurire y'umubiri kandi bishobora gutera cataract, kwangirika kw'impyiko n'umwijima, ndetse no kwangirika k'umuhondo. Ibikoresho bitandukanye bya PAH biotransformation nka diols, epoxides, quinones na free radicals (cations) byagaragaye ko bikora ADN adducts. Byagaragaye ko uduce duto tw’amaraso duhindagurika duhindura uburyo bwo gukora ADN, mu gihe uduce duto tudahindagurika dushobora gukuraho ADN (cyane cyane adenine ndetse rimwe na rimwe na guanine); byombi bishobora gutera amakosa atuma habaho impinduka (Schweigert et al. 2001). Byongeye kandi, quinones (benzo-/pan-) zishobora gukora ubwoko bwa ogisijeni bukora neza (ROS), bigatera kwangirika gukomeye kwa ADN n’izindi macromolecules, bityo bigira ingaruka ku mikorere/ubuzima bw’ingingo (Ewa na Danuta 2017). Guhura kenshi n’ibice bike bya pyrene, biphenyl na naphthalene byagaragaye ko bitera kanseri mu nyamaswa zakorewe igerageza (Diggs et al. 2012). Bitewe n’uburozi bwazo bwica, gusukura/gukuraho izi PAH ahantu zangiritse/zanduye ni ikintu cy’ingenzi.
Uburyo butandukanye bwo mu mubiri n’ubutabire bwakoreshejwe mu gukuraho PAH ahantu/ibidukikije byanduye. Uburyo nko gutwika, gukuraho chlorine, oxidation ya UV, gufunga, no gukuramo solvent bufite ingaruka mbi nyinshi, harimo kurema ibintu birimo uburozi, imiterere y’imikorere, ibibazo by’umutekano n’amategeko, imikorere mike, ndetse n’igiciro kinini. Ariko, kwangirika kwa mikorobe (byitwa bioremediation) ni ubundi buryo bwiza bwo gukoresha mikorobe mu buryo bw’imiti cyangwa koloni. Ugereranije n’uburyo bwo mu mubiri n’ubutabire, ubu buryo ni bwiza ku bidukikije, ntibutera ingaruka mbi, buhendutse, kandi burambye. Bioremediation ishobora gukorerwa ahantu hagize ikibazo (aho kiri) cyangwa ahantu hateguwe byihariye (aho kiri) bityo ikaba ifatwa nk'uburyo burambye bwo kuvura kurusha uburyo gakondo bwo mu mubiri n’ubutabire (Juhasz na Naidu, 2000; Andreoni na Gianfreda, 2007; Megharaj et al., 2011; Phale et al., 2020; Sarkar et al., 2020).
Gusobanukirwa intambwe za mikorobe zikoreshwa mu kwangiza imyanda ihumanya ikirere bigira ingaruka zikomeye mu bya siyansi no mu bukungu ku bidukikije no mu kubungabunga ibidukikije. Garama 2.1 × 1018 za karuboni (C) zibikwa mu myanda n'ibintu bikomoka ku bimera (ni ukuvuga peteroli, gaze karemano, na makara, ni ukuvuga ibikomoka ku mabuye y'agaciro) ku isi yose, bikagira uruhare runini mu mikorere ya karuboni ku isi. Ariko, inganda zigenda ziyongera vuba, gucukura ibikomoka ku mabuye y'agaciro, n'ibikorwa by'abantu birimo kugabanya ibigega bya karuboni, bisohora garama 5.5 × 1015 za karuboni karemano (nk'imyanda) mu kirere buri mwaka (Gonzalez-Gaya et al., 2019). Inyinshi muri iyi karuboni karemano yinjira mu bidukikije byo ku isi no mu nyanja binyuze mu gusya, gutwara no gusohora amazi. Byongeye kandi, imyanda mishya ikomoka ku bikomoka ku mabuye y'agaciro, nka pulasitiki, pulasitiki na pulasitiki ihamye (phthalate na isomers zazo), yangiza cyane ibidukikije byo mu mazi, ubutaka n'ibyo mu mazi ndetse n'ibinyabuzima byabyo, bityo bikongera ibyago byo kuzamuka kw'ikirere ku isi. Ubwoko butandukanye bwa microplastics, nanoplastics, ibice bya pulasitiki hamwe n'ibikomoka kuri monomer byazo byangiza bikomoka kuri polyethylene terephthalate (PET) byakusanyije mu Nyanja ya Pasifika hagati ya Amerika ya Ruguru na Aziya y'Amajyepfo y'Uburasirazuba, birema "Great Pacific Garbage Patch", byangiza ibinyabuzima byo mu mazi (Newell et al., 2020). Ubushakashatsi bwa siyansi bwagaragaje ko bidashoboka gukuraho imyanda nk'iyo hakoreshejwe uburyo ubwo aribwo bwose bufatika cyangwa bwa shimi. Muri uru rwego, microorganisms zigira akamaro cyane ni izishobora guhindura imyanda muri oxidative dioxide de carbone, ingufu za chemical n'ibindi bintu bitari uburozi amaherezo byinjira mu zindi ntungamubiri (H, O, N, S, P, Fe, nibindi). Bityo, gusobanukirwa imiterere y'ibinyabuzima bya microflora y'ubutare bwa aromatike n'uburyo bwo kugenzura ibidukikije ni ingenzi mu gusuzuma uruziga rwa microflora de carbone, ingengo y'imari ya carbone yuzuye n'ingaruka z'ikirere mu gihe kizaza. Bitewe n'uko hakenewe cyane gukuraho ibyo bintu mu bidukikije, havutse inganda zitandukanye zibanda ku ikoranabuhanga risukuye. Ubundi buryo, guha agaciro imyanda/imyanda ikomoka mu nganda ikusanyirizwa mu rusobe rw’ibinyabuzima (ni ukuvuga uburyo bwo gufata imyanda mu buryo bw’ubukungu) bifatwa nk'imwe mu nkingi z'ubukungu buzenguruka n'intego z'iterambere rirambye (Close et al., 2012). Kubwibyo, gusobanukirwa imiterere y'ingufu, enzymes na genetike by'ibi bishobora kwangirika ni ingenzi cyane kugira ngo ikurwaho neza kandi ivugururwe neza ry'imyanda ihumura.
Mu bintu byinshi bihumanya ikirere, twita cyane ku bintu bihumanya ikirere bifite uburemere buke nka naphthalene na naphthalene zasimbuwe. Ibi bintu ni bimwe mu bigize ibikomoka kuri peteroli, amarangi y'imyenda, ibikomoka ku bicuruzwa, imiti yica udukoko (udukoko tw'inyenzi n'udukoko twica udukoko), plastike na tannins bityo bikaba bikwirakwira mu bidukikije byinshi (Preuss et al., 2003). Raporo ziherutse zigaragaza ubwiyongere bw'ingano ya naphthalene mu myanda y'amazi, mu butaka bwo munsi y'ubutaka no mu butaka bwo munsi y'ubutaka, mu turere twa vadose no mu nkengero z'imigezi, bigaragaza ko ikora neza mu bidukikije (Duttagupta et al., 2019, 2020). Imbonerahamwe ya 2 igaragaza imiterere ya fizikiki, ikoreshwa ryayo n'ingaruka ku buzima bwa naphthalene n'ibiyikomokaho. Ugereranyije n'izindi PAH zifite uburemere bwinshi, naphthalene n'ibiyikomokaho ntabwo bikunda amazi menshi, bishonga cyane mu mazi kandi bikwirakwira cyane mu bidukikije, bityo bikunze gukoreshwa nk'icyitegererezo cyo kwiga metabolisme, imiterere y'uturemangingo n'uburyo metabolisme ya PAH. Umubare munini w’udukoko dushobora guhindura naphthalene n’ibiyikomokaho, kandi amakuru arambuye araboneka ku nzira zabyo zo guhindura imikorere y’umubiri, enzymes n’imikorere y’amategeko (Mallick et al., 2011; Phale et al., 2019, 2020). Byongeye kandi, naphthalene n’ibiyikomokaho byagenwe nk’ibintu by’icyitegererezo byo gusuzuma ihumana ry’ibidukikije bitewe n’ubwinshi bwabyo n’uburyo biboneka. Ikigo gishinzwe kurengera ibidukikije muri Amerika kivuga ko ikigereranyo cya naphthalene ari 5.19 μg kuri metero kibe uvuye ku mwotsi w’itabi, ahanini uturutse ku gutwika kutuzuye, na 7.8 kugeza kuri 46 μg uturutse ku mwotsi wo ku ruhande, mu gihe guhura na creosote na naphthalene byikubye inshuro 100 kugeza ku 10.000 (Preuss et al. 2003). By’umwihariko, naphthalene byagaragaye ko ifite uburozi bw’ubuhumekero mu moko, mu turere, no mu gitsina ndetse n’ingaruka za kanseri. Hashingiwe ku bushakashatsi bw’inyamaswa, Ikigo Mpuzamahanga Gishinzwe Ubushakashatsi ku Kanseri (IARC) cyashyize naphthalene mu cyiciro cya "kanseri ishobora gutera umuntu" (Itsinda rya 2B)1. Kubona naphthalene zasimbuwe, cyane cyane iyo umuntu ahumeka cyangwa ahawe mu kanwa, bitera kwangirika kw'imitsi y'ibihaha kandi byongera umubare w'ibibyimba by'ibihaha mu mbeba n'imbeba (Gahunda y'Igihugu y'Uburozi bw'Ibihaha 2). Ingaruka zikomeye zirimo isesemi, kuruka, kubabara mu nda, impiswi, kuribwa umutwe, urujijo, ibyuya byinshi, umuriro, tachycardia, nibindi. Ku rundi ruhande, umuti wica udukoko witwa carbamate carbaryl (1-naphthyl N-methylcarbamate) wavuzweho uburozi ku nyamaswa zo mu mazi, inyamaswa zo mu mazi, inzuki z'ubuki n'abantu kandi byagaragaye ko ubuza acetylcholinesterase gutera paralysis (Smulders et al., 2003; Bulen na Distel, 2011). Kubwibyo, gusobanukirwa uburyo bwo kwangirika kwa mikorobe, kugenzura imiterere y'uturemangingo, imikorere ya enzymatic n'uturemangingo ni ingenzi mu guteza imbere ingamba zo kuvura indwara mu bidukikije byanduye.
Imbonerahamwe ya 2. Amakuru arambuye ku miterere ya fizikiki, imikoreshereze, uburyo bwo kumenya n'indwara zifitanye isano na naphthalene n'ibiyikomokaho.
Mu bice byanduye, imyanda ihumanya ikirere ishobora gutera ingaruka zitandukanye ku mikorobe z’ibidukikije (umuryango), nko guhinduka kw’amazi mu ruhu, kwinjira mu ruhu, kubyimba kw’ibinure mu ruhu, guhungabanya ihererekanya ry’ingufu (electron transport chain/proton motivation force), n’imikorere ya poroteyine zifitanye isano na ruhu (Sikkema et al., 1995). Byongeye kandi, bimwe mu bintu bishongesha nk’ibinyabutabire na quinones bitanga ubwoko bwa ogisijeni bukora (ROS) kandi bigatera ADN na poroteyine (Penning et al., 1999). Bityo, ubwinshi bw’ibi bintu mu rusobe rw’ibinyabuzima butera igitutu ku miterere y’udukoko kugira ngo tubashe kwangiza neza ku rwego rutandukanye rw’umubiri, harimo no kutwaka/gutwara, guhinduranya imiterere y’uturemangingo, guhuza/gukoresha, no gushyira hamwe.
Ubushakashatsi bwakozwe ku mushinga wa Ribosomal Database Project-II (RDP-II) bwagaragaje ko ubwoko bwose bwa bagiteri 926 bwakuwe mu miterere cyangwa mu miterere y’ibinyabuzima byandujwe na naphthalene cyangwa ibikomoka kuri yo. Itsinda rya Proteobacteria ryari rifite umubare munini w’ibiyigize (n = 755), rigakurikirwa na Firmicutes (52), Bacteroidetes (43), Actinobacteria (39), Tenericutes (10), na bagiteri zitarashyirwa mu byiciro (8) (Ishusho ya 2). Ibihagarariye γ-Proteobacteria (Pseudomonadales na Xanthomonadales) byiganje mu matsinda yose ya Gram-negative afite G+C nyinshi (54%), mu gihe Clostridiales na Bacillales (30%) byari amatsinda ya Gram-negative afite G+C nkeya. Pseudomonas (umubare munini, amoko 338) byavuzwe ko ishobora kwangiza naphthalene n'ibikomoka kuri methyl mu buryo butandukanye bwanduye (ikarafuriya y'amakara, peteroli, peteroli, ibitaka, amavuta yamenetse, amazi yanduye, imyanda y'ibinyabuzima n'aho imyanda ijugunywa) ndetse no mu buryo budahinduka (ubutaka, imigezi, ibishanga n'amazi yo munsi y'ubutaka) (Ishusho ya 2). Byongeye kandi, inyigo zo kongera ubwinshi bw'ibinyabuzima n'isesengura rya metagenomike muri bimwe muri utu turere byagaragaje ko ubwoko bwa Legionella na Clostridium butari buhingwa bushobora kuba bufite ubushobozi bwo kwangiza, bigaragaza ko ari ngombwa ko ubwo bwoko bwa bagiteri buhingwa kugira ngo bwige inzira nshya n'uburyo butandukanye bwo guhindura imikorere y'ibinyabuzima.
Ishusho ya 2. Ubudasa bw'uturemangingo n'uburyo bagiteri zikwirakwira mu bidukikije byandujwe na naphthalene n'ibikomoka kuri naphthalene.
Mu dukoko dutandukanye twangiza hydrocarbon, inyinshi muri two zishobora kwangiza naphthalene nk'isoko yonyine ya karuboni n'ingufu. Urukurikirane rw'ibintu bigira uruhare mu ihinduka rya naphthalene rwasobanuwe kuri Pseudomonas sp. (ubwoko: NCIB 9816-4, G7, AK-5, PMD-1 na CSV86), Pseudomonas stutzeri AN10, Pseudomonas fluorescens PC20 n'ubundi bwoko (ND6 na AS1) (Mahajan et al., 1994; Resnick et al., 1996; Annweiler et al., 2000; Basu et al., 2003; Dennis na Zylstra, 2004; Sota et al., 2006; Metabolisme itangizwa na dioxygenese igizwe n'ibice byinshi [naphthalene dioxygenease (NDO), ring hydroxylating dioxygenease] itera oxidation y'imwe mu mpeta zihumura za naphthalene ikoresheje ogisijeni ya molekile nk'indi substrate, igahindura naphthalene cis-naphthalenediol (Ishusho ya 3). Cis-dihydrodiol ihindurwamo 1,2-dihydroxynaphthalene binyuze dehydrogenase. Dioxygenese ifata impeta, 1,2-dihydroxynaphthalene dioxygenase (12DHNDO), ihindura 1,2-dihydroxynaphthalene muri aside 2-hydroxychromene-2-carboxylic. Isomerization ya cis-trans ya enzimatike ikora trans-o-hydroxybenzylidenepyruvate, igakurwamo na hydratase aldolase ikaba salicylic aldehyde na pyruvate. Aside organic pyruvate ni yo ya mbere C3 ikomoka ku gashishwa ka naphthalene karubone ikerekeza mu nzira yo hagati ya karubone. Byongeye kandi, salicylaldehyde dehydrogenase ishingiye kuri NAD+ ihindura salicylaldehyde ikaba aside salicylic. Metabolism muri iki cyiciro yitwa "inzira yo hejuru" yo kwangirika kwa naphthalene. Iyi nzira ikunze kugaragara muri bagiteri nyinshi za naphthalene. Ariko, hari ibintu bike bidasanzwe; urugero, muri Bacillus hamburgii 2 ikunda ikirere, kwangirika kwa naphthalene bitangirwa na naphthalene. 2,3-dioxygenase kugirango ikore 2,3-dihydroxynaphthalene (Annweiler et al., 2000).
Ishusho ya 3. Inzira za naphthalene, methylnaphthalene, aside naphthoic, na carbaryl yononekara. Imibare ifite uruziga igaragaza enzymes zishinzwe guhindura naphthalene n'ibiyikomokaho mu buryo bukurikira. 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.
Bitewe n'imiterere y'ibinyabuzima n'imiterere yabyo, aside salicylic ivamo irushaho guhindurwa binyuze mu nzira ya catechol hakoreshejwe salicylate 1-hydroxylase (S1H) cyangwa binyuze mu nzira ya gentisate hakoreshejwe salicylate 5-hydroxylase (S5H) (Ishusho ya 3). Kubera ko aside salicylic ari yo ikoreshwa cyane mu mikorere ya naphthalene (inzira yo hejuru), intambwe ziva kuri aside salicylic zijya kuri TCA intermediate zikunze kwitwa inzira yo hasi, kandi gene zigizwe na operon imwe. Ni ibisanzwe kubona ko gene ziri mu nzira yo hejuru ya operon (nah) na operon yo hasi ya operon (sal) zigenzurwa n'ibintu bisanzwe bigenzurwa; urugero, NahR na aside salicylic bikora nk'ibitera imbaraga, bigatuma operon zombi zishobora guhindurwa burundu naphthalene (Phale et al., 2019, 2020).
Byongeye kandi, catechol igabanywamo kabiri na 2-hydroxymuconate semialdehyde binyuze mu nzira ya meta na catechol 2,3-dioxygenase (C23DO) (Yen et al., 1988) hanyuma ikongera gushonga na 2-hydroxymuconate semialdehyde hydrolase kugira ngo ikore 2-hydroxypent-2,4-dienoic aside. 2-hydroxypent-2,4-dienoate ihindurwamo pyruvate na acetaldehyde binyuze kuri hydratase (2-oxopent-4-enoate hydratase) na aldolase (4-hydroxy-2-oxopentanoate aldolase) hanyuma ikinjira mu nzira ya karuboni yo hagati (Ishusho ya 3). Ubundi buryo, catechol igabanywamo kabiri na cis,cis-muconate binyuze mu nzira ya ortho na catechol 1,2-oxygenase (C12DO). Muconate cycloisomerase, muconolactone isomerase, na β-ketoadipate-nollactone hydrolase bihindura cis, cis-muconate muri 3-oxoadipate, byinjira mu nzira ya karuboni yo hagati binyuze muri succinyl-CoA na acetyl-CoA (Nozaki et al., 1968) (Ishusho ya 3).
Mu nzira ya gentisate (2,5-dihydroxybenzoate), impeta ihumura icibwamo na gentisate 1,2-dioxygenase (GDO) kugira ngo ikore maleylpyruvate. Iki gicuruzwa gishobora guhindurwamo hydrolyze mu buryo butaziguye kugira ngo gikore pyruvate na malate, cyangwa gishobora guhindurwamo isomer kugira ngo gikore fumarylpyruvate, hanyuma kigahindurwamo hydrolyze mu gukora pyruvate na fumarate (Larkin na Day, 1986). Guhitamo inzira yindi byagaragaye muri bagiteri za Gram-negative na Gram-positive ku rwego rwa biochemical na gene (Morawski et al., 1997; Whyte et al., 1997). Bagiteri za Gram-negative (Pseudomonas) zikunda gukoresha aside salicylic, itera metabolism ya naphthalene, ikayikuramo katekoli ikoresheje salicylate 1-hydroxylase (Gibson na Subramanian, 1984). Ku rundi ruhande, muri bagiteri za Gram-positive (Rhodococcus), salicylate 5-hydroxylase ihindura aside salicylic muri aside gentisic, mu gihe aside salicylic itagira ingaruka ku ihindurwa rya genes za naphthalene (Grund et al., 1992) (Ishusho ya 3).
Byaravuzwe ko ubwoko nka Pseudomonas CSV86, Oceanobacterium NCE312, Marinhomonas naphthotrophicus, Sphingomonas paucimobilis 2322, Vibrio cyclotrophus, Pseudomonas fluorescens LP6a, ubwoko bwa Pseudomonas na Mycobacterium bushobora kwangiza monomethylnaphthalene cyangwa dimethylnaphthalene (Dean-Raymond na Bartha, 1975; Cane na Williams, 1982; Mahajan et al., 1994; Dutta et al., 1998; Hedlund et al., 1999). Muri bwo, inzira ya 1-methylnaphthalene na 2-methylnaphthalene yo kwangirika kwa Pseudomonas sp. CSV86 yizwe neza ku rwego rwa biochemical na enzyme (Mahajan et al., 1994). 1-Methylnaphthalene itunganywa binyuze mu nzira ebyiri. Ubwa mbere, impeta ihumura iba hydroxylated (uruziga rudasimbuwe rwa methylnaphthalene) kugira ngo ikore cis-1,2-dihydroxy-1,2-dihydro-8-methylnaphthalene, ikongera guhindurwa methyl salicylate na methylcatechol, hanyuma ikinjira mu nzira yo hagati ya karuboni nyuma yo gucika kw'uruziga (Ishusho ya 3). Iyi nzira yitwa "inzira yo gusiga karuboni". Mu nzira ya kabiri "yo gusiga uburozi", itsinda rya methyl rishobora guhindurwa hydroxylated na NDO kugira ngo rikore 1-hydroxymethylnaphthalene, ikongera guhindurwa 1-naphthoic acid hanyuma igasohorwa mu buryo bw'imikurire nk'umusaruro udapfa. Ubushakashatsi bwagaragaje ko ubwoko bwa CSV86 budashobora gukura kuri aside 1 na 2-naphthoic nk'isoko yonyine ya karuboni n'ingufu, byemeza inzira yayo yo gusiga uburozi (Mahajan et al., 1994; Basu et al., 2003). Muri 2-methylnaphthalene, itsinda rya methyl rinyura muri hydroxylation na hydroxylase kugira ngo rikore 2-hydroxymethylnaphthalene. Byongeye kandi, impeta idasimbuwe y'impeta ya naphthalene inyura muri hydroxylation kugira ngo ikore dihydrodiol, ihinduka oxidized muri 4-hydroxymethylcatechol mu ruhererekane rw'ibikorwa bya enzyme-catalyzed reactions hanyuma ikinjira mu nzira yo hagati ya karuboni binyuze mu nzira ya meta-ring cleavage. Mu buryo nk'ubwo, S. paucimobilis 2322 yavuzwe ko ikoresha NDO muri hydroxylate 2-methylnaphthalene, irushaho oxidized kugira ngo ikore methyl salicylate na methylcatechol (Dutta et al., 1998).
Aside naphthoic (yasimbuwe/itasimbuwe) ni ibintu biva mu guhumanya/guhindura imiterere y’ibinyabuzima bikomoka ku ihumana rya methylnaphthalene, phenanthrene na anthracene bigashyirwa mu buryo bwo gukwirakwiza ibidukikije. Byaravuzwe ko Stenotrophomonas maltophilia CSV89 ishobora guhindura aside 1-naphthoic nk'isoko ya karuboni (Phale et al., 1995). Ihinduka ry’ibintu ritangirana na dihydroxylation y’impeta ihumura neza igakora 1,2-dihydroxy-8-carboxynaphthalene. Diol ivamo ihinduka catechol binyuze kuri 2-hydroxy-3-carboxybenzylidenepyruvate, aside 3-formylsalicylic, aside 2-hydroxyisophthalic na aside salicylic hanyuma ikinjira mu nzira ya karuboni yo hagati binyuze mu nzira ya meta-ring cleavage (Ishusho ya 3).
Carbaryl ni umuti wica udukoko wa naphthyl carbamate. Kuva mu mpinduramatwara y’ibidukikije mu Buhinde mu myaka ya 1970, ikoreshwa ry’ifumbire mvaruganda n’imiti yica udukoko byatumye habaho kwiyongera kw’imyuka ihumanya ikirere ya polycyclic aromatic hydrocarbon (PAH) iva mu buhinzi butari ahantu hanini (Pingali, 2012; Duttagupta et al., 2020). Ubutaka bwose bugera kuri 55% (hegitari 85,722,000) mu Buhinde buterwa imiti yica udukoko. Mu myaka itanu ishize (2015–2020), urwego rw’ubuhinzi mu Buhinde rwakoresheje impuzandengo ya toni 55,000 kugeza 60,000 z’imiti yica udukoko buri mwaka (Ishami rishinzwe Amakoperative n’Imibereho Myiza y’Abahinzi, Minisiteri y’Ubuhinzi, Guverinoma y’Ubuhinde, Kanama 2020). Mu turere twa Gangetic two mu majyaruguru no hagati (intara zifite abaturage benshi n’ubucucike bw’abaturage), ikoreshwa ry’imiti yica udukoko ku bihingwa rikwirakwiriye cyane, aho imiti yica udukoko yiganjemo. Carbaryl (1-naphthyl-N-methylcarbamate) ni umuti wica udukoko wa carbamate ufite uburozi buri hagati y’uburozi busanzwe cyangwa uburozi bwinshi ukoreshwa mu buhinzi bw’Ubuhinde ku kigero cya toni 100–110. Ukunze kugurishwa ku izina ry’ubucuruzi rya Sevin kandi ukoreshwa mu kurwanya udukoko (udusimba, imiserebanya, inda, udusimba, ibicurane n’utundi dukoko twinshi two mu gasozi) bigira ingaruka ku bihingwa bitandukanye (ibigori, soya, ipamba, imbuto n’imboga). Udukoko tumwe na tumwe nka Pseudomonas (NCIB 12042, 12043, C4, C5, C6, C7, Pseudomonas putida XWY-1), Rhodococcus (NCIB 12038), Sphingobacterium spp. (CF06), Burkholderia (C3), Micrococcus na Arthrobacter nabyo bishobora gukoreshwa mu kurwanya utundi dukoko. Byaratangajwe ko RC100 ishobora kwangiza carbaryl (Larkin na Day, 1986; Chapalamadugu na Chaudhry, 1991; Hayatsu et al., 1999; Swetha na Phale, 2005; Trivedi et al., 2017). Inzira yo kwangirika kwa carbaryl yizwe cyane ku rwego rwa biochemical, enzyme na genetiki mu butaka bwa Pseudomonas sp. Strains C4, C5 na C6 (Swetha na Phale, 2005; Trivedi et al., 2016) (Ishusho ya 3). Inzira ya metabolisme itangirira kuri hydrolysis ya ester bond na carbaryl hydrolase (CH) kugira ngo habeho 1-naphthol, methylamine na dioxyde de carbone. Hanyuma 1-naphthol ihindurwa 1,2-dihydroxynaphthalene na 1-naphthol hydroxylase (1-NH), ikomeza gukoreshwa binyuze mu nzira yo hagati ya karuboni binyuze muri salicylate na gentisate. Hari bagiteri zimwe na zimwe zigabanya karuboni zavuzwe ko ziyihinduramo aside salicylic binyuze mu gucamo impeta ya catechol ortho (Larkin na Day, 1986; Chapalamadugu na Chaudhry, 1991). Ikigaragara ni uko bagiteri zigabanya naphthalene ahanini zihindura aside salicylic binyuze muri catechol, mu gihe bagiteri zigabanya karuboni zikunda gukoresha aside salicylic binyuze mu nzira ya gentisate.
Aside naphthalenesulfonike/aside disulfonike hamwe n’ibikomoka kuri aside naphthylaminesulfonike bishobora gukoreshwa nk'ibice by'ingenzi mu gukora irangi rya azo, ibintu bitose, ibikwirakwiza amazi, nibindi. Nubwo ibi bintu bifite uburozi buke ku bantu, isuzuma rya cytotoxicity ryagaragaje ko byica amafi, daphnia n'ibimera (Greim et al., 1994). Abahagarariye ubwoko bwa Pseudomonas (ubwoko bwa A3, C22) byavuzwe ko batangira metabolism binyuze mu gukurura kabiri hydroxylation y'impeta ihumura irimo itsinda rya aside sulfonic kugira ngo ikore dihydrodiol, ihinduka 1,2-dihydroxynaphthalene binyuze mu gucikamo kw'itsinda rya sulfite (Brilon et al., 1981). 1,2-dihydroxynaphthalene ivamo ihinduka binyuze mu nzira ya naphthalene ya kera, ni ukuvuga inzira ya catechol cyangwa gentisate (Ishusho ya 4). Byagaragaye ko aside aminonaphthalenesulfonique na aside hydroxynaphthalenesulfonique bishobora kwangirika burundu hakoreshejwe uburyo bwa bagiteri buvanze hamwe n’inzira zuzuzanya za katabolic (Nortemann et al., 1986). Byagaragaye ko kimwe mu bigize ihuriro gikuraho aside aminonaphthalenesulfonique cyangwa aside hydroxynaphthalenesulfonique binyuze muri 1,2-dioxygenation, mu gihe aminosalicylate cyangwa hydroxysalicylate birekurwa mu buryo bw’imikurire nk’ikintu kidakora neza hanyuma bigafatwa n’ibindi bigize ihuriro. Aside Naphthalenedisulfonique ni polarique ariko ntiyoroherwa neza bityo ikaba ishobora kwangirika binyuze mu nzira zitandukanye. Kureka kwa mbere kwa sulfurization bibaho mu gihe cyo guhindura imiterere y’impeta ya aromatique n’itsinda rya aside sulfonique; Gukuraho sulfuri kwa kabiri bibaho mu gihe cy’ihindagurika rya aside 5-sulfosalicylic na aside salicylic 5-hydroxylase kugira ngo bikore aside gentisic, yinjira mu nzira yo hagati ya karuboni (Brilon et al., 1981) (Ishusho ya 4). Enzymes zishinzwe kwangirika kwa naphthalene nazo zishinzwe metabolism ya naphthalene sulfonate (Brilon et al., 1981; Keck et al., 2006).
Ishusho ya 4. Inzira za metabolike zituma naphthalene sulfonate yangirika. Imibare iri mu ruziga igaragaza imisemburo ikora metabolike ya naphthyl sulfonate, isa/imeze nk'imisemburo ivugwa ku ISHUSHO YA 3.
PAH nkeya za molekile (LMW-PAHs) ziragabanuka, ntizikunda amazi kandi ntizishonga neza, bityo ntizishobora kwangirika/kwangirika bisanzwe. Ariko, mikorobe zo mu mwuka zishobora kuyikora binyuze mu kwinjiza ogisijeni ya molekile (O2). Izi enzymes ahanini ziri mu itsinda rya oxidoreductases kandi zishobora gukora ibikorwa bitandukanye nko kugabanya hydroxylation ya aromatike (mono- cyangwa dihydroxylation), kugabanya hydrogenation no gucika kw'impeta ya aromatike. Ibikomoka kuri izi actives biri mu rwego rwo hejuru rwa oxidation kandi byoroshye guhindura imikorere binyuze mu nzira ya karuboni yo hagati (Phale et al., 2020). Bivugwa ko enzymes ziri mu nzira yo kwangirika zishobora kwangirika. Imikorere y'izi enzymes iba iri hasi cyane cyangwa ntoya iyo uturemangingo dukurira ku masoko yoroshye ya karuboni nka glucose cyangwa aside organic. Imbonerahamwe ya 3 igaragaza muri make enzymes zitandukanye (oxygenases, hydrolases, dehydrogenases, oxidases, nibindi) zigira uruhare mu mikorere ya naphthalene n'ibiyikomokaho.
Imbonerahamwe ya 3. Imiterere y'ibinyabutabire bya enzymes zituma naphthalene n'ibiyikomokaho byangirika.
Ubushakashatsi bwa radioisotope (18O2) bwagaragaje ko gushyira O2 mu mpande zihumura hakoreshejwe ogisijeni ari intambwe y'ingenzi cyane mu gutuma ikintu kirushaho kubora (Hayaishi et al., 1955; Mason et al., 1955). Kwinjizwa kwa atome imwe ya ogisijeni (O) iva muri ogisijeni ya ogisijeni (O2) mu gice cy'ubutaka bitangirwa na monooxygenases zo mu bwoko bwa endogenous cyangwa exogenous (zitwa kandi hydroxylases). Indi atome ya ogisijeni igabanuka ikaba amazi. Monooxygenases zo mu bwoko bwa exogenous zigabanya flavin hamwe na NADH cyangwa NADPH, mu gihe muri endomonooxygenases flavin igabanuka bitewe n'igice cy'ubutaka. Aho hydroxylation iherereye hatuma habaho ubudasa mu mikorere y'ibicuruzwa. Urugero, salicylate 1-hydroxylase hydroxylates salicylic acid ku mwanya wa C1, igakora catechol. Ku rundi ruhande, salicylate 5-hydroxylase ifite ibice byinshi (irimo reductase, ferredoxin, na oxygenase subunits) hydroxylates salicylic acid ku mwanya wa C5, ikora aside gentisic (Yamamoto et al., 1965).
Dioxygeneases zishyira atome ebyiri za O2 muri substrate. Bitewe n'ibikomoka ku bimera, zigabanyijemo dioxygeneases zikora ring hydroxylating na dioxygeneases zikora ring cleaving. Dioxygeneases zikora ring hydroxylating zihindura substrate za aromatic mo cis-dihydrodiols (urugero: naphthalene) kandi zikwirakwira muri bagiteri. Kugeza ubu, byagaragaye ko ibinyabuzima birimo dioxygeneases zikora ring hydroxylating zishobora gukura ku masoko atandukanye ya karuboni aromatic, kandi izi enzymes zishyirwa mu byiciro bya NDO (naphthalene), toluene dioxygenase (TDO, toluene), na biphenyl dioxygenase (BPDO, biphenyl). NDO na BPDO byombi bishobora gukurura oxidation ebyiri n'umuyoboro wa hydroxylation y'uturemangingo dutandukanye twa polycyclic aromatic hydrocarbons (toluene, nitrotoluene, xylene, ethylbenzene, naphthalene, biphenyl, fluorene, indole, methylnaphthalene, naphthalenesulfonate, phenanthrene, anthracene, acetophenone, nibindi) (Boyd na Sheldrake, 1998; Phale et al., 2020). NDO ni sisitemu y'ibice byinshi igizwe na oxidoreductase, ferredoxin, n'igice cy'umwuka kirimo oxygenase (Gibson na Subramanian, 1984; Resnick et al., 1996). Igice cya catalytic cya NDO kigizwe n'igice kinini cya α n'igice gito cya β gishyizwe mu buryo bwa α3β3. NDO iri mu muryango munini w’uturemangingo twa ogisijeni kandi α-subunit yayo irimo ahantu ha Rieske [2Fe-2S] hamwe n’icyuma kitagira ingirabuzimafatizo kimwe, kigena imiterere yihariye ya NDO (Parales et al., 1998). Ubusanzwe, mu gihe kimwe cya katalytic, electron ebyiri zivuye mu kugabanuka kwa nucleotide ya pyridine zimurirwa kuri iyoni ya Fe (II) iri ahantu hakora binyuze muri reductase, ferredoxin n’ahantu ha Rieske. Uturemangingo duto dutuma ogisijeni ya molekile ikora, ari nabyo bisabwa kugira ngo substrate dihydroxylation iboneke (Ferraro et al., 2005). Kugeza ubu, ni NDO nke gusa zatunganyijwe kandi zigaragazwa mu buryo burambuye n’ubwoko butandukanye kandi uburyo bwo kugenzura inzira zigize ingaruka ku kwangirika kwa naphthalene bwizweho mu buryo burambuye (Resnick et al., 1996; Parales et al., 1998; Karlsson et al., 2003). Diooxygenase zigabanya impeta (enzymes zigabanya impeta cyangwa ortho-ring na enzymes zigabanya impeta cyangwa meta-ring) zigira uruhare ku bintu birungo bya hydroxylated. Urugero, diooxygenase igabanya impeta ni catechol-1,2-dioxygenase, mu gihe diooxygenase igabanya impeta ari catechol-2,3-dioxygenase (Kojima et al., 1961; Nozaki et al., 1968). Uretse ogisijeni zitandukanye, hari kandi dehydrogenase zitandukanye zituma dihydrodiols zigabanya impeta, alcool na aldehydes zigabanuka kandi zigakoresha NAD+/NADP+ nk'ibikoresho bitanga electron, ari na bimwe mu bintu by'ingenzi bigira uruhare mu mikorere y'umubiri (Gibson na Subramanian, 1984; Shaw na Harayama, 1990; Fahle et al., 2020).
Enzymes nka hydrolases (esterases, amidases) ni ubwoko bwa kabiri bw'ingenzi bwa enzymes zikoresha amazi kugira ngo zice imigozi ya covalent kandi zigaragaze umwihariko wa substrate. Carbaryl hydrolase n'izindi hydrolases bifatwa nk'ibice bya periplasm (transmembrane) mu bigize bagiteri za Gram-negative (Kamini et al., 2018). Carbaryl ifite isano ya amide na ester; bityo, ishobora gushonga na esterase cyangwa amidase kugira ngo ikore 1-naphthol. Carbaryl muri Rhizobium rhizobium strain AC10023 na Arthrobacter strain RC100 byavuzwe ko bikora nka esterase na amidase, uko bikurikirana. Carbaryl muri Arthrobacter strain RC100 nayo ikora nka amidase. Byagaragaye ko RC100 ikoresha hydrolyze imiti ine yo mu bwoko bwa N-methylcarbamate nka carbaryl, methomyl, aside mefenamic na XMC (Hayaatsu et al., 2001). Byatangajwe ko CH muri Pseudomonas sp. C5pp ishobora gukora kuri carbaryl (ibikorwa 100%) na 1-naphthyl acetate (ibikorwa 36%), ariko ntabwo ikora kuri 1-naphthylacetamide, bigaragaza ko ari esterase (Trivesi et al., 2016).
Inyigo za biochemical, imiterere ya enzymes, n'isesengura rya genetike byagaragaje ko genes za naphthalene zo kwangirika zigizwe n'ibice bibiri bigenzura cyangwa "operon": nah ("inzira yo hejuru", guhindura naphthalene muri aside salicylic) na sal ("inzira yo hasi", guhindura aside salicylic mu nzira yo hagati ya karuboni binyuze muri catechol). Aside salicylic n'ibindi biyigize bishobora gukora nk'ibitera imbaraga (Shamsuzzaman na Barnsley, 1974). Iyo hari glucose cyangwa aside organic, operon irahagarikwa. Ishusho ya 5 igaragaza imiterere yuzuye ya genetike yo kwangirika kwa naphthalene (mu buryo bwa operon). Imodoka nyinshi/ubwoko bwa nah gene (ndo/pah/dox) zasobanuwe kandi zasanze zifite imiterere yo hejuru (90%) mu moko yose ya Pseudomonas (Abbasian et al., 2016). Genes z'inzira yo hejuru ya naphthalene muri rusange zateguwe mu buryo bwumvikana nk'uko bigaragara ku Ishusho ya 5A. Hari indi gene, nahQ, yavuzweho kugira uruhare mu mikorere ya naphthalene kandi ubusanzwe yari iri hagati ya nahC na nahE, ariko imikorere yayo nyayo iracyari hafi gusobanurwa. Mu buryo nk'ubwo, gene ya nahY, ishinzwe chemotaxis iterwa na naphthalene, yabonetse ku mpera ya nah operon muri bamwe mu bagize urwego. Muri Ralstonia sp., gene ya U2 ikora glutathione S-transferase (gsh) yagaragaye ko iri hagati ya nahAa na nahAb ariko ntabwo yagize ingaruka ku mikoreshereze ya naphthalene (Zylstra et al., 1997).
Ishusho ya 5. Imiterere n'ubutandukanye byagaragaye mu gihe naphthalene yangiritse mu moko ya bagiteri; (A) Inzira ya naphthalene yo hejuru, imikorere ya naphthalene ihinduka aside salicylic; (B) Inzira ya naphthalene yo hasi, aside salicylic binyuze muri catechol ijya mu nzira ya karuboni yo hagati; (C) aside salicylic inyuze muri gentisate ijya mu nzira ya karuboni yo hagati.
"Inzira yo hasi" (sal operon) ubusanzwe igizwe nahGTHINLMOKJ kandi igahindura salicylate muri pyruvate na acetaldehyde binyuze mu nzira ya catechol metaring cleavage. Gene ya nahG (ishyira salicylate hydroxylase) byagaragaye ko iguma ku mpera ya operon (Ishusho ya 5B). Ugereranije n'izindi naphthalene-degrading types, muri P. putida CSV86 nah na sal operons zirafatanye kandi zifitanye isano ya hafi cyane (hafi 7.5 kb). Muri zimwe muri bagiteri za Gram-negative, nka Ralstonia sp. U2, Polaromonas naphthalenivorans CJ2, na P. putida AK5, naphthalene ihinduka metabolite ya karuboni hagati binyuze mu nzira ya gentisate (mu buryo bwa sgp/nag operon). Kaseti ya gene ikunze kugaragara mu buryo bwa nagAaGHabAcAdBFCQEDJI, aho nagR (ishyiraho uburyo bwa LysR-type regulator) iherereye ku mpera yo hejuru (Ishusho ya 5C).
Karuboni yinjira mu gice cyo hagati cya karuboni binyuze mu mikorere ya 1-naphthol, 1,2-dihydroxynaphthalene, aside salicylic, na aside gentisic (Ishusho ya 3). Hashingiwe ku bushakashatsi bwa genetiki n'impinduka mu mikorere, byatekerejwe ko iyi nzira igabanywamo "izamuka" (ihinduka rya karuboni muri aside salicylic), "hagati" (ihinduka rya aside salicylic muri aside gentisic), na "imanuka" (ihinduka rya aside gentisic muri intermediates ya central carbon pathway) (Singh et al., 2013). Isesengura rya genomike rya C5pp (supercontig A, 76.3 kb) ryagaragaje ko gene ya mcbACBDEF igira uruhare mu guhindura carbaryl muri aside salicylic, hagakurikiraho mcbIJKL mu guhindura aside salicylic muri aside gentisic, na mcbOQP mu guhindura aside gentisic muri intermediates ya karuboni yo hagati (fumarate na pyruvate, Trivedi et al., 2016) (Ishusho ya 6).
Byaravuzwe ko enzymes zigira uruhare mu kwangirika kwa hydrocarbons zihumura neza (harimo naphthalene na aside salicylic) zishobora guterwa n'ibintu bifitanye isano kandi bigakumirwa n'amasoko yoroshye ya karuboni nka glucose cyangwa aside organic (Shingler, 2003; Phale et al., 2019, 2020). Mu nzira zitandukanye za naphthalene n'ibikomoka kuri yo, imiterere y'igenzura rya naphthalene na carbaryl yakozweho ubushakashatsi ku rugero runaka. Kuri naphthalene, genes mu nzira zo hejuru n'izo hasi zigenzurwa na NahR, urwego rwa LysR-type trans-acting positive regulator. Irakenewe kugira ngo gene ya nah ishyirwemo na aside salicylic hamwe n'imikorere yayo yo ku rwego rwo hejuru (Yen na Gunsalus, 1982). Byongeye kandi, ubushakashatsi bwagaragaje ko integrative host factor (IHF) na XylR (sigma 54-dependent transcriptional regulator) nabyo ari ingenzi mu gutuma genes zikora transcriptional mu metabolism ya naphthalene (Ramos et al., 1997). Ubushakashatsi bwagaragaje ko enzymes zo mu nzira yo gufungura catechol meta-ring, ari zo catechol 2,3-dioxygenase, ziterwa iyo hari naphthalene cyangwa aside salicylic (Basu et al., 2006). Ubushakashatsi bwagaragaje ko enzymes zo mu nzira yo gufungura catechol ortho-ring, ari zo catechol 1,2-dioxygenase, ziterwa iyo hari aside benzoic na cis, cis-muconate (Parsek et al., 1994; Tover et al., 2001).
Muri C5pp, gene eshanu, mcbG, mcbH, mcbN, mcbR na mcbS, zishyiraho amabwiriza agenga umuryango wa LysR/TetR w’abagenzura uburyo bwo kwandika inyandiko bashinzwe kugenzura iyangirika rya karubori. Gene ya mcbG ifitanye isano rya hafi na PhnS igenzura ubwoko bwa LysR (58% aside amine identity) igira uruhare mu mikorere ya phenanthrene muri Burkholderia RP00725 (Trivesi et al., 2016). Gene ya mcbH yagaragaye ko igira uruhare mu nzira yo hagati (guhindura aside salicylic muri aside gentisic) kandi ikaba iri mu igenzura uburyo bwo kwandika ubwoko bwa LysR NagR/DntR/NahR muri Pseudomonas na Burkholderia. Abagize uyu muryango bavuzwe ko bemera aside salicylic nk'ikintu cyihariye gituma gene zangirika ziyongera. Ku rundi ruhande, jeni eshatu, mcbN, mcbR na mcbS, zigize LysR na TetR type transcriptional regulators, zabonetse mu nzira yo hasi (metabolites za genzisate-central carbon pathway).
Muri prokaryote, inzira zo kohereza gene zitambitse (gufata, guhanahana, cyangwa kwimura) binyuze muri plasmide, transposons, prophages, genomic islands, na integrative conjugative elements (ICE) ni zo mpamvu zikomeye zitera plasticity muri genome za bagiteri, biganisha ku kwiyongera cyangwa gutakaza imikorere/imiterere yihariye. Bituma bagiteri zishobora kwihuza vuba n’imimerere itandukanye y’ibidukikije, bigatanga inyungu zishobora guhinduka ku mubiri w’umuntu, nko kwangirika kw’ibinyabutabire bihumura. Impinduka mu mikorere ya metabolic akenshi zigerwaho binyuze mu gutunganya neza operons zo kwangirika, uburyo bwazo bwo kuzigenzura, n’imiterere ya enzyme, byoroshya kwangirika k’ubwoko bwinshi bw’ibinyabutabire bihumura (Nojiri et al., 2004; Phale et al., 2019, 2020). Kasete za gene zo kwangirika kwa naphthalene zabonetse ziherereye ku bintu bitandukanye bigenda nka plasmide (bihuza n’ibitahuza), transposons, genome, ICEs, n’uruvange rw’ubwoko butandukanye bwa bagiteri (Ishusho ya 5). Muri Pseudomonas G7, nah na sal operon za plasmid NAH7 zanditswe mu cyerekezo kimwe kandi ni bimwe mu bigize transposon ifite inenge isaba transposase Tn4653 kugira ngo ikoreshwe (Sota et al., 2006). Muri Pseudomonas strain NCIB9816-4, gene yabonetse kuri conjugative plasmid pDTG1 nk'ama-operon abiri (hafi 15 kb hagati) yanditswe mu byerekezo bitandukanye (Dennis na Zylstra, 2004). Muri Pseudomonas putida strain AK5, non-conjugative plasmid pAK5 ishyiraho enzyme ishinzwe kwangirika kwa naphthalene binyuze mu nzira ya gentisate (Izmalkova et al., 2013). Muri Pseudomonas strain PMD-1, nah operon iherereye kuri chromosome, mu gihe sal operon iherereye kuri plasmid pMWD-1 ihuza (Zuniga et al., 1981). Ariko, muri Pseudomonas stutzeri AN10, genes zose za naphthalene degradation (nah na sal operon) ziherereye kuri chromosome kandi birashoboka ko ziboneka binyuze mu guhinduranya, gusubiramo, no gusubiza hamwe (Bosch et al., 2000). Muri Pseudomonas sp. CSV86, nah na sal operon ziherereye muri genome mu buryo bwa ICE (ICECSV86). Imiterere irinzwe na tRNAGly hagakurikiraho gusubiramo mu buryo butaziguye bigaragaza ahantu ho gusubiranamo/gufatana (attR na attL) hamwe n'integrase isa na phage iherereye ku mpera zombi za tRNAGly, bityo ikaba isa n'igice cya ICEclc (ICEclcB13 muri Pseudomonas knackmusii yo kwangirika kwa chlorocatechol). Byaravuzwe ko genes kuri ICE zishobora kwimurirwa mu buryo bwo guhuza hamwe n'umuvuduko muto cyane wo kwimurira (10-8), bityo zigahita zimurira imiterere yo kwangirika ku muntu wakiriye (Basu na Phale, 2008; Phale et al., 2019).
Inyinshi mu ngirabuzimafatizo zituma carbaryl yangirika ziherereye kuri plasmide. Arthrobacter sp. RC100 irimo plasmide eshatu (pRC1, pRC2 na pRC300) muri zo plasmide ebyiri zihuza, pRC1 na pRC2, zishyiraho enzymes zihindura carbaryl muri gentisate. Ku rundi ruhande, enzymes zigize uruhare mu guhindura gentisate muri metabolite ya karuboni yo hagati ziherereye kuri chromosome (Hayaatsu et al., 1999). Bagiteri zo mu bwoko bwa Rhizobium. Strain AC100, ikoreshwa mu guhindura carbaryl muri 1-naphthol, irimo plasmide pAC200, itwara gene ya cehA ikora CH nk'igice cya transposon ya Tnceh ikikijwe n'ibice bisa n'ibice (istA na istB) (Hashimoto et al., 2002). Muri CF06 ya Sphingomonas, gene ya carbaryl degradation bikekwa ko iboneka muri plasmidi eshanu: pCF01, pCF02, pCF03, pCF04, na pCF05. Ubushobozi bwa ADN bw'izi plasmidi buri hejuru, bigaragaza ko hariho igikorwa cyo kwikuba kwa gene (Feng et al., 1997). Mu bwoko bwa carbaryl-degrading symbiont bugizwe n'ubwoko bubiri bwa Pseudomonas, strain 50581 irimo plasmidi pCD1 (50 kb) ikora gene ya mcd carbaryl hydrolase, mu gihe plasmidi ya conjugative muri strain 50552 ikora enzyme ya 1-naphthol-degrading (Chapalamadugu na Chaudhry, 1991). Muri strain ya Achromobacter WM111, gene ya mcd furadan hydrolase iherereye kuri plasmidi ya 100 kb (pPDL11). Iyi gene byagaragaye ko iboneka kuri plasmide zitandukanye (100, 105, 115 cyangwa 124 kb) muri bagiteri zitandukanye zituruka mu turere dutandukanye tw’isi (Parekh et al., 1995). Muri Pseudomonas sp. C5pp, gene zose zituma carbaryl yangirika ziherereye muri genome ifite 76.3 kb y’uruhererekane (Trivesi et al., 2016). Isesengura rya genome (6.15 Mb) ryagaragaje ko hari MGE 42 na GEI 36, muri zo MGE 17 zari muri supercontig A (76.3 kb) zifite impuzandengo ya G+C idafite ingano (54–60 mol%), bigaragaza ko hashobora kubaho impinduka mu buryo butambitse (Trivesi et al., 2016). P. putida XWY-1 igaragaza imiterere nk'iyo ya genes zigabanya carbaryl, ariko izi genes ziherereye kuri plasmide (Zhu et al., 2019).
Uretse ubushobozi bwa metabolisme ku rwego rwa biochemical na genome, mikorobe zigaragaza izindi miterere cyangwa ibisubizo nka chemotaxis, imiterere yo guhindura ubuso bw'uturemangingo, gushyira ibice mu bice, ikoreshwa ryihariye, ikorwa rya biosurfactant, nibindi, bibafasha gutunganya neza imyanda ihumanya ikirere mu bidukikije byanduye (Ishusho ya 7).
Ishusho ya 7. Ingamba zitandukanye zo gusubiza uturemangingo tw’uturemangingo tw’udukoko twangiza hydrocarbon kugira ngo tubore neza ibintu bihumanya ikirere.
Uburyo bwo kurwanya imyuka ihumanya ikirere bufatwa nk'ibintu byongera kwangirika kw'imyanda ihumanya ikirere mu duce tw'ibinyabuzima twanduye cyane. (2002) byagaragaje ko imyuka ihumanya ikirere ya Pseudomonas sp. G7 kuri naphthalene yongereye umuvuduko wo kwangirika kwa naphthalene mu buryo bw'amazi. Ubwoko bw'inyamaswa bwa G7 bwangizaga naphthalene vuba cyane kurusha ubwoko bw'ibinyabuzima bidakoresha chemotaxis. Poroteyine ya NahY (aside amine 538 ifite topology ya membrane) byagaragaye ko ihujwe na genes za metacleavage pathway kuri plasmid ya NAH7, kandi kimwe na transducers za chemotaxis, iyi poroteyine isa nkaho ikora nk'umusemburo wa chemoreceptor wo kwangirika kwa naphthalene (Grimm na Harwood 1997). Ubundi bushakashatsi bwakozwe na Hansel et al. (2009) bwagaragaje ko poroteyine ari chemotactic, ariko igipimo cyayo cyo kwangirika kiri hejuru. (2011) yagaragaje uburyo Pseudomonas (P. putida) ikora ku buryo bwa chemotactic kuri naphthalene ikora, aho ikwirakwira ry’ingufu ryatumye naphthalene ikomeza gutembera mu turemangingo, ibyo bikaba byaragenzuraga uburyo chemotactic ikora ku turemangingo. Abashakashatsi bakoresheje iyi myitwarire ya chemotactic mu gushushanya mikorobe zakongera umuvuduko wo kwangirika. Ubushakashatsi bwagaragaje ko inzira za chemosensory zigenga izindi mikorere y’uturemangingo nko kwigabanyamo uturemangingo, kugenzura imikorere y’uturemangingo, no gukora biofilm, bityo bigafasha mu kugenzura umuvuduko wo kwangirika. Ariko, gukoresha iyi miterere (chemotaxis) kugira ngo ivugururwe neza bibangamirwa n’imbogamizi nyinshi. Imbogamizi zikomeye ni izi: (a) receptors zitandukanye za paralogous zibona ibintu bimwe/ligands; (b) kubaho kwa receptors zindi, ni ukuvuga, energie tropism; (c) itandukaniro rikomeye mu bice by’ibyiyumviro by’umuryango umwe wa receptor; na (d) kubura amakuru kuri poroteyine zikomeye za bagiteri (Ortega et al., 2017; Martin-Mora et al., 2018). Hari igihe, kubora kw'ibinyabuzima bya hydrocarboni bihumura bitanga metabolite nyinshi/intermediates, zishobora kuba chemotactic ku itsinda rimwe rya bagiteri ariko zigatera ishozi ku zindi, birushaho kugorana inzira. Kugira ngo tumenye imikoranire ya ligands (hydrocarboni zihumura neza) n'ibikoresho bya chemical, twubatse poroteyine zihuza sensor (PcaY, McfR, na NahY) duhuza sensor na signaling domains za Pseudomonas putida na Escherichia coli, zigamije receptors za aside ahumura, intermediates za TCA, na naphthalene, uko bikurikirana (Luu et al., 2019).
Bitewe n’ingaruka za naphthalene n’izindi polycyclic aromatic hydrocarbons (PAHs), imiterere y’urukiramende rwa bagiteri n’ubusugire bwa microorganisms bihinduka cyane. Ubushakashatsi bwagaragaje ko naphthalene ibangamira imikoranire y’urukiramende rwa acyl binyuze mu mikoranire ya hydrophobic, bityo byongera kubyimba no kugenda kw’urukiramende (Sikkema et al., 1995). Kugira ngo hirindwe iyi ngaruka mbi, bagiteri zigenzura imiterere y’urukiramende binyuze mu guhindura ikigereranyo n’imiterere ya aside irike hagati ya aside irike ya iso/anteiso branched-chain na isomerizing cis-unsaturated acids muri trans-isomers zijyanye nayo (Heipieper na de Bont, 1994). Muri Pseudomonas stutzeri yakuwe mu miti ya naphthalene, ikigereranyo cya aside irike yuzuye cyariyongereye kuva kuri 1.1 kugera kuri 2.1, mu gihe muri Pseudomonas JS150 iki kigereranyo cyariyongereye kuva kuri 7.5 kugera kuri 12.0 (Mrozik et al., 2004). Ubwo yakuraga kuri naphthalene, uturemangingo twa Achromobacter KA 3-5 twagaragaje ukwishyira hamwe kw'uturemangingo dukikije kristu za naphthalene ndetse no kugabanuka k'umuvuduko w'uturemangingo (kuva kuri -22.5 kugeza kuri -2.5 mV) hamwe no guhumeka kwa cytoplasm no kuvanga, bigaragaza impinduka mu miterere y'uturemangingo n'imiterere y'ubuso bw'uturemangingo (Mohapatra et al., 2019). Nubwo impinduka mu turemangingo/ubuso zifitanye isano itaziguye no gufata neza imyanda ihumanya ikirere, ingamba zijyanye na bioengineering ntizanonosowe neza. Gukoresha imiterere y'uturemangingo ntabwo byigeze bikoreshwa cyane mu kunoza imikorere y'ibinyabuzima (Volke na Nikel, 2018). Gukuraho uturemangingo bigira ingaruka ku kwigabanya kw'uturemangingo bitera impinduka mu miterere y'uturemangingo. Gukuraho uturemangingo bigira ingaruka ku kwigabanya kw'uturemangingo bitera impinduka mu miterere y'uturemangingo. Muri Bacillus subtilis, poroteyine ya septum y'uturemangingo SepF byagaragaye ko igira uruhare mu ishingwa rya septum kandi ikenewe mu ntambwe zikurikiraho zo kwigabanya kw'uturemangingo, ariko si gene y'ingenzi. Gusiba kwa jeni zikora peptide glycan hydrolases muri Bacillus subtilis byatumye uturemangingo twiyongera, umuvuduko wo gukura wiyongera, ndetse n'ubushobozi bwo gukora enzymes bwiyongera (Cui et al., 2018).
Hatanzwe uburyo bwo gushyira hamwe inzira yo kwangirika kwa carbaryl kugira ngo hagerwe ku kwangirika neza k'ubwoko bwa Pseudomonas C5pp na C7 (Kamini et al., 2018). Hatanzwe igitekerezo ko carbaryl ijyanwa mu mwanya wa periplasmic binyuze muri septum yo hanze ya membrane cyangwa binyuze muri porins zishobora gukwirakwira. CH ni enzyme ya periplasmic itera hydrolysis ya carbaryl kuri 1-naphthol, ikaba ihamye, idakunda amazi kandi ifite uburozi bwinshi. CH iboneka muri periplasm kandi ifite ubushobozi buke bwa carbaryl, bityo ikagenzura ikorwa rya 1-naphthol, bityo ikarinda kwirundanya kwayo mu turemangingo no kugabanya uburozi bwayo ku turemangingo (Kamini et al., 2018). 1-naphthol ivamo ijyanwa muri cytoplasm mu membrane y'imbere binyuze mu kugabanya no gukwirakwira, hanyuma ikavanwa muri hydroxylated kuri 1,2-dihydroxynaphthalene na enzyme ifite 1NH kugira ngo ikomeze gukora neza mu nzira ya karuboni yo hagati.
Nubwo mikorobe zifite ubushobozi bwo kwangiza amasoko ya karuboni ya xenobiotic, imiterere y’ikoreshwa ryazo (ni ukuvuga gukoresha cyane amasoko ya karuboni yoroshye kuruta ayoroshye) ni imbogamizi ikomeye ku kwangirika kw’ibinyabuzima. Kuba hariho kandi ikoreshwa ry’amasoko ya karuboni yoroshye rigabanya imiterere ya za jeni zikora imisemburo yangiza amasoko ya karuboni yoroshye/atari meza nka PAH. Urugero rwizwe neza ni uko iyo glucose na lactose bihujwe na Escherichia coli, glucose ikoreshwa neza kurusha lactose (Jacob na Monod, 1965). Byaragaragaye ko Pseudomonas yangiza ubwoko butandukanye bwa PAH n’ibinyabutabire bya xenobiotic nk’isoko ya karuboni. Inzira yo gukoresha amasoko ya karuboni muri Pseudomonas ni aside organic > glucose > ibinyabutabire bihumura neza (Hylemon na Phibbs, 1972; Collier et al., 1996). Ariko, hari itandukaniro. Igishimishije ni uko Pseudomonas sp. CSV86 igaragaza imiterere yihariye y’urwego rw’ibanze ikoresha cyane cyane hydrocarboni zihumura neza (aside benzoic, naphthalene, nibindi) aho gukoresha glucose kandi igahuza hydrocarboni zihumura neza hamwe na aside organic (Basu et al., 2006). Muri iyi bagiteri, genes zo kwangiza no gutwara hydrocarboni zihumura neza ntizigabanywa nubwo haba hari isoko ya kabiri ya karuboni nka glucose cyangwa aside organic. Iyo yakuriye muri glucose na hydrocarboni zihumura neza, byagaragaye ko genes zo gutwara glucose na metabolisme zagabanijwe, hydrocarboni zihumura neza zakoreshejwe mu cyiciro cya mbere cy’igenzura, naho glucose ikoreshwa mu cyiciro cya kabiri cy’igenzura (Basu et al., 2006; Choudhary et al., 2017). Ku rundi ruhande, kuba hariho aside organice ntacyo byagizeho ingaruka ku mikorere ya hydrocarboni zihumura neza, bityo iyi bagiteri yitezweho kuba ubwoko bushobora gukoreshwa mu bushakashatsi ku ibora ry’ibinyabuzima (Phale et al., 2020).
Birazwi neza ko guhindura imiterere y’ibinyabuzima muri hydrocarbon bishobora gutera stress ya oxidative no kwiyongera kwa enzymes za antioxidant mu tunyangingo duto. Kudakora neza kwa naphthalene haba mu tunyangingo tudakora neza ndetse no mu gihe hari ibintu by’uburozi bituma habaho ubwoko bwa ogisijeni ikora neza (ROS) (Kang et al. 2006). Kubera ko enzymes za naphthalene zirimo amatsinda ya fer-sulfur, iyo stress ya oxidative, icyuma kiri muri heme na poroteyine za fer-sulfur zizahinduka oxidative, bigatuma poroteyine idakora. Ferredoxin-NADP+ reductase (Fpr), hamwe na superoxide dismutase (SOD), bihuza redox reaction isubira inyuma hagati ya NADP+/NADPH na molekile ebyiri za ferredoxin cyangwa flavodoxin, bityo bigakuraho ROS no gusubiza ikigo cya fer-sulfur iyo stress ya oxidative (Li et al. 2006). Byaratangajwe ko Fpr na SodA (SOD) muri Pseudomonas bishobora guterwa n’umuvuduko wa ogisijeni, kandi kwiyongera kwa SOD na catalase byagaragaye mu moko ane ya Pseudomonas (O1, W1, As1, na G1) mu gihe cyo gukura munsi y’imiterere ya naphthalene (Kang et al., 2006). Ubushakashatsi bwagaragaje ko kongeramo antioxydants nka aside ascorbic cyangwa ferrous iron (Fe2+) bishobora kongera umuvuduko wo gukura kwa naphthalene. Ubwo Rhodococcus erythropolis yakuraga mu buryo bwa naphthalene, imiterere ya genes za cytochrome P450 zifitanye isano na oxidative stress irimo sodA (Fe/Mn superoxide dismutase), sodC (Cu/Zn superoxide dismutase), na recA byongerewe (Sazykin et al., 2019). Isesengura rya poroteyine rigereranya ingano y’uturemangingo twa Pseudomonas twakuwe muri naphthalene ryagaragaje ko kongera poroteyine zitandukanye zijyanye n’ingaruka z’umuvuduko wa oxidative ari ingamba yo guhangana n’umuvuduko (Herbst et al., 2013).
Udukoko twavuzwe ko dukora biosurfactants binyuze mu bikorwa by’isoko rya karubone idakoresha amazi. Izi surfactants ni ibintu bikora ku buso bwa amfifili bishobora gukora aggregates ku mazi y’amavuta cyangwa aho amazi y’umwuka ahurira. Ibi bitera gushonga kw’ibintu bidafite aho bihuriye kandi byoroshya uburyo hydrocarbons zihumura neza, bigatuma bibora neza (Rahman et al., 2002). Kubera iyo miterere, biosurfactants zikoreshwa cyane mu nganda zitandukanye. Kongeramo chemical surfactants cyangwa biosurfactants mu miterere ya bagiteri bishobora kongera imikorere n’umuvuduko wo kwangirika kwa hydrocarbon. Muri biosurfactants, rhamnolipides zikorwa na Pseudomonas aeruginosa zakoreweho ubushakashatsi bwinshi kandi zigaragazwa (Hisatsuka et al., 1971; Rahman et al., 2002). Byongeye kandi, ubundi bwoko bwa biosurfactants burimo lipopeptides (mucins ziva kuri Pseudomonas fluorescens), emulsifier 378 (ziva kuri Pseudomonas fluorescens) (Rosenberg na Ron, 1999), lipide za trehalose disaccharide ziva kuri Rhodococcus (Ramdahl, 1985), lichenin ziva kuri Bacillus (Saraswathy na Hallberg, 2002), na surfactant ziva kuri Bacillus subtilis (Siegmund na Wagner, 1991) na Bacillus amyloliquefaciens (Zhi et al., 2017). Izi surfactants zikomeye zagaragaye ko zigabanya ubushyuhe bw'ubuso kuva kuri 72 dynes/cm kugeza kuri munsi ya 30 dynes/cm, bigatuma hydrocarbon ibasha kwinjiza neza. Byaravuzwe ko Pseudomonas, Bacillus, Rhodococcus, Burkholderia n'izindi bagiteri zishobora gukora biosurfactants zitandukanye za rhamnolipide na glycolipide iyo zihinzwe muri naphthalene na methylnaphthalene (Kanga et al., 1997; Puntus et al., 2005). Pseudomonas maltophilia CSV89 ishobora gukora biosurfactant yo hanze ya extracellular Biosur-Pm iyo ihinzwe ku bintu bihumura nka aside naphthoic (Phale et al., 1995). Ikinetiki yo gukora Biosur-Pm yagaragaje ko gukora kwayo ari inzira ishingiye ku gukura no ku pH. Byagaragaye ko ingano ya Biosur-Pm ikorwa n'uturemangingo kuri pH neutral yari hejuru ugereranyije n'iyo kuri pH 8.5. Uturemangingo twahinzwe kuri pH 8.5 twari dufite hydrophobic cyane kandi twari dufite affinity nyinshi ku bintu bihumura na aliphatic kuruta utwo duremangingo twahinzwe kuri pH 7.0. Muri Rhodococcus spp. N6, igipimo cyo hejuru cya karuboni kuri azote (C:N) hamwe n'ubudahangarwa bw'icyuma ni ibintu byiza cyane mu gukora biosurfactants zo hanze y'umubiri (Mutalik et al., 2008). Hakozwe igerageza ryo kunoza imikorere ya biosurfactants (surfactins) binyuze mu kunoza ubwoko n'imikorere ya fermentation. Ariko, igipimo cya surfactant mu buryo bw'ubuhinzi ni gito (1.0 g/L), ibi bikaba ari imbogamizi ku musaruro munini (Jiao et al., 2017; Wu et al., 2019). Kubwibyo, uburyo bwo gukora genetiki bwakoreshejwe kugira ngo bunoze imikorere yayo. Ariko, guhindura imiterere yayo biragoye bitewe n'ingano nini ya operon (∼25 kb) n'uburyo bugoye bwo gukora biosurfactants muri sisitemu yo kumenya imiterere ya quorum (Jiao et al., 2017; Wu et al., 2019). Impinduka nyinshi mu buhanga bw’uturemangingo zakozwe muri bagiteri ya Bacillus, zigamije cyane cyane kongera umusaruro wa surfactin hashingiwe ku musaruro ukomoka kuri surfactin (srfA operon), hagashyirwa hejuru cyane poroteyine yo kohereza surfactin muri YerP hamwe n’ibintu bigenzura ComX na PhrC (Jiao et al., 2017). Ariko, ubu buryo bw’ubuhanga bw’uturemangingo bwageze ku mpinduka imwe cyangwa nke gusa mu buhanga bw’uturemangingo kandi ntiburagera ku musaruro w’ubucuruzi. Kubwibyo, ni ngombwa kwiga uburyo bwo kunoza ubumenyi bushingiye ku bumenyi.
Inyigo zo kwangirika kwa PAH zikorwa ahanini mu buryo busanzwe bwo muri laboratwari. Ariko, ahantu handuye cyangwa ahantu handuye, ibintu byinshi bitaba ibyo mu mubiri (ubushyuhe, pH, ogisijeni, kuboneka kw'intungamubiri, kuba ibintu bikomoka ku binyabuzima, izindi xenobiotics, gukumira umusaruro wa nyuma, nibindi) byagaragaye ko bigira ingaruka ku bushobozi bwo kwangirika kwa mikorobe.
Ubushyuhe bugira ingaruka zikomeye ku kubora kwa PAH. Uko ubushyuhe bwiyongera, ubwinshi bwa ogisijeni ishongeshejwe buragabanuka, ibyo bikaba bigira ingaruka ku mikorere ya mikorobe zo mu mubiri, kuko zikenera ogisijeni ya molekile nk'imwe mu ngingo zituma ogisijeni ikora ibikorwa byo kubora cyangwa gucana kw'impeta. Akenshi bikunze kugaragara ko ubushyuhe bwinshi buhindura PAH z'ababyeyi mo ibintu bihumanya cyane, bityo bikabuza kubora kwa ogisijeni (Muller et al., 1998).
Byagaragaye ko ahantu henshi handuye PAH hari pH ikabije, nko ahantu handuye aside mine (pH 1-4) n'ahantu haterwa gaze karemano/amakara handuye alkaline leachate (pH 8-12). Ibi bishobora kugira ingaruka zikomeye ku ibora ry'ibinyabuzima. Kubwibyo, mbere yo gukoresha mikorobe mu gutunganya ibintu, ni byiza guhindura pH wongeramo imiti ikwiye (ifite ubushobozi bwo kugabanya ogisijeni buri hagati kugeza kuri make cyane) nka ammonium sulfate cyangwa ammonium nitrate ku butaka bwa alkaline cyangwa liming hamwe na calcium carbonate cyangwa magnesium carbonate ku hantu hafite aside (Bowlen et al. 1995; Gupta na Sar 2020).
Gutanga ogisijeni mu gace kagizweho ingaruka ni cyo kintu kigabanya umuvuduko wa PAH mu kubora kw'ibinyabuzima. Bitewe n'imiterere y'ibidukikije, ibikorwa byo gutunganya ogisijeni mu gace runaka akenshi bisaba ko ogisijeni ijyanwa mu bintu byo hanze (guhinga, gutanga umwuka mwinshi, no kongeramo imiti) (Pardieck et al., 1992). Odenkranz et al. (1996) bagaragaje ko kongeramo peroxide ya magnesium (ikintu gitanga ogisijeni) mu mazi yanduye bishobora gutuma BTEX ikora neza. Ubundi bushakashatsi bwakorewe ku igabanuka rya phenol na BTEX mu mazi yanduye binyuze mu gutera nitrati ya sodium no kubaka amariba yo gukuramo kugira ngo haboneke uburyo bwo gutunganya ogisijeni neza (Bewley na Webb, 2001).