Malembe aaya ncibeela chamutwe wamakani wakuvwuntauzya "Nzila zyakusumpula zintu zipona antoomwe anzila zyakubelesya zintu zipona zyakabambwa kale". Langa zibalo zyoonse zili kkumi azibili
Ma molecule masyoonto aa polycyclic aromatic hydrocarbons (PAHs) mbuli naphthalene antoomwe a substituted naphthalenes (methylnaphthalene, naphthoic acid, 1-naphthyl-N-methylcarbamate, azimwi) alabelesyegwa kapati mumakwebo aasiyene-siyene alimwi alajaya ma gene, mutagenic alimwi/naa carcinogenic kuzilenge. Aaya ma organic compounds (SOCs) naakuti xenobiotics abonwa kuti ngamasofweede aakusaanguna alimwi aleta ntenda mpati kubusena bwanyika yoonse abuumi bwabantu boonse. Bupati bwamilimo yabantu (nkokuti kupanga muya wamalasha, kusalazya mafuta, kuzwa kwamyootokala alimwi akubelesya mubulimi) mbobupa kuti kube kuvwula, kuyooba alimwi akutolwa kwazintu eezyi zijanika koonse-koonse alimwi zitamani. Mukuyungizya kunzila zyakusilisya/kugwisya, nzila zitanyonganyi busena zilimbuli bioremediation, eezyo zibelesya tuzunda tukonzya kunyonyoona POCs naakuti kuzisandula kuba zintu zitakwe tombe, zyakazwa mbuli nzila imwi iikwabilidwe, iitaduli alimwi iikonzya kugwasya. Misyobo yamabacteria yiindene-indene iili mumulongo wa Proteobacteria (Pseudomonas, Pseudomonas, Comamonas, Burkholderia, a Neosphingobacterium), Firmicutes (Bacillus a Paenibacillus), antoomwe a Actinobacteria (Rhodococcus a Arthrobacter) mubulongo yakatondezya kuti ilakonzya kunyonyoona zintu zisiyene-siyene. ziiyo zya metabolic, genomics, antoomwe a metagenomic analysis zitugwasya kumvwisisisya buyumuyumu bwa catabolic akusiyana kuliwo mubukkale oobu buuba, oobo bukonzya kubelesegwa kunembo kubola kwabuumi. Kubako kwa PAHs kwachiindi chilamfu kwapa kuti kube nzila zipya zyakubola kwiinda mukutola ma gene kubelesya zintu zya gene zilimbuli ma plasmids, transposons, bacteriophages, genomic islands, antoomwe a integrative conjugative elements. Bbayoloji yazyintu zipona antoomwe akubamba ma gene aabantu bamwi naakuti bantu bamwi (consortia) zilakonzya kugwasya kuti kube kubambulula kwa PAHs ooku kwiinda mukubelesya nzila zibeleka antoomwe. Mukulanga-langa ooku, tulabikkila maanu kunzila zyaandeene zyakusanduka kwazyakulya akusiyana, bupange bwama gene akusiyana, alimwi akusanduka kwamaseelo/kucinca kwa naphthalene alimwi amabacteria aanyonganya naphthalene aabikkwa mubusena bwawo. Eezi ziyoopa makani aajatikizya zyintu zipona mumuunda akuchita kuti zintu zibe kabotu.
Kusumpuka kwakufwambaana kwamakwebo (mafuta, bulimi, misamu, madayi aamasani, zyakulisamika, azimwi) kwagwasya kuzwidilila kwabuvubi bwanyika yoonse akusumpula buumi bwabantu. Ooku kuyaambele kupati kwapa kuti kube kupanga mweelwe mupati wazintu zipangwa abantu (SOCs), eezyo zibelesyegwa kupanga zintu zisiyene. Eeyi misamu yakunze naakuti SOCs ibikkilizya ma polycyclic aromatic hydrocarbons (PAHs), misamu iijaya tuzunda, misamu iijaya nsaku, misamu iikola, madayi, misamu iikola, ma organophosphates, misamu iikwabilila mulilo, misamu iikonzya kupya, azimwi. Zizwa mumulengalenga, mumaanzi amunyika ooko nkozikonzya kunyonganya zintu zyiingi, zyintu zikonzya kuchitika kuzilenge zyaandeene-andeene kwiinda mukuchincha zintu zipona antoomwe abupange bwabantu (Petrie et al., 2015; Bernhardt et al., 2017; Sarkar et al., 2020). Zintu zyiingi zisofweede zilijisi nguzu zyakunyonyoona zintu zyiingi zipona/masena aajisi zintu zipona (nkokuti ma coral reefs, Arctic/Antarctic ice sheets, madambwa aamumalundu malamfu, lwizi lwaansi, azimwi) (Jones 2010; Beyer et al. 2020; Nordborg et al. 2020). Buvwuntauzi bwa geomicrobiological bwakacitwa ino-ino bwakatondezya kuti kubikkwa kwa zintu zipona (nkokuti zintu zisofweede) azintu zizwa kulinzizyo atalaa zintu zipangidwe (nkokuti busena bwakayakidwe) (nkokuti masena aazilengwa leza azintu zyakabambwa a granite, mabwe, zisamu abutale) zifwambaanya kunyonyooka kwazyo (Gadd Liu et al. 2017; 2018). Milimu yabantu ilakonzya kuyungizya akubija kunyonyooka kwazintu zipona zyazikozyanyo amayake kwiinda mukusofwaazya muuya akusanduka kwachiimo chamulengalenga (Liu et al. 2020). Eezyi zintu zisofweede zilaswaangana abwaanda bwamaanzi mumulengalenga mpawo zilakkala abwaanda, mpawo zipa kuti zintu zinyonyooke. Kubola kwazintu zipona nchintu chizyibidwe loko kuti nkucinca kutali kabotu mukuboneka abukkale bwazintu zyakachitika akaambo kazintu zipona eezyo zipa kuti zikwabililwe (Pochon a Jaton, 1967). Kuyungizya kubeleka kwama microbial (kusanduka kwazintu) kwa misamu eeyi kulakonzya kucesya kusyomeka kwabupange, kugwasya kwakukwabilila alimwi abulemu bwazilengwa (Gadd, 2017; Liu et al., 2018). Kulubazu lumwi, muziindi zimwi, kuzyibila kwatunyama tusyoonto-syoonto alimwi akuvwiila kuzintu eezyi kwajanwa kuti kulagwasya nkaambo zipanga ma biofilm alimwi azintu zimwi zikwabilila zicesya mweelwe wakubola/kubola (Martino, 2016). Aboobo, kuyaambele kwanzila zigwasya zyakukwabilila zintu zyamabwe, nsimbi azisamu ziyandika kumvwisisisya nzila zipati zijatikizyidwe mumulimo ooyu. Kweezyanisya azintu zicitika muzilengwa leza (zintu zicitika munyika, mulilo wamusyokwe, kupya kwamalundu, kusanduka kwazisyango alimwi amabacteria), milimo yabantu ipa kuti kube kuzwa kwamaanzi manji aa polycyclic aromatic hydrocarbons (PAHs) alimwi a organic carbon iimbi (OC) muzilenge zyamunyika. Ma PAH manji abelesyegwa mubulimi (misamu iijaya tuzunda amisamu iijaya tuzunda mbuli DDT, atrazine, carbaryl, pentachlorophenol, azimwi), mumakkampani (mafuta aatakwe busani, bulongo bwamafuta, mapulasitiki aazwa mumafuta, ma PCB, mapulasitiki, ma detergent, ma infectant, fumigants, zintu zikwabilila tununkilizyo), azintu zikwabilila tununkilizyo), (mafuta aakukwabilila zuba, misamu iijaya tuzunda, misamu iitanda tuzunda alimwi a polycyclic musks) alimwi azilwanyo (zintu zikonzya kubboloka mbuli 2,4,6-TNT) nzintu zikonzya kunyonganya buumi bwanyika (Srogi, 2007; Vamsee-Krishna a Phale, 2008; Petrie et al., 2015). Mulongo ooyu ulakonzya kuyungizigwa kuti ubikkilizye zintu zizwa mumafuta (mafuta, mafuta, ma asphaltenes), ma bioplastics aajisi ma molecule manji, alimwi ama ionic liquids (Amde et al., 2015). Chibeela 1 chilabikkilizya zintu zisofweede zisiyene-siyene akubelesya kwazyo mumakwebo aasiyene-siyene. Muminyaka misyoonto yainda, kuzwa kwabantu kwazintu zipona zitazungaani, antoomwe a carbon dioxide alimwi amwi muya uusofweede, zyakatalika kuyaambele (Dvorak et al., 2017). Nokuba boobo, ntenda zyabantu zilainda loko ntenda zyazilengwa leza. Mukuyungizya, twakajana kuti mweelwe wa SOCs ulazumanana mumasena miingi alimwi wakajanwa kuti nzintu zisofwaazya zilikuzwa antangalala zyeendelezya zintu zipona (Chitondeezyo 1). Mbunga zikwabilila busena zilimbuli United States Environmental Protection Agency (USEPA) zyakabikkilizya bunji bwazintu eezyi zisofwaazya mumulongo wazintu nzyobeelede kuchita akaambo kakuti zilakonzya kujaya maseelo, kujaya ma gene, kuchincha, akujaya kkansa. Aboobo, milawu miyumu yakusowa tombe anzila zigwasya zyakugwisya tombe kuzwa mumasena aasofweede zilayandikana. Nzila zyaandeene zyakusilisya zilimbuli pyrolysis, oxidative thermal treatment, airation airation, filling, incineration, azimwi tazibeleki pe mpawo ziladula mpawo zipa kuti kube zintu zibola, zikola alimwi zikatazya kusilika. Alubo luzibo lwanyika yoonse lwazyintu zizingulukide, tuzunda tukonzya kunyonyoona zintu eezyi zisofwaazya azintu zizwa kulinzizyo (mbuli halogenated, nitro, alkyl a/naa methyl) tulayungizya kubikkilwa maanu (Fennell et al., 2004; Haritash and Kaushik, 2009; Phale et al., 2002; Sarkar et al., 2020; Sarkar et al., 2020; Schwanemann abamwi, 2020). Kubelesya tuzunda ootu tusyoonto-syoonto tulikke naa muzilengwa zisanganizidwe (makoloni) mukugwisya zintu zisofwaazya tununkilizyo kuli bubotu mumakani aakukwabilila busena, muulo, kubeleka kabotu, kubeleka kabotu, alimwi akuzumanana. Basikuvwuntauzya alimwi bali mukulanga-langa kuswaanganya nzila zyama microbial anzila zya electrochemical redox, nkokuti bioelectrochemical systems (BES), mbuli nzila mbotu yakugwisya tombe (Huang et al., 2011). Busongo bwa BES bwakakwela bantu banji akaambo kakubeleka kabotu, tabuyandiki mali manji, bulakwabilila busena, bulabeleka kabotu, zibelesyo zikonzya kweendelana azintu zipona, alimwi akukonzya kujana zintu ziyandisi (nkokuti, magesi, mafuta, alimwi amakemikalu) (Pant et al., 2012; Nazari et al., 2020). Kuboola kwakubelesya nzila zyiingi zyakutobelezya ma genome antoomwe azibelesyo/nzila zya omics kwapa buvwubi bwamakani mapya aatala akweendelezya ma gene, ma proteomics, antoomwe a fluxomics zyakusanduka kwa tuzunda tusyoonto-syoonto tujaya. Kuswaanganya zibelesyo eezi antoomwe a systems biology kwayungizya kumvwisisisya kwesu kwakusala akubamba kabotu nzila zya catabolic muzilenge zipona (nkokuti, nzila zya metabolic) kuchitila kuti kube kubola kwazintu zipona. Kuti tupange nzila zigwasya zyakuponya zintu zipona kubelesya tuzunda tusyoonto-syoonto, tweelede kumvwisisisya nguzu zyabuumi, kusiyana kwazyakulya, bupange bwama gene, antoomwe abukkale (autoecology/synecology) bwatunyama tusyoonto-syoonto.
Cifwanikiso. 1. Nkozizwa anzila zya low-molecular PAHs kwiinda mumasena aasiyene-siyene azintu zisiyene-siyene zijatikizya buumi. Mizila iiteteete yiiminina kujatana aakati kazyintu zipona.
Mukulanga-langa ooku, twasola kupandulula mubufwiifwi data aatala akunyonyooka kwa PAHs zitakatazyi zilimbuli naphthalene antoomwe a naphthalenes zyakanjililwa mubusena bwabacteria zisiyene-siyene zijatikizya nzila zyakusanduka kwazyakulya akusiyana, ma enzymes aajatikizyidwe mukunyonyooka, bupange bwama gene/zintu zili mukati akusiyana, nsandulo zyamaseelo azintu zisiyene-siyene zya bioremediation. Kumvwisisisya zyeelelo zya biochemical antoomwe a molecule kuyoogwasya mukuziba misyobo yeelede ya host alimwi akuyungizya nzila zyabo zya genetic engineering kutegwa kugwasyigwe kabotu ku bioremediation ya zintu zisofwaazya zili boobu. Eezi ziyoogwasya mukubamba nzila zyakutalisya mbungano zyamabacteria aabusena bumwi kutegwa kube kugwasya kusilika zintu zipona.
Kubako kwamweelwe mupati wazintu zikola alimwi ziyoosya (kuzuzikizya mulawu wa Huckel 4n + 2π electrons, n = 1, 2, 3, ...) kuletela ntenda mpati kuzyintu zyamuchilengwaleza mbuli muuya, bulongo, mabwe, alimwi amaanzi aali atalaa nyika alimwi aali ansi (Puglisi et al., 2007). Eezyi zintu zilijisi zibeela zya benzene zyomwe (monocyclic) naa zibeela zya benzene zinji (polycyclic) zibikkidwe munzila iili mumulongo, mumbali naa muzibeela alimwi zitondezya kukkalikana (kukkalilila/kutakkalilila) mubusena akaambo kanguzu zyakuvwundauka zitali kabotu alimwi akubula nguzu (kubula nguzu), eezyo zinga zyapandululwa akaambo kakutayandi maanzi alimwi akuceya kwazyo. Eelyo ringi ya aromatic niyanjililwa busena a methyl (-CH3), carboxyl (-COOH), hydroxyl (-OH), naa sulfonate (-HSO3) tubunga, ilakkalilila, ilaa luyando lupati ku macromolecules, alimwi ilaa bioaccumulative muzibeela zyabuumi (Seo et al., 2009; Phale et al., 2020). Amwi ma polycyclic aromatic hydrocarbons (LMWAHs), mbuli naphthalene azintu zizwa mulinjiyo [methylnaphthalene, naphthoic acid, naphthalenesulfonate, antoomwe a 1-naphthyl N-methylcarbamate (carbaryl)], akabikkilizyigwa mumulongo wazintu zipona ziyandikana kapati a US Environmental Agency zyintu zikonzya kujaya ma gene, mutagenic, alimwi/naa carcinogenic (Cerniglia, 1984). Kugwisigwa kwamusyobo ooyu wa NM-PAHs mubusena bulakonzya kupa kuti zintu eezyi zivwule muzibeela zyoonse zyazyakulya, aboobo zilakonzya kunyonganya buumi bwazilenge (Binkova et al., 2000; Srogi, 2007; Quinn et al., 2009).
Nkozizwa anzila zya PAHs kuya kuzilenge zipona zili kwiinda mukulonga akuswaangana aakati kazyintu zyamunyika zisiyene mbuli bulongo, maanzi aansi, maanzi aali atala, zilyo amulengalenga (Arey a Atkinson, 2003). Cifwanikiso 1 chitondeezya kujatana akwaabila kwa PAHs zyama molecule masyoonto zisiyene-siyene muzilenge zyamunyika anzila zyabo zyakunjila muzintu zipona/bantu. Ma PAHs alabikkwa atala akaambo kakusofwaala kwamuya alimwi akulonga (kuzungaana) kwamuya uuzwa mumyootokala, muya uuzwa mumafakitali (kupanga muya wamalasha, kuumpa alimwi akupanga coke) alimwi akubikkwa kwawo. Milimu yamakkampani mbuli kupanga zisani zyakupanga, madayi amapenti; kubamba nkuni; kubamba raba; milimu yakupanga sementi; kupanga misamu iijaya tuzunda; alimwi akubelesya mubulimi nzintu zipati zipa kuti kube PAHs munyika amumaanzi (Bamforth a Singleton, 2005; Wick et al., 2011). Ziiyo zyakatondezya kuti bulongo bwamuminzi, afwaafwi amigwagwa mipati, amuminzi mipati bulakonzya kuciswa akaambo kakuzwa mumaanda aapanga magesi, kupya mumaanda, muya amumigwagwa, alimwi amilimo yakuyaka (Suman et al., 2016). (2008) wakatondezya kuti ma PAHs mumuse afwaafwi amigwagwa muNew Orleans, Louisiana, USA akali kusika ku 7189 μg/kg, mpawo mubusena butakkwene, akali buyo 2404 μg/kg. Mbubwenya buyo, mweelwe wa PAH uusika ku 300 μg/kg wakaambwa mumasena aali afwaafwi amasena aakupanga muya wamalasha mumadolopo manji aaku Amelika (Kanaly a Harayama, 2000; Bamforth a Singleton, 2005). Bulongo buzwa mumadolopo aasiyene-siyene aakuIndia mbuli Delhi (Sharma et al., 2008), Agra (Dubey et al., 2014), Mumbai (Kulkarni a Venkataraman, 2000) antoomwe a Visakhapatnam (Kulkarni et al., 2014) bwakaambwa kuti bulijisi bunji bwa PAH. Zintu zijisi tununkilizyo zilafwambaana kunjila mubulongo, zintu zipona alimwi abulongo, aboobo ziba zintu zipati zipa kuti kaboni kaboni muzilenge zipona (Srogi, 2007; Peng et al., 2008). Nkozyaka zipati zya PAHs muzilenge zyamumaanzi mvwula (mvwula iiteteete/iyumu alimwi abusi bwamaanzi), kuzwa kwamaanzi mumadolopo, kuzwa kwamaanzi aasofweede, kuzuzya maanzi aansi azimwi (Srogi, 2007). Kuyeeyelwa kuti aali 80% ya PAHs muzilenge zyamulwizi zizwa kumvwula, kuseluka, akusowa tombe (Motelay-Massei et al., 2006; Srogi, 2007). Bunji bwa PAHs mumaanzi aali atalaa nyika naakuti mumaanzi azwa mumasena aakusowela tombe alazuza mumaanzi aansi, eezi zileta ntenda mpati kubuumi bwabantu nkaambo bantu bainda ku 70% baku South a Southeast Asia banywa maanzi aansi (Duttagupta et al., 2019). Buvwuntauzi bwakachitika ino-ino aba Duttagupta abamwi. (2020) yamulonga (32) amaanzi aansi (235) kuzwa kuWest Bengal, India, yakajana kuti 53% yabantu bakkala mumadolopo a 44% yabantu bakkala kumyuunzi (boonse antoomwe bali 20 miliyoni) balakonzya kujatikizyigwa a naphthalene (4.9–10.6 μg/L) azintu zizwa kulinjiyo. Kusiyana kwakubelesya nyika akuyungizya kugwisigwa kwamaanzi aansi zilabonwa kuti nzintu zipati zyeendelezya kweenda (kuyungizya) kwa PAHs zili ansi anyika. Kuzwa kwamaanzi aabulimi, kuzwa kwamaanzi aasofweede aamuminzi amumafakitali, alimwi akuzwa kwamaanzi aasofweede/tombe zyakajanika kuti zilajatikizyigwa ama PAHs mumilonga amumasena aali munsi-munsi. Mvwula iizwa mumulengalenga ilayungizya busofwaazi bwa PAH. Bunji bwa PAHs antoomwe azintu zizwa kuli nzizyo (51 zyoonse antoomwe) zyakaambwa mumilonga/masena aajisi maanzi nyika yoonse, mbuli Mulonga wa Fraser, Mulonga wa Louan, Mulonga wa Denso, Mulonga wa Missouri, Mulonga wa Anacostia, Mulonga wa Ebro, a Mulonga wa Delaware (Yunker et al., 2002; Motelay-Massei et al., 2006; Li et al., 2006; 2010; Amoako abamwi, 2011; Kim abamwi, 2018). Mumulonga wa Ganges, naphthalene antoomwe a phenanthrene zyakajanwa kuti nzizyo zipati (zyakajanika mu 70% yazitondezyo) (Duttagupta et al., 2019). Kwiindawo, ziiyo zyakatondezya kuti kubikka chlorine mumaanzi aakunwa kulakonzya kuleta kupanga ma PAHs aajisi oxygen a chlorine (Manoli a Samara, 1999). Ma PAH alabungika muzilyo, michelo amuzisyu akaambo kakutolwa azisyango kuzwa kubulongo busofweede, maanzi aali ansi anyika alimwi amvwula (Fismes et al., 2002). Zilenge zyiingi zyamumaanzi zilimbuli nswi, nswi, nswi, nswi zyamaanzi zilasofwaazyigwa a PAHs kwiinda mukulya chakulya chisofweede amaanzi aalwizi, antoomwe anyama alukanda (Mackay a Fraser, 2000). Nzila zyakujika/kubamba mbuli kujika, kuyoka, kufweba, kujika, kuyuma, kujika akujika amalasha zilakonzya kuleta mweelwe mupati wa PAHs muzyakulya. Eezi zyeendelana akusala kwachibelesyo chakufweba, bunji bwa phenolic/aromatic hydrocarbon, nzila yakujika, musyobo wachintu chakupya, bunji bwamaanzi, muuya wa oxygen akupya (Guillén et al., 2000; Gomes et al., 2013). Ma polycyclic aromatic hydrocarbons (PAHs) aalo akajanwa mumukaka mubunji bwiindene-indene (0.75–2.1 mg/L) (Girelli et al., 2014). Kuyoboloka kwa PAHs muzyakulya akwalo kuyeeme abukkale bwazyakulya, mpawo ntenda zyabo zyeendelana amilimo yamubili, milimo ya metabolic, kunywa, kwaabila akwaabila kwamubili (Mechini et al., 2011).
Bulowa abubi bwa polycyclic aromatic hydrocarbons (PAHs) zyakazibidwe kwachiindi chilamfu (Cherniglia, 1984). Ma molecule masyoonto aa polycyclic aromatic hydrocarbons (LMW-PAHs) (ma ringi obile kusikila kuli otatwe) alakonzya kuswaangana ama macromolecules aasiyene-siyene mbuli DNA, RNA ama puloteni mpawo alakonzya kuleta kkansa (Santarelli et al., 2008). Akaambo kakuti taziyandi maanzi, zilaandaanizyigwa aama lipid membranes. Mubantu, ma cytochrome P450 monooxygenases apanga PAHs kuba ma epoxides, amwi aayo alabeleka loko (nkokuti, baediol epoxide) mpawo alakonzya kusololela kukusanduka kwamaseelo aali kabotu kuba ma epoxide (Marston et al., 2001). Mukuyungizya, zintu zizwa mukusanduka kwa PAHs zilimbuli quinones, phenols, epoxides, diols, azimwi zili abuti kwiinda zintu zizyala. Amwi ma PAH azintu zili akati-akati zyabo zilakonzya kunyonganya mahomoni ama enzyme aasiyene-siyene mumubili, aboobo zilakonzya kunyonganya kukomena, nsinga zyamukati, nzila zyakuzyala azyakukwabilila mubili (Swetha a Phale, 2005; Vamsee-Krishna et al., 2006; Oostingh et al., 2008). Kukkala kwachiindi chifwiifwi kuma PAHs aajisi ma molecule masyoonto kwakaambwa kuti kuletela kutabeleka kabotu kwamapunga akuzimba kwamapunga mubantu bachiswa bulwazi bwakufwundilila mpawo akuyungizya ntenda yakuciswa kkansa yachikkumba, yamapunga, yachikumba ayamumatumbu (Olsson et al., 2010; Diggs et al., 2011). Buvwuntauzi bwabanyama bwakatondezya kuti kujatikizyigwa a PAH kulakonzya kuba azintu zitali kabotu kukuzyala akukomena alimwi kulakonzya kuleta bulwazi bwameso, kunyonyooka kwa nswi achibindi, antoomwe akuciswa kwa jaundice. Zintu zisiyene-siyene zyakusanduka kwa PAH zilimbuli diols, epoxides, quinones antoomwe ama free radicals (cations) zyakatondezyegwa kuti zilakonzya kupanga DNA adducts. Ziyungizyo zitazungaani zyakatondezyegwa kuti zilakonzya kuchincha nzila zyakupanga DNA, mpawo zintu zitazungaani zilakonzya kugwisya DNA (ikapati kuba adenine mpawo chimwi chiindi kuba guanine); zyoonse zyobile zilakonzya kuleta tumpenda tusololela kukusanduka (Schweigert et al. 2001). Kuyungizya waawo, ma quinones (benzo-/pan-) alakonzya kupanga musyobo wa oxygen (ROS), akupa kuti DNA azimwi zipati zinyonyooke, aboobo zilakonzya kunyonganya mulimo wanyama (Ewa a Danuta 2017). Kukkala kwachiindi chilamfu mubunji bwa pyrene, biphenyl a naphthalene kwakaambwa kuti kuletela kkansa kubanyama bakali kusunkwa (Diggs et al. 2012). Akaambo kakujaya kwazyo, kusalazya/kugwisya ma PAH aaya kuzwa mumasena aajatikizyidwe/asofweede nchintu chiyandikana.
Nzila zyaandeene zyakumubili azyamakemikalu zyakabelesegwa kugwisya ma PAH kuzwa mumasena/masena aasofweede. Nzila zilimbuli kutenta, kugwisya chlorine, UV oxidation, kubamba, akugwisya solvent zilijisi mapenzi manji, kubikkilizya akupanga zintu zikola, buyumuyumu bwanzila, lukwabililo atwaambo twamilawu, kutabeleka kabotu, amali manji. Nokuba boobo, kunyonyooka kwatuzunda tusyoonto-syoonto (kwiitwa kuti bioremediation) ninzila imwi iisyomya iijatikizya kubelesya tuzunda tusyoonto-syoonto munzila yazilengwa zisalala naa zibeela zyabantu. Kweezyanisya anzila zyakumubili azyamakemikalu, eeyi nzila tainyonganyi busena, tainyonganyi pe, taibelesyi mali manji, alimwi ilakonzya kuzumanana. Kubambulula zintu zipona kulakonzya kuchitwa kubusena bujisi ntenda (mubusena) naakuti kubusena bwakabambilwa (anze) aboobo kubonwa kuti ninzila yakubambulula iikonzya kuzumanana kwiinda nzila zyakale zyakumubili azyamakemikalu (Juhasz a Naidu, 2000; Andreoni a Gianfreda, 2007; Megharaj et al., 2011; Phale et al., 2020; Phale et al., 2020). Sarkar abamwi, 2020).
Kumvwisisisya ntaamu zyakusanduka kwatuzunda zijatikizyidwe mukunyonyoona zintu zisofweede zijisi tununkilizyo tujisi mpindu mpati mumakani aasayaansi alimwi aabuvwubi mukukwabilila buumi bwazilenge alimwi abusena. Kuyeeyelwa kuti 2.1×1018 grams ya kaboni (C) ilayobolwa muma sediments amuzintu zipona (nkokuti, mafuta, muya uuzwa kuzilenge, amalasha, nkokuti, mafuta aazwa kuzintu zyakasyaalila) nyika yoonse, akugwasya kapati mukweendeenda kwa kaboni munyika yoonse. Nokuba boobo, kufwambaana kwakupanga zintu, kugwisya mafuta aakaindi, amilimo yabantu zili mukumana mabwe aaya aajisi kaboni, akugusya 5.5×1015 g ya kaboni (mbuli zintu zisofwaazya) mumulengalenga amwaka (Gonzalez-Gaya et al., 2019). Bunji bwa kaboni eeyi ilanjila munyika amulwizi kwiinda mukuseluka, kweenzyegwa, akuzwa kwamaanzi. Mukuyungizya, zintu zipya zisofwaazya zizwa kumafuta aakaindi, zilimbuli mapulasitiki, mapulasitiki alimwi azintu ziyumya mapulasitiki (ma phthalates alimwi ama isomers), zisofwaazya kapati zintu zili mulwizi, bulongo alimwi azilenge zyamumaanzi alimwi azintu zipona, aboobo ziyungizya ntenda zyakucinca kwamvwula munyika yoonse. Misyobo yiindene-indene yamapulasitiki masyoonto, mapulasitiki masyoonto, zibeela zyamapulasitiki azintu zyabo zikola zizwa ku polyethylene terephthalate (PET) zyabungene mu Lwizi lwa Pacific akati ka North America a Southeast Asia, akupanga “Great Pacific Garbage Patch”, akunyonganya buumi bwamulwizi (Newell et al., 2020). Ziiyo zyasayaansi zyakatondezya kuti tazikonzeki pe kugwisya zintu zisofweede/zisofweede ziliboobu kwiinda mukubelesya nzila zili zyoonse zyakumubili naa zyamakemikalu. Muchiimo eechi, tuzunda tugwasya kapati ntotukonzya kusandula zintu zisofweede kuti zibe carbon dioxide, manguzu aamakemikalu azintu zimwi zitakwe tombe zikonzya kunjila munzila zimwi zyakuzungaana kwazyakulya (H, O, N, S, P, Fe, azimwi). Aboobo, kumvwisisisya buumi bwama microbial bwakupanga zintu zisofweede antoomwe akweendelezya busena bwazyo nchintu chipati loko mukulanga-langa nzila zya microbial carbon cycle, net carbon budget alimwi antenda zyakumbele zyakucinca kwachiindi. Akaambo kakuyandika kwakufwambaana kugwisya misamu iili boobu mubusena, makwebo aasiyene-siyene aabikkila maano kuzyintu zisalala akazwa. Munzila imwi, kusumpula tombe lyamakkampani/makemikalu aasofweede aabungene muzilenge (nkokuti nzila ya tombe kuya kubuvubi) ilabonwa kuti ninzila imwi yabuvubi bwakuzyunguluka antoomwe amakanze aakusumpula zintu (Close et al., 2012). Aboobo, kumvwisisisya nzila zya metabolic, enzymatic antoomwe a genetic zyazintu zikonzya kunyonyooka nchintu chipati loko mukugwisya akubambulula zintu zisofweede.
Akati kazintu zyiingi zisofweede, tulabikkila maanu kapati ku PAHs zitakwe ma molecule masyoonto zilimbuli naphthalene antoomwe a naphthalenes. Eezyi zintu nzintu zipati zipangwa mumafuta aazwa ku petroleum, madayi aamasani, zintu zibelesyegwa abantu, misamu iijaya tuzunda (ma mothballs alimwi ama insect repellents), ma plasticizers alimwi ama tannins aboobo zilidumide mumasena manji (Preuss et al., 2003). Malipooti alino-lino atondezya kuyoboloka kwa naphthalene mumaanzi, mumaanzi aansi amubulongo buli munsi-munsi, mumasena aajisi mavadose amumilonga, kutondeezya kuyoboloka kwayo mubusena (Duttagupta et al., 2019, 2020). Chikozyano 2 chilapandulula bube bwa physicochemical, kubelesegwa antoomwe akuchisa kwa naphthalene azintu zizwa mulinjiyo. Kweezyanisya azimwi PAHs zijisi ma molecule manji, naphthalene azintu zizwa kulinjiyo taziyandi maanzi pe, zilakonzya kunywika mumaanzi alimwi zilamwaika mumasena manji, aboobo zilabelesyegwa mbuli nzila zyakuyiisya nzila zyakusanduka kwazyakulya, zyakuzyalwa antoomwe akusiyana kwazyakulya zya PAHs. Mweelwe mupati watuzunda tusyoonto-syoonto tulakonzya kusanduka naphthalene azintu zizwa kulinjiyo, alimwi makani manji alajanika kujatikizya nzila zyabo zyakusandula, ma enzyme alimwi anzila zyakweendelezya (Mallick et al., 2011; Phale et al., 2019, 2020). Mukuyungizya, naphthalene azintu zizwa kulinjiyo zyakasalwa kuti zibe zibelesyo zyakusaanguna zyakupima kusofwaala kwabusena akaambo kakuvwula kwazyo alimwi akujanika kwazyo. Bbunga lyakukwabilila busena lyaku Amelika liyeeyela kuti mweelwe waakati-kati wa naphthalene uli 5.19 μg a cubic meter kuzwa kubusi bwa tombwe, ikapati kuzwa kukupya kutazulide, alimwi a 7.8 kusikila ku 46 μg kuzwa ku busi bwamumbali, kakuli kujatikizyigwa a creosote a naphthalene kuli 100 kusikila ku 1000, 000s (Preer et al. 2003). Naphthalene ikapati yakajanika kuti ilakonzya kuleta ntenda yakuyoya iijatikizya musyobo, busena, abwaamu. Kwiinda mukuvwuntauzya kwabanyama, mbunga yanyika yoonse iilanganya kkansa (IARC) yakabikka naphthalene kuti "ilakonzya kuleta kkansa kubantu" (Kabunga 2B)1. Kuswaangana a naphthalenes, ikapati kwiinda mukuyoya naakuti kubelesya mumulomo, kuletela kuchisa kwamapunga akuyungizya kuzimba kwamapunga mu mbeba amu mbeba (National Toxicology Program 2). Zitondeezyo zipati zibikkilizya kufwubaala, kuluka, kuchisa mwida, kuzwa bulo, kuchisa mutwe, kunyongana, kuzuza kapati, kupya mubili, kutyompwa kwamoyo, azimwi. Kulubazu lumwi, musamu uujaya tuzunda uutegwa carbamate carbaryl (1-naphthyl N-methylcarbamate) wakaambwa kuti ulajaya tunyama tupona mumaanzi, abantu alimwi kwakatondezyegwa kuti kulakonzya kulesya acetylcholinesterase kupa kuti muntu aleke kubeleka (Smulders et al., 2003; Bulen a Distel, 2011). Aboobo, kumvwisisisya nzila zyakunyonyooka kwatuzunda, kweendelezya kwamajini, nsandulo zyama enzyme antoomwe amaseelo nchintu chipati loko mukupanga nzila zyakubambulula zintu zipona mumasena aasofweede.
Chikozyano 2. Makani aajatikizya bube bwa physicochemical, kubelesyegwa, nzila zyakuzyiba alimwi amalwazi aajatikizyidwe aa naphthalene azintu zizwa kulinjiyo.
Mumasena aasofweede, zintu zisofweede zitayandi maanzi alimwi aziyandisya mafuta zilakonzya kuleta ntenda ziindene-indene kumaseelo aali mubusena oobo (bantu), mbuli kucinca kwamaanzi aamubili, kunjila kwamubili, kuzimba kwamafuta, kunyongana kwanguzu (nketani yakutola ma electron/nguzu zya protoni), alimwi akubeleka kwamapulotini aajatikizyidwe mumubili (Sikkema et al. 1995). Mukuyungizya, zimwi zintu zikonzya kunywika mbuli ma catechols ama quinones zipa kuti kube oxygen species (ROS) mpawo zipanga ma adducts a DNA ama proteins (Penning et al., 1999). Aboobo, bunji bwazintu ziliboobu muzilenge zyamunyika zipa kuti tuzunda tube aanguzu zyakusofwaazya kabotu mumasena aasiyene-siyene, kubikkilizya akubweza/kutola, kusanduka kwamukati kamaseelo, kuswaangana/kubelesya, akwaandaanya.
Kuvwuntauzya kwa Ribosomal Database Project-II (RDP-II) kwakatondeezya kuti mweelwe wamabacteria aali 926 wakagwisigwa kuzwa kuzyintu zyakali kubelesegwa naakuti kuvwuzya zyakali kusofwaazigwa a naphthalene naakuti zintu zizwa kulinjiyo. Kabunga ka Proteobacteria kakalijisi mweelwe mupati wabaiminizi (n = 755), katobela Firmicutes (52), Bacteroidetes (43), Actinobacteria (39), Tenericutes (10), antoomwe amabacteria aatabikkidwe (8) (Mufanikisyo 2). Baiminizi ba γ-Proteobacteria (Pseudomonadales a Xanthomonadales) bakali kweendelezya nkamu zyoonse zya Gram-negative zili a G+C mpati (54%), mpawo Clostridiales a Bacillales (30%) zyakali nkamu zya Gram-positive zili a G+C nsyoonto. Pseudomonas (mweelwe mupati, misyobo iili 338) yakaambwa kuti ilakonzya kunyonyoona naphthalene antoomwe azintu zizwa mulinjiyo muzintu zisofweede (malasha, mafuta, mafuta aatakwe busani, bulongo, mafuta aakali kutila, maanzi aasofweede, tombe lyazilenge alimwi amasena aakusowela tombe) alimwi amuzilenge zitakwe ntenda (maanzi aamumilonga, tombe, alimwi amaanzi aansi) (Chitondeezyo 2). Kwiindawo, ziiyo zyakuyungizya akuvwuntauzya kwa metagenomic mumasena amwi aaya zyakayubununa kuti misyobo ya Legionella a Clostridium iitakomene ilakonzya kuba anguzu zyakubola, kutondezya mbokuyandika kukomezya ma bacteria aaya kuti twiiye nzila zipya akusiyana kwa metabolic.
Cifwanikiso. 2. Kusiyana kwazibeela antoomwe akumwaika kwazyintu zipona mumasena aasofweede a naphthalene azintu zizwa ku naphthalene.
Akati kazyintu zisiyene-siyene zinyonyoona ma hydrocarbon, bunji bwazyo zilakonzya kunyonyoona naphthalene mbuli nzila ilikke ya carbon anguzu. Kutobelana kwazintu zijatikizyidwe mukusanduka kwa naphthalene kwakapandululwa kuli Pseudomonas sp. (misyobo: NCIB 9816-4, G7, AK-5, PMD-1 a CSV86), Pseudomonas stutzeri AN10, Pseudomonas fluorescens PC20 amisyobo imwi (ND6 a AS1) (Mahajan et al., 1994; Resnick et al., 1996; Anweer et al., 2000; al., 2003; Dennis a Zylstra, 2004; Sota et al., 2006; Kusanduka kwamubili kutalisyigwa a multicomponent dioxygenase [naphthalene dioxygenase (NDO), ring hydroxylating dioxygenase] eeyo iipa kuti chimwi chaziingi zya naphthalene zyeendelezyegwe. substrate, kusandula naphthalene kuba cis-naphthalenediol (Chitondeezyo 3 Cis-dihydrodiol yakasandulwa kuba 1,2-dihydroxynaphthalene kwiinda mukubelesya dehydrogenase A ring-cleaving dioxygenase, 1,2-dihydroxynaphthalene dioxygenase (12-dihydroxynaphthalene dioxygenase (12-dihydrooxynaphthalene), kusanduka kuba DHNDO. 2-hydroxychromene-2-carboxylic acid isomerization ya enzymatic ilapanga trans-o-hydroxybenzylidenepyruvate, eeyo iikosolwa a hydratase aldolase kuba salicylic aldehyde a pyruvate I organic acid pyruvate yakali C3 yakusaanguna kuzwa ku naphone mpawo yakanjila mukati ka carbon. Mukuyungizya, NAD+-dependent salicylaldehyde isandula salicylaldehyde aciindi eeci yiitwa kuti "nzila yakujulu" yakunyonyooka kwa naphthalene. hamburgii 2, kunyonyooka kwa naphthalene kutalisyigwa a naphthalene 2,3-dioxygenase kuti ipange 2,3-dihydroxynaphthalene (Annweiler et al., 2000).
Cifwanikiso 3. Nzila zyakunyonyooka kwa naphthalene, methylnaphthalene, naphthoic acid, antoomwe a carbaryl. Myeelwe iizungulukide yiiminina ma enzymes ajisi mukuli wakusandula naphthalene azintu zizwa mulinjiyo kuti zibe zintu zitobela. 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.
Kwiinda muchilengwa eechi abubambe bwazyo, salicylic acid iizwa mulinjiyo ilayungizigwa kwiinda munzila ya catechol kubelesya salicylate 1-hydroxylase (S1H) naakuti kwiinda munzila ya gentisate kubelesya salicylate 5-hydroxylase (S5H) (Mufanikisyo 3). Mbwaanga salicylic acid ninzila mpati iili akati-akati mukusanduka kwa naphthalene (nzila yakujulu), ntaamu kuzwa ku salicylic acid kuya ku TCA akati-kati kanjikanji zyaambwa kuti ninzila yaansi, mpawo ma genes abamba mu operan yomwe. Nchintu chidumide kubona kuti ma genes aali munzila yakujulu (nah) amunzila yamunsi (sal) zyeendelezyegwa azintu zyeendelezya; mbuli chitondeezyo, NahR a salicylic acid zibeleka mbuli zintu zipa kuti kube kuchincha, kupa kuti ma operon oonse obile asandule naphthalene (Phale et al., 2019, 2020).
Mukuyungizya, catechol ilaandaanizyigwa kuba 2-hydroxymuconate semialdehyde kwiinda munzila ya meta kwiinda mukubelesya catechol 2,3-dioxygenase (C23DO) (Yen et al., 1988) mpawo ilayungizyigwa hydrolyzed a 2-hydroxymuconate semialdehyde hydrolase kuti ipange 2-hydroxymuconate semialdehyde hydrolase, 2-4-dipenenoid acid. 2-hydroxypent-2,4-dienoate mpawo yasandulwa kuba pyruvate a acetaldehyde a hydratase (2-oxopent-4-enoate hydratase) antoomwe a aldolase (4-hydroxy-2-oxopentanoate aldolase) mpawo yanjila munzila ya carbon (Mufanikisyo 3). Nikunyina, catechol ilaandaanizyigwa kuba cis, cis-muconate kwiinda munzila ya ortho kwiinda mukubelesya catechol 1,2-oxygenase (C12DO). Muconate cycloisomerase, muconolactone isomerase, antoomwe a β-ketoadipate-nollactone hydrolase zisandula cis, cis-muconate kuba 3-oxoadipate, eeyo iinjila munzila ya carbon kwiinda mu succinyl-CoA a acetyl-CoA (Nozaki et al., 1968) (Figure 33).
Munzila ya gentisate (2,5-dihydroxybenzoate), nsinga zya aromatic zilaandaanizyigwa a gentisate 1,2-dioxygenase (GDO) kuti zipange maleylpyruvate. Eechi chintu chilakonzya kusanduka kuba pyruvate a malate, naakuti chilakonzya kusanduka kuba fumarylpyruvate, eecho chikonzya kusanduka kuba pyruvate a fumarate (Larkin a Day, 1986). Kusala kwanzila imbi kwakabonwa mulizyoonse zyobile Gram-negative a Gram-positive bacteria kuma biochemical a genetic levels (Morawski et al., 1997; Whyte et al., 1997). Mabakiteliya aatali kabotu (Pseudomonas) alayanda kubelesya asidi ya salicylic, eeyo iipa kuti naphthalene ichite kabotu, akuyigwisya kuba catechol kubelesya salicylate 1-hydroxylase (Gibson a Subramanian, 1984). Kulubazu lumwi, mu Gram-positive bacteria (Rhodococcus), salicylate 5-hydroxylase isandula salicylic acid kuba gentisic acid, mpawo salicylic acid taijisi nguzu zyakusandula ma naphthalene genes (Grund et al., 1992) (Figure 3).
Kwakaambwa kuti misyobo iili mbuli Pseudomonas CSV86, Oceanobacterium NCE312, Marinhomonas naphthotrophicus, Sphingomonas paucimobilis 2322, Vibrio cyclotrophus, Pseudomonas fluorescens LP6a, Pseudomonas antoomwe a Mycobacterium misyobo ya moncicome dimethylnaphthalene (Dean-Raymond a Bartha, 1975; Cane a Williams, 1982; Mahajan abamwi, 1994; Dutta abamwi, 1998; Hedlund abamwi, 1999). Akati kabo, 1-methylnaphthalene antoomwe a 2-methylnaphthalene nzila yakunyonyooka ya Pseudomonas sp. CSV86 yakavwuntauzyigwa kabotu-kabotu kuma biochemical a enzymatic levels (Mahajan et al., 1994). 1-Methylnaphthalene ilasanduka kwiinda munzila zyobile. Chakusanguna, nkomo iijisi tununkilizyo ilasandulwa (nkomo iitakwe busena bwa methylnaphthalene) kuti ipange cis-1,2-dihydroxy-1,2-dihydro-8-methylnaphthalene, eeyo iyungizigwa kuba methyl salicylate a methylcatechol, mpawo yanjila munzila yakati kaboni nikwakainda nkomo (Mufanikisyo 3). Eeyi nzila yiitwa kuti "nzila yakujana kaboni". Munzila yabili "yakugwisya toxin", nkamu ya methyl ilakonzya kusandulwa a NDO kuti ipange 1-hydroxymethylnaphthalene, eeyo iyungizigwa kusika ku 1-naphthoic acid mpawo yazwa mubusena bwakukomezya mbuli chintu chifwide. Ziiyo zyakatondezya kuti musyobo wa CSV86 taukonzyi kukomena a 1- a 2-naphthoic acid mbuli kaboni anguzu, eezi zisinizya nzila yakugwisya tombe (Mahajan et al., 1994; Basu et al., 2003). Mu 2-methylnaphthalene, kabunga ka methyl kalainda mu hydroxylation a hydroxylase kuchita 2-hydroxymethylnaphthalene. Mukuyungizya, nkomo iitakwe busena ya nsingo ya naphthalene ilanjila mu nsingo ya hydroxylation kuti ipange dihydrodiol, eeyo iisanduka kuba 4-hydroxymethylcatechol mumulongo wazyintu zichitwa a enzyme mpawo injila munzila ya carbon akati kwiinda munzila ya meta-ring cleavage. Mbubwenya buyo, S. paucimobilis 2322 yakaambwa kuti ibelesya NDO kuchita hydroxylate 2-methylnaphthalene, eeyo yakayungizigwa kupanga methyl salicylate a methylcatechol (Dutta et al., 1998).
Naphthoic acids (yakanjililwa/iitabikkidwe) nzintu zizwa mukugwisya toxini/kusanduka kwabuumi zipangwa muchiindi chakunyonyooka kwa methylnaphthalene, phenanthrene antoomwe a anthracene mpawo zyazwa mubusena bwakukomezya. Kwakaambwa kuti bulongo bwa Stenotrophomonas maltophilia CSV89 bulakonzya kuchincha 1-naphthoic acid mbuli nzila yakujana kaboni (Phale et al., 1995). Kusanduka kwamubili kutalikila akugwisigwa kwamaanzi aamununkilizyo kuti kube 1,2-dihydroxy-8-carboxynaphthalene. Diol iizwa muli 2-hydroxy-3-carboxybenzylidenepyruvate, 3-formylsalicylic acid, 2-hydroxyisophthalic acid antoomwe a salicylic acid mpawo yanjila munzila ya carbon kwiinda munzila ya meta-ring cleavage (Chitondeezyo 3).
Carbaryl musamu uujaya tuzunda. Kuzwa muchiindi cha Green Revolution muIndia muma 1970, kubelesya manyolo amisamu iijaya tuzunda kwapa kuti kube kuyungizigwa kwa polycyclic aromatic hydrocarbon (PAH) emissions kuzwa kuzintu zitali zyabulimi (Pingali, 2012; Duttagupta et al., 2020). Kuyeeyelwa kuti 55% (85,722,000 hectares) yanyika yoonse yabulimi muIndia ilasanzyigwa amisamu iijaya tuzunda. Muminyaka yosanwe yainda (2015-2020), chibaka chabulimi chakuIndia chakabelesya mweelwe waakati-kati wamatani aasika ku 55,000 kusikila ku 60,000 amisamu iijaya tuzunda amwaka (Dipatimenti iilanganya zyakubelekela antoomwe abasikulima, Mutabi wabulimi, Mfulumende yakuIndia, mwezi waChiloba 2020). Kunyika aakati-kati ka Gangetic plains (zisi zili abantu banji loko), kubelesya misamu iijaya tuzunda kuzilyo kulivwulide, amisamu iijaya tuzunda njiivwulide. Carbaryl (1-naphthyl-N-methylcarbamate) nchintu chilamfu, chili aakati-akati kusikila kuli chiimo chibi loko chakujaya tuzunda chibelesyegwa mubulimi bwakuIndia amweelwe wa 100-110 matani. Chiindi chiingi chisambalwa azina lyakuti Sevin mpo chibelesyegwa kulwana tuzunda (ma aphid, nswi zyamulilo, nkumba, nkumba, nkumba azimwi zijaya tuzunda) zijaya zilyo zyiingi (maila, soya, kkotoni, michelo azisyu). Tumwi tuzunda tulimbuli Pseudomonas (NCIB 12042, 12043, C4, C5, C6, C7, Pseudomonas putida XWY-1), Rhodococcus (NCIB 12038), Sphingobacterium spp. (CF06), Burkholderia (C3), Micrococcus antoomwe a Arthrobacter zilakonzya kubelesegwa kujaya tuzunda tumwi. Kwakaambwa kuti RC100 ilakonzya kunyonyoona carbaryl (Larkin a Day, 1986; Chapalamadugu a Chaudhry, 1991; Hayatsu abamwi, 1999; Swetha a Phale, 2005; Trivedi abamwi, 2017). Nzila yakunyonyooka kwa carbaryl yakavwuntauzyigwa kapati kuma biochemical, enzymatic alimwi ama genetic mubulongo bwa Pseudomonas sp. Misyobo C4, C5 a C6 (Swetha a Phale, 2005; Trivedi abamwi, 2016) (Fig. 3). Nzila yakucinca kwamubili italika akusanduka kwa ester bond a carbaryl hydrolase (CH) kuti ipange 1-naphthol, methylamine antoomwe a carbon dioxide. 1-naphthol mpawo yasandulwa kuba 1,2-dihydroxynaphthalene a 1-naphthol hydroxylase (1-NH), eeyo iiyungizigwa kwiinda munzila ya carbon kwiinda mu salicylate a gentisate. Amwi ma bacteria anyonganya carbaryl akaambwa kuti alakonzya kuchisandula kuba salicylic acid kwiinda mukukosola catechol ortho ring (Larkin a Day, 1986; Chapalamadugu a Chaudhry, 1991). Chazibinkene, mabakiteriya anyonganya naphthalene asandula salicylic acid kwiinda mukubelesya catechol, mpawo mabakiteriya anyonganya carbaryl alayanda kusanduka salicylic acid kwiinda munzila ya gentisate.
Naphthalenesulfonic acid/disulfonic acid antoomwe a naphthylaminesulfonic acid zilakonzya kubelesegwa mbuli zintu zili akati-akati mukupanga madayi aa azo, zintu zitontola, zintu zimwaika, azimwi. Nikuba kuti eezyi zintu tazikwe ntenda kubantu, buvwuntauzi bwakujaya nswi zyakatondezya kuti zilakonzya kujaya nswi, daphim, algae, et. 1994). Bamwi baiminizi bamusyobo wa Pseudomonas (misyobo ya A3, C22) bakaambwa kuti batalika kusanduka kwazyakulya kwiinda mukubelesya nzila zyobile zya hydroxylation ya aromatic ring iijisi kabunga ka sulfonic acid kuti bapange dihydrodiol, eeyo isandulwa kuba 1,2-dihydroxynaphthalene kwiinda mukukosola kwa sulfite group (B rilon et al., 1918). I 1,2-dihydroxynaphthalene iizwa mumo ilasanduka kwiinda munzila ya naphthalene, nkokuti, nzila ya catechol naa gentisate (Mufanikisyo 4). Kwakatondezyegwa kuti aminonaphthalenesulfonic acid antoomwe a hydroxynaphthalenesulfonic acid zilakonzya kunyonyoonwa akusanganizigwa kwamabacteria antoomwe anzila zyakugwasyanya (Nortemann et al., 1986). Kwakatondezyegwa kuti umwi wabaabo bali mukabunga ulagwisya aminonaphthalenesulfonic acid naa hydroxynaphthalenesulfonic acid kwiinda mukubelesya 1,2-dioxygenation, kakuli aminosalicylate naa hydroxysalicylate ilagwisigwa mubusena bwakukomezya mbuli chintu chifwide mpawo ilatolwa abamwi bali mukabunga. Naphthalenedisulfonic acid ilakonzya kubola pesi taikonzyi kubola pe aboobo ilakonzya kusanduka kwiinda munzila zisiyene. Kugwisigwa kwasulfur kutaanzi kulachitika muchiindi chakusala kwa dihydroxylation ya aromatic ring antoomwe a sulfonic acid group; kugwisya sulfurization kwabili kulachitika muchiindi cha hydroxylation ya 5-sulfosalicylic acid a salicylic acid 5-hydroxylase kuti ipange gentisic acid, eeyo iinjila munzila ya kaboni (Brilon et al., 1981) (Chitondeezyo 4). Ma enzymes achita kuti naphthalene inyonyoonwe aalo ngaachita kuti naphthalene sulfonate ibeleke (Brilon et al., 1981; Keck et al., 2006).
Cifwanikiso 4. Nzila zyakusanduka kwa naphthalene sulfonate. Myeelwe iili mukati kazikobela yiiminina ma enzymes ajisi mukuli wakusanduka kwa naphthyl sulfonate, aakozyenye/akozyenye ama enzymes aapanduludwe mu FIG. 3.
Ma PAHs aalema ma molecule masyoonto (LMW-PAHs) alakonzya kucheya, taayandi maanzi alimwi taali kabotu kunywika pe, aboobo taakonzyi kunyonyooka pe. Nokuba boobo, tuzunda tusyoonto-syoonto tulakonzya kuchisandula kwiinda mukunyona muya uutegwa oxygen (O2). Aaya ma enzymes kanjikanji ali mukabunga ka oxidoreductases mpawo alakonzya kuchita zintu zyiingi mbuli aromatic ring hydroxylation (mono- naa dihydroxylation), dehydrogenation antoomwe a aromatic ring cleavage. Zintu zizwa mukusanduka ooku zilimuchiimo chakuyungizya muya mpawo zilakonzya kusanduka kwiinda munzila ya kaboni (Phale et al., 2020). Ma enzymes munzila yakubola akaambwa kuti alakonzya kuchitwa. Ikubeleka kwama enzyme aaya nkusyoonto loko naakuti nkutabikkilwi maanu aawo maselu naakomenena azintu zitakatazyi mbuli glucose naakuti organic acids. Chikozyano cha 3 chilapandulula ma enzyme aasiyene (ma oxygenases, ma hydrolases, ma dehydrogenases, ma oxidases, azimwi) aajatikizyidwe mukusanduka kwa naphthalene azintu zizwa kulinjiyo.
Chikozyano 3. Zitondeezyo zya biochemical zya enzymes zipa kuti naphthalene azintu zizwa mulinjiyo zinyonyooke.
Ziiyo zya Radioisotope (18O2) zyakatondezya kuti kubikka O2 muma molecule muzibeela zya aromatic rings kwiinda mukubelesya oxygenases nintaamu iiyandika kapati mukupa kuti musamu uzumanane kunyonyooka (Hayaishi et al., 1955; Mason et al., 1955). Kubikkilizya atomu yomwe ya oxygen (O) kuzwa ku molecule oxygen (O2) mukati ka substrate kutalisyigwa a endogenous naa exogenous monooxygenases (alimwi yiitwa kuti hydroxylases). Imwi atomu ya oxygen ilasanduka kuba maanzi. Ma monooxygenases aazwa anze alacesya flavin a NADH naa NADPH, mpawo muma endomonooxygenases flavin alacesyegwa a substrate. Busena bwa hydroxylation bupa kuti kube kusiyana kwakupanga zintu. Mbuli chitondeezyo, salicylate 1-hydroxylase ila hydroxylate salicylic acid aawo C1, mpawo ipanga catechol. Kulubazu lumwi, multicomponent salicylate 5-hydroxylase (iijisi reductase, ferredoxin, antoomwe a oxygenase subunits) ila hydroxylate salicylic acid a C5 position, akupanga gentisic acid (Yamamoto et al., 1965).
Ma dioxygenases abikkilizya ma atomu obile aa O2 mukati kachintu. Kwiinda muzintu zyakabambwa, zilaandaanizyigwa kuba ma ring hydroxylating dioxygenases antoomwe ama ring cleaving dioxygenases. Ma ring hydroxylating dioxygenases asandula zintu zijisi tununkilizyo kuba cis-dihydrodiols (nkokuti, naphthalene) mpawo zilajanika kapati akati kamabacteria. Kusikila sunu, kwakatondezyegwa kuti zintu zipona zijisi ma ring hydroxylating dioxygenases zilakonzya kukomena mumasena aasiyene-siyene aajisi kaboni, mpawo ma enzymes aaya abikkwa muzibeela zya NDO (naphthalene), toluene dioxygenase (TDO, toluene), antoomwe a biphenyl dioxygenase (BPDO, biphenyl). Zyoonse zyobile NDO a BPDO zilakonzya kuchita kuti kube kusanduka kwabili antoomwe akusanduka kwama hydroxylation aa polycyclic aromatic hydrocarbons (toluene, nitrotoluene, xylene, ethylbenzene, naphthalene, biphenyl, fluorene, indole, methylnaphthalene, naphthalene, phenthalenethrenethrenethreneth, acetophenone, azimwi) (Boyd a Sheldrake, 1998; Phale abamwi, 2020). NDO ninzila iijisi zibeela zyiingi iijisi oxidoreductase, ferredoxin, antoomwe a active site-containing oxygenase component (Gibson a Subramanian, 1984; Resnick et al., 1996). Cibeela ca NDO cipa kuti zintu zibe kabotu cijisi cibeela cipati ca α alimwi acibeela cisyoonto ca β cibikkidwe munzila ya α3β3. NDO ili mumpuli mpati yama oxygenases mpawo α-subunit yayo ilijisi Rieske site [2Fe-2S] antoomwe a mononuclear non-heme iron, eeyo iipa kuti NDO ibeleke kabotu (Parales et al., 1998). Mbukubede, muchiindi chomwe chakuchita zintu, ma electron obile kuzwa kukucheya kwa pyridine nucleotide alatolwa ku Fe(II) ion mubusena bubeleka kwiinda mu reductase, a ferredoxin antoomwe a Rieske site. Ikucheya kweelene kupa kuti muya uutegwa oxygen ubeleke, ooyo uuyandikana kuti kube dihydroxygen (Ferraro et al., 2005). Kusikila sunu, ma NDO masyoonto buyo ngakasalazyigwa akuzyibwa kabotu-kabotu kuzwa kumisyobo yiindene-indene alimwi kweendelezya kwama gene kwanzila zijatikizyidwe mukunyonyooka kwa naphthalene kwakavwuntauzyigwa munzila iimvwika (Resnick et al., 1996; Parales et al., 1998; Karlsson et al., 2003). Ring-cleaving dioxygenases (endo- naa ortho-ring-cleaving enzymes antoomwe a exodiol- naa meta-ring-cleaving enzymes) zibeleka aa hydroxylated aromatic compounds. Mbuli chitondeezyo, ortho-ring-cleaving dioxygenase ni catechol-1,2-dioxygenase, mpawo meta-ring-cleaving dioxygenase ni catechol-2,3-dioxygenase (Kojima et al., 1961; Nozaki et al., 1968). Mukuyungizya kuma oxygenases aasiyene, kuli a dehydrogenases aasiyene-siyene ajisi mukuli wakugusya ma-hydrogenation aa aromatic dihydrodiols, alcohols a aldehydes akubelesya NAD+/NADP+ mbuli zibelesyo zitambula ma electron, eezyo zili zimwi zyama enzymes aayandika kapati aajatikizyidwe mu metabolism (Gibson a Subramanian, 1984; Shaw a Farayama, 1909; Farayama, et al., et al. 2020).
Ma enzymes mbuli ma hydrolases (esterases, amidases) ngamusyobo wabili uuyandika kapati wama enzymes abelesya maanzi kutyolaula ma covalent bonds mpawo atondezya bube bwa substrate bupati. Carbaryl hydrolase azimwi hydrolases zilabonwa kuti nzibeela zya periplasm (transmembrane) muzibeela zya Gram-negative bacteria (Kamini et al., 2018). Carbaryl ilijisi amide antoomwe a ester; aboobo, ilakonzya kuyobolwa a esterase naa amidase kuti ipange 1-naphthol. Carbaryl mu Rhizobium rhizobium strain AC10023 antoomwe a Arthrobacter strain RC100 zyaambwa kuti zibeleka mbuli esterase antoomwe amidase, kweendelana. Carbaryl mu Arthrobacter strain RC100 alimwi ibelekela mbuli amidase. RC100 yakatondezyegwa kuti ilakonzya kujaya misamu yone ya N-methylcarbamate mbuli carbaryl, methomyl, mefenamic acid a XMC (Hayaatsu et al., 2001). Kwakaambwa kuti CH mu Pseudomonas sp. C5pp ilakonzya kubeleka a carbaryl (100% activity) a 1-naphthyl acetate (36% activity), pesi kutali a 1-naphthylacetamide, kutondezya kuti ni esterase (Trivedi et al., 2016).
Ziiyo zya biochemical, nzila zyakweendelezya ma enzyme, akuvwuntauzya kwama gene zyakatondezya kuti ma genes aakunyonyooka kwa naphthalene ajisi zibeela zyobile zikonzya kweendelezya naakuti "operons": nah ("nzila yakujulu", kusandula naphthalene kuba salicylic acid) antoomwe a sal ("nzila yakuselemuka", kusandula salicylic acid kuba nzila ya carbon catecholic). Asidi ya salicylic azimwi zikozyenye anjiyo zilakonzya kubeleka mbuli zintu zipa kuti kube (Shamsuzzaman a Barnsley, 1974). Mukubako kwa glucose naa ma organic acids, operon ilalesyegwa. Cifwanikiso 5 chitondeezya bubambe boonse bwakunyonyooka kwa naphthalene (muchiimo cha operon). Zibeela zyiingi zya nah gene (ndo/pah/dox) zyakapandululwa mpawo zyakajanwa kuti zilijisi kweendelana kupati (90%) akati kamisyobo yoonse ya Pseudomonas (Abbasian et al., 2016). Ma genes aa naphthalene upstream pathway akalibikkidwe munzila yakuzuminana mbuli mbokutondezyedwe achifanikisyo 5A. Aimbi gene, nahQ, yakaambwa kuti ilajatikizyidwe mukusanduka kwa naphthalene alimwi kanjikanji yakali akati ka nahC a nahE, asi mulimo wayo wini tiiwakazyibwa pe. Mbubwenya buyo, nahY gene, iilanganya naphthalene-sensitive chemotaxis, yakajanwa kumamanino aa nah operon muli zimwi zizo. Mu Ralstonia sp., U2 gene iibamba glutathione S-transferase (gsh) yakajanwa kuti ili akati ka nahAa a nahAb pesi tiiyakajatikizya bube bwakubelesya naphthalene (Zylstra et al., 1997).
Cifwanikiso 5. Bubambe bwamajini akusiyana kwakabonwa muchiindi chakunyonyooka kwa naphthalene akati kamisyobo yamabacteria; (A) Nzila ya naphthalene yakujulu, kusanduka kwa naphthalene kuya ku salicylic acid; (B) Nzila ya naphthalene, salicylic acid kwiinda mu catechol kuya kunzila ya kaboni iili akati; (C) salicylic acid kwiinda mu gentisate kuya kunzila ya kaboni iili akati.
"Nzila yamunsi" (sal operon) kanjikanji ilaa nahGTHINLMOKJ mpawo isandula salicylate kuba pyruvate antoomwe a acetaldehyde kwiinda munzila ya catechol metaring cleavage. I nahG gene (ilembedwe salicylate hydroxylase) yakajanika kuti yakakwabililwa kumamanino aa operon (Fig. 5B). Kweezyanisya amisyobo imwi iinyonganya naphthalene, mu P. putida CSV86 nah a sal operons zili antoomwe mpawo zili afwaafwi loko (aasika ku 7.5 kb). Kuli zimwi zilwazi zya Gram-negative, zilimbuli Ralstonia sp. U2, Polaromonas naphthalenivorans CJ2, antoomwe a P. putida AK5, naphthalene ilasanduka mbuli kaboni metabolite kwiinda munzila ya gentisate (muchiimo cha sgp/nag operon). Ikkaseti yamajini kanjikanji itondezyegwa muchikozyano cha nagAaGHAbAcAdBFCQEDJI, aawo nagR (iibamba LysR-type regulator) mpoili kumamanino (Chitondeezyo 5C).
Carbaryl ilanjila mukati ka carbon cycle kwiinda mukusanduka kwa 1-naphthol, 1,2-dihydroxynaphthalene, salicylic acid, antoomwe a gentisic acid (Mufanikisyo 3). Kwiinda muziiyo zyama genetic a metabolic, kwakayeeyelwa kuti nzila eeyi yaandaanizyigwe mu "kujulu" (kusanduka kwa carbaryl kuba salicylic acid), "akati" (kusanduka kwa salicylic acid kuya ku gentisic acid), antoomwe a "ansi" (kusanduka kwa gentisic acid kuba akati-kati ka carbon) (Sing et al., 2013). Kuvwuntauzya kwa C5pp (supercontig A, 76.3 kb) kwakatondeezya kuti mcbACBDEF gene ilajatikizyidwe mukusandula kwa carbaryl kuba salicylic acid, mpawo mcbIJKL mukusandula salicylic acid kuba gentisic acid, mpawo mcbOQP mukusandula kwa central genmaratisic acid ( pyruvate, Trivedi abamwi, 2016) (Mufanikisyo 6).
Kwakaambwa kuti ma enzymes aajatikizyidwe mukunyonyooka kwa aromatic hydrocarbons (kubikkilizya a naphthalene a salicylic acid) alakonzya kuchitwa azintu eezyo mpawo alesyegwa azintu zitakatazyi mbuli glucose naa organic acids (Shingler, 2003; Phale et al., 2019, 2020). Akati kanzila zyaandeene zyakusanduka kwa naphthalene azintu zizwa kulinjiyo, kweendelezya kwa naphthalene a carbaryl zyakavwuntauzyigwa kusikila mpocigolela. Ku naphthalene, ma genes aali munzila zyoonse zyobile zyakujulu azyaansi zyeendelezyegwa a NahR, LysR-type trans-acting positive regulator. Nchintu chiyandikana mukutalisya nah gene a salicylic acid mpawo akutondezya kwayo kwaatala (Yen a Gunsalus, 1982). Kuya kunembo, ziiyo zyakatondezya kuti integrative host factor (IHF) antoomwe a XylR (sigma 54-dependent transcriptional regulator) azyalo zilayandika kapati mukuchita kuti ma genes abelesegwe mu naphthalene metabolism (Ramos et al., 1997). ziiyo zyakatondezya kuti ma enzymes aa catechol meta-ring opening pathway, nkokuti catechol 2,3-dioxygenase, alachitwa kuti kakuli naphthalene antoomwe a salicylic acid (Basu et al., 2006). Ziiyo zyakatondezya kuti ma enzymes aa catechol ortho-ring opening pathway, nkokuti catechol 1,2-dioxygenase, alachitwa kuti kakuli benzoic acid a cis,cis-muconate (Parsek et al., 1994; Tover et al., 2001).
Mumusyobo wa C5pp, ma genes osanwe, mcbG, mcbH, mcbN, mcbR antoomwe a mcbS, abamba zyeendelezyo zili mumukwasyi wa LysR/TetR wazyeendelezyo zyakulemba zyeendelezya kunyonyooka kwa carbaryl. Ijini mcbG yakajanwa kuti ilaswaangene kapati a LysR-type regulator PhnS (58% amino acid identity) iijatikizyidwe mu phenanthrene metabolism mu Burkholderia RP00725 (Trivedi et al., 2016). Ijini lya mcbH lyakajanwa kuti lilajatikizyidwe munzila yaakati-kati (kusandula salicylic acid kuya ku gentisic acid) mpo lili mu LysR-type transcriptional regulator NagR/DntR/NahR mu Pseudomonas a Burkholderia. Bamwi bamukwasyi ooyu bakaambwa kuti balizyi kuti salicylic acid nchintu chikonzya kuchita kuti ma genes anyonganye. Kulubazu lumwi, ma genes otatwe, mcbN, mcbR antoomwe a mcbS, aali mu LysR a TetR type transcriptional regulators, akajanwa munzila yakuselemuka (gentisate-central carbon pathway metabolites).
Muma prokaryotes, nzila zyakutola ma gene (kujana, kuchincha, naa kutola) kwiinda muma plasmids, transposons, prophages, genomic islands, antoomwe a integrative conjugative elements (ICE) nzintu zipati zipa kuti ma genomes aabacteria abe abubotu, mpawo eezi zipa kuti kube kuyungizya naa kusweekelwa kwamilimo imwi. Zipa kuti mabakiteriya afwambaane kuzyibila mubukkale bwiindene-indene, kupa bubotu bwakuzyibila kumuntu uutambula, mbuli kunyonyooka kwazintu zijisi tununkilizyo. Kuchincha kwazyakulya mumubili kanjikanji kulachitika kwiinda mukubamba kabotu ma operons aakunyonyooka, nzila zyabo zyakweendelezya, anzila zya enzyme, eezyo zigwasya kunyonyooka kwamisyobo myiingi yama aromatic compounds (Nojiri et al., 2004; Phale et al., 2019, 2020). Ma kaseti aama gene aakunyonyooka kwa naphthalene akajanwa kuti ali muzintu zyiingi zyeenda mbuli ma plasmids (aaswaangana aayo aataswaangani), ma transposons, ma genomes, ma ICEs, antoomwe akusanganya misyobo ya bacteria yiindene-indene (Chitondeezyo 5). Mu Pseudomonas G7, nah a sal operons zya plasmid NAH7 zilalembedwe munzila yomwe mpawo nzibeela zya transposon iitabeleki kabotu iiyandika transposase Tn4653 kuti ichite (Sota et al., 2006). Mu Pseudomonas strain NCIB9816-4, gene yakajanwa a conjugative plasmid pDTG1 mbuli ma operon obile (aasika ku 15 kb kuzwa kuli umwi) alo aakalembwa munzila ziimpene (Dennis a Zylstra, 2004). Mu Pseudomonas putida strain AK5, non-conjugative plasmid pAK5 ilaa enzyme iilanganya kunyonyooka kwa naphthalene kwiinda munzila ya gentisate (Izmalkova et al., 2013). Mu Pseudomonas strain PMD-1, nah operon ili aa chromosome, mpawo sal operon ili a conjugative plasmid pMWD-1 (Zuniga et al., 1981). Nokuba boobo, mu Pseudomonas stutzeri AN10, ma genes woonse aakunyonyooka kwa naphthalene (nah a sal operons) alajanwa aa chromosome mpawo alakonzya kujanwa kwiinda mukucinca, kusanganya alimwi, akubamba alimwi (Bosch et al., 2000). Mu Pseudomonas sp. CSV86, nah a sal operons zili mu genome muchimo cha ICE (ICECSV86). Bupange bulikwabilidwe a tRNAGly kutobelwa akwiinduluka kutondezya masena aakuswaangana/kujatana (attR a attL) antoomwe a phage-like integrase iijanika kumamanino oonse obile aa tRNAGly, aboobo bupange bulikozyenye a ICEclc element (ICEclcB13 mu Pseudomonas knackmusii mukunyonyoona chlorocatechol). Kwakaambwa kuti ma genes aali mu ICE alakonzya kuchinchwa kwiinda mukuswaangana achiindi chifwiifwi loko (10-8), aboobo akutola bube bwakunyonyooka kuli yooyo uutambula (Basu a Phale, 2008; Phale et al., 2019).
Bunji bwama genes achita kuti carbaryl inyonyoonwe ajanika muma plasmids. Arthrobacter sp. RC100 ilijisi ma plasmid otatwe (pRC1, pRC2 antoomwe a pRC300) oomo ma plasmid obile aajatikizyidwe, pRC1 antoomwe a pRC2, ajisi ma enzyme aasandula carbaryl kuba gentisate. Kulubazu lumwi, ma enzyme aajatikizyidwe mukusanduka kwa gentisate kuba ma carbon metabolites aakati-kati alajanwa aa chromosome (Hayaatsu et al., 1999). Mabacteria aamusyobo wa Rhizobium. Musyobo wa AC100, uubelesyegwa kusandula carbaryl kuba 1-naphthol, ulijisi plasmid pAC200, eeyo iijisi cehA gene iibamba CH mbuli cibeela ca Tnceh transposon izingulukidwe azintu zilimbuli zintu (istA a istB) (Hashimoto et al., 2002). Mu Sphingomonas strain CF06, carbaryl degradation gene ilasyomwa kuti ili muma plasmid osanwe: pCF01, pCF02, pCF03, pCF04, antoomwe a pCF05. Kukozyanya kwa DNA kwama plasmid aaya kulisumpukide, kutondezya kubako kwachitika chakuvwula kwama gene (Feng et al., 1997). Mumusyobo wa carbaryl-degrading symbiont uupangidwe amisyobo yobile ya Pseudomonas, musyobo 50581 ulijisi conjugative plasmid pCD1 (50 kb) iibamba mcd carbaryl hydrolase gene, mpawo conjugative plasmid mumusyobo 50552 ilijisi 1-naphthode-engrading engrameme (Cpadu Chaudhry, 1991). Mu Achromobacter strain WM111, mcd furadan hydrolase gene ili a 100 kb plasmid (pPDL11). Eeyi gene yakatondezyegwa kuti ilajanika muma plasmid aasiyene (100, 105, 115 naa 124 kb) muma bacteria aasiyene kuzwa mumasena aasiyene (Parekh et al., 1995). Mu Pseudomonas sp. C5pp, ma gene woonse ajisi mukuli wakumana kwa carbaryl ali mu genome iijisi 76.3 kb ya sequence (Trivedi et al., 2016). Kuvwuntauzya kwa genome (6.15 Mb) kwakayubununa kubako kwa 42 MGEs antoomwe a 36 GEIs, oomo 17 MGEs mozyakali mu supercontig A (76.3 kb) a G+C iitakwe mweelwe (54–60 mol%), kutondeezya kuti kuli zyintu zikonzya kuchitika zyakucinca kwama gene (Trivedi et al., 2016). P. putida XWY-1 itondezya bubambe bukozyenye bwama genes aakonzya kunyonyoona carbaryl, pesi aaya ma genes ali a plasmid (Zhu et al., 2019).
Mukuyungizya kukubeleka obotu kwa metabolic kuma biochemical a genomic levels, microorganisms azyalo zitondezya bube bumbi naakuti nsandulo mbuli chemotaxis, bube bwakucinca maseelo, kwaandaanya, kubelesya kabotu, kupanga biosurfactant, azimwi, eezyo zibagwasya kuti bachinche kabotu zintu zisofweede mumasena aasofweede (Chitondeezyo 7).
Cifwanikiso 7. Nzila zyakuvwiila zyamaseelo ziindene-indene zyabacteria zibotu zikonzya kunyonyoona ma hydrocarbon kutegwa zintu zisofweede zyeende kabotu.
Ikwiingula kwa chemotactic kulabonwa kuti nzintu zisumpula kunyonyooka kwa zintu zisofweede muzilenge zisofweede ziindene-indene. (2002) bakatondezya kuti chemotaxis ya Pseudomonas sp. G7 ku naphthalene yakayungizya mweelwe wakunyonyooka kwa naphthalene mumaanzi. Musyobo wamusyobo wa G7 wakanyonyoona naphthalene chakufwambaana kwiinda musyobo uutakwe chemotaxis. Ma protein aa NahY (538 amino acids a membrane topology) akajanwa kuti alembedwe antoomwe ama metacleavage pathway genes aa NAH7 plasmid, alimwi mbuli chemotaxis transducers, eeyi protein ilibonya kuti ibelekela mbuli chemoreceptor ya naphthalene degradation (Grimm a Harwood 1997). Alubo buvwuntauzi bumbi bwakacitwa aHansel abamwi. (2009) bakatondezya kuti ma protein alakonzya kukwabilila misamu, asi kubola kwawo kulisumpukide. (2011) bakatondezya nsandulo ya chemotactic ya Pseudomonas (P. putida) ku gaseous naphthalene, oomo gas phase diffusion yakapa kuti naphthalene izumanane kweenda mumaseelo, eeyo yakali kweendelezya chemotactic response yamaseelo. Basikuvwuntauzya bakabelesya nzila eeyi ya chemotactic kupanga ma microbes aakonzya kuyungizya mweelwe wakunyonyooka. Ziiyo zyakatondezya kuti nzila zya chemosensory azyalo zyeendelezya milimu imwi yamaseelo mbuli kwaandaana kwamaseelo, kweendelezya kweenda kwamaseelo, akupanga biofilm, aboobo zilagwasya kweendelezya mweelwe wakunyonyooka. Nokuba boobo, kubelesya eechi chintu (chemotaxis) kuti chibelesyegwe kabotu kulalesyegwa azintu zyiingi. Zisinkilizyo zipati nzizyeezyi: (a) zibelesyo ziindene-indene zitambula zintu zyeendelana; (b) kubako kwazyintu zimbi zitambula, nkokuti, nguzu zyakutontola; (c) kwiimpana kupati kwakutobelana kwazibeela zyakumvwa zyamukwasyi omwe; a (d) kubula makani aatala amapulotini mapati aamabacteria (Ortega et al., 2017; Martin-Mora et al., 2018). Chimwi chiindi, kunyonyooka kwa ma hydrocarbons aanunkilila kupa kuti kube zintu zyiingi zipangwa mumubili, eezyo zinga zyaba chemotactic kuli kabunga kamwi kamabacteria pesi kazisesemya kuli bamwi, akupa kuti nzila eeyi ikatazye. Kuziba kujatana kwa ligands (aromatic hydrocarbons) antoomwe a chemical receptors, twakapanga ma hybrid sensor proteins (PcaY, McfR, a NahY) kwiinda mukuswaanganya sensor a signaling domains zya Pseudomonas putida antoomwe a Escherichia coli, eezyo zilanga zibeela zya aromatic acids, TCA intermediates, naphalene, antoomwe, antoomwe (Luthalene et al. 2019).
Mukuyungwa kwa naphthalene azimwi zintu zijisi tununkilizyo tunji (PAHs), bupange bwachikumba chabacteria akusyomeka kwatuzunda tusyoonto-syoonto zilachincha kapati. Ziiyo zyakatondezya kuti naphthalene ilanyonganya kuswaangana kwa acyl chain kwiinda mukuswaangana kwa hydrophobic, aboobo ikuyungizya kuzimba akuzuza kwa membrane (Sikkema et al., 1995). Kulwisya penzi eeli, mabakiteliya alalanganya kweenda kwa membrane kwiinda mukuchincha mweelwe amafuta aakati ka iso/anteiso branched-chain fatty acids a isomerizing cis-unsaturated fatty acids kuba trans-isomers (Heipieper a de Bont, 1994). Mu Pseudomonas stutzeri yakakomezyegwa a naphthalene, mweelwe wamafuta aazwide kusikila kuli uutakwe busani wakatanta kuzwa ku 1.1 kusika ku 2.1, mpawo mu Pseudomonas JS150 mweelwe ooyu wakatanta kuzwa ku 7.5 kusika ku 12.0 (Mrozik et al., 2004). Nibakakomezyegwa a naphthalene, maselu aa Achromobacter KAs 3-5 akatondezya kubungana kwamaseelo kuzinguluka mabwe aa naphthalene akucheya kwanguzu zyaatala amaseelo (kuzwa ku -22.5 kusika ku -2.5 mV) antoomwe akuvwundauka kwa cytoplasmic a vacuolization, kutondezya kucinca kwabupange bwamaseelo alimwi abube bwamaseelo (Mohapatra et al., 2019). Nikuba kuti kuchincha kwamaseelo/aatala kulaswaangene akutambula kabotu zintu zisofweede, nzila zyakupanga zintu zipona tazinabambululwa pe. Kuchincha chiimo chamaseelo takubelesyegwi pe kugwasya nzila zyabuumi (Volke a Nikel, 2018). Kugwisigwa kwama genes aajatikizya kwaandaana kwamaseelo kuletela kuchincha kwabupange bwamaseelo. Kugwisigwa kwama genes aajatikizya kwaandaana kwamaseelo kuletela kuchincha kwabupange bwamaseelo. Mu Bacillus subtilis, ma protein aa septum aa SepF atondezyedwe kuti alajatikizyigwa mukupanga septum alimwi alayandika muzibeela zitobela zyakwaandaana kwa maselu, pesi teensi gene iiyandika pe. Kugwisigwa kwama genes ajisi peptide glycan hydrolases mu Bacillus subtilis kwakapa kuti maseelo akomene, akuyungizya mweelwe wakukomena, akusumpula nguzu zyakupanga ma enzyme (Cui et al., 2018).
Kupambukanya nzila yakunyonyooka kwa carbaryl kwakayeeyelwa kuti kube kunyonyooka kubotu kwa Pseudomonas strains C5pp a C7 (Kamini et al., 2018). Kwakayeeyelwa kuti carbaryl ilatolwa mubusena bwa periplasmic kwiinda muchibaka chaanze cha membrane alimwi/naa kwiinda mu diffusible porins. CH ni enzyme iili mukati ka plassmic iipa kuti carbaryl ibe 1-naphthol, eeyo iitazungaani, iitayandi maanzi alimwi ilakola. CH ili mukati ka periplasm mpawo ilaa luyando lusyoonto ku carbaryl, aboobo ilakonzya kweendelezya kupanga kwa 1-naphthol, aboobo ilakonzya kulesya kuyoboloka kwayo mumaseelo akucesya ntenda yayo kumaseelo (Kamini et al., 2018). I 1-naphthol iizwa mulinjiyo ilatolwa mu cytoplasm kwiinda mukati ka membrane kwiinda mukwaandaanya a/naa kumwaika, mpawo ilasandulwa kuba 1,2-dihydroxynaphthalene a high-affinity enzyme 1NH kuti iyungizye kusanduka munzila ya kaboni.
Nikuba kuti tuzunda tusyoonto-syoonto tulijisi nguzu zyakunyonyoona zyintu zipa kuti kube kaboni, bupange bwakubelesya kwazyo (nkokuti, kubelesya zintu zitakatazyi kwiinda zintu zikatazya kumvwisisisya) nchintu chipati chisinkila kubola. Kubako akubelesya nzila zitakatazyi zyakujana kaboni kulacesya ma genes aabamba ma enzymes aakonzya kunyonyoona zintu zitayandiki kapati zya kaboni mbuli PAHs. Chikozyano chakavwuntauzyigwa kabotu nzakuti eelyo glucose a lactose nozipegwa antoomwe ku Escherichia coli, glucose ilabelesyegwa kabotu kwiinda lactose (Jacob a Monod, 1965). Pseudomonas yakaambwa kuti ilakonzya kunyonyoona ma PAHs a xenobiotic mbuli nzila zyakujana kaboni. Bweendelezi bwakubelesya kaboni mu Pseudomonas mbwakuti asidi zipona > glucose > zintu zijisi tununkilizyo (Hylemon a Phibbs, 1972; Collier abamwi, 1996). Nokuba boobo, kuli chintu chiliimpene. Chakugambya, Pseudomonas sp. CSV86 itondezya bupange bulibedelede bwalo bubelesya ma aromatic hydrocarbons (benzoic acid, naphthalene, azimwi) kwiinda glucose mpawo ilasandula ma aromatic hydrocarbons ama organic acids (Basu et al., 2006). Muchilwazi eechi, ma genes aakunyonyooka akutola ma hydrocarbons aanunkilila taakwe naalilwa pe nikuba kuti kuli kaboni yabili mbuli glucose naa organic acids. Eelyo nozyakali kukomezyegwa mu glucose a aromatic hydrocarbons, kwakabonwa kuti ma genes aakunyamuna glucose a metabolism akacheya, ma aromatic hydrocarbons akabelesegwa muchibeela chakusanguna, mpawo glucose yakabelesegwa muchibeela cabili (Basu et al., 2006; Choudhary et al., 2017). Kulubazu lumwi, kujanika kwa asidi zyabuumi tiikwakachizya kutondezyegwa kwa aromatic hydrocarbon metabolism, aboobo eeyi bacterium ilalangilwa kuba musyobo uukonzya kubelesegwa muziiyo zyakubola (Phale et al., 2020).
Chilizyibidwe kuti kusanduka kwa hydrocarbon kulakonzya kuleta penzi lyakukataazikana akuyungizya ma enzyme aakwabilila mubili muzintu zipona. Kubola kwa naphthalene kutabeleki kabotu mumaseelo aatazungaani alimwi akujanika kwazintu zikola kupa kuti kube musyobo wa oxygen uukonzya kubeleka (ROS) (Kang et al. 2006). Mbwaanga ma enzymes anyonganya naphthalene ajisi tubunga twa iron-sulfur, muchiindi chakukatazyigwa kwa oxidative, iron iili mu heme ama protein aa iron-sulfur ziyoosanduka, eezi zipa kuti ma protein aleke kubeleka. Ferredoxin-NADP+ reductase (Fpr), antoomwe a superoxide dismutase (SOD), ilagwasya kujokela musyule akati ka NADP+/NADPH ama molecule obile aa ferredoxin naa flavodoxin, aboobo ilagwisya ROS akubukulusya busena bwa iron-sulfur muchiindi cha oxidative stress (Li et al. 2006). Kwakaambwa kuti zyoonse zyobile Fpr a SodA (SOD) mu Pseudomonas zilakonzya kuchitwa akukatazyigwa kwa oxidative, mpawo kuyungizigwa kwa SOD a catalase kwakabonwa mumisyobo yone ya Pseudomonas (O1, W1, As1, a G1) muchiindi chakukomena mubukkale bwakuyungizya naphthalene (Kang et al., 2006). Ziiyo zyakatondezya kuti kuyungizya ma antioxidants mbuli ascorbic acid naa ferrous iron (Fe2+) kulakonzya kuyungizya kukomena kwa naphthalene. Eelyo Rhodococcus erythropolis niyakakomena mu naphthalene medium, kulembwa kwa oxidative stress-related cytochrome P450 genes kubikkilizya a sodA (Fe/Mn superoxide dismutase), sodC (Cu/Zn superoxide dismutase), antoomwe a recA kwakayungizyigwa (Sazykin et al., 2019). Kweezyanisya mweelwe wama proteomic aamaseelo aa Pseudomonas aakakomezyegwa mu naphthalene kwakatondezya kuti kuyungizya ma protein aasiyene-siyene aajatikizya kwiingula kwakukatazyigwa kwa oxidative ninzila yakulwana penzi lyakukataazikana (Herbst et al., 2013).
Tunyama tusyoonto-syoonto twakaambwa kuti tulapanga zintu zipona kwiinda mukubelesya zintu zitayandi maanzi. Aaya ma surfactants ngama amphiphilic surface active compounds aakonzya kupanga ma aggregates aawo mafuta amaanzi naa muuya amaanzi. Eezi zisumpula kusanduka kwakubeja akugwasya kunjila kwama haayidirokaboni aanunkilila, eezi zipa kuti kube kunyonyooka kubotu (Rahman et al., 2002). Akaambo kazyintu eezyi, ma biosurfactants alabelesyegwa kapati mumakwebo aasiyene. Kuyungizya misamu iikola naakuti misamu iikola kumisela yamabacteria kulakonzya kuyungizya bubotu akufwambaana kwakubola kwa hydrocarbon. Akati kazyintu zipona, ma rhamnolipids azwa ku Pseudomonas aeruginosa akavwuntauzyigwa akuzyibwa (Hisatsuka et al., 1971; Rahman et al., 2002). Kuyungizya waawo, imwi misyobo ya biosurfactants ibikkilizya ma lipopeptides (mucins kuzwa ku Pseudomonas fluorescens), emulsifier 378 (kuzwa ku Pseudomonas fluorescens) (Rosenberg a Ron, 1999), trehalose disaccharide lipids kuzwa ku Rhodococcus (Ramdahl, 1988), lipopeptides kuzwa ku Hallinberg a Sacillus, 2002), antoomwe a surfactant kuzwa ku Bacillus subtilis (Siegmund a Wagner, 1991) antoomwe a Bacillus amyloliquefaciens (Zhi et al., 2017). Aaya ma surfactants aanguzu atondezyedwe kuti alacesya buyumuyumu bwaatala kuzwa ku 72 dynes/cm kusika kunsi a 30 dynes/cm, kupa kuti kube kunjila kabotu kwa hydrocarbon. Kwakaambwa kuti Pseudomonas, Bacillus, Rhodococcus, Burkholderia azimwi misyobo yabacteria ilakonzya kupanga ma rhamnolipid a glycolipid-based biosurfactants kuti yakomena mu naphthalene a methylnaphthalene media (Kanga et al., 1997; Puntus et al., 2005). Pseudomonas maltophilia CSV89 ilakonzya kupanga biosurfactant Biosur-Pm iizwa kunze lyamaselu kuti yakomezyegwa azintu zijisi tununkilizyo mbuli naphthoic acid (Phale et al., 1995). Ikweenda kwa Biosur-Pm kwakatondezya kuti kupanga kwayo nchintu chiyeeme akukomena antoomwe a pH. Kwakajanika kuti mweelwe wa Biosur-Pm wakapangwa amaseelo aa pH ya neutral wakali mupati kwiinda pH 8.5. Maseelo akakomenena a pH 8.5 taakwe naakali kuyanda maanzi alimwi akalijisi luyando lupati kuzintu zijisi tununkilizyo alimwi a aliphatic kwiinda maseelo aakakomezyegwa a pH 7.0. Mu Rhodococcus spp. N6, mweelwe mupati wa carbon ku nitrogen (C:N) antoomwe akucheya kwa iron nzintu zibotu zyakupanga zintu zipona zili kunze lyamaseelo (Mutalik et al., 2008). Kusoleka kwakachitwa kuti kube kusumpula kupanga zintu zipona (surfactins) kwiinda mukusumpula misyobo akubila. Nokuba boobo, mweelwe wa surfactant mubusena bwakukomezya uli munsi (1.0 g/L), eezi zileta buyumuyumu mukupanga zintu zyiingi (Jiao et al., 2017; Wu et al., 2019). Aboobo, nzila zyakucinca ma gene zyakabelesegwa kusumpula nzila zyakupanga zintu. Nokuba boobo, kuchincha kwayo kwabunjiniyela nkuyumu akaambo kabupati bwa operon (∼25 kb) antoomwe akweendelezya kwakupanga zintu zipona (Jiao et al., 2017; Wu et al., 2019). Mweelwe wakucinca kwama genetic engineering kwakachitwa mu Bacillus bacteria, ikapati kugaminina kuyungizya kupanga surfactin kwiinda mukubikka busena bwa promoter (srfA operon), ikuyungizya surfactin export protein YerP antoomwe a regulatory factors ComX a PhrC (Jiao et al., 2017). Nokuba boobo, nzila eezyi zyakucinca ma gene zyakazwidilila buyo mukucinca komwe naakuti zisyoonto zyama gene mpawo tazinasika pe mumakwebo. Aboobo, kwiiya kunembo kwanzila zyakusumpula zyeendelana aluzibo kulayandika.
ziiyo zyakubola kwa PAH zichitwa kapati muchiimo che laboratory. Nokuba boobo, mumasena aasofweede naakuti mumasena aasofweede, zintu zyiingi zipona (kupya, pH, muya mubotu, kujanika kwazyakulya, kujanika kwazyakulya, zimwi zintu zipona, kukasya zintu zipangwa kumamanino, azimwi) zyakatondezyegwa kuti zilakonzya kuchincha akusungwaazya nguzu zyakubola zyatuzunda.
Kupya kulakonzya kuchita obotu kukunyonyooka kwa PAH. Mbuli kupya mbokuyaabukomena, muya uusofweede ulaselemuka, eecho chijatikizya kusanduka kwatunyama tusyoonto-syoonto, nkaambo tuyanda muya mubotu mbuli chimwi chazibelesyo zya oxygenases zichita hydroxylation naakuti ring cleavage reactions. Chiindi chiingi chilabonwa kuti kupya kupati kusandula ma PAHs kuba zintu zikola, aboobo kulesya kubola (Muller et al., 1998).
Kwakazyibwa kuti masena manji aasofweede a PAH alijisi pH mpati loko, mbuli masena aasofweede aamugoti wa asidi (pH 1-4) antoomwe amasena aasofweede aamuwo/malasha aasofweede a alkaline leachate (pH 8-12). Eezyi zintu zilakonzya kunyonganya kapati nzila yakubola kwazintu zipona. Aboobo, katutanabelesya tuzunda tusyoonto-syoonto mukubamba zintu zipona, ncibotu kuti uchinche pH kwiinda mukuyungizya makemikalu (aakonzya kugwisya muya) mbuli ammonium sulfate naa ammonium nitrate mubulongo bujisi alkaline naa kubamba liming a calcium carbonate naa magnesium carbonate mubulongo bwa acid (Bowpta et al 195; Gulen and Gulen et al. 1995). Sar 2020).
Kunjila kwa muya mubotu kubusena bujatikizyidwe ncecintu cipa kuti PAH itanyonyooki. Akaambo kabukkale bwa redox bwabusena, nzila zyakubambulula zintu zipona kanjikanji ziyandika kunjizya muya uuzwa anze (kulima, kugwisya muya, akuyungizya misamu) (Pardieck et al., 1992). Odenkranz abamwi. (1996) wakatondezya kuti kuyungizya magnesium peroxide (musamu uugwisya muya mubotu) kumulonga uusofweede ulakonzya kugwasya BTEX. Alubo buvwuntauzi bumbi bwakavwuntauzya kunyonyooka kwa phenol a BTEX mubusena bwamaanzi aasofweede kwiinda mukunjizya sodium nitrate akuyaka zikala zyakugwisya zintu kutegwa kube kubambulula zintu zipona (Bewley a Webb, 2001).